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HARVARD UNIVERSITY 






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Library of the 






Museum of 






Comparative Zoology 





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BREVIORA 



MUSEUM OF COMPARATIVE ZOOLOGY 
Harvard University 



Numbers 231-264 
1965-1967 



CAMBRIDGE, MASS.. U.S.A. 
1967 



Edited 

By 

Nelda E. Wright 



CONTENTS 

BREVIORA 

Museum of Comparative Zoology 

Numbers 231-264 

1965 

No. 231. A new anole (Sauria. Iguanidae) from Puerto Rico. By 
Ernest E. Williams, Juan A. Rivero and Richard 
Thomas. 18 pp. October 29. 

No. 232. Hispaniolan giant anoles (Sauria, Iguanidae): new data 
and a new subspecies. By Ernest E. Williams. 7 pp. 
October 29. 

No. 233. South American Anolis (Sauria, Iguanidae): two new 
species of the punctatus group. By Ernest E. Williams. 
15 pp. October 29. 

No. 234. Studies on neotropical Pompilidae (Hymenoptera). I. 
The genus Agenioideus Ashmead in South America. 
By Howard E. Evans. 7 pp. October 29. 

No. 235. A new salamander of the genus Chiropterothton (Cau- 
data: Plethodontidae) from Mexico. By George B. 
Rabb. 8 pp. November 19. 

No. 236. Variation in the number of marginal tooth positions in 
three species of iguanid lizards. By Clayton E. Ray. 
15 pp. December 10. 

No. 237. A new species of the ant genus Dacetinops from Sarawak. 
By Robert W. Taylor. 4 pp. December 15. 



1966 

No. 238. An evaluation of Jamaican Dromicus (Serpentes, Colu- 
bridae) with the description of a new species. By 
Donald W. Buden. 10 pp. February 25. 



No. 239. South American anoles: Anolis biporcatus and Anolis 
fraseri (Sauria, Iguanidae) compared. By Ernest E. 
Williams. 14 pp. February 25. 

No. 240. Gynmothorax galetae, a new moray eel from the Atlantic 
Coast of Panama. By Ira Rubinoff. 4 pp. February 

25. 

No. 241. Avocettinops ycinoi, a new nemichthyid eel from the 
southern Indian Ocean. By Giles W. Mead and Ira 
Rubinoff. 6 pp. February 25. 

No. 242. The supposed "sponge spicules" of Merrill, 1895, from 
the Lower Cretaceous (Albian) of Texas. By William 
A. S. Sarjeant. 15 pp. February 25. 

No. 243. Quaternary fish fossils from west of Lake Rudolf, Kenya. 
By Keith Stewart Thomson. 10 pp. April 29. 

No. 244. A new species of Ashmimella from West Texas (Mol- 
lusca: Pulmonata). By W. J. Clench and W. B. Mil- 
ler. 6 pp. April 29. 

No. 245. Notes and descriptions of new Urocoptidae from Cuba 
and Hispaniola (MoUusca: Pulmonata). By William 
J. Clench. 14 pp. April 29. 

No. 246. Pseudanthessius procwrens n.sp., a cyclopoid copepod 
associated with a cidarid echinoid in Madagascar. By 
Arthur G. Humes. 14 pp. May 3. 

No. 247. The Chariares (Argentina) Triassic reptile fauna. I. In- 
troduction. By Alfred Sherwood Romer. 14 pp. May 
3. 

No. 248. A Triassic ammonite from the Hindubagh region, Baluch- 
istan, West Pakistan. By Bernhard Kummel. 5 pp. 
July 29. 

No. 249. Additional notes on the amphisbaenids of greater Puerto 
Rico. By Richard Thomas. 23 pp. July 29. 

No. 250. The brain of the emu Dromaeus novaehollandiae. II. 
Anatomy of the principal nerve cell ganglia and tracts. 
By Stanley Cobb. 27 pp. November 4. 



No. 25 1 . Chronological survey of the tetrapod-bearing Triassic of 
Argentina. By J. F. Bonaparte. 13 pp. November 4. 

No. 252. The Chafiares (Argentina) Triassic reptile fauna. II. 
Sketch of the geology of the Rio Chanares-Rio Gualo 
region. By Alfred Sherwood Romer and James A. Jen- 
sen. 20 pp. November 4. 

No. 253. A new Hispaniolan g^cko. By Richard Thomas. 5 pp. 
November 4. 

No. 254. Preliminary descriptions of new Honey-Eaters (Aves. 
Meliphagidae ) . By Finn Salomonsen. 12 pp. Novem- 
ber 4. 

No. 255. A revision of the fossil selenodont artiodactyls from the 
Middle Miocene Thomas Farm, Gilchrist County, 
Florida. By Vincent Joseph Maglio. 27 pp. Decem- 
ber 6. 



1967 

No. 256. Anolis chocorum, a new pimctatus-\\\:.Q anole from Dari- 
en, Panama (Sauria, Iguanidae). By Ernest E. Wil- 
liams and William E. Duellman. 12 pp. February 3. 

No. 257. A review of the Clark Fork vertebrate fauna. By Roger 
C. Wood. 30 pp. February 3. 

No. 258. Biology of the parthenogenetic fungus beetle Cis juscipes 
Mellie (Coleoptera: Ciidae). By John F. Lawrence. 
14 pp. February 3. 

No. 259. Expanding the palpi of male spiders. By William A. 
Shear. 27 pp. February 3. 

No. 260. Monograph of the genus Spiroceramiis (Mollusca: Pul- 
monata: Urocoptidae). By William J. Clench. 10 pp. 
February 3. 

No. 261. The monticola group of the lizard genus Anolis in His- 
paniola. By Richard Thomas and Albert Schwartz. 
27 pp. March 31. 



No. 262. A phylogenetic survey of molluscan shell matrix proteins. 
By Michael T. Ghiselin, Egon T. Degens, Derek W. 
Spencer, and Robert H. Parker. 35 pp. March 31. 

No. 263. The hydroid of Vcmnuccia forbesii (Anthomedusae, 
Tubulariidae). By Anita Brinckmann-Voss. 10 pp. 
March 31. 

No. 264. The Chanares (Argentina) Triassic reptile fauna. III. 
Two new gomphodonts, Massetognathus pascuali and 
M. teruggii. By Alfred Sherwood Romer. 25 pp. 
April 6. 



INDEX OF AUTHORS 
BREVIORA 

Museum of Comparative Zoology 
Numbers 231-264 

1965-1967 

Bonaparte, J. F 25 1 

Brinckmann-Voss, Anita 263 

BuDEN, Donald W 238 

Clench, William J 244, 245, 260 

Cobb, Stanley 250 

Degens, Egon T 262 

DuELLMAN, William E 256 

Evans, Howard E 234 

Ghiselin, Michael T 262 

Humes, Arthur G 246 

Jensen, James A 252 

Kummel, Bernhard 248 

Lawrence, John F 258 

Maglio, Vincent Joseph 255 

Mead, Giles W 241 

Miller, W. B 244 

Parker, Robert H 262 

Rabb, George B 235 

Ray, Clayton E 236 

RivERo, Juan A 231 

RoMER, Alfred Sherwood 247, 252, 264 



RuBiNOFF, Ira 240, 241 

Salomonsen, Finn 254 

Sarjeant, William A. S 242 

Schwartz, Albert 261 

Shear, William A 259 

Spencer, Derek W 262 

Taylor, Robert W 237 

Thomas, Richard 231, 249, 253, 261 

Thomson, Keith Stewart 243 

Williams, Ernest E 231, 232, 233, 239, 256 

Wood, Roger C 257 



BREVIORA 

Mmseiiim of Comparative Zoology 

Cambridge, Mass. Octobeu 29, 19G5 Number 231 

A NEW ANGLE (SAURIA, IGUANIDAE) FROM 
PUERTO RICO 

By Ebxest E. Williams, 
Juan A. Rivero and Richard Thomas 

PART I. DESCRIPTION 

By Ernest E. Williams 
Museum of Comparative Zoology 

and Juan A. Rivero 

University of Puerto Eieo, Mayaguez 



On June 18, 1963, Juan Rivero was collecting at Cerro La 
Punta at an approximate elevation of 1200 meters when his wife 
and son, who had separated from the rest of the party, shouted 
that they had what appeared to be a new species of lizard. The 
animal was first seen in the axilla of one of the outer leaves of 
the bromeliad Vriesia (Thecodacfyllion) sintenisii, which al- 
though usually epiphytic, was growing abundantly, together 
with Guzmania herteroniana, on the forest floor alongside the 
road. Although the cloud forest is fairly heavy in this area, 
trees and tree ferns had been cut for about 10 or 15 meters 
along the margin of the road, leaving an open although some- 
what shaded strip on each side. It was in the marginal area 
between the forest and the cleared area that the new anole was 
collected. 

When capture of the specimen was attempted, it jumped out 
of the bromeliad and climbed a small bush nearby. The move- 
ment was descril)ed as slow, but the observers are not certain 
if it walked, jumped or crawled. Search for other specimens in 
and out of bromeliads was fruitless. 



2 BREVIORA No. 231 

On July 9, Mr. Francis Rolle collected a second specimen at 
the Maricao Reserve Forest. The animal was at the entrance of 
a hole on a partially rotten buttress of a tree at about 1.8 meters 
from the ground, and 600 to 750 mm from a bromeliad. Mr. 
Rolle used a flj^ swatter to stun the animal (which later died 
enroute) and its behavior could not be observed. Except for its 
lighter color, this specimen does not differ materially from the 
type. 

The first specimen was kept in the laboratory for several days, 
but although it was eating well (DrosopMla) and appeared to 
be in good condition it was eventually preserved in view of the 
danger that it might die and decay over a week end when it 
was not under observation. The animal expanded the dewlap 
on one occasion, and when given the opportunity to move on the 
floor, it always did so by jumping, or, if forced to move about 
until tired, by moving the hind limbs simultaneously, as if it 
were swimming. A 16 mm black and white film forms part of 
the type material filed at Museum of Comparative Zoology. 

During its first few days in captivity, the new anole slept by 
lying flat on one of the side panels of the aquarium which served 
to hold it. All four legs were on the glass, but the tail, on the 
floor of the aquarium, apparently served to prop the animal 
upwards. Later, it slept on a twig, in typical Anolis fashion. 
It vvas never seen to enter a small bromeliad that was provided 
inside the small tank, but its tail sometimes encircled twigs or 
small branches with its tip, although the animal was never seen 
to hang from its tail. 

No further specimens were obtained until early 1965 when 
Richard Thomas and later he and Albert Schwartz collected the 
new species in numbers at four localities in the mountains of 
southern Puerto Rico. Thanks to this material more is known 
both of the habitat and distribution of the new species. 

The material of the new species has been divided among a 
number of institutions : the Museum of Comparative Zoology 
(MCZ), the American Museum of Natural History (AMNH), 
the Carnegie Museum (CM), the Museum of Zoology University 
of Michigan (UMMZ), and the United States National Museum 
(USNM). A number of the Thomas specimens have been re- 
tained in the Albert Schwartz collection (ASFS) or the Richard 
Thomas collection (RT). 

In allusion to its long concealment from scientific record we 
call the new species by the Latin adjective vv^hich means "hid- 
den ' ' : 



1965 



NEW ANGLE FROM PUERTO RICO 




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4 BREVIORA No. 231 

Anolis occultus new species 

Holotypc. I\ICZ 80303, adult female, on Rd. 143, midway be- 
tween Cerro La Punta (1338 m) and Cerro Maravilla (1183 m), 
Puerto Rico, J. Rivero coll., 16 June, 1963. 

Paratypes. MCZ 83735, male, Maricao Reserve Forest, Fran- 
cis Rolle coll., 9 July, 1963; ASFS-V5489-91, V5494, AMNH 
94560-61, 13.7 Tim N Sahana Grande, 2800 feet (850 meters), 
A. Schwartz, Richard Thomas coll., 25 February, 1965; MCZ 
83661-63, same locality, Richard Thomas coll., 28 February, 
1965; A^F^-Y 6196-91] 10.6 km S8E Villa Perez, 3400 feet 
(1040 meters), Richard Thomas coll., 27 March, 1965; MCZ 
83664-67, same locality, Richard Thomas coll., 28 March 1965; 
MCZ 83656, 18.6 km NE Guayama, 2000 feet (610 meters), Rich- 
ard Thomas coll., 31 January, 1965; MCZ 83657-59, ASFS- 
V4891-92, V4901, CM 40691-92, UMMZ 126021-22, USNM 
157107-08, 20.9 km NNE Guayama, 2300 feet (700 meters), 
Richard Thomas coll., 2 February 1965; MCZ 83660, same lo- 
cality, Richard Thomas coll., 5 February, 1965 ; ASFS-V5017-19, 
same locality, Richard Thomas coll., 3 February, 1965. ASFS- 
V6662-65, RT 1332, 13.7 km S Palmer, Richard Thomas coll. 29 
July 1965 ; ASFS-V6670-71, same locality, Richard Thomas coll. 
30 July 1965. 

Diagnosis. An Anolis distinguished from all others by the 
wholly granular supraciliary margin, with no enlarged or elon- 
gate scales, and by the extreme reduction of the canthus, of 
which only the two posterior scales can be said to be differenti- 
ated from surrounding scales, as well as by the dorsal scales of 
the tail w^hich are very small and smooth. 

Description (paratype variation in parentheses). Head: Nar- 
row, elongate. Head scales small, smooth, ca. 13 (9-13) scales 
across snout between the hardly developed second canthals. A 
very shallow frontal depression. Nostril oval, nasal scale sep- 
arated from rostral by one small round scale. 

Supraorbital semicircles weakly developed, separated by 2 
(2-4) scales, supraocular area with enlarged scales medially, 
grading into granules laterally. Entire supraciliary margin 
granular: neither any elongate supraciliaries nor any series of 
enlarged squarish supraciliary scales. Canthal ridge barely ap- 
parent, its posterior inception indicated by two slightly enlarged 
squarish scales, hardly larger than the adjacent dorsal snout 
scales. First two canthals continued by slightly differentiated 
rectangular scales to a point below the naris. Loreal rows 4 (2-6, 



1965 



NEW ANGLE FROM PUERTO RICO 



usually 4), all scales subequal. Supratemporal scales subgranu- 
lar, flattened, grading- upward into irregularly enlarged scales 
surrounding interparietal. Interparietal round, small, larger 
or smaller than ear, separated /rom the supraorbital semicircles 
by 3 (2-6, usually 4) scales. Ear small, subround, placed far 
ventrally, directly behind the commissure of the mouth. 

Suboculars in contact with supralabials, anteriorly grading 
into loreals, posteriorly grading into supratempoials. Ten to 
eleven supralabials to the center of the eye, the posterior supra- 
labials very low and small. 




Fig. 2. Anolis occultus new species. Dorsal, lateral and ventral views of 
head of type, MCZ S0303. 



6 BREVIORA No. 231 

Mentals small, about as deep as wide, in contact with 4 (4-6, 
usually 4) scales between the infralabials. Several rows of throat 
scales medial to infralabials and posterior to mental somewhat 
enlarged, grading into granular scales on center of throat ; all 
throat scales smooth. 

Trunk : Middorsal scales smooth, flat, not larger than flank 
scales. Ventrals larger than dorsals, smooth, round, juxtaposed, 
in transverse rows. 

Gular fan : Large, present in both sexes and well developed 
even in juveniles, lateral margins inset ("slotted") in general 
skin of neck, all scales granular, smaller than throat scales, 
much smaller than ventrals, edge scales especially small, lateral 
scales small but well developed in well separated rows ( $ ) or 
these scales weakly developed, or almost absent { S ). 

Limbs and digits : Limbs short, flattened dorsoventrally, tibial 
length not exceeding distance snout-to-middle-of-eye. About 16 
(14-20. usually 16) lamellae under phalanges ii and iii of fourth 
toe. Scales of limbs smooth, much smaller than ventrals, supra- 
digital scales smooth. 

Tail : Round, no dorsal crest. No enlarged postanals in males. 
Scales on dorsal surface very small, smooth, imbricate. Scales 
behind vent smooth, ca. 4-6 ventral rows on distal half of tail, 
enlarged and keeled. Verticils indistinct, but apparently 11 
dorsal granules above, 5 keeled scales below, per verticil. 

Color. In the living type the head was gray, with sparse, dark 
mottles ; the eyelids of a lighter, yellowish gray color ; general 
color of dorsum gray, with dark vermiculations and a light 
bluish cast on the neck region ; flanks above the shoulders yel- 
lowish green, with black longitudinal spots or vermiculations; 
rest of flanks yellowish gray turning greenish towards the two 
extremities; an ochraceous stain on the dorsum, on the midline, 





Fig. 3. Scales around nostril compared in Anolis occultus (right) MCZ 
83661 and Anolis evermanni (left) MCZ 61223. 



1965 



NEW ANGLE FROM PUERTO RICO 



between the anterior limbs; tAvo small, round and well defined 
yellow spots on each side of the tail base, just below the sacral 
bones. Below, white, except for some speckling- on the throat and 
tail. The animal could change' colors easily, becoming darker or 
lighter with ease. 

The preserved type is of a darker gray color, with lighter reti- 
culation and spotting; there is a light colored frontal band in 
front of the eyes and a couple of light spots at the base of the 
tail. Venter and one-third of tail whitish with dark spots and 
specks, throat with dark areas along the margin of the lip and 
speckles and spots in the center. 

Size (in mm). Holotype : snout-vent, 34; tail 36. Largest 
specimen : snout-vent, 42. 

DISCUSSION 

Anolis occultus is far more distinct from other Puerto Rican 
anoles than a mere tabulation of conventional scale counts and 
characters (Tables 1 and 2) would indicate. It is a rather 






Fig. 4. Tail of A. occultus. Top: MCZ 83666, a verticil showing typical 
squamation. ILiddle : ASFS-V 5490, uiiregencrated tail tip. Bottom: ASFS-Y 
5489, regenerated tail tip. 



8 BREVIORA No. 231 

variable species and therefore approaches or overlaps other 
species in varying ways (e.g. gundlachi in certain characters, 
evermanni in others). It may be helpful to indicate the many 
ways in which it differs from all other Puerto Rican species: 
It is smaller in maximum size and has a longer head with usually 
smaller head scales. The nasal scale has not fused with the pre- 
nasal scale (Fig. 3). The absence of a differentiated supraocular 
disk and of elongate supraciliaries and differentiated post-supra- 
ciliary rows is very striking (Fig. 2). The mentals are remark- 
ably narrow (as if in occult us the apparent first infralabials had 
segmented oft" from formerly wider mentals). There are no 
differentiated sublabials ; these are present and large in all other 
Puerto Rican species. Enlarged postanal scales are absent in 
males. The tail is weakly if at all compressed and without trace 
or hint of any dorsal crest (Fig. 4) ; the dorsal caudal scales are 
not keeled or enlarged but finely granular and smooth. The 
dorsal limb scales are granular and smooth, not keeled, and the 
supradigital scales are also smooth. The fingers, including the 
distal phalanx, are notably short. The dewlap is sparsely scaled 
laterally and the lateral skin tucks under the well scaled skin 
of the throat to give a "slotted" appearance (Fig. 1). (See 
remarks by Thomas, below.) 

It is in fact necessary to go far afield to find species genuinely 
comparable with A. occultus. The suite of living species that 
requires comparison with occultus is the same that required 
comparison with the recently described Anolis in amber — 
Anolis electrum from the Miocene ambers of Chiapas, Mexico 
(Lazell, 1965) — A. fuscoawmtus, A. chloris, A. macidiventris, 
etc. 

The Miocene A7wlis, represented only by its skin, not its skele- 
ton, was remarkable for its small, quite uniform body and limb 
squamation. In this respect at least — A. electrum is unfortun- 
ately not complete and many characters are not available — ■ the 
Miocene anole and A. occultus are very similar, and electrum 
might well be, of all known species, the one closest to occultus. 
Of the living species which Lazell compared with electrum, the 
one closest in totality of features to occultus is maculiventris of 
the Colombian Choco. As Table 3 shows, in all scale counts there 
is at least overlap of extremes. Certain differences between 
maculiventris and occultus are those associated with the shorter, 
less tapered head and smaller absolute and relative scale size in 
maculiventris. Thus, in the latter feature, in a larger animal 
(snout-vent length 42 mm), the body scales are smaller, scales 
around interparietal smaller, gulars smaller and ventrals much 



1965 NEW ANGLE FROM PUERTO RICO 9 

smaller. Other differences are the presence of a small supra- 
ciliary, the wide mental, the unicarinate dorsal limb scales, the 
nmlticarinate siipradigital scales, the nacrower and slenderer 
digits and the larger caudal scales. The prenasal scale is fre- 
quently but not invariably fused with the nasal. The dewlap is 
not "slotted." 

It is not at all probal^le that occult us is close to maculiventris, 
but the necessity of comparison with a mainland species geo- 
graphically so distant emphasizes the very isolated position of 
occultus in the West Indies. Resemblance to maculiventris and 
to electrum may indicate preservation in occultus to a greater 
or lesser degree of a primitive anoline squamation pattern. This, 
however, is a point which will receive attention in a separate 
paper in which these and the osteological characters of this 
strange new Puerto Rican anole will be analyzed at length and 
their implications assessed. 

No detailed discussion of the distribution of occultus in Puerto 
Rico is possible. It has been collected at the two extremes of 
the Cordillera Central and with all probability^ will be found 
wherever the proper habitat exists all along this range. It has 
been recorded also by very recent collections (July 29-30, 1965) 
on El Yunque. All present records are above 600 meters. 

One comment more must be made. The discovery of so distinct 
a species in an island thought to be well known herpetologically 
and in which the anoles have received special attention must give 
us pause. As will appear from the observations reported below 
by Thomas, A. occultus is a creature of the canopy. We may 
well be ignorant of many another species of the canopy on the 
islands and on the mainland. 

ACKNOWLEDGMENTS 

We are indebted to Mr. Richard Thomas and Dr. Albert 
Schwartz for the privilege of examining the more than 30 
specimens collected by Mr. Thomas, and to Mr. Francis Rolle for 
the gift of the specimen collected by him. Permission to quote 
certain numerical data on Puerto Rican Anolis from an unpub- 
lished thesis has been granted by Dr. A. S. Rand. The illustra- 
tions are by Joshua Clark. This study has been supported by 
National Science Foundation Grant GB-2444. 

LITEEATUKE CITED 
Lazell, J. D. Jr. 

1965. An Anolis (Sauria, Iguanidae) in amber. Jour. Paleout. 
39: 379-382. 



10 BREVIORA No. 231 

PART II. FIELD OBSERVATIONS ON 
ANOLIS OCCULTUS WILLIAMS AND RIVERO 

By Richard Thomas 

10,000 SW 84th Street 
Miami, Florida 33143 

Having been fortunate enough to collect the first substantial 
series of the newly described Anolis occultiis, I take the oppor- 
tunity to present those observations on habits, habitat and color 
repertory which were made incidentally to collecting the speci- 
mens. I am indebted to Dr. Albert Schwartz, without whose 
support the collections would not have been made, and Dr. 
Ernest E. Williams whose advice was to be alert for a strange 
and new little anole in the forests of Puerto Rico. 

All but two specimens of occult us collected by me were taken 
at night while they slept. They invariably slept on dead or 
leafless vines and twigs. Although the sleeping sites were usually 
associated with a viny or bushy tangle, frequently individuals 
slept on single branches which projected beyond the main mass, 
or on pendant pieces of vine. While sleeping (Fig. 5) the head 




Fig. 5. Typical sleeping posture of Anolis occultus. 

was almost always towards the distal part of a twig or vine 
(where this could be determined). The forelimbs were flexed 
and the hindlimbs extended along the sleeping surface only 
slightly flexed. At times the fifth toe was positioned against the 
tail on either side, as if to hold it in place (a posture seen in 
some "grass anoles"). The tail was curled loosely about the vine 
or twig to a varying extent, sometimes completely encircling it. 
This characteristic prehensility of the tail enabled specimens of 
occultus to be identified at a glance even at a distance that would 



1965 NEW ANGLE FROM PUERTO RICO 11 

not ordinarily allow positive identification of so smajl a lizard. 
No specimens were ever observed sleeping on leafy portions of 
plants, whether green or dried, despite the abundance of ferns 
and grasses, and shrubs, at all the localities. 

When grasped, these lizards characteristically held tightly 
to the sleeping surface, and it was frequently simpler to break 
off the vine or twig on either end of the specimen than to risk 
injuring it by pulling it off. Typically, they showed great toler- 
ance to shaking and disturbance of their sleeping sites. When 
dislodged with a stick they might fall a short distance but 
adroitly catch themselves on the next object in their path, simul- 
taneously assuming the sleeping posture. On two occasions speci- 
mens were disturbed sufficiently (by approach of the collector 
with flashlight, not by being shaken) that they suddenly released 
their hold and dropped. One was recovered when it was found 
resting immobile in a tangle of vegetation on the ground. 

Williams and Rivero have commented on the slow movement 
reported by the collectors of the first specimen of this species. 
It is my observation that movement was "not slow but had a 
desultory aspect, not a hasty scampering characteristic of so 
many anoles" (field notes on specimen collected during day). 
The relatively short limbs of this species probably account for 
this apparent lack of haste or a leaping mode of progression. 
I have seen specimens of this lizard confined in a plastic bag 
make short hopping movements, but in the field evasion took 
place as a fast walk from stem to stem. AVhen progressing over 
a uniform surface (along a smooth stick, for instance), an almost 
salamander-like crawl is employed. 

When caught or handled, A. occultus may emit a rather per- 
sistent squeaking during its struggles to escape. On two occa- 
sions captive specimens were observed displaying the dewlap. 
In the admittedly artificial environment of a plastic bag filled 
with a coil of dead vines, the specimens displayed from a hori- 
zontal position with little flexure of the head and neck. 

The dewlap of this species, when retracted, appears to fit into 
a longitudinal slot in the throat and anterior chest. This is a 
characteristic of living specimens and is not an artifact of 
hardening in preservative. This "slot" (which is not evident 
when the dewlap is extended) is in the form of a distinct in- 
vagination of skin on either side of the retracted dewlap ; the 
folds on each side meet posteriorly and the entire structure is 



12 



BREVIOEA 



No. 231 




Fig. 6. Diagrammatic cross-section of a situation similar to those encoun- 
tered at localities 2 and 4. The dotted lines indicate the approximate zones 
where occultus was seen; the vertical line equals an approximate height of 
six feet. 



elongate and U-shaped. The posterior part of the dewlap in- 
serts beneath the "U" portion of the fold. The feature is pres- 
ent in even the smallest specimens. 

In life the eyes are chameleon-like : ' ' very protuberant and 
capable of considerable movement in all directions ; some degree 
of independent movement was evident" (field notes). 

Locality 1 : 18.6 km NXE Guayama, Puerto Rico, 2000 feet 
(610 meters), 1 specimen. The first specimen of Anolis occultus 
collected by me was found on a broad-leafed green plant growing 
in an open, sunny spot along the side of a path through montane 
rain forest. The specimen was on the upper surface of a leaf 
and moved to the under surface of an adjacent leaf when ap- 
proached. Subsequent search at this locality both day and night 
yielded no other specimens. 



1965 NEW ANGLE FROM PUERTO RICO 13 

Locality 2: 20.9 km NNE Guayama, 2300 feet (700 liieters), 
16 specimens. At this locality there was a path cut into a 
forested hillside (Fig. 2). The forest grew close to the sides of 
the path and in places overhung it. Downhill from the path 
grew what may be called the forest proper, which in places 
leveled off after a short drop. Above the path grew a dense 
second growth of generally smaller trees, ferns and viny tangles. 
A. occultus was found sleeping at night in tangles of dead (or 
leafless) vines and twigs along both sides of the path, four to 
ten feet above the ground (i.e. above the path on the uphill side, 
which includes the height of the cut-bank, or above the slope 
directly below the specimen on the downhill side). 

All but one specimen taken at this locality were collected at 
night while they slept. Specimens invariably slept on dead or 
leafless vines and twigs. 

An attempt to collect occultus in the forest below the path 
failed. The apparently preferred sleeping places were abundant 
enough, but all these were well lielow the canopy of the forest 
and not exposed by breaks in the forest. 

A brief stop was made at this locality during the daytime. 
One specimen was seen resting on the underside of the tip of a 
dead fern leaf, where it had possibly retreated at my approach. 

Locality 3 : 13.7 km N Sabana Grande, 2800 feet (850 meters), 
9 specimens. This locality was a region of montane rain forest 
of the same general composition as the first two localities but 
with a lower canopy. Specimens of occultus were found along 
paths through the forest but not of the 'cut bank' sort reported 
for the second locality. Specimens were definitely associated 
with an opening in the canopy. Where the forest canopy com- 
pletely closed over the paths, occultus was not to be found, de- 
spite the apparent abundance of the required viny tangles. 
The forest in this area was relatively low, so that there was not 
so large a space between the canopy and the ground. Specimens 
were found sleeping in similar situations to those of the second 
locality at heights above the ground of approximately four to 
ten feet. Three specimens were seen which were not collected 
due to the height at which they were sleeping (about 15 feet). 
One of these remained in place until the dead vine on which it 
rested was broken by being rather violently shaken. 

Locality 4: 10.6 km SSE Villa Perez, 3400 feet (1040 meters), 
8 specimens. Here specimens were collected in a situation very 
similar to the second locality. A path cut into a hillside wound 



14 BREVIORA No. 231 

through about two kilometers of montane forest. Specimens 
were collected at night as they slept. All were taken in an area 
where the downhill woods were high enough to partially or, in 
spots, completely enclose the path. Even under intensive search, 
specimens were not found in areas where the forest had been 
cleared below the path or the vegetation was low and sparse, 
even though the vegetation above the cut bank was of the proper 
type and density. The lizards slept at heights of about five to 
twelve feet above the ground. 

A visit was made to the Dona Juana Insular Forest, where no 
specimens of occultus were seen. The forest at this locality was 
very high and the canopy consequently out of easy access or 
sight. Likewise, the Toro Negro Insular Forest yielded no speci- 
mens, no doubt because of the lack of enough accessible situations 
in which the species is most easily collected.^ 

From the foregoing observations it is seen that A. occultus 
prefers as its sleeping site (and this should be a fairly good in- 
dication of its general habitat preference, though slightly more 
restricted) dead or bare twigs and vines in areas of close forest 
in the proximity of small breaks in the canopy. Normally, of 
course, the canopy is not accessible to the collector. It appears 
to be only in certain situations (breaks in the forest which allow, 
in effect, a descent of the canopy to a low level) that these 
lizards are accessible. It seems not improbable that occultus 
is normally a lizard of the canopy, demanding a thicket of 
bare vines and twigs among the foliage with readily available 
exposure to the sky. The possibility that it might be simply a 
clearing-edge anole is not indicated by my collecting experience. 
Specimens were not found along the edges of forest adjacent to 
large clearings or wide roads. 

At the first locality two other species of anoles were collected, 
A. gundlachi and A. evermanni. Both of these lizards are essen- 
tially ' ' tree anoles ' ' of forested regions ; the former is primarily 
a species of deep shade, the latter is often found on herbaceous 
plants such as Heliconia and Musa, but it is by no means re- 
stricted to such situations. At the second locality specimens of 



1 Since tliis manuscript went to press, another visit was made to Puerto 
Rico where seven more specimens of A. occultus were collected in the El 
Yunque region. All were taken at night along a path on a forested hillside, 
a situation very similar to that described for localities 2 and 4. One speci- 
men was found sleeping on the edge of a vertically oriented green leaf. 



1965 NEW ANGLE FROM PUERTO RICO 15 

I 

A. cristatcllus and A. krugi were collected; giindlachi and ever- 
mnnni were also seen there. A. h'rugi is an anole of bushes and 
grass and is sometimes almost terrestrial in habits. Its sleeping 
site preferences are less restricted than those of occultns. It may 
sleep on green or dead plants, leaves or stems, and is much less 
particular about how it positions itself. At locality 3 only A. 
fiundlachi was collected in addition to occult us. At locality 4, 
crisiatellus and krugi were taken ; gundlachi was seen. Krugi 
was very common there and during the day was often seen run- 
ning through the grass along the more open parts of the path. 
In addition, a juvenile specimen of an as yet undetermined 
species of large anole was collected at this locality. 

COLOR REPERTORY 

Anolis occult us possesses a well developed and complex dis- 
ruptive pattern which is variously manifested in different color 
stages. In its complete development, the pattern consists of the 
following elements : a dark cephalic figure or interocular tri- 
angle, which may be solid or hollow ; a pattern of dark radiating 
eye lines; four zones of transverse banding on the body (scapu- 
lar, dorsal, lumbar, and sacral) which may be manifested as 
either very hazy, indistinct dark l)ands or as bands with a 
sharply defined, sinuous, dark anterior edge and a posteriorly 
fading zone of dark pigment; a lumbar spot (occasionally 
paired), which is present in the lumbar band but is frequently 
evident when the band is not and is perhaps the most constant 
pattern element ; a fine reticulum of dark lines, which frequently 
appears as faint small ocelli. The venter is light, frequently with 
some stippling and a distinct but irregular zone of juncture with 
the dorsal coloration. The transverse body banding continues 
onto the tail as small dark chevrons. 

The colorations displayed by this lizard may be characterized 
as follows : 

Unicolor: Varies from gray through olive-brown, olive, yellow- 
green to a dirty orange color. Pattern elements are minimal in 
this range of phases but parts of the major elements may be 
present ; the lumbar spot is usually present. Axillary and in- 
guinal areas may be a dull or bright yellow ; a j'ellow edge to the 
lumbar spot may be present. This phase is the predominant one 
seen in specimens in the wild, which are usually green in the 
davtime and grav or brown at night. 



16 BREVIORA No. 231 

Lichcnate: The pattern is boldly developed and black or dark 
gray ; the ground color is off-white or very light gray. This pre- 
sents a very complex, striking and disruptive pattern ; it is per- 
haps a "fright" pattern as it is often seen in freshly collected 
or killed specimens. 

Intermediate: This category covers a wide range of effects in- 
termediate in various combinations between the other two. The 
pattern may be well developed and the ground color various 
shades as seen in the unicolor phase ; the pattern may be frag- 
mented, moderately or poorly developed with various shades of 
ground color, commonly brown, yellow-brown or gray. A fre- 
quent variant is almost uniformly reticulate above with especially 
prominent dark diagonal lines in the neck and scapular region. 

Dewlap color: The dewlaps were frequently noted as being 
pinkish gra.y. An individual whose dewlap was color-noted 
(ASFS V4890) with the Maerz and Paul Dictionary of Color 
(1950) was keyed to Plate 7E7. The posterior edge of the dewlap 
was rusty. This coloration was typical of the other specimens. 

LITEEATURE CITED 

Maerz, A. and M. Rea PxVUL 

1950. A dictionary of color. New York, McGraw-Hill Book Co., 
pp. i-vii, 1-23, 137-208, 56 pis. 

(Received 29 June 1965.) 



1965 



NEW ANGLE FROM PUERTO RICO 



17 




18 



BREVIORA 



No. 231 



Table 1 
Scale characters in Puerto Rican Anolis 

scales across labials 

S-V length (mm) lamellap snout at to center 

Anolia adult cf 4th toe second canthal loreals of eye 

occultvs 34 14^18 9^ 3 2^6 8^3~ 

cuvieri 126 31-33 8-9 6 7-8 

evervmnni 60-70 25-30 4-6 4-6 5-7 

stratulus 44-48 22-24 4-7 3-6 5-8 

cristatellus 65-74 22-25 4-7 5-7 5-7 

gundlachi 58-69 18-22 7-10 6-10 5-8 

Tcrugi 45-55 21-23 4-7 6-7 6-8 

pulchellns 45-50 19-21 4-5 3-5 5-7 

poncensis 43-46 18-21 4-6 3-5 5-6 



Anolis 



Scale characters in Puerto Rican Anolis 



scales interparietal 
between from middorsal rows 

semicircles semicircles enlarged 



ventrals 



tail crest 



occultus 


2-4 


2-6 







8j 


— 


cuvieri 


3 


3 




1 


pi 


+ ! 


evermanni 


0-1 


1-4 







si 


( + ) 


stratulus 


0-1 


0-3 







si 


+ 


cristatellus 


0-1 


1-4 




2 


s/k, i 


+ ! 


gundlachi 


1-3 


3-8 




2 


ki 


+ ! 


Icrugi 


0-2 


1-5 




4-6 


ki 


( + ) 


pulchelhi^ 


0-2 


1-5 


ca 


12-18 


ki 


( + ) 


poncensis 


0-1 


1-2 


ca 


16-20 


ki 


( + ) 



s — smooth ; k — keeled ; i = imbricate ; j =z juxtaposed ; I z= strong; ( ) = weak. 



Table 3 
Scale character comparison 



scales across snout at second canthal 

scales bordering rostral posteriorly 

scales between semicircles 

loreal rows 

interparietal from semicircles 

labials to center of eye 

scales bordering mental between 

infralabials 
fourth toe lamellae 



'uliventris 


occultu 


9-16 


9-13 


6-9 


5-9 


2-4 


2-4 


6-9 


2-6 


5-10 


2-6 


6-9 


8-13 


5-8 


4-6 


5-18 


14-18 



B R E V I O R A 

Mosceum of Coimparsitive Zoology 

Cambridge, Mass. October 29, 1965 Number 232 



HISPANIOLAN GIANT ANGLES (SAURIA, IGUANIDAE): 
NEW DATA AND A NEW SUBSPECIES 

By Ernest E. Williams 

Recent collections of Hispaniolan giant anoles {Anolis ricordii) 
by Albert Schwartz and his co-workers, and smaller numbers of 
specimens procured by Luc and George Whiteman for the Museum 
of Comparative Zoology, fill in many of the gaps in the distribu- 
tional record (Williams, 1962) and permit formal description of a 
new race. 

This study has been supported by National Science Foundation 
Grant GB 2444. The material of the new race has been divided at 
Dr. Schwartz's direction between the Museum of Comparative 
Zoology (MCZ), the American Museum of Natural History 
(AMNH), and his own collection (ASFS). A single specimen 
from the collection of Donald W. Buden (DWB) has been examined 
through the courtesy of Dr. Schwartz. 

The new material confirms the taxonomic utility of the scale 
characters previously employed in the analysis of geographic 
variation (Williams, 1962). However, although Dr. Schwartz has 
generously made available field notes on colors in life, variation in 
color and pattern is such, and there is such a repertoire available 
to any single individual, that the hesitations and cautions expressed 
in the 1962 paper on this subject seem still very pertinent. The 
attempt has been made to use color data as fully as possible, but it 
will be apparent in the following pages that I have succeeded only 
to a limited degree. While I have described below a new subspecies 
primarily on color pattern, I do so with the explicit admission that 
only crass differences in pattern or pattern repertoire seem at all 
usable. 

It is easiest to confront the new data in terms of the major 
distinguishable populations — i.e. the subspecies. I have therefore 
begun with the nominate race: 



2 BREVIORA No. 232 

Anolis ricordii ricordii Dumcril and Bibron 

New Records: Haiti. Departcmcnt de VOuest. Lancerouelle near 
Mirebalais, MVVj 69404; AOrebalais, MCZ 68479; Saltrou, MCZ 
69405. 

Dominican Republic. Monte Cristi Province. Laguna de Salo- 
dillo, 7 km SE Pepillo Salcedo, ASFS V 1470; 1 km W Copey, 
ASFS V 1269, V 1411-12, V 1470. 

The new specimens indicate that this race, in addition to 
occupying all of Haiti north of the Cul de Sac Plain, extends into 
the Dominican Republic in the iVIonte Cristi region. Hence the 
suggestion (Williams, 1962) that Ti-Guinen just west of Cape 
Haitien on the north coast of Haiti might be an intergrade area is 
in error. The extent of black on the head, nape, and shoulders of 
ricordii ricordii is apparently individually variable to a greater 
extent than I then realized and is perhaps less evident in the live 
animal than in the preserved animals. The presence of intense 
black patches on the anterior body and head is still a strongly 
marked characteristic of the males of this subspecies and is strongly 
correlated with the very low nape and dorsal crest scales, as well 
as with a high number of scales across the snout at the second 
canthal. 

One character not previously available has been the color of the 
dewlap. The dewlap in the Monte Cristi specimens is described as 
ranging through pale peach and brown, pale peach, very pale 
peach speckled with brown proximally to pale yellowish gray. 

Despite the new collections one eml)arrassment remains. No 
certain int(^rgrades between the two strikingly different forms 
ricordii and baleatus are yet known. In ^'iew of the erratic occur- 
rence of these giant anoles and the usual difficulty of collecting a 
series (see, however, below for Camp Perrin in southwest Haiti), 
it is not completely surprising that this should still be true. How- 
ever, the area in which intergrades may occur is being narrowed; 
on the north coast of the Dominican Republic between Monte 
Cristi and Santiago and in the center of Hispaniola between 
Mirebalais (MCZ 68479, 69404) and Santiago. This still leaves a 
very wide area of ignorance. 

A single specimen from Saltrou (MCZ 69405) narrows the 
geographic gap between ricordii ricordii and r. barahonae; how- 
ever, it does nothing to narrow the character gap. This is a male 
extremely heavity marked with intense black not only on nape and 
back of head but also on the flanks — more heavily than any 
other specimen; in squamation also it is quite typical of ricordii. 

Dr. Schwartz (pers. comm.) reports taking the ^Nlonte Cristi 
series in vines at night in a wooded area about a small cattle pond 



1965 HISPANIOLAN GIANT ANGLES 3 

between Copey and Pepillo Salcedo — an 0/ther\vise arid region. 
He states that in general ricordii can be captured with moderate 
regularity while sleeping at night in vin}^ tangles, especially where 
there are dense "mats" or "curtains" of vines under a canopy. 
This is a usually reliable technique which, however, sometimes 
fails; there may be areas in which the species just does not occur. 

Anolis kicordii baleatus Cope 

New records: Dominican Republic: La Vega Province. 4 km SW 
El Rio, ASFS X 8558: 0.3 mi (0.5 km) E El Rio, ASFS X 8114; 
12.8 km NW Bonao, ra. 1200 feet (360 m), ASFS V 4317. Puerto 
Plata Province. 11 km SE Sosua, ASFS V 1717. Duarte Province. 
5.6 km SW San Francisco de Alacoris, DWB 271 ; ca. 4 km NE 
Ponton (Rio Cuaba), ASFS \' 2987. Samana Province. 6 km E 
Sanchez, ASFS V 1904. Peravia Province. 1.1 mi (1.8 km) S San 
Jose de Ocoa, 1400 feet (425 m), ASFS V 723. El Seibo Province. 
3.5 mi (5.8 km) S Sabana de la Mar, ASFS X 7877; 2.1 mi (3.5 km) 
N El Valle, ASFS V 7861-62; 3 km N El Valle, ASFS V 3157-58. 
La Romana Province. 0.7 mi (1.2 km) W Higuey, ASFS V 854-55; 
1 mi (1.7 km) W Higuey, ASFS V 1038; 2.5 km NW Boca de 
Yuma, ASFS V 1136; 0.5 mi (0.8 km) NW Boca de Yuma, ASFS 
V 961-62. 

The new specimens of baleatus add no critical localities; only the 
eastern Dominican Republic is represented. No qualification of the 
characters previously reported for this race is recjuired; the 
squamation characters show a very limited variability. 

Again, the colors of dewlap and chin may be added as new 
characters. The dewlap is described as "orange" (adult cf), 
"dark orange" (adult cf), "very bright orange" (adult d^), "dull 
grayish orange" (adult 9 ), "dirty orange" (adult 9 ), "orange 
brown" (adult 9), "pale grayish orange" (juvenile cf), "dewlap 
skin charcoal, scales yellow-green" (ju\'enile 9 ). The chin of the 
male from southeast of So.^ua, Puerto Plata Province, with a bright 
orange dewlap is described as having the chin bright orange also. 
Other males from El Seibo Province, La Romana Province, and 
La Vega Province are described as having the chins some shade or 
other of green, vaiying from dark green through mottlings to 
yellow green. The significance of these differences is at present 
quite obscure; it would be particularly important to know the 
individual powers of color change. 

Anolls ricordii barahoxae Williams 

New records: Dominican Republic. Barahona Province. 8 km 
SE Las Auyamas, 2600 feet (7880 m), ASFS X 9676. Pedernales 



4 BREVIORA No. 232 

Province. 13.1 mi. (21.8 km) SW Enriquillo, ASFS V 4422. 

One new specimen from 8 km SE Las Auyamas, Barahona 
Province, which is quite near Polo, is almost topotypic and adds 
no new information. As in typical specimens, the dorsum has 
obscure, irregular, dark blotching. The lower flanks are boldly 
blotched black on white. Blotching here is usual but is not usually 
so bold. 

A second — ASFS V 4422 — from 13.1 miles (21.8 km) SW of 
Enricjuillo, Pedernales, is typical in squamation but peculiar in 
having very distinct small light spots on the flanks. The color in 
life of this specimen is given by Schwartz in his field notes as: 
''Dorsal ground color brown to grayish, with white (faintly 
bluish) dark-edged ocelli. Venter white with gray mottling and 
stippling. Dcivlap pale yellow, pink along outer edge. Head light 
brown. Soles of hands and feet pale yellow." 

It will be recalled that it was a specimen from Enriquillo 
(AMNH 51241) that caused some hesitation when barahonae was 
first described. In AAINH 51241 the pattern was thought to be 
obscure banding: the present specimen clearly shows spots tending 
to be vertically aligned — a condition which is easily transformed 
into vertical banding. It is possible that the ricordii populations 
in the vicinity of Enriquillo consistently show a distinctive pattern 
though characteristically barahonae in scjuamation. 

Anolis ricordii leberi new subspecies' 

Holotype: MCZ 80935, adult male from Camp Perrin, Haiti, 
native-collected for Albert Schwartz, 26 July 1962. 

Paratypes: MCZ 80936-42, 83982, A:\INH 93713-21, ASFS X 
3033 35, 3038-39, 3041-42, same data as type; MCZ 80943-53, 
AAINH 93722 36, ASFS X 3182, same data as type except 
collected 28 July 1962. 

Diagnosis: A subspecies of ricordii resembling barahonae in the 
size of the scales of the snout (4-6 across snout between second 
canthals) and in the slender, tapering, but small scales of the 
nuchal crest, but differing from barahonae in the higher and larger 
dorsal crest scales and in the presence in adults of both sexes of a 
pattern of bold alternating black and light lines on nape and 
flanks, the black hues more or less broken into and complicated b3' 
light cross-banding. (One or two specimens have this pattern very 
much reduced and juveniles are irregularly spotted, without light 
or black lines.) Dewlap in life is a rich yellow. 

'Named for David C. Leber who has prepared accurate and beautiful water 
color portraits from life of this and many otl.er West Indian anoles. 



1965 



HISPANIOLAN GIANT AXOLES 




a 



6 BREVIORA No. 232 

Comments. Almost all specimens show the broken, linear pattern 
quite clearly; probably all adults would do so in the darker phases 
of their pattern repertoire. 

So large a series from a single locality is quite extraordinary for 
any of the giant anoles. Albert Schwartz (pers. comm.) confesses 
that none were seen by himself and coworkers during the period in 
which this splendid sample was collected by local people. A. S. 
Rand {in Williams, 1962) has commented on the shyness and 
difficulty of catching the single individual of this same species 
which he saw in 1960. Obviously only the diligence of the local 
Haitians has permitted this glimpse of the real numbers of these 
anoles which, as inhabitants of a tree crown habitat, usually elude 
the general herpetological collector. 

From the point of view of the study of the geographic variation 
to which subspecific names are expected to call attention, a sample 
from one locality, however large, is not ideal. 

The single specimen from outside the Camp Perrin region which 
is referred to leheri — AICZ 38277, collected by P. J. Darlington, 
Jr., discussed in Williams, 1962 — has not been made a paratype 
of the new taxon. It is a juvenile which in squamation completely 
agrees with lebcri. However, while in its spotted, non-lineate 
pattern it is roughly like the single topotypic juvenile available 
(AICZ 83982), the pattern agreement is not striking enough and 
the preservation (showing an odd purplish tone not seen in any 
other specimen) is too different for confident assignment of the 
animal. 

The Darlington specimen is from Tardieu in the Massif de la 
Hotte, not far west of Camp Perrin, but this portion of the south- 
west peninsula of Haiti is a region of Hispaniola in which sharp 
character changes occur in other anoles (e.g. distichus, cybotes) 
within very short distances. It is a region also which, except for 
the road connecting Les Ca^^es and Jeremie on which Camp Perrin 
lies, is largely herpetologically unknown. It cannot be cavalierly 
assumed that onlj^ one ricordii race will be native to this area. 
Perhaps no surprises await us, but in the W^est Indies sharp faunal 
changes are not unusual and casual generalizations are never 
safely made. 

REFERENCE ClTtt) 

Williams, E.E. 

1962. Notes on Hispaniolan herpetology. 6. The giant anoles. Breviora, 
Mus. Comp. Zool., No. 155: 1-15. 

(Received 29 June 1965) 



1965 



HISPAXIOLAN GIANT ANGLES 




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BREVIORA 

Miiseiuiim of Comparative Zoology 

Cambridge, Mass. October 29, 1965 Number 233 

SOUTH AMERICAN ANOLIS (SAURIA, IGUANIDAE): 
TWO NEW SPECIES OF THE PUNCTATUS GROUP 

By Ernest E. Williams 



Anolis pundatus Daudin is the earliest discovered, most widely 
distributed, and morphologically the central member, of a group 
of forest anoles in South America. The males of pundatus (and of 
its race boulengeri O'Shaughnessy) have the rostral swollen, pro- 
jecting strongly beyond the lower Up. This is the first adumbration 
of a condition which becomes weirdly emphasized in three species 
which can be collectively called the proboscis anoles: A. laevis 
Cope, A. phyllorhinus Myers and Carvalho, and A. proboscis 
Peters and Orces. The strange rostral appendages in these three 
anoles are probably confined to males; at least all known specimens 
of these species are males. 

I here associate with this strange group of anoles, which lengthen 
the outline of the head by a swelling or soft protuberance, two 
undescribed species that elongate the bony frame of the snout 
itself. I associate these two with pundatus and its allies because 
of their general similarity in squamation despite the conspicuous 
difference in trend. This resemblance may be superficial and con- 
vergent, but I find the hypothesis of relationship useful as a 
provisional organization of the data. It is above all desirable at 
this moment to arrange and put in order the welter of Anolis 
species — even if somewhat artificially — and this is all the more 
desirable when still further species must be named. 

The two species here described are, for South American species, 
remarkably distinct. It is usual in South American anoles to be 
uncertain whether a distinctive population is an unrecognized 
species or not. No such question exists in these cases. The first 
species I describe has such strong differences from all described 
species that I name it : 



BREVIORA 



No. 233 



Anolis dissimilis new species 

Holotype: CNHAF 81369, an adult cf, Itahuania, upper Rio 
Madre de Dios, Madre de Dios Province, Peru, Kalinowski coll., 
15 October 1954. 

Diagnosis: Close to Anolis proboscis Peters and Orces, A. 
phyUorhinus Myers and Carvalho, A. laevis (Cope), and A. punc- 
tatus Daudin. From all of these the new species differs by the 
absence of any trace of swelling on the snout, by having the 
ventrals in oblique rather than transverse rows, by its more 
slender body and longer, narrower head (measured to interparietal 
scale 1.5 times tibia, ca. 3^ snout- vent length). 

Description. Head: All head scales wrinkled or striate, moderate 
to large posteriorly, small anteriorly. Seven scales across head 




Fig. 1. Aywlis dissimilis new species. Type, CNHM 81369. 



'For abbreviations see p. 11. 



1965 ANOLIS PUNCTATUS GROUP 3 

between second canthals. No frontal ridges but a shallow frontal 
depression. Five scales bordering rostral posteriorly. Rostral 
partly divided. Nasal scale anterior to canthal ridge, separated 
from rostral by one scale. Six scales between supranasals. 

Supraorbital semicircles broadly in contact, posteriorly partly 
separated from, anteriorly in contact with the supraocular disks, 
which consist of approximately nine enlarged scales, the antero- 
medial one much the largest, the remainder grading posteriorly 
into granules that become smaller toward the supraciliary margin, 
anteriorly grading more gradually into the moderately enlarged 
scales that are in contact with the single supraciliary. The latter 
short, continued posteriorly by undifferentiated granules, anteri- 
orly fused on both sides with the first canthal. Canthus distinct, 
canthal scales nme, the second the longest, diminishing gradually 
forward. Loreal rows five, the lowest row distinctly the largest. 
Temporal and supratemporal scales subgranular, smooth, grading 
into the large flat but wrinkled scales that surround the inter- 
parietal. Interparietal very much larger than the very small ear, 
in contact with the supraorbital semicircles. 

Suboculars in contact with supralabials, continued behind the 
eye by granules only, anteriorly in contact with the canthal ridge. 
Eleven supralabials to the center of the eye. 

Mentals distinctly longer than wide, in contact with two 
elongate throat scales. Sublabials large, wide, four in contact with 
infralabials on one side, five on the other. Central throat scales 
elongate, swollen, not keeled. 

Trunk: Middorsal scales feebly keeled, distinctly larger than 
flank granules but grading into them very gradually. 

Ventrals larger than dorsals, swollen, smooth, subcycloid, 
imbricate or subimbricate, arranged in oblique rows. 

Gular fan: Fan very large, extending well back on belly, scales 
narrow, smooth, much longer than ventrals, arranged in close-set 
lines. 

Limbs and digits: Hand and foot scales multicarinate. Largest 
arm and leg scales unicarinate, smaller than ventrals. About 17 
lamellae under phalanges 2 and 3 of fourth toe. 

Tail: Tail distinctly compressed, surmounted by a crest of 
enlarged keeled scales, very uniform in size, that give it a very 
serrate upper border. Verticils not evident. Lateral caudal scales 
irregular in size, smaller. Ventral surface of tail covered by two 
rows of keeled, more elongate scales, smaller than crest scales. 

Color: As preserved, essentially uniform dark above, lighter 
below. No evident pattern. 



4 BREVIORA No. 233 

Size: Snout-vent length 56 mm. 

The second species which requires description is most easily 
recognizable by a very narrow black line extending down the 
middle of the back. It is therefore described as: 

Anolis nigrolineatus, new species 

Holotype: MCZ 38940, Machalai, El Oro Province, Ecuador. 
Luis A. Perez coll. 

Paratype: USNM 12280, Guayaquil, Ecuador. No collector 
listed. 

Diagnosis: Similar to A . punctatus Daudin but differing in color 
(lighter and with a black midvertebral line and faint dark markings 
on flanks and limbs) and in squamation (fewer scales in contact 
with the rostral, fewer lamellae under fourth toe). 

Description. (Paratype differences in parentheses.) Head: 
Head scales moderate, posteriorly smooth, fiat, anteriorly weakly 
keeled; 10 (8) scales across snout between second canthals; frontal 
depression distinct ; 5 (7) scales border rostral posteriorly ; anterior 
nasal scale in contact with rostral ; 6 (5) scales between supranasals. 

Supraorbital semicircles separated medially by 2 (1) scales 
from each other on each side and by one row of granules from the 
well-defined supraocular disk of 10-13 flat scales. One or two 
elongate supraciliaries bordered medially by polygonal scales and 
continued posteriorly by granules. Canthus indistinct anteriorly, 
not forming a single continuous row; 6 loreal rows, subequal. 

Temporals and supratemporals both subgranular but the supra- 
temporals longer and grading gradually into the somewhat enlarged 
scales surrounding the interparietal, which is larger than the ear 
and separated from the supraorbital semicircles by 3 to 4 scales. 
Scales posterior to interparietal grading very gradually into nape 
scales. 

Suboculars weakly keeled, in contact with supralabials, anteri- 
orly separated from canthal ridge by 4 (3) scales, posteriorly grad- 
ing into temporals; 10-11 (8-9) supralabials to center of eye. 

Mentals wider than long, deeply indented by first sublabials; 
2-3 sublabials in contact with inf ralabials ; 4 gular scales in a 



^The exact locality of collection is in doubt. All specimens in the Perez 
collection come from El Oro Province and most from the vicinity of Machala 
but exact data were not kept. It would be preferable to make the USNM 
specimen the type of this very distinct species except that it is not as well 
preserved and the locality is not — with any probabiHty —- more exact. 



1965 



ANOLIS PUNCTATUS GROUP 




Fig. 2. Anolis nigrolineatus new species. Type, MCZ 38940. 



6 BREVIORA No. 233 

gentle forward arc in contact with mentals between sublabials. 
Gular scales smallest medially grading laterally into sublabials. 

Trunk: Two middorsal rows slightly but distinctly enlarged, 
swollen on nape, grading into flank granules. Ventrals larger, 
smooth, imbricate. 

Gular Jan: Very large, extending posteriorly onto more than 
the first third of the belly, rather densely scaled, the scales at the 
edge at least the size of the ventrals; smooth, lateral scales 
smaller. 

Limbs: Scales on limbs unicarinate, the largest on hind limb 
larger than ventrals. Supradigital scales multicarinate. Eighteen 
lamellae under phalanges ii and iii of fourth toe. 

Tail: Compressed, without verticils or dorsal crest. Greatly 
enlarged postanals present. 

Color: As preserved, light brown with faint and broken streaking 
or reticulation. A black middorsal line two scales wide. Top of 
head including area around interparietal, but not supratemporals, 
blackish also. A dark postocular spot. Dewlap light at edge but 
with a long black spot at base. 

Comparative material examined: A. punctafus (including the sub- 
species boulengeri) : Specimens from many localities in Brasil, 
Bolivia, Peru, Ecuador and the Guianas. 

.4. laevis: Type only, ANSP 11368, between Moyabamba and 
Balsa Puerto on the Rio Huallaga, E. Peru. 

A. phyllorhinus: [Type from Borba, lower Rio Madeira, Ama- 
zonas, Brasil, not seen], DZ 7118, Rio Tapajoz, Jacareacanga, 
Para, Brasil. 

A. proboscis: Type only, MCZ 54800, vicinity of Cunuco, N.W. 
of Mindo, on S. bank of Rio Mindo, Pichincha, Ecuador. 

Definition of the Anolis punctatus species group 

Reference of two new species to a "punctatus''^ group is an 
obligation to define the group. I attempt this after rather than 
before describing the new species because of the characteristic 
difficulties which attend all definitions of species groups within 
Anolis based on externals. This is a genus pervaded with con- 
vergence and parallel adaptation to such an extent that only those 
species series which are also obvious superspecies are clear, discrete 
units. Just beyond the level of the superspecies it is often not 
difficult to assemble larger clumps of species that seem evidently 
related. But here the problem of exclusion rather than inclusion 
enters. Each added species leads insensibly toward species still 
more remote from the core species of the putative group. 



1965 AXOLIS PUNCTATUS GROUP 7 

This situation is well exemplified by the pimclatus group as here 
seen. A definition can be proposed but there will be multiple 
exceptions to certain characters, and certain fringe species which 
could as well be included or excluded. 

A tentative definition (with exceptions) follows: Species of 
moderate size (but nigrolineatus is rather small, and laevis rela- 
tively large). The snout is produced either by bony structure, 
swelling, or soft proboscis. The color is probably green in life 
(usually purples and blues in alcohol), perhaps an exception in 
nigrolineatus. The ear is small and rather ventrally placed (on the 
level of mouth). The nostril is separated by a single prenasal scale 
from the rostral. The head scales are flat, pavimentose (even in 
boulengeri, which has keeled ventrals and keeling of the dorsal 
scales). The loreal rows are few (as few as two in laevis, as many as 
seven in punclatus. There are no or only a few scales between the 
supraorbital semicircles (0-2). The interparietal, larger than the 
ear, is of moderate size (large in laevis, dissimilis and proboscis) and 
separated from the semicircles by to 4 scales. Suboculars broadly 
in contact with supralabials. Mental deep, not wide (widest in 
nigrolineatus). Well developed sublabials present. Dewlap large, 
scales in rows narrowly separated b}^ naked skin (scales not in 
rows in proboscis). Middorsals not or not appreciably larger than 
flank scales (a dorsal crest in proboscis). Ventrals smooth (keeled 
in boulengeri which, howe\'er, intergrades with smooth-scaled 
punctaius), squarish, transverse. Tail more or less compressed with 
double row of scales dorsally (a single crest in dissimilis and 
proboscis) . 

This is an extensive list of similarities. However, the differences 
between species, emphasized above by the exceptions, are as 
striking as the similarities. They are of many sorts and it is 
natural to inquire whether the differences are less important than 
the common characters. It will be useful, therefore, to examine the 
differences in some detail. 

1. Snout differences. A. nigrolineatus is in this regard not very 
different from many anoles not closely related to it; the bony 
structure of the snout itself has been stretched into a tapering, 
])luntly pointed structure. A. dissimilis carries the condition of 
nigrolineatus to an extreme. In contrast, the bony snout of A. 
punctatus is very little modified, but the rostral scale is swollen, 
protuberant. A. laevis has this scale produced into a broad-based 
flexible appendage. A. phyllorhinus has a narrow flexible appendage 
above the triangular rostral scale, this appendage having small 
granular scales. A. proboscis is very simjiar in the general con- 
formation of the area but the scales on the flexible proboscis are 



8 BREVIORA No. 233 

elongate. The differences here imply, as we have already sug- 
gested, that no single linear series can be envisioned; at least two 
are required. It is, however, possible to suppose that there has 
been radiation from a central type — perhaps punctatus, perhaps 
an ancestor of punctatus. 

2. Interparietal size. The interparietal is large and in direct 
contact with the semicircles in so many species of diverse relation- 
ships, and the interparietal may so often differ in size and in 
distance from the semicircles in closely related species that this 
character is probably of minimal systematic value above the 
species or superspecies level. 

3. Dewlap squamation. Again a character subject to much 
parallel modification and often different within a superspecies. 
A. proboscis is anomalous among the compared species in having a 
rather uniform squamation of the dewlap rather than scales in 
distinct, separated rows, but this is probably of no major sig- 
nificance. 

4. Dorsal crest. A. proboscis is again very peculiar in having a 
dorsal crest of strongly enlarged sub-triangular scales. Such a crest 
is known in several West Indian so-called giant anoles but is very 
unusual in mainland species. It does, however, occur in some 
Guatemalan A . pentaprion — whether as an anomaly or a popula- 
tion character is unknown. Special though this feature seems in 
A. proboscis, it is hard to regard this as more significant than the 
extraordinary proboscis — so like that of phyllorhinus in which 
there is no hint of a dorsal crest. 

5. Tail. The difference between a tail with two rows of scales 
dorsally and one with a pronounced single crest is a very obvious 
one. Schmidt (1939), in describing Anolis barkeri, and also Myers 
and Carvalho (1945), in describing A. phyllorhinus, have made 
much of the double-rowed condition — a feature which does 
appear to be unusual in anoles. There is usually in Anolis only a 
single row, whether or not this is produced into a crest. The 
systematic value of the double row is, however, much diminished 
by just the case in which Schmidt first used it: A. barkeri is a 
Mexican species which on osteological grounds (Etheridge, 1959) 
belongs to a very different section of the genus from the South 
American species in which this peculiarity is otherwise known. 

In my judgment these differences, though disturbing at first 
glance, do not provide serious difficulty for a concept which unites 
all these species as a unit group. There are strong cross re- 
semblances between species that on other characters would be 
separated. Thus, dissimilis shares with proboscis the character, 



1965 ANOLIS PUNCTATUS GROUP 9 

unusual in South America, of a crested tail, but in the nasal 
appendage and snout structure proboscis resembles phyllorhinus 
and is very different from dissimilis. 

More awkward for the desiderate goal of taxonomic clarity are 
the species I have described as "fringe species." These are: 
transversalis (including buckleyi); the solitarius-tigrinus super- 
species ; jacare. 

All of these have a double row of scales dorsally on the tail, all 
have smooth ventrals, few loreal rows, pavimentose dorsal head 
scales, few or no rows between the supraorbital semicircles, sub- 
oculars broadly in contact with supralabials, mental deep, not 
wide, well developed sublabials. 

However, the ear is rather large, the color is complex with much 
cross-barring and spotting. The species of the solitarius-tigrinus 
series are all of small size (40-50 mm snout- vent length), jacare and 
transversalis are of moderate size. One peculiar feature which 
unites this set of species but is untrue or unknown for all those I 
have referred to the punctatus group (untrue for punctatus, un- 
known for the others) is the presence of black pigment in the 
female dewlap, and its absence in the male structure (which is also 
somewhat better developed). 

To include these species in the punctatus group would seem to 
enlarge it too much; yet a considerable degree of affinity seems 
probable. 

An evolutionary perspective for the punctatus group 

The first described proboscis anoles — A . laevis Cope (Rio 
Huallaga, Peru) and ^4. phyllorhinus Myers and Carvalho (lower 
Rio Madeira, Brasil) were Amazonian. This is also true of A. 
dissimilis (on the upper Rio Madre de Dios). But A. proboscis 
Peters is from Pichincha Province in Ecuador, west of the Andean 
water shed, and A. nigrolineatus is from the Pacific lowlands near 
Guayaquil. A. punctatus — the central species of this putative 
complex — is much more widespread than any of the other species 
occurring in Amazonia, the Guianas and the forests of eastern 
Brasil, but neither it nor its western race (with keeled ventrals) — 
boulengeri — ever transgresses into the Trans- Andean Province. 

There is, thus, in this species group — if it is a reality — no 
special geographic pattern except that of being clearly and wholly 
South American. 

That this group is part of a wider autochthonous South American 
section of Anolis has been demonstrated by Richard Etheridge 
(1959). The species of the punctatus group, all examined radio- 
graphically by him, and a wider Icircle of forms which include such 



10 BREVIORA No. 233 

species as bocttgeri, chloris, fasciatus, fraseri, jrenatus, insignis, 
jacare, latifrons, microtus, minis, nasojrontalis, peraccae, pseudoH- 
grinus, solitarius, squamulatus, tigrinus, transversalis, ventrimacu- 
latus, are all characterized by possession of posterior caudal 
vertebrae without transverse processes and without autotomy 
septa, by the possession of four parasternal chevrons attached to 
the ribs and by having the latei-al arms of the interclavicle diver- 
gent from the proximal parts of the clavicles. This is an assemblage 
of characters that Etheridge has demonstrated to be quite distinc- 
tive, and geographically quite coherent, occurring in species of 
mainland South America (but not those of the West Indies or 
]\Ialpelo Island) and also in two or three species — jrenatus, 
insignis and microtus — present in extreme southern Central 
America. 

This is a very varied series in everything but these distinctive 
skeletal characters. These anoles are very different in size (includ- 
ing both dwarfs and giants), and in squamation (Table 2). The 
series, therefore, has every appearance of being an old assemblage 
which has had the time to diversify and which has exploited its 
opportunities. 

The autochthonous South American section of Anolis shares 
South America with a group clearly not autochthonous but with 
its stronghold and center of origin to the north in Central America 
and Mexico. Though the latter group is clearly an invader from the 
north, it has reached every part of the total range of Anolis in 
South America. It is amazing that the ranges of these two groups 
of divergent history should be so closely coterminous in mainland 
South America. 

This invader group is distinguished on Etheridge's osteological 
characters by having caudal vertebrae with caudal autotomy septa, 
with transverse processes which are inclined forward, and with an 
interclavicle the lateral arms of which are in contact with the 
clavicles. Osteologically, therefore, they are quite distinct from 
the old South American anoles. In squamation, as Table 2 shows, 
there is broad overlap. It is, therefore, impossible on external 
characters to make a separation of the two groups. Indeed, 
species belonging to the two groups have sometimes been confused 
with one another, and, in other cases, while the species characters 
permit ready separation, it will still be impossible on externals to 
allocate the species to group other than randomly. 

Yet in bias and trend the two groups do differ. This too is shown 
in Table 2. In toe lamellae the bias of the alpha group is to higher 
numbers, that of the beta group to lower numbers. This character 



1965 ANOLIS PUNCTATUS GROUP 11 

(and probabl}^ more obscurely some of the others) is a reflection 
of an ecological bias in the two groups: the alpha group includes 
more deep forest, highly arboreal species, the beta group more 
species of open country — ground, grassland, or bush. 

In ecology, as in so much else, there is strong overlap, but the 
bias or trend is clear. At the extreme of the beta series is an anole 
that actually lives in or at least takes refuge in holes in the ground 
(Ruthven, 1922), that in fact has abandoned wholly its arboreal 
heritage and with it the clinging hairs on the toe lamellae so 
characteristic of all other anoles. This anole, though only the 
extreme of its series, is customarily placed in a distinct genus; it is 
Tropidodactylus onca. 

At the opposing extreme in the alpha series are probably to be 
placed the proboscis anoles — which again might be placed in a 
genus apart did they not seem to achiev^e their distinctive rostral 
structure in different ways. Neighbors to these in the extreme 
wing of the alpha series are pundatus and, if I interpret matters 
rightly, the two new species that are described in this paper. 

Acknowledgments: I am deeply grateful to Dr. Gustavo Orces-V. 
for allowing me to examine his Ecuadorian A nolis — material of 
extraordinary value — and for donating the type of A. nigro- 
lineatus to the Museum of Comparative Zoology. Dr. Doris 
Cochran of the United States National Museum (USNM), 
Dr. Robert Inger of the Chicago Natural History Museum 
(CNHM), Dr. James E. Bohlke of the Academy of Natural 
Sciences, Philadelphia (ANSP), and Dr. Paulo Vanzohni, De- 
partamento de Zoologia, Sao Paulo (DZ) have generously allowed 
study of material under their care. James D. Lazell, Jr. prepared 
the figures of A. nigrolineatus and Nicholas Strekalovsky those of 
A. dissimilis. Dr. Mary Willson prepared the map. This series of 
studies on South American and other Anolis is supported by Nat- 
ional Science Foundation Grant GB 2444. 

literati:re cited 

Cope, E. D. 

1875. Report on the reptiles brought by Professor James Orton from the 
middle upper Amazon and western Peru. Jour. Acad. Nat. Sci. 
Philadelphia, (2) 8: 159-188. 
Etheridge, R. 

1959. The relationships of the anoles (Reptilia: Sauria: Iguanidae): 
an interpretation based on skeletal morpholog3\ University 
Microfilms, Ann Arbor. 236 pp. 
Myers, G. S. and A. Carvalho 

1945. A strange new leaf-nosed lizard of the genus Anolis from Ama- 
zonia. Bol. Mus. Nac, Rio de Janeiro, N. S. Zool., No. 43: 1-14. 



12 



BREVIORA 



No. 233 



Peters, J. A. and G. Orc6s-V. 

1956. A third leaf-nosed species of the lizard genus Anolis from South 
America. Breviora, No. 62: 1-8. 
Schmidt, K. P. 

1939. A new lizard from Mexico with a note on the genus Nor ops. 
Zool. Ser. Field Mus. Nat. Hist., 29: 7-10. 
Rtjthven, a. 

1922. The amphibians and reptiles of the Sierra Nevada de Santa 
Marta, Colombia. Misc. Publ. Mus. Zool. Univ. Michigan, 8: 
1-69. 

(Received 26 July 1965.) 



80' 

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♦ A. proboscis 
©A dissimilis 

200 mi, 

1 ' 



Fig. 3. Map of the distribution of pundatus group anoles except A. punc- 
tatus itself. The exact limits of the latter's very wide range are not known. 



1965 



ANOLIS PUNCTATUS GROUP 



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No. 233 





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1965 



ANTOLIS PUNCTATUS GROUP 



15 



number of toe lamellae 
scales across snout 
scales between semicircles 
dorsal scale rows enlarged 



Table 2 

Character range in South American anoles 

alpha anoles beta anoles 

14-30 (mode 18-21) 10-27 (mode 14-18) 

4-25 7-20 

0-5 (7 *) 0-4 

0-2 0-12 



* Though several alphas range up to five scales between the supraorbital 
semicircles, the maximum reported liere occurs in two exceptional specimens 
of A. princeps Boulenger (= ? frenatus Cope). 



R E V I O R A 

Mmseiuimni of Comnipsirsitive Zoology 

Cambridge, ]\Iass. October 29, 1965 Number 234 

STUDIES ON NEOTROPICAL POMPILIDAE 
(HYMENOPTERA) 

I. THE GENUS AGENIOIDEUS ASHMEAD 
IN SOUTH AMERICA 

By Howard E. Evans 



The rich pompihd fauna of the neotropics has unfortunately been 
subjected to a good deal of bad taxonomy. The reasons are the 
usual ones: workers have been content to erect new species and 
genera without having seen the types of described species, which 
are widely scattered throughout Europe and North and South 
America; each worker has tended to use his own system of classi- 
fication; and many parts of the neotropics remain very inade- 
quately collected. The two papers of Nathan Banks on the South 
American Pompilidae [Bull. Mus. Comp. Zool., 96: 311-525 
(1946), and 99: 371-486 (1947)] are useful, but suffer from all three 
of these deficiencies as well as from Banks' failure to cite references 
and his failure to cover quite a number of described species. 

It will be many years before the taxonomy of the neotropical 
Pompilidae can be brought to a reasonably high level. In the mean- 
time, I hope to publish a series of short papers covering such 
segments of the fauna as I am able to work out to my satisfaction. 
In addition to the specimens which Banks studied, I have seen 
much additional material from southeastern Brazil collected by 
Fritz Plaumann, several collections made in Chile and Peru by 
Luis Pena, and the material from a recent trip to Peru, Chile, and 
Argentina by C. C. Porter of Harvard University. I shall also 
make use of other material as available, including specimens from 
my own collecting in Mexico, Central America, and the West 
Indies. A review of the subfamily Pompilinae in Mexico and 
Central America is being published elsewhere (Alem. Amer. Ent. 
Soc, in press). 



2 BREVIORA No. 234 

The genus Agenioidcus is primarily characteristic of the warmer 
parts of the Holarctic region. There is considerable structural 
diversity within the genus, but the group nevertheless holds 
together well on the basis of wing venation, the weak development 
of the pulvillar pad and comb, and other features which I outhned 
in 1950 (Trans. Amer. Ent. Soc, 75: 189). One of the four North 
American species is confined to Mexico, two others range from 
southern United States into Mexico, and another (humilis Cresson) 
ranges all the way from southern Canada to Panama. 

Banks did not regard Agenioideus as occurring in South America. 
However, three of the four species which he included in Serico- 
pompilus do, in fact, belong in Agenioideus. The fourth species, 
exilis Banks, has only two submarginal cells and other features 
characteristic of the genus Euplaniceps, to which genus this species 
is here reassigned. To the best of my knowledge, Sericopompilus 
does not occur in South America. The three species of Agenioideus 
(which I regard as only two) are closely related and somewhat 
divergent from the North American Agenioideus, although still 
falling readily within that genus. Although the females lack a 
tarsal comb, as in the subgenus Gymnochares, the over-all resem- 
blance is perhaps greatest to Agenioideus sensu stricto. A new sub- 
genus, Enbanksia, is here proposed for this group. This group may 
represent the descendants of a stock of Agenioideus which succeeded 
in entering South America sometime during the Tertiary and which 
underwent a small radiation there. 

A review of this small group seems justified at this time not only 
to clarify the correct generic position of these species, but also to 
properly characterize the male sex. Of the males assigned by 
Banks to South American Sericopompilus, only one (the allotype 
of accoleus Banks) properly belongs with this group. Also, Banks 
made several errors of fact which should be corrected; for example, 
he stated that the clypeus of the type of accoleus is "fully three and 
one-half times as broad as long" when in fact it measures 2.4 X as 
broad as long. He also did not have sufficient material to appreci- 
ate the variation in some of the characters used, for example in the 
dentition of the claws. 

Enbanksia new subgenus 

Type species. — Sericopompilus accoleus Banks, 1947. 

Subgeneric characters. — Small wasps (4-9 mm), the females 
(and males of one species) with banded wings, both sexes with the 
tibial spurs entirely white and with a white spot near the base of 
the hind tibiae; males with the apical abdominal tergite white; 



1965 AGENIOIDEUS IN SOUTH AMERICA 3 

integument smooth and polished, except the front sometimes 
micropunctuate; body virtually without erect setae but extensively 
clothed with silvery pubescence, which is especially conspicuous 
on the posterior slope of the propodeum. Clypeus wider than lower 
face, truncate, with a slightly raised apical rim; antennae relatively 
short for the genus, third segment in the female not equal to the 
upper interocular distance, in the male not much if any longer than 
second segment; ocelli in a broad, flat triangle. Thoracic dorsum 
forming a rather smooth arc except the scutellum and metanotal 
disc prominent, somewhat compressed; pronotum sloping smoothly 
in front, its posterior margin broadly angulate or subangulate; 
postnotum somewhat polished, constricted on the midline; propo- 
deal slope smooth and even; middle and hind tibiae strongly 
spinose, but the front tarsus without a comb, the apical tarsal seg- 
ments not spined beneath. Claws dentate, the tooth sometimes 
close to the outer ray, such that the claws appear almost bifid ; fore 
tarsal claws of male alike, both dentate. Wing venation similar to 
that of other Agenioideus (see Evans, 1950, Trans. Amer. Ent. 
Soc, 75: 190, fig. 58). Abdominal segments showing no strong 
tendency to telescope; male subgenital plate moderately com- 
pressed, simple (Figs. 3, 4); male genitalia without basal booklets, 
but with some stout setae arising from a lobe at the base of the 
digitus (Figs. 1,2). 

Remarks. — This subgenus is named for N. Banks, who described 
all its presently known components. 

KEY TO SPECIES AND SUBSPECIES 

Females 

1. Antennae short, third segment 2.0-3.5 X as long as thick, equal to 

.35-.45 X the upper interocular distance; wings with only one strong 
band, sometimes with a weak second band over the basal vein; legs be- 
yond the basal part of the femora rufous 1. minutus (Banks) 

Antennae longer, third segment 4.5-5.0 X as long as thick, equal to 
.65-.S0 X the upper interocular distance; wings strongly twice-binled; 
legs fuscous, at least in large part "2 

2. Pronotum and mesoscutum ferruginous, except the former with somi p dj 

yellow markings; front relatively narrow, middle interocular distance 

.56 X transfacial distance 2a. accjleus accohus (Baaks) 

Pronotum and mesoscutum fuscous, e.xcept the former with pale yellow 
markings on the collar and often along the posterior margin; front 
broader, middle interocular distance .59-.61 X transfacial distance. . . . 
2b. accoleus lucanus (Banks) 



BREVIORA 



No. 234 



Males 

1. Wings clear hyaline, with a whitish bloom; legs rufous beyond basal parts 

of femora; third antennal segment wider than long; subgenital plate 

narrowly truncate apically (Fig. 3) 1. minutus (Banks) 

Wings banded; legs mostly fuscous; third antennal segment longer than 
wide; subgenital plate subacute apically (Fig. 4) 2 

2. Prouotum, mesoscutum, scutellum, metanotum, and upper part of meso- 

pleura ferruginous; middle interocular distance .59 X transfacial dis- 
tance 2a. accoleus accoleus (Banks) 

Thorax black; middle interocular distance .62-. 66 X transfacial dis- 
tance 2b. accoleus lucanus (Banks) 

1. Agenioideus (Enbanksia) minutus (Banks) new combination 

Sericopompilus ininutus Banks, 1947, Bull. Mus. Comp. Zool. Harvard, 
99:435. [Type: 9, BRAZIL: Tres Lagoas, Matto Grosso, 6-10 Dec. 
(Cornell Univ. Exped.) (Cornell Univ.)] 

Female. — Length 5.0-6.5 mm; fore wing 4.6-6.0 mm. Black, 
except as follows: pronotum with pale yellow markings on the 
collar and along the posterior margin: clypeus and mandibles 
rufo-testaceous, the latter darker apically; antennae ruf ©-testa- 
ceous except second and apical few segments usually somewhat 
infuscated; legs beyond the trochanters (or at least beyond the 




Fig. 1. 
aspect. 
Fig. 2. 
Fig. 3. 
Fig. 4. 



Male genitalia of Agenioideus (Enbanksia) minutus (Banks), ventral 

Same of A.{E.) accoleus lucanus (Banks), left side omitted. 
Tip of subgenital plate of male A.{E.) minutus (Banks). 
Subgenital plate of male A.(E.) accoleus lucanus (Banks). 



1965 AGENIOIDEUS IN SOUTH AMERICA 5 

middle of the femora) bright ruf o-castaneous ; fore wings with a 
strong brown band from the marginal cell through the outer 
discoidal cell, some specimens weakly tinged with brown around 
the basal vein. Clypeus 2.9 X as wide as high. Front broad, 
middle interocular distance .65-.69 X transfacial distance; upper 
interocular distance .88-.94 X lower interocular distance; 
POL : OOL = 4:3. Third antennal segment three only slightly 
if at all longer than four, measuring 2.0-3.5 X as long as thick, 
.35-.45 X upper interocular distance. Propodeum somewhat more 
strongly convex and abdomen stouter than in the species which 
follows. All claws weakly dentate. 

Male. — Length 5-6 mm; fore wing 4-5 mm. Black; pronotum 
marked with pale yellow on the collar and in a broad band along 
the posterior margin ; coxae, trochanters, and basal parts of femora 
black, legs otherwise ferruginous except tarsi dusky; apical half of 
mandibles rufo-testaceous ; antennae dark brown except scape pale 
beneath; wings clear hyaline, with a whitish bloom, veins and 
stigma brown. Clypeus 3 X as wide as high, truncate. Head 
broad, vertex forming a strong arc above the eye tops; front 
broad, middle interocular distance .67-.69 X transfacial distance; 
upper and lower interocular distances subequal, but middle inter- 
ocular distance 1.2-1.3 X upper interocular distance; POL: OOL = 
7:4. Third antennal segment very small, slightly wider than long, 
not longer than segment two or more than half the length of four. 
Postnotum sUghtly shorter than metanotum ; propodeum in profile 
forming a somewhat higher arc than in the following species. Sub- 
genital plate narrowly truncate apically (Fig. 3) ; genitalia as 
shown in Figure 1. 

Distribution. — Southern Brazil, Paraguay, and eastern Peru. 

Specimens examined. — S 9 9, 3cf'cf. BRAZIL: 7 9 9,1 d". 
Nova Teutonia, Santa Catarina, Nov.-Feb. (F. Plaumann) 
[Mus. Comp. ZooL, Cornell Univ., Coll. G. R. Ferguson]; 1 9, 
Tres Lagoas, Alatto Grosso [type, Cornell Univ.]. PARAGUAY: 
1 d", Caacupe, Oct. 25, 1955 (F. Schade) [Coll. G. R. Ferguson] 
PERU: 1 d", Avispas, Madre de Dios, 400 meters, Sept. 10-30' 
1962 (L. Pena) [Mus. Comp. Zool.]. 

2a. Agenioideus (Enbanksia) accoleus accoleus (Banks) new 
combination 

Sericopom.pilus accoleus Banks, 1947, Bull. Mus. Comp. Zool. Harvard, 
99: 433. [Type: 9, BRAZIL: Maracajii, Matto Grosso, Apr.-May 1937 
(G. Fairchild) (Mus. Comp. Zool.)] 



6 BREVIORA No. 234 

Female. — Length 8.5 mm; fore wing 8.7 mm. Dark brownish- 
fuscous except as follows: mandibles and clypeus pale ferruginous, 
the latter with yellow blotching on each side; basal two antennal 
segments pale ferruginous, the remainder brownish ; pronotum and 
mesoscutum ferruginous, except the collar marked with pale yellow 
and the posterior pronotal margin indistinctly marked with 
yellowish; front tibiae and tarsi testaceous; wings hyaline, fore 
wing with a strong brown band across the basal vein and a broader 
band across the wing at the marginal cell. Clypeus 2.4 X as wide 
as high. Front relatively narrow, middle interocular distance 
.56 X transfacial distance; upper interocular distance .80 X lower 
interocular distance; POL : OOL = 8:5. Third antennal segment 
5 X as long as thick, .8 as long as the upper interocular distance. 
Slope of propodeum low and even, median line somewhat impressed. 
Claws strongly dentate, the inner ray acute, in the front tarsi the 
two rays rather close together, subparallel. 

Male. — Length 7 mm; fore wing 6.5 mm. Dark brownish- 
fuscous except pronotal collar marked with pale yellow, remainder 
of pronotum ferruginous; mesoscutum, scutellum, metanotum, and 
upper two-thirds of the mesopleura also ferruginous; antennae 
brown except scape whitish below, flagellum testaceous below ; legs 
brown except for the usual spot on the hind tibiae and the pale 
spurs; fore wings with a dark band at the marginal cell as in the 
female, but with only a weak, narrow infuscation at the basal vein. 
Clypeus 2.5 X as wide as high. Front rather narrow, middle inter- 
ocular distance .59 X transfacial distance; upper interocular dis- 
tance very slightly exceeding lower interocular ; POL : OOL = 7 :5 ; 
vertex weakly humped at the ocellar triangle. Third antennal seg- 
ment rather short although longer than second, measuring about 
L2 X as long as thick. Postnotum nearly as long as metanotum; 
slope of propodeum very low and even. Terminalia as described 
under accoleus lucanus. 

Distribution — Brazil (states of Matto Grosso and Sao Paulo) . 

Specimens examined. — Only the type and allotype, the latter 
from Campinas, Sao Paulo, March 1924 (F. X. Williams) [Mus. 
Comp. Zool.]. 

2b. Agenioideus (Enbanksia) accoleus lucanus (Banks) new 
status, new combination 

Sericopompilus lucanus Banks, 1947, Bull. AIus. Comp. Zool., Harvard, 
99: 434. [Type: 9, BRAZIL: Nova Teutonia, Santa Catarina, 25 Jan. 
1939 (F. Plaumann) (Mus. Comp. Zool.)j 



1965 AGENIOIDEUS IN SOUTH AMERICA 7 

Female. — Length 6.5-8.5 mm; fore wing 6.5-9.0 mm. Black, 
except as follows: mandibles and clypeus pale ferruginous, the 
latter sometimes with obscure yellowish blotching, sometimes 
infuscated on the upper half; basal 2.0-2.5 antennal segments 
testaceous; pronotum marked with pale yellow on the collar and 
usually with a more or less complete yellowish band along the 
posterior margin; legs fuscous, front tibiae and tarsi somewhat 
paler than the remainder; wings banded as in the nominate sub- 
species. Middle interocular distance .59-. 61 X transfacial dis- 
tance; upper interocular distance .82-.86 X lower interocular. 
Third antennal segment 4.5-5.0 X as long as thick, equal to .65- 
.75 X upper interocular distance. Claws of front tarsus variable, 
the inner ray close to the outer ray or somewhat removed from it. 
Other features as described for a. accoleus. 

Male. — Length 6.0-6.5 mm; fore wing 5.0-5.5 mm. Dark 
brownish-fuscous except mouthparts and clypeus rufo-testaceous, 
basal two antennal segments and under side of segment three (and 
sometimes four) testaceous, front legs beyond the trochanters and 
also middle tibiae sometimes rufo-testaceous; wings banded as in 
female but the band over the basal vein weak. Aliddle interocular 
distance .62-.66 X transfacial distance; upper and lower inter- 
ocular distances subequal. Third antennal segment 1.3-1.6 X as 
long as thick, sometimes barely shorter than fourth segment. Sub- 
genital plate subacute apically (Fig. 4). Genitalia differing from 
those of minutus in having both the aedoeagus and the digiti more 
broadly expanded apically (Fig. 2). 

Distributioyi — Southern Brazil (Santa Catarina). 

Specimens examined. — 13 9 9 , 4 6^ cf , all from Nova Teutonia, 
Santa Catarina, Oct.-March (F. Plaumann) [AIus. Comp. Zool., 
Coll. G. R. Ferguson]. 

(Received 14 June 1965) 



BREVIORA 

Museiuim of Comparative Zoology 

Cambridge, Mass. November 19, 1965 Number 235 



A NEW SALAMANDER OF THE GENUS 

CHIROPTEROTRITON (GAUD ATA: PLETHODONTIDAE) 

FRO.AI MEXIGO. 

By GeorCxE B. Rabb 

Chicago Zoological Park, Brookfield, Illinois 

As Martin (1958:12) has noted, the karst caves of the Sierra 
Madre Oriental of Mexico are important amphibian habitats. Most 
of the salamander species found in such caves are widespread forms, 
but at least one described species from San Luis Potosi, Chiro- 
pterotriton mosaueri, is probably restricted to caves. Another 
species apparently in this category is described below. For it, I 
propose the name : 

Ghiropterotriton magnipes sp. nov. 
Figs. 1, 2 

Holotype. — Museum of Gomparative Zoology 30607, a young 
adult male collected in the Gueva de Potrerillos, about 2 km WSW 
of Ahuacatlan, which is approximately 8 km SW of Xilitla, San 
Luis Potosi, Mexico, on 18 November 1955 by Alejandro Villa- 
lobos F. 

Parafypes. — MGZ 44033, GNHM 142457, and four uncata- 
logued specimens from the type locality in the collection of the 
Instituto Politecnico Nacional, Mexico, collected on 20 January 
1952 by F. Bonet; UMMZ 125423 (GH 9878), collected at the tvpe 
locality in July 1964 by Kraig Adler; GNHM 142458, UMMZ 
125179, and three uncatalogued males in the Instituto Politecnico 
collection from Gueva del Madrono, near Laguna Golorada, 
IMunicipio de Landa de Alatamoros, Queretaro, Alexico, collected 
on 16 January 1952 by F. Bonet. 

Diagnosis — A species distinguished from all others in the genus 
by its very fully webbed feet, large size (adults 40-60 mm snout- 
vent length) and greater number of teeth (average of 79 maxillary- 
premaxillary in adult males). 



2 BREVIORA No. 235 

Description of the holotype. — General habitus long and de- 
pressed. Tail relatively short (slightly less than snout- vent length), 
rounded in cross section. Basal constriction of tail not indicated. 
Limbs relatively long; adpressed limbs overlap by two or more 
costal spaces. Feet broad ; toes and fingers joined by thick webbing. 
On foot, webbing indented between digits 2-3, 3-4, and 4-5 to 
level of distal end of penultimate phalanx of digits 3 and 4 (Fig. 1). 
Terminal pads, subtending digits 2-5 on foot and 2-4 on hand, 
present but not prominently developed, palmar surfaces of hand 
and foot otherwise smooth. Contours of digits not prominent 
dorsally, except for pigmentless areas marking some joints. Head 
flat and wide, eyes large and bulging. Prominent subocular groove 
extending in an arc from anterior corner of eye to position beneath 
posterior corner of eye. Postocular and other cephalic and nuchal 
grooves weakly developed. Viewed from above, snout truncate. 
Labial protuberances small. Total number of maxillary-premaxil- 
lary teeth 80; 64 countable mandibular teeth (dentary broken on 
left side near symphysis). Vomerine teeth in slightly arched single 
rows numbering 13 on each side. Teeth on posterior vomerine 
shelf beneath parasphenoid 93, arranged in two obovate patches 
slightly separated in midline. Skin rather thin. Mental gland not 
prominent, although marked by less pigment than rest of chin. 
Cloacal papillae well developed. Two symmetrical pockets with 
slit-like openings in dorsal posterior cloacal wall. Testis unilobate, 
pigmentless. Color of body in alcohol uniform dark lavender- 
brown above, somewhat lighter on ventral surfaces and feet. 
Palmar surfaces pigmentless. Post-iliac gland weakly indicated by 
light area. Anterior ends of ceratohyals blade-like, relatively 
narrow. Transverse processes of first caudal vertebra extend 
anteriorly to midpoint of preceding postsacral vertebra (see Fig. 2). 
Distal tarsal 5 large, articulating with fibulare and centrale 2-3; 
distal tarsal 4 accordingly not in contact with fibulare. Terminal 
phalanges with T-shaped distal ends. Measurements in mm, fol- 
lowing techniques in Rabb (1958): Total length, 78; snout-vent 
length, 40.2; head width, 8.0; eye width, 3.2; nostril width, 0.4; 
snout-eye length, 1.8; snout-angle of jaw length, 7.1; arm length, 
13.8; leg length 14.5; hind foot width, 6.5. 

Variation. — The holotype is the smallest of the specimens stud- 
ied. The others are variously shrunken by preservative, but the 
measurements are generally comparable (Table 1). Sexual dimor- 
phism is apparent in the longer snout of the larger males, and 
perhaps in their relatively wider heads. Females evidently reach 
a larger size than the males, since three of the four available are 



1965 NEW CHIROPTEROTRITON 3 

longer than the oldest and largest male. The tail is proportion- 
ately longer in males. The average number of vomerine and 
maxillary-premaxillary teeth is greater in the females, possibly 
reflecting the larger size and probably greater age of the females. 

There is no marked difference detectable in the numbers of teeth 
in the three age classes represented among the males, although 
the high vomerine count of the largest male suggests a slight 
increase with age. The two larger males from Cueva de Potrerillos 
have some melanophores in the covering of the testes, but there is 
no such pigmentation in those from Cueva del Madrono. The 
number of glands in the mental cluster ranged from about 240 to 
390 in the three males examined. 

There is slight variation in the amount of webbing of the feet, 
but the average arrangement is that described for the type. 

Habitat. — The type locality was well described by Bonet (1953). 
The cave is at an altitude of about 1300 m, and apparently in what 
was originally a cloud forest zone. According to the field notes of 
Dr. Villalobos, the topography of the cave in 1955 was considerably 
changed from that seen by Bonet in 1952. 

Bonet (op. dt.) also provided data on the Cueva del Madroiio, 
which is at an altitude of 1810 m in mixed pine-oak woods con- 
taining madroiio (Arbutus) and Gary a. The stalagmites in the 
cave were actively growing in 1952. 

Judging from Bonet's account, Chiropterotriton magnipes was 
common in both the Cueva de Potrerillos and the Cueva del 
Madrono along with various cavernicolous arthropods. However, 
C. magnipes is not the only cave-inhabiting salamander in the 
region. A specimen collected by Dr. Villalobos at Cueva de la 
Hoya, which is only 3 km from the Cueva de Potrerillos, is refer- 
able to C. arboreus (MCZ 30605). 

Relationships and comparisons. — The genus Chiropterotriton is 
composed of a group of Central American boletoglossal pletho- 
dontids that has a distinctive, relatively delicate, general habitus 
but few diagnostic characteristics. Taylor (1944) defined the 
group on the basis of the partial webbing of the feet. Tanner (1952) 
described the throat anatomy in various species, and Rabb (1956) 
described the skull and other anatomical features of the apparently 
most primitive species of the group. David Wake (in litt.) has 
informed me that the tarsal arrangement mentioned for C. magnipes 
distinguishes many members of the genus from the species groups 
in Pseudoeurycea. Although the foot shape is like that of certain 
species of Bolitoglossa, C. magnipes clearly belongs to the genus 



4 BREVIORA No. 235 

Chiropterotriton in respect to general habitus, tarsal pattern, sub- 
lingual fold, form of the ceratohyal, and such features of its 
skeletal anatomy as can be determined from X-rays and one 
cleared and stained specimen (CNHAI 142458). Wake and Brame 
(1963) indicated that fully-webbed conditions of the feet developed 
several times within the genus Bolitoglossa. The occurrence of a 
similar condition in a species of Chiropterotriton lends indirect 
support to their idea. 

The feet of Chiropterotriton magnipes are unique in the genus in 
fullness of webbing and pad-like structure. The closest approach 
to the condition in C. magnipes is in an unnamed form from El 
Chihue, Tamaulipas (illustrated in Rabb, 1958, pi. II). 

In proportions, C. magnipes differs from other Chiropterotriton 
in having a somewhat wider head (average, 19.1 per cent of snout- 
vent length in males). In relation to head length, C. magnipes has 
a large eye (average percentage, 35.5, cf cf ; 37.9, 9 9), although 
the proportion is matched in C. multidentatus from Rancho del 
Cielo, Tamaulipas and El Chico, Hidalgo, and in the short-snouted 
species C. hromeliacia. In adults of most forms, this proportion is 
29 to 33 per cent. 

The tooth count averages in C. magnipes exceed the maxima 
for all other forms. However, the size of individual teeth in 
C. magnipes is small. The largest premaxillary teeth in males 
project about 0.25 mm from the gum, and the ordinary maxillary 
and dentary teeth are about half this size. 

The foot structure and long limbs of Chiropterotriton magnipes 
indicate scansorial habits. The lack of testis pigmentation and 
the large eye suggest that the species is truly troglodytic and not 
a casual cave inhabitant. In these respects the species is slightly 
more specialized than the cave-dwelling C. mosaueri and the 
Tamaulipan Chiropterotriton with a large foot. Presumably its 
closest relationships are with these and other large, arboreal or 
scansorial northern Mexican species of the genus (C 7nultidentatus, 
C. arboreus). 

Acknowledgements. — I am grateful to Dr. E. E. Williams for the 
opportunity to describe this salamander. Dr. Alejandro Villa- 
lobos F. of the Instituto de Biologia, Universidad Autonoma de 
Mexico, has kindly supplied data on the type locahty. Dr. F. Bonet 
of the Instituto Politecnico Nacional, Escuela Nacional de Ciencias 
Biologicas (IPN), loaned his series of C. magnipes for study and 
graciously allowed deposit of specimens in major U. S. collections. 
George Zug sent information on species in the University of 
Michigan Museum of Zoology (UMMZ). X-rays were made with 



1965 NEW CHIROPTEROTRITON 5 

the beryllium-window machine at the University of ^Michigan by 
Kraig Adler. The prints are the work of Raymond Simpson. 
Abbreviations besides those indicated above are: MCZ, Museum 
of Comparative Zoology, and CNHAI, Chicago Natural History 
Museum. 

LITERATURE CITED 

BONET, F. 

1953. Espeleologia Mexicana. Cuevas de la Sierra Madre Oriental en la 
region de Xilitla. Bol. Univ. Nac. Aut. Mexico Inst. Geol., 
57: vi+96pp. 

Martin, P. S. 

1958. A biogeograpln- of reptiles and amphibians in the Gomez Farias 
Region, Tamaulipas, Mexico. Misc. Fubl. Mus. Zool. Univ. 
Michigan, 101: 1-102, 7 pis. 

Rabb, G. B. 

1956. A new plethodontid salamander from Nuevo Leon, Mexico. 

Fieldiana: Zoology, 39: 11-20. 
1958. On certain Mexican salamanders of the plethodontid genus 

Chiropterotriton. Occ. Papers Mus. Zool. Univ. Michigan, No. 

587: 1-37, 3 pis. 
Tanner, W. W. 

1952. A comparative study of the throat musculature of the Pletho- 

dontidae of Mexico and Central America. Univ. Kansas Sci. Bull., 

34: 583-677. 
Taylor, E. H. 

1944. The genera of plethodont salamanders in Mexico. Ft. I. Univ. 

Kansas Sci. Bull., 30: 189-232. 
Wake, D. B. and A. H. Brame, Jr. 

1963. The status of the plethodontid salamander genera Bolitoglossa and 

Magnadigita. Copeia, 1963: 382-387. 



BREVIORA 



No. 233 



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1965 



NEW CHIROPTEROTRITON 




Fig. 1. Chiropierotriton magnipes sp. nov., holotype MCZ 30607. X 2.1. 




No. 235 



^'^^fe** 



BREVIORA 

Miiseitanii of Compsirsitive Zoology 

Cambridge, Mass. December 10, 1965 Number 236 



VARIATION IN THE NUMBER OF 

MARGINAL TOOTH POSITIONS 

IN THREE SPECIES OF IGUANID LIZARDS 

By 

Claytox E. Ray 

U. S. National Museum 

Although tooth counts (more accurately, the number of tooth 
positions) are customarily given in the description of fossil lizards, 
their possible taxonomic value is generally unassessable for want 
of quantitative data on adequate samples of modern lizards. 
Further, the possible developmental and adaptive imphcations of 
tooth number have been inadequately explored. Edmund (1960, 
p. 66) notes an apparent lack of correlation between wave length 
in tooth replacement and number of tooth positions in iguanids, but 
the condition in juveniles of species with long wave lengths, such as 
Ctenosaura pecfinata with 11 teeth per wave, would be especially 
interesting. Hotton (1955, p. 97) in a studj^ of adaptive relations 
of dentition to diet in iguanids, including C. similis, states regard- 
ing tooth number only that there are "slight tendencies toward 
fewer teeth in the maxillary row ... in smaller individuals." 

The availability of adequate, though not impeccable, samples 
of cranial material (assembled for other purposes) representing 
Ctenosaura similis, C. pedinata, and Anolis carolinensis suggested 
the desirability of characterizing these samples statistically, in 
order to compare ontogenetic and taxonomic variability in number 
of tooth positions, and strength of correlation between number of 
tooth positions and length of tooth row among two closely related 
taxa and one distantly related taxon. 



2 BREVIORA No. 236 

MATERIAL 

Ctcnosanra similis: Well-preserved, dissociated elements repre- 
senting no less than 223 individuals, collected from deposits in the 
ruins of Mayapan (20°37'48"N, 89°27'42"E), Yucatan, during 
expeditions (1950-1956) of the Department of Archaeology, 
Carnegie Institution of Washington. Among the more than 6000 
bones identified from these deposits, over a third pertain to 
C. similis, the only lizard represented (Pollock and Ray, 1957). 
The material utilized in the present study undoubtedly dates 
almost entirely from the period of major occupation of the city, 
the JMaya Resurgence, estimated (and in part confirmed through 
radiocarbon dating) to have extended from about 1200 through 
1450 A.D. Thus the sample was accumulated during an interval 
of perhaps 250 years or more. The specimens probably were 
secured within close proximity of the city. The sample is undoubt- 
edly biased in favor of large individuals. Early juveniles are absent 
altogether. Although there is certainly high representation of left 
and right sides of single individuals among the dissociated maxillae 
and dentaries, the correspondence is far from complete and can be 
demonstrated in no given instance. The material was made avail- 
able for study and was deposited in the Museum of Comparative 
Zoology at Harvard College through the courtesy of Dr. H. E. D. 
Pollock. 

Ctenosaura pcctinafa: Cleaned skulls of 17 modern specimens 
collected over many years at widespread localities through the 
range of this nominal species. Specimens are housed in the col- 
lections of the Museum of Comparative Zoology, American Mus- 
eum of Natural History, Chicago Natural History ^luseum, and 
University of Michigan Museum of Zoology, and were made avail- 
able by the curators of those collections. 

Anolis carolinensis: Macerated skulls of 58 individuals collected 
during 1955 in the vicinity of Gainesville, Alachua County, Florida. 
This sample, like those of Ctenosaura, does not include very young 
indi\'iduals, and thus is not ideal for the study of growth phe- 
nomena. In other respects it is for statistical purposes the most 
desirably constituted sample of the three. The specimens were 
made available by Dr. Walter Auffenberg and are deposited in the 
Museum of Comparative Zoology. 

METHODS 

The following symbols represent the statistics calculated in this 
study : 

N = the number of individuals in the sample. 



1965 TOOTH VARIATION IN IGUANID LIZARDS 3 

X = the mean of x, here the straight -Hue length of tooth row, 

maxillary or dentary. 
y = the mean of y, here the number of teeth or tooth posi- 
tions per maxilla, dentary, or premaxilla (fused pre- 
maxillae). 
Sx = the standard deviation of x. 
Sy = the standard deviation of y. 
Vx = the coefficient of variation of x. 
Vy = the coefficient of variation of y. 
ORx = the observed range of x. 
ORy = the observed range of y. 

r = the coefficient of correlation, here between x and y. 
a = the growth ratio. 
Ca = the standard error of a. 
b = the initial growth index. 

Sd = the standard deviation of the diagonal distances of points 
from the reduced major axis; a measure of absolute dis- 
persion of points about that axis. 
Dd = the coefficient of relative dispersion about the reduced 
major axis. 
z = statistic computed in testing for significant difference in 
slope of growth lines. 
[Most of these are the familiar statistics of univariate and bivari- 
ate anal^'sis dealt with in standard statistical handbooks. However, 
Imbrie's paper (1956) is especially valuable regarding the basis for 
utilization of the reduced major axis rather than regression lines, 
as well as for a lucid interpretation of all of the statistics used 
herein. 

Premaxillae: The fused premaxillae were treated as a single 
median element. Only the number of tooth po.sitions was deter- 
mined for this element, as adecjuate means of measuring the (short) 
length of the tooth row, especially in Anolis, were not at hand when 
the samples were assembled. Thus only the standard uni\'ariate 
statistics are presented (Table 1, Figure 1). 

Maxillae and dentaries: All of the statistics listed above were 
computed for both left and right maxillae and dentaries of the two 
samples of Ctcnosaura, and for the right side only of Anolis (Tables 
2, 3, and 4). As a check on symmetry, the frequency distribution 
for length of tooth row and number of tooth positions was in 
every case represented l)y histograms, one example of which is 
presented here (Fig. 2). Scatter diagrams of the two variates were 
prepared in each case, from which it was determined that the dis- 
tribution of points approximates a straight line closely enough to 



4 BREVIORA No. 236 

permit representation of the growth line by the ecjuation: 
y = ax + b. The scatter diagrams for all elements of the right 
side are presented in Figures 3 and 4. 

RESULTS 

Premaxillae: As might be anticipated, the two species of 
Ctenosaura are very similar, in fact essentially identical, in statis- 
tical characteristics of tooth number, and differ considerably from 
Anolis carolinensis (Table 1). Actually, these representatives of 
the two genera differ much more strikingly than the statistics alone 
reveal. For practical purposes it can be said that the variate will 
never assume an odd \'alue in Anolis^, whereas it does so in 70 per 
cent or more of the Ctenosaura sampled (Fig. 1). The tooth 
count in an Anolis premaxilla more likely than not will be 10, and 
in Ctenosaura 7. The fact that premaxillary tooth counts in Anolis 
can assume only even values might suggest that the coefficient of 
variation (Vy) for Anolis is artificially high. However, if only one 
side of the fused element is considered, with tooth counts of 3-6, 
Vy is not altered. The V for tooth counts here and in the maxillae 
and dentaries is of little interest except that it reflects rather low 
variation in tooth number over the broad size range represented 
in the samples. 

Maxillae and dentaries: The frequency distribution for number 
of tooth positions is in general approximately symmetrical (Fig. 
2B), although it is skewed considerably to the right in the right 
dentary of Anolis. The length of tooth row in no case approaches 
a normal curve, and a polymodal frequency distribution is sug- 
gested for Ctenosaura similis (Fig. 2A) and Anolis, particularly the 
former; N is not adequate to suggest a pattern in C. pectinata. It 
is tempting to suppose that broadly overlapping frequency dis- 
tributions of different sexual or age classes or both are responsible 
for the poly modality, analogous to that demonstrated by Klaul)er 
(1937, fig. 4) for Crotalus v. viridis, but this could not be tested on 
the basis of samples at hand. 

The number of tooth positions per se is not very useful in dis- 
tinguishing the three taxa tested. It is of no value in separating 
the two species of Ctenosaura, and its power of resolution is very 
low even between taxa as remote as Ctenosaura and Anolis. Tooth 



^ Reference to "Anolis" and to "Ctenosaura" is to be understood as a shrrt- 
hand device applying only to the samples tested here, and in no case to either 
genus as a whole. 



1965 TOOTH VARIATION IN IGUANID LIZARDS 5 

count alone in an isolated fossil specimen would })e of taxonomic 
interest only if it lay within the portion of the Ctenosaura dis- 
tribution well above the OR of Ayiolis. The sample of Anolis is 
separable from those of Ctenosaura on the basis of absolute length 
of toothrow, which reflects the valid biological distinction that 
Ctenosaura is a much larger lizard, although the true population 
parameters unquestionably overlap. The two samples of Ctenosaura 
are not clearly distinguishable on the basis of any statistic pre- 
sented in Tables 3 and 4. The relatively high values of Vx, Vy, sj, 
and Dd in Ctenosaura pectinata reflect principally its greater ORx 
but may result in part also from geographic variation, not a factor 
in the other two samples. 

The most interesting features of the relation between length of 
tooth row and num])er of tooth positions in the samples studied 
are the relative strength of correlation between the two variates 
and the slope of their growth hue-. The two samples of Ctenosaura 
are essentially identical in these features. Both samples exhibit 
a strong correlation between the variates, both visually (Figs. 3A, 
B, and 4A, B) and statistically (minimum value of r = .83). The 
probability is greater than .05 that the minor deviations in slope 
of the growth lines are attributable to chance, for the values of z 
are in every case less than 1.96 (Table 4). 

Anolis, on the other hand, differs radically from Ctenosaura 
both in strength of correlation and in slope of growth line. Cor- 
relation is not visually detectable (except in a mildly suggestive 
linear clustering of points within the wide scatter) in Figures 3C 
and 4C, and is only weakly indicated by the correlation coefficient. 
However, the probability of obtaining a calculated r as great as 
.24 if the sample was drawn from a population in which p = is 
less than .10, or, if, as is reasonable on developmental grounds, it 
is assumed that r is necessarily positive, less than .05 (Simpson, 
Roe, and Lewontin, 1960, appendix table V, p. 426). The slope of 
the groAvth line is very steep in Anolis, very gentle in Ctenosaura. 
The difference in slope is statistically highly significant (Table 4), 
but biologically means only that the teeth of Anolis are con- 
siderably smaller than those of Ctenosaura. Thus, more teeth per 
unit increase in length of tooth row can be accommodated in 
Anolis than in Ctenosaura. 



- Strictly speaking, the line is not purely a "growth" line as the scatter of 
points on which it is based undoubtedly represents in part non-ontogenetic 
variation. Designation of the line as a growth line rests on the assumption 
that regression in individuals of different sizes is the same as in individuals of 
different ages. 



6 BREVIORA No. 236 

Although so few samples pro^'ide the basis for no more than 
suggested interpretations, it may be supposed that a difference such 
as that observed in correlation between number of tooth positions 
and length of tooth row has adaptive significance. The most 
fundamental adaptive distinction relating to dentition between 
Anolis and Ctenosaura is in diet; Anolis is a predatory insect 
eater, and Ctenosaura an extreme vegetarian. In Ctenosaura the 
maxillary and dentary tooth rows constitute continuous dental 
palisades in which the broadly expanded, strongly cusped crowns 
(more pronounced posteriorly) succeed one another in close 
order, and tend to overlap, an arrangement apparently advan- 
tageous in leaf-chopping, whereas in Anolis the tooth rows con- 
stitute open series in which the broadly based, apically narrow 
teeth with weak cusps are separated by variable gaps, possibly 
adaptive in the apprehension and puncturing of insect prey (Fig. 5). 
If in a strict vegetarian there is selective advantage in maintaining 
a continuous tooth row, it would be reasonable to expect a con- 
sistent addition of tooth positions with increasing size, i.e., a 
strong positive correlation between the two phenomena. The 
slope of the growth line would depend in part on the size of addi- 
tional teeth and on the degree of increase in size of successive 
replacement teeth. In a form such as Anolis carolinensis with no 
apparent selection pressure toward a closed tooth row, and perhaps 
with positive selection for an open tooth row, one might expect 
only a loose correlation between number of tooth positions and 
length of tooth row. Obviously these suggestions require testing 
on broader ontogenetic series of the taxa studied, and on similar 
series of other taxa. 

I wish to thank Drs. Thomas Frazzetta, Nicholas Hotton, 
George Simpson, and Ernest Williams for reading the manuscript. 

LITERATURE CITED 

Edmund, A. G. 

1960. Tooth replacement phenomena in the lower vertebrates. Roy. 
Ontario Mus., Life Sci. Div., Contrib. No. .52: 1-190, .5,8 figs. 
Hotton, Nicholas, III 

1955. A survey of adaptive relationships of dentition to diet in the North 
American Iguanidae. Am. Midland Nat., 53: 88-114. 

Imbrie, John 

1956. Biometrical methods in the study of invertebrate fossils Hull. .^m. 
Mus. Nat. Hist., 108: 211-252, 10 figs. 

Klauber, L. AI. 

1937. A statistical study of the rattlesnakes. IV. The growth of the 
rattlesnake. Occ. Pap. San Diego Soc. Nat. Hist., 3: ]-5(), 6 figs. 



1965 



TOOTH VARIATION IN IGUANID LIZARDS 



Pollock, H. E. D., and C. E. Ray 

19")7. Notes on vertebrate animal remains from Maj-apan. Carnegie 
Inst. Washington, Dept. Archaeol., Current Repts., 41: 638-656. 
Simpson, G. G., Anne Roe, and R. C. Lewontin 

1960. Quantitative zoology. Harcourt, Brace and Co., New York, 
vii + 440 pp., rev. ed. 

(Received 24 August 1965) 



Table 1. Univariate statistical characterization of tooth count (y) 
in the fused premaxillae of three iguanid Hzards. 









A nolis 


Statistic 


Cicnosaura si mil is 


Ctenosaura pectinata 


rarnlinensis 


N 


74 


17 


58 


y 


7.05 =t .07 


7. IS ± .17 


9.72 ± .21 


Sy 


.63 ± .05 


.70 ± .12 


1.60 ± .15 


Vy 


8.94 ± .73 


9.75 ± 1.67 


16.40 ± 1.52 


ORy 


6-9 


6 9 


6-12 



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10 BREVIORA No. 236 



Table 4. Values of the statistic z in tests for significant difference 
in slope of pairs of growth Hnes. 

Maxilla Dentaiy 

C. siinilis 

left -right .22 - .84 

('. peiiiiiata 

left -right - .317 - .08 

C. similis - C. peclinata 

left -left 1.12 - 1.28 

right - right .52 - 1.02 

(' . similis - Atiolis 

right - right 5.83 5.28 

('. pertinala - Anolis 

right - right 5.82 4.70 



1965 



TOOTH VARIATION IN IGUANID LIZARDS 



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in the fused premaxillao of (A) ('Irnnnaiini .s('w///.s, (B) ('. pcrlinata, and 
(C) Anolis carolinrnsis. 




20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 
LENGTH OF TOOTH ROW (MM ) 



22 24 26 28 30 32 34 36 38 

TOOTH COUNT 



Fig. 2. Histograms illustiating the frequency distribution of (A) length 
of tooth row and (li) tooth counts in the riglit dentary of ('(enc.sanra sinnlis. 



12 



BREVIORA 



No. 236 



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(C) Arwlis Caroline nsis. The broken line represents the reduced major axis; 
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1965 



TOOTH VARIATION IN IGUANID LIZARDS 



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14 



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R E V I O R A 

Museiam of Comparative Zoology 

Cambridge, AIass. December 15, 1965 Number 237 



A NEW SPECIES OF THE ANT GENUS DACETINOPS 
FROZ^I SARAWAKi 

By Robert W. Taylor 

Biological Laboratories, Harvard University 

The singular myrmicine genus Dacetinops Brown and Wilson 
was discovered only ten years ago. The original series included 
twelve workers and three dealate females collected by Edward O. 
Wilson in rain forest at the lower Busu River, near Lae, N. E. 
New Guinea. These specimens were described as D. cibdella by 
W. L. Brown and E. 0. Wilson (1957), and a larva collected with 
them was characterized by G. C. and J. Wheeler (1957). 

A second species, described below, has recently come to hand 
among material collected in Sarawak by Professor A. E. and 
Mrs. Eleanor Em^erson. This significant and unexpected record 
robs Dacetinops of its apparent status as a New Guinean endemic. 
In the hght of this discovery it would not be surprising if the genus 
was found to be more widespread in the Indo-Australian area. 

Dacetinops concinna new species 

Type locality. SARAWAK: Kapit District, Third Division, 
Nanga Tekalit Camp, 1° 38' N, 113° 35' E, January 29, 1963, 
A. E. and E. Emerson. 

The holotype and paratypes were taken together in a dead log 
on the floor of virgin upland rain forest at an elevation of about 
180 meters. Nanga TekaUt was a remote field camp estabhshed by 
the Chicago Natural History JNIuseum, situated about 70 miles 
above Kapit on the Batang Balleh — Sungei JNIengiong River 
system in western Sarawak. 



1 Research supported by U. S. National Science Foundation Grant No. 
GB 1634. 



2 BREVIORA No. 237 

Type deposition. Holotype, two paratypes, and fragments of a 
third, all workers, deposited in the Museum of Comparative 
Zoology (type No. 31171). 

Description. The 3 intact type specimens have the following 
dimensions (holotype cited first). The parameters of measurement 
and the abbreviations are those of Brown (1953); measurements 
are given in millimeters: TL 4.8, 4.5, 5.0; HL 1.15, 1.12, 1.22; 
HW 1.02, 1.01, 1.09; CI 88, 90, 89; ML 0.35, 0.34, 0.38; MI 32, 
30, 31; WL 1.47, 1.42, 1.57; petiole L 0.49, 0.45, 0.52; index of 
cephalic depression 59, 60, 59. 

Form of head and mandibles as shown in Figure 2. Mandibles 
longer and more slender than in D. cibdeUa; outer borders distinctly 
concave; inner borders with about 15 very small, obtuse, back- 
wardly directed teeth; apical tooth largest, more acute than 
remainder, which increase slightly in size towards the base. 
Antennal funiculus slightly incrassate, lacking a segmentally 
differentiated club. Palpal formula: maxillary 2, labial 2, struc- 
tural details of palopmeres as in cibdella (Brown and Wilson, 1957, 
fig. 2). Antennal scrobes as in cibdeUa; longitudinal dividing 
carinae feeble; upper scrobe margins more broadly lamellate, 
almost completely obscuring posterior parts of scrobes in frontal 
view. Eyes proportionately larger and more convex, maximum 
diameter in holotype about 0.17 mm. Occipital lobes well de- 
veloped. 

Mesosomal profile (Figure 1) indented at metanotal area. Pro- 
notal dorsum not margined anteriorly, submarginate at the sides. 
Propodeal structure and dentition as in cibdella, bases of teeth 
separated by sHghtly less than twice their length. Propodeal 
spiracles directed posterolaterally, placed well back on sides of 
propodeum, not as far back as in cibdella. Petiolar node longer 
than in cibdella, its dorsal profile more broadly and evenly arched. 
Postpetiole less transverse, subcircular in dorsal view. Gastric 
structure generally as in cibdella; first segment relatively narrow, 
elliptical in dorsal view, about 0.8 X as broad as long. 

Yellowish white spongiform material developed on petiole, 
postpetiole and gaster, its distribution as in cibdella, the festoons 
less massive; tergal postpetiolar blocks separated from sternal 
mass on each side. 

General sculpture consisting of very coarse longitudinal costula- 
tion, with a tendency towards reticulation due to the costular 
crests being shghtly irregular in outline, and to the development of 
weak transverse ribs in the inter-costal grooves. Reticulation more 
distinct on posterior part of frons, occipital lobes, and especially 



I 



1965 



ANT GENUS DACETINOPS 




Dacetinops concinna sp. n., holotype worker. Fig. 1, mesosoma, petiole and 
postpetiole in lateral view. Fig. 2, head in frontal view; sculpturation omitted, 
and only hairs nearest the periphery shown. 



on front of pronotum and postgenal areas of cranium ; these parts 
also with a finely shagreened mierosculpture superimposed on the 
costulae, which are generally smooth and shining elsewhere. 
About 10 costulae cross the cranium at its broadest point. Costae 
of mesosomal dorsum less cleanly cut than in cibdeUa; 4 distinct 
median ones along pronotal dorsum, with weak traces of another 
at the edge on each side. Sides of mesosoma coarsely and some- 
what irregularly rugose. 

Propodeal declivity smooth and shining, with several weak 
transverse carinae. Petiole and postpetiole moderately coarsely 
rugo-reticulate. First gastric segment moderately shining, with 
fine reticulate mierosculpture; its basal half with about 10 sharp 



4 BREVIORA No. 237 

straight longitudinal costae, the median four or five strongest, 
those at extreme sides almost vestigial and more closely spaced. 

Mandibles smooth and shining. Antennae very finely sha- 
greened. Antennal scrobes and coxae coarsely shagreened. Re- 
maining leg segments finely shagreened, and with fine irregular 
longitudinal striae. 

Long Avhitish body hairs extensively developed, proportionately 
longer than in cibdeUa (0.12-0.65 mm) but similarly distributed; 
many on head (particularly its margins), mesosoma and nodes 
occupying positions homologous to those of cibdeUa. Shorter erect 
to suberect hairs on legs and gaster, finer subadpressed ones form- 
ing a pilosity on underside of head, anterior faces of fore-coxae and 
tip of gaster. 

Color almost exactly as in cibdeUa, deep reddish brown, mandi- 
bles, antennae, and legs dark yellowish brown. 

Dismission. D. concinna is readily distinguished from D. cibdeUa 
by numerous characters, most of which are apparent on comparison 
of the published descriptions and figures of the two species. None- 
theless these ants constitute a compact and clearly delimited 
genus, and the generic diagnosis of Brown and Wilson requires only 
slight modification at this point. The statements concerning the 
antennal club structure, the mesosomal profile, and the mandibular 
shape and sculpturation need qualification, but these are minor 
matters. 

REFERENCES 

Brown, W. L., Jr. 

1953. Revisionary studies in the ant tribe Dacetini. Amer. Midland 
Natur., 50 (1): 1-137 (pp. 7-15). 
Brown, W. L., Jr., and E. O. Wilson 

1957. Dacetinops, a new ant genus from New Guinea. Breviora, No. 
77: 1-7. 
Wheeler, G. C., and J. Wheeler 

1957. The larva of the ant genus Dacetinops Brown and Wilson. Brevi- 
ora, No. 78: 1-4. 

(Received 15 September, 1965) 



BREVIORA 

Mmseiuim of Conraparsitive Zoology 

Cambridge, AIass. February 25, 1966 Number 238 



AN EVALUATION OF JAMAICAN DBOMICUS 

(SERPENTES, COLUBRIDAE) 

WITH THE DESCRIPTION OF A NEW SPECIES 

By Donald W. Buden 

Department of Biological Sciences 
University of Idaho, Moscow, Idaho 

Grant (1938) evaluated the populations of Dromicus on the 
island of Jamaica and distinguished two species, D. callilaemus 
(Gosse, 1851), and D. funereus (Cope, 1862). His review was 
based upon the examination of only seventeen individuals, sev- 
eral of which lacked adequate locality data. The original de- 
scriptions incorporated data from only two specimens of D. 
callilaemus and three specimens of Z>. funereus. 

Recently, much more material has been made available for 
study. A long series of Dromicus collected by Schwartz and 
others during the summer of 1961 together with material bor- 
roAved from several museums (a total of 227 specimens) has been 
utilized in the present study. The examination of a large num- 
ber of individuals has made possible an analysis of the popula- 
tions as well as an accumulation of data heretofore not present 
in the literature. A re-evaluation of the classification is necessary. 
The population regarded by Grant (1938) as D. funereus actually 
consists of at least two forms. Present evidence indicates that 
these forms are best treated as distinct species. 

All measurements are in millimeters. The methods used in 
obtaining scale counts have been taken from Dowding (1951), 
and the terminology in regard to hemipenial descriptions has 
been derived mainly from Dowling and Savage (I960). Several 
paratypes of the new species have been deposited in the Ameri- 
can Museum of Natural History (AMNH), and the University 
of Illinois Museum of Natural History (UIMNH), as well as 
in other collections. 



2 BREVIORA No. 238 

Dromicus callilaemfs (Gosse, 1851) 

Natrix callilaema Gossl', 1851, Naturalist 's Sojourn in Jamaica, p. 384. 
Type locality: Bluctields, Jamaica. 

Since only two of the three (fide Boulenger, 1894:143) syn- 
types of D. callilaemus were examined for the present stndy, it 
seems expedient to designate one of these specimens as a lecto- 
type, thus eliminating the possibility of confusion should the 
third specimen prove to be representative of a form other than 
Z>. callilaemus. 

Description of the lectotype: British Museum (Natural His- 
tory) 1946.1.5.90, Bluefields, Westmoreland Parish, Jamaica ; 
juvenile, total length 157 ; tail length 57 ; ventrals 136 ; paired 
subcaudals 112 ; supralabials 7/7 ; infralabials 9/9 ; prefrontals 
2 ; loreals 1/1 ; preoculars 1/1 ; postoculars 2/2 ; temporals 1 + 
2/1 -|- 2 ; dorsals smooth, dorsal row formula 19-19-17. Inter- 
nasal as long as broad, internasal suture considerably shorter 
than prefrontal suture ; frontal longer than broad, much broader 
than supraoculars, about as long as its distance to the tip of the 
snout. A dark cap on the frontal scale, accentuated by darker 
pigmentation on the anterior portions of the supraoculars and 
the posteromedial borders of the prefrontals; cap narrowing 
posteriorly along the parietal sutures, diffusing into darker 
coloration on the neck; dorsum pale except for darker pigmen- 
tation in the area of the head and neck, pattern consisting of 
two rows of dark paramedian punctations on a light ground 
color and a dark lateral stripe incorporating scale rows 3, 4, 
and 5 ; venter pale except for dark mottling in the area of the 
chin and throat; dark pigmentation more concentrated along 
the sutures of the infralabials, supralabials and the anterior 
chin shields. Because of its small size the specimen could not 
be sexed ; the subcauclal count, however, correlates with those 
of males from other localities. 

Distril)utio7i: Jamaica, apparently island-wide. 

Redefinition of species: A relatively small, slender, brick red 
snake with a distinct head pattern; ventrals 133-141 in males, 
130-140 in females ; anal divided ; subcaudals paired, 106-122 
in males, 92-109 in females; ventrals + subcaudals 242-262 in 
males and 231-242 in females; largest male 445, largest female 
465 ; ratio of tail length to snout-vent length 0.57-0.75 in males 
and 0.50-0.73 in females ; dorsal pattern usually consisting of 
three indistinct stripes, the lateralmost found on scale rows 



1966 



JAMAICAN DROMICUS 



3, 4 and 5, occasionally restricted to 3 and 4 or 4 and 5 ; pigmen- 
tation of stripes darker than ground color; venter pale, un- 
marked except for dark stippling or mottling in area of chin 
and throat, anterior scales often with light centers (Fig. 2, left) ; 
labials pale but mottled, flecked or stippled with darker pigmen- 
tation ; a narrow dark bar extending from naris to the anterior 
border of the eye, bar continuing as a broad postocular stripe 
(Fig. 1, top) ; a dark cap on the frontal scale variously di- 
lated to include portions of the supraoculars, prefrontals and 





Fig. 1. Top: Dromicus callilaemus, lateral view of head, MCZ 69070, 
from Mona, St. Andrew Parish, Jamaica. Bottom: Dromicus funereus, lat- 
eral view of head, ASFS 15922, from Christiana, Manchester Parish, Jamaica. 



nasals; cap usually narrowing posteriorly along the parietal 
sutures and continuing as a middorsal stripe, at times diffusing 
into the darker ground color of the neck ; prefrontals, nasals, and 
supraoculars often with a dark spot; rostral pale, usually with 
a dark inverted U-shaped bar ; pattern more pronounced in juven- 
iles; young specimens also with many flecks or punctations 
on a lighter dorsum ; supralabials 7/7 ; inf ralabials 9/9 ; pre- 
frontals 2 ; loreals usually present 1/1 ; preoculars 1/1 ; postocu- 
lars 2/2 ; temporals 1 + 2/1 + 2 ; dorsals smooth, dorsal row 
formula 19-19-17. 



BREVIORA 



No. 238 



Hemipenis (based upon the incompletely everted left organ 
of MCZ 69078) as follows: organ slightly bilobed, extending to 
the level of the ninth or tenth subeaudal; sulcus spermaticus 
bifurcating at the level of the third subeaudal ; large spines 
near basal and midportions of the organ, remainder of organ 
covered with small spines. 

The faded condition of the specimens examined precludes a 
detailed color description. The following comments have been 
appended from Barbour (1910) : "All the Kingston specimens 
are red when adult with scattered marks on the sides of the 
head and throat. The young have a blackish stripe along the 
body and are spotted with dark brown. The throat and head are 
heavily marked with brown. The ground color, however, is the 
same as in the full-grown examples, viz., brick red." 

Variation: Two specimens (USNM 108531 and MCZ 73761) 
have 8/9 and 5/8 inf ralabials, respectively ; MCZ 7374 lacks both 
loreal scales, and MCZ 69073 maintains only the right loreal, 
considerably reduced in size; USNM 102650 has a dorsal row 
formula 19-17-17 ; temporal scale counts are generally 1 + 2/1 
+ 2, but these scales may be variously fused or divided. 

Minor variation in color and pattern intensity is frequent. 
The head and dorsal pattern may be reduced, nearly absent, or 
somewhat obscured by darker pigmentation. This condition 





Fig. 2. Left: Dromicus callilaemus, ventral view of head, MCZ 52255, 
from Galloway Gap, St. Andrew Parish, Jamaica. Bight: Drovucus funereus, 
ventral view of head, ASFS 15922, from Christiana, Manchester Parish, 
Jamaica. 



1966 JAMAICAN DROMICUS 5 

occurs in several specimens from Montego Bay, Kingston, and 
Port Antonio. 

Specimens examined: Jamaica, Westmoreland Parish, Blue- 
fields, two (BMNH 1946.1.5.90 [lectotype], 1946.1.5.91 [para- 
lectotype] ) ; St. James Parish, Montego Bay, two (MCZ 52254, 
USNM 42881), Greenwood, one (SMIJ) ; St. Elizabeth Parish, 
Malvern, one (SMIJ) ; Manchester Parish, Mandeville, one 
(UMMZ 85948) ; St. Andrew Parish, Kingston and immediate 
vicinity, 47 (CM 33100, 33112; MCZ 7374, 18387, 44207-15, 
44903-04, 52255, 53147, 66893, 69072-81, 69181, 73761; USNM 
36662-63, 108079-80, 102650, 38947-48; UMMZ 85949, 85951; 
SMIJ [8 specimens] ) ; St. Thomas Parish, Arntully, two (USNM 
79637-38) ; Portland Parish, Boston Bay, two (USNM 108530- 
31), Windsor, one (USNM 117675). 

Dromicus funebeus (Cope, 1862) 

Alsophis funereus Cope, 1862, Proe. Acad. Nat. Sci. Philadelphia, p. 77. 
Type locality : ' ' Jamaica. ' ' 

In view of the fact that the only specimen of D. funereus cited 
by number, by Cope, has been lost, as indicated by Stejneger in 
Grant (1938:85), the designation of a lectotype from the two 
extant syntypes is in order. 

Description of the lectotype: USNM 12372, adult male; total 
length 378 ; tail length 140 ; ventrals 134 ; anal divided ; paired 
subcaudals 90 ; supralabials 7/7 ; inf ralabials 9/9 ; prefrontals 2 ; 
loreal 1/1 ; preoculars 1/1 ; postoculars 2/2 ; temporals 1 + 2/1 
+ 2 ; dorsals smooth, dorsal row formula 19-19-17 ; internasals as 
long as broad ; internasal suture slightly shorter than prefrontal 
suture ; frontal slightly longer than broad, about twice as broad 
as supraoculars, about as long as its distance from the tip of the 
snout ; loreal small, somewhat square ; pigmentation dark, speci- 
men appears somewhat faded ; no discernible pattern. 

Distribution: Western Jamaica, eastward as far as Bog Walk 
and Port Maria. 

Definition: A relatively small, stout, brownish black snake, 
generall}^ lacking a conspicuous head or dorsal pattern ; ventrals 
123-139 in males and 128-140 in females; anal divided; sub- 
caudals paired, 78-91 in males, 62-78 in females ; ventrals + 
subcaudals 203-224 in males and 192-218 in females ; largest male 
479, largest female 462 ; ratio of tail length to snout-vent length 
0.43-0.59 in males, and 0.38-0.47 in females; tail length/snout- 
vent length ratio somewhat lower in juveniles than in adults ; 



BREVIORA 



No. 288 



dorsum generally dark, unpatterued ; venter usually dark, similar 
in intensity to the dorsum, paler anteriorly in area of chin and 
throat with a suffusion of darker pigmentation (Figs. 2 and 4) ; 
juveniles often exhibiting a dorsal pattern of longitudinal 
stripes; head with a dark cap generally darker than the sur- 
rounding ground color, and venter darker than ground color 
of dorsum ; supralabials 7/7 ; inf ralabials 9/9 ; prefrontals 2 ; 
loreals 1/1 ; preoculars 1/1 ; postoculars 2/2 ; temporals 1 + 2/1 
+ 2 ; dorsals smooth, dorsal row formula 19-19-17. 

An account of the hemipenial configuration based upon the 
examination of the right extruded organ of ASPS 15172 is as 
follows : a single relatively slender, clavate organ extending to 
the level of the eighth subcaudal ; sulcus spermaticus bifurcating 
on the apical surface with each branch terminating on opposite 
sides of a fleshy rowel (Fig. 3, left) ; organ virtually nude 
from the base to the level of the seventh subcaudal, thence be- 
coming spinose to the apex, the spines generally small though 
somewhat larger on the non-sulcate surface. 

Variation: Anomalies in numbers of certain head scales were 
occasionally encountered ; scales involved were : supralabials, 





:■■•<» 







Fig. 3. Left: DromicuJi funereus, hemipenis, view of sulcate surface, 
ASFS 15172, from 1 mi. S Reading, St. James, Jamaica. Eight: Dromicus 
polylepis, hemipenis, view of sulcate surface, ASFS 1353G, from Port 
Antonio, Portland Parish, Jamaica. 



1966 JA.MAICAX DROMICIJS 7 

iufralabials, prefrontals, preoculars and postoeulars. Color varia- 
tions appear negligible. ASFS 15991 and 15994 were noted as 
being partially erythrystic in life. Juveniles differ considerably 
from adults in possessing a relatively well marked dorsal pattern 
on a light ground color. 

Specimens examined: Jamaica, St. James Parish, Montego 
Bay, two (USNll 108337-38), 5 mi. W Montego Bay, seven (MCZ 
44897-900; UMMZ 85941-43), 1 mi. S Reading, 59 (ASFS 
14743, 14932, 15060-63, 15066-67, 15146-48, 15159-77, 15264-92) ; 
St. Elizabeth Parish, Balaclava, two (USNM 73276-77) ; Trelawny 
Parish, Quick Step, one (USNIM 117676), Windsor, two (USNM 
83719-20) ; 3Ianchester Parish, Christiana, 39 (ASFS 15827-29, 
15916-25, 15982-16007), Mandeville, 12 (MCZ 13294-96, 44895-96. 
+ two untagged specimens); USNM 31096, 108240-41; UMMZ 
85946-47) ; Williamsfield, one (USNM 108533) ; Clarendon Par- 
ish, Grantham, eight (ASFS 16091-98); St. Ann. Parish, at 
mouth of Roaring River, one (MCZ 28092); St. Mary Parish, 
4 mi. S Port Maria, two (MCZ 44901-02) ; St. Catherine Parish. 
Bog Walk, two (UMMZ 85944-45) ; Jamaica (no other locality 
designation) one (USNM 5780, paratype). 

Dromicus polylepis new species 

Holotype: MCZ 81020, an adult male, from Port Antonio. 
Portland Parish, Jamaica, collected 19 June 1961 by native 
Jamaican collector. Original number ASFS 13122. 

Paratypcs: Portland Parish, Jamaica, all from Port Antonio, 
as follows : AMNH 94171-75, native collector, 22 June 1961 ; 
AMNH 94176, USNM 152589, native collector, 23 June 1961; 
USNM 152590-92, native collector, 26 June 1961; UIMNH 
56911-14, native collector, 27 June 1961 ; ASFS 13533-36, native 
collector, 28 June 1961 ; ASFS 13606-09, native collector, 30 June 
1961 ; MCZ 81021, native collector, 29 June 1961 ; MCZ 7363-64 
plus two untagged specimens (field tags 39 and 66), A. E. 
Wright, no date. 

Associated specimens: Portland Parish, Port Antonio, one 
(field tagged specimen, number 65, duplicate of series MCZ 
7363-64); St. Andrew Parish, Constant Spring, one (SMIJ). 
Cross Roads, one (SMIJ). 

Distribution: Eastern Jamaica: known only from Portland 
Parish, Port Antonio, and St. Andrew Parish, Constant Spring 
and Cross Roads. 



8 BREVIORA No. 238 

Diagnosis: A relatively small (largest male 455 mm, largest 
female 412 mm), slender (juveniles) to stout (adults), brownish 
black snake, usually lacking any conspicuous pattern; ventrals 
+ subcaudals 230-245 in males, and 222-237 in females ; liemi- 
penis single, somewhat bulbous, and heavily spinose. 

Description of holotype: An adult male, total length 387, tail 
length 150 ; ventrals 137 ; anal divided ; paired subcaudals 106 ; 
head scalation typical of a generalized colubrid; internasals as 
long as broad, internasal suture slightly shorter than prefrontal 
suture ; frontal longer than broad, broader than supraoculars, as 
long as its distance to the tip of the snout ; loreal nearly square ; 
supralabials 7/7 ; inf ralabials 9/9 ; prefrontals 2 ; loreals 1/1 ; 
preoculars 1/1 ; postoculars 2/2 ; temporals 1 + 2/1 + 2 ; dorsals 
smooth, dorsal row formula 19-17-17 ; dorsum dark brown to black 
with a pair of light brown lines of blotches, lower two or three 
scale rows paler ; venter black, suffused with brown anteriorly. 

Variation: Ventrals 136-141 in males, and 137-146 in females; 
paired subcaudals 92-106 in males, 76-93 in females ; ventrals + 
subcaudals 230-245 in males, 222-237 in females ; ratio of tail 
length to snout-vent length 0.44-0.63 in males, and 0.42-0.54 in 
females ; supralabials 7/7 ; inf ralabials 9/9 ; prefrontals 2 ; tem- 
porals 1 + 2/1 + 2 ; a dorsal row formula of 19-19-17, occasionally 
19-17-17 ; slight deviation from the normal number of head scales 
usually involving supralabials, infralabials, and postoculars. 

An account of the hemipenial configuration based upon the ex- 
amination of the right everted organ of ASFS 13536 (Fig. 3, 
right) is as follows: organ single, relatively stout and somewhat 
bulbous ; apex extending to the level of the eighth or ninth suIj- 
caudals; sulcus spermaticus bifurcating at a distance from the 
base approximately equivalent to three-fifths the total length of 
the organ, each branch terminating in a fleshy apical papilla : 
a series of relatively large spines extending along either side of 
the unbranched portion of the sulcus, diverging laterally just 
proximal to the bifurcation of the sulcus and becoming confluent 
on the non-sulcate surface, forming four or five distinct rows 
at the level of the sixth and seventh subcaudals ; spines some- 
what reduced in size and number near the base of the organ ; 
remainder of organ covered with small undifferentiated spines ; 
a depression lateral to each apical papilla forming a fleshy ridge 
connecting the non-sulcate and sulcate surfaces. 

Dorsum generally dark, unpatterned ; venter usually dark, 
similar in intensity to the dorsum, paler anteriorly in area of 



1966 JAMAICAN DROMICUS 9 

chin and throat with a suffusion of darker pigmentation; varia- 
tions in color intensity negligible, young specimens tending to 
be somewhat lighter. 

Comparisons: Although quite similar to funereus in colora- 
tion, polylepis may be distinguished on the basis of the total 
number of underbody scales and the configuration of the hemi- 
penis. The ventral plus subcaudal counts of polylepis (230-245 in 
males, 222-237 in females) are more than those of funereus (203- 
224 in males, 192-218 in females) in specimens of the same sex, 
and the hemipenis is stout, bulbous, and heavily spined as op- 
posed to the relatively slender, clavate, lightly spined and 
rowelled organ of funereus. In addition, males of polylepis have 
a higher subcaudal count (92-106) than males of funereus (78- 
91). There is no indication of overlap in distribution between 
the populations of these two species. Occurring sympatrically 
with callilaemus in eastern Jamaica, polylepis may be distin- 
guished from the latter by its dark coloration and lack of con- 
spicuous head and dorsal patterns. In addition to differences in 
the structure of the hemipenis, specimens of polylepis generally 
exhibit a lower subcaudal count (males 92-106, females 76-93) 
than specimens of callilaemus (males 106-122, females 92-109). 
The single-structured hemipenis of polylepis distinguishes it 
from the remaining forms of West Indian Dromicus, all of which 
are characterized by a bilobed organ. 

Remarks: D. p)olylepis is the third species of Dromicus de- 
scribed from Jamaica. Except for this instance, not more than 
one species of Dromicus occurs on any one island in the West 
Indies. In general morphology, coloration, and pattern, polylepis 
is strikingly similar to funereus, and the two may be regarded 
as sibling species. In addition to D. polylepis, a lizard (Anolis 
grahami aquarum Underw'ood and Williams, 1959), and a hum- 
mingbird {Trochilus polytmus scitulus Brewster and Bangs; cf. 
Bond, 1956) have undergone differentiation in extreme eastern 
Jamaica, although in neither of these cases has the differentia- 
tion reached the status of species. 

ACKNOWLEDGEMENTS 

I wish to express my thanks to Dr. Albert Schwartz for use 
of specimens in the Albert Schwartz Field Series (ASFS), for 
obtaining the loan of the remainder of the specimens examined, 
and especially for his assistance in the laboratory; to Kichard 
Thomas for his assistance in the laboratory as well as for the 



10 BREVIORA No. 238 

illustrations of the hemipenes of funereus and polylcpis; and to 
Ronald F. Klinikowski for the illustrations of the lateral and 
ventral head views of callilaemus and funereus. 

For the loan of comparative material I wish to extend thanks 
to the following : James C. Bohlke and Edmond V. Malnate, 
Academy of Natural Sciences of Philadelphia; Miss Alice G. C. 
Grandison, British Museum (Natural History) (BMNH) ; Neil 
D. Richmond, Carnegie Museum (CM) ; Ernest E. Williams, 
Museum of Comparative Zoology (MCZ) ; C. Bernard Lewis, Sci- 
ence Museum, Institute of Jamaica (SMIJ) ; Charles F. Walker 
and George R. Zug, Museum of Zoology, University of Michigan 
(UMMZ), and Doris M. Cochran and James A. Peters, United 
States National Museum (USNM). 

LITERATURE CITED 

Barbour, Thomas 

1910. Notes ou the lierpetology of Jamaica. Bull. Miis. Conip. Zool., 
52(15) :273-301. 
Bond, James 

1956. CTiecklist of birds of the West Indies. Acad. Nat. Sei. Philadel- 
phia, ed. 4, ix + 214 pp. 
Boulenger, G. E. 

1894. Catalogue of the snakes in the British Museiun (Natural His- 
tory). London, vol. 2, xiii + 382 pp. 
DOWLING, Herndon G. 

1951. A proposed standard system of counting ventrals in snakes. 
British Jour. Herpetology, 1(5) :97-99. 

, and Jay M. Savage 

1960. A guide to the snake hemipenis: a survey of basic structure and 
systematic characteristics. Zoologica, 45(2) :17-28. 
Grant, Chapman 

1938. The Jamaican Dromicus funereus re-established. Copeia, 1938:83- 
86. 
Underwood, Garth, and Ernest Williams 

1959. The anoline lizards of Jamaica. Bull. Inst. Jamaica, No. 9:1-48. 

(Received 26 July 1965.) 



\y 



BREVIORA 

escemm of Comparative Zoology 



Cambridge, Mass. February 25, 1966 Number 230 

SOUTH AMERICxVX ANGLES: 

AN0LI8 BIPORCATUS AND ANOLIS FRA8EBI 

(SAURIA, IGUANIDAE) COMPARED 

By Ernest E. Williams 

INTRODUCTION 

Confusion of the two species Anolis hiporcatus Wiegmann 1834 
and Anolis fraseri Giinther 1859 first occurred in the original 
description of A. fraseri by Giinther. One of the syntypes of 
Anolis fraseri Giinther 1859 from western Ecuador was in fact 
a specimen of Anolis hiporcatus Wiegmann 1834, a species first 
described from Mexico. Very unfortunately, it was this syntype 
which was sent to Paris for examination and which Bocourt 
(1873, pi. 15, fig. 12) figured as A. fraseri. The confusion of 
these two species which began thus has continued till the present. 

Boulenger (1885) did, indeed, correct Giinther 's error and 
remove the hiporcatus specimen from the syntype series, placing 
it in "copii" {=copci Bocourt 1873, a synonym of hiporcatus'^) . 
He was, however, apparenth' quite convinced that "copii'' and 
fraseri were closely related ; he placed them side by side in his 
catalogue and a collation of his two descriptions shovrs only 
minor differences. This belief in a close relationship continues 
to the present day: L. C. Stuart (1955) in discussing the affinities 
of Central American hiporcatus has suggested that fraseri is the 
South American repr(^sentative of hiporcatus. 

A contrary view that A. hiporcatus and .1. f)-asc)i are not at 
all close is also currently held. This opinion is implicit in Dunn's 
1937 discussion of mainland giant anoles of the latifrons group. 
A. fraseri is placed with these, and hiporcatus or its synonym 
copei goes quite unmentioned. Most recently a more explicit 
denial of close relationship has Ijeen propounded : R. Etheridge. 

1 See Stuart (1948) for discussion of the usage of the name biporcatii.^ Wio^mann 
rather than copei Bocourt. The question is one of the identity of the Berlin type 
of hiporcatus. 



2 BREVIORA No. 239 

in a doctoral thesis (1960) at the University of Michigan, has 
placed A. fraseri and A. hiporcatus in separate major sections 
of the genus Anolis, the first in his alpha series, the second 
in his beta series. (The primary distinction is the presence or 
absence of transverse processes on certain of the caudal verte- 
brae, an apparently trivial point which, however, has some very 
striking" zoogeographic and other correlations.) 

A. BIPORCATUS IN SOUTH AMERICA 

Central to the confusion of hiporcatus and fi-aseri has been 
the question of the presence of hiporcatus itself in South America. 
This has been so poorly documented in the literature that doubt 
of the authenticity of the South American records has existed. 
Thus, while Burt and Burt (1933) list the range of hiporcatus 
as "western Ecuador northward into Central America," Smith 
and Taylor (1950) give, instead, "Chiapas to Panama." 

Boulenger, of course, had already listed one of the Giinther 
syntypes from western Ecuador as the hiporcatus synonym 
"copii," but the absence of further records in the literature has 
probably progressively diminished for many workers the im- 
portance which has been attached to this single specimen. Thus, 
Barbour (1934), while mentioning the Giinther syntype in the 
synonymy of copei, totally ignored it in his discussion of the 
range, which he regarded as extending only to Panama. Further- 
more, the description from the Santa Marta Mountains of 
Colombia by Ruthven (1916) of Anolis solifer, regarded by him 
as a relative of copei, may have seemed to make the occurrence 
of hiporcatus itself in South America still more improbable. 

It is a measure of our ignorance of the Anolis of South America 
that confusion so extreme could exist regarding one of the larger 
and more distinct forms occurring there. 

It is only because I have been able to see a quite unusual 
amount of material (some of it only recently collected) that 
it is possible to bring a measure of clarity into this muddled 
situation. I have been privileged to examine the material re- 
cently collected by James A. Peters (JAP) in Ecuador, as well 
as material in the American Museum of Natural History 
(AMNH), the Academy of Natural Sciences, Philadelphia 
(ANSP), the British Museum (BM), the Chicago Natural His- 
tory ]\Iuseum (CNHM), the Instituto La Salle, Bogota, Colombia 
(ILS). the Institut Royal des Sciences Naturelles, Brussels 
(IRSN), the Escuela Polytecnica, Quito, Ecuador (Orces), the 
Universidad Central de Venezuela (UCV), the University of 



1966 ANOLIS BIPORCATUS AND ANOLIS FRASERI 3 

Michigan Museum of Zoology (UMMZ), the United States Na- 
tional Museum (USNM), the Vienna Museum (VM), and the 
Berlin ]\Iuseum (ZMB). To the collectors and curators who 
have so generously loaned material I am deeply indebted. Dr. 
Fred Medem graciously donated Colombian material collected 
by him. National Science Foundation Grant GB 2444 supported 
this study. 

On the basis of this newly examined material — so much more 
abundant than anything seen before — plus the collections of the 
Museum of Comparative Zoology, I have been able to come to the 
following conclusions. 

1. Anolis hiporcatus Wiegmann does occur in both Colombia 
and Ecuador and also in Venezuela. 

2. Anolis solifer Ruthven must be considered a synonym of 
A. hiporcatus Wiegmann. 

3. A. hiporcatus Wiegmann is at least in a broad sense sym- 
patric with A. frascri but the sympatric populations appear not 
to belong to the nominate race. 

4. A. hiporcatus belongs to a quite different group from A. 
fraseri, the relationships of which are, as Dunn and Etheridge 
have suggested, to endemic South American anoles. 

COMPARISON OF A. BIPORCATUS AND A. FRASERI. 

The characters of these two species — so often confused — are 
best compared in tabular fashion ( Table 1 ) . In many characters 
there is wide overlap. If in any numerical character the range 




Fig. 1. Anolis fraseri, MCZ 70227. Dorsal view of head. 



RREVIORA 



No. 239 



Table 1 



South American hiporcatiis 

Head scales, at least aiiterioi-ly, 
keeled. 



2-3 scales between circumnasal 
and rostral. 

Supraorbital semicircles sepa- 
rated by 0-3 scales. 

Supraocular scales keeled. 

2-3 more or less elongate supra- 
ciliary scales followed by a dou- 
ble series of scales wliicli become 
smaller posteriorly and merge 
into granules like those medial 
to the series. 

6-9 loreal rows. 

Interparietal small (but some- 
times larger than the ear which 
is usually very small). 

Subocular scales usually com- 
pletely and always at least 
partially separated from the 
supralabials by a row of inter- 
vening scales. 



fraseri 



Head scales never heeled, usu- 
ally flat, pavementose, hut some- 
times swollen or wrinkled. 

1-2 scales between circumnasal 
and rostral. 

Supraorbital semicircles sepa- 
rated hy 2-4 scales. 

Supraocular scales smooth. 

Usually no elongate supracili- 
ary scales; instead, the whole 
supraciliary margin of round 
or squarish scales distinctly 
larger than the granules medial 
to them. 

4-9 loreal rows. 

Interparietal larger (larger 
than the ear which is moderate 
in size). 

Suhocular scales broadly in con- 
tact with supralabials. 



7-12 supralabials to center of 6-9 supralabials to center of 
eye. eye. 

Dewlap scales strongly keeled, Dewlap scales smooth, small, 



almost as large as ventrals, 
rather uniformly distributed on 
the small fan. 

Ventral scales strongly keeled, 
mueronate. 

22-26 lamellae under phalanges 
ii and iii of 4th toe. 



almost granular, grouped in 
rows which are separated by 
naked skin. 

Ventral scales .smooth or weak- 
ly keeled. 

20-24 lamellae under phalanges 
ii and iii of 4th toe. 



19(i0 ANOLIS BIPORCATUS AND ANOLIS FRASERI 5 

ill one species completely overlaps tlie known range of the other. 
I have omitted this character. The most striking differences are 
emphasized hy italicizing the pertinent characters for fraseri. 
Both are large, short-legged, stout-bodied, relatively short- 
headed forms. The similarities are indeed amazing since the 
two species are, according to Etheridge, osteologically in totally 
different sections of the genus. Tiie similarities would be even 
greater if Mexican and other northern biporcatus were com- 
pared with fraseri since the northern animals tend to have 3-4 
scales between the supraorbital semicircles (c/. Stuart, 1955). 
Overlapping variation in many features, as shown in Table 1, 
is characteristic of many sympatrie species in Ayiolis and is one 
of the several reasons why this genus is described as "difficult." 
As in all such cases, the recognition of the valid species depends 
upon recognition of the constant (or almost constant) associa- 
tion of characters, however trivial. Smooth ur keeled scales, sul)- 
oculars in contact with or separated from supralabials, inter- 
parietal large or small are none of them characters universally 
useful in Anolis. The distinctive supraciliaries of fraseri are 
more useful because a substantial morphological gap between 
this condition and that shown by hiporcatiis will always distin- 
guish these two species. (The fraseri supraciliaries occur, how- 
ever, also and in a more exaggerated degree in latifrons; the 
condition is not unique to fraseri.) 




F'liX- -■ Anulis bipoicciiH.s iiarraiiril tt.s, new subspecies, ^ICZ 78935. Dorsal 
view of liead of type. 



6 BREVIORA No. 239 

INTRASPECIES VARIATION: A. BIPORCATUS 

While I have not endeavored to examine all specimens of A. 
hiporcatus from all parts of its range, I have sampled the 
northern populations (Panama, Costa Rica, Guatemala, Mexico), 
and I have had the especial advantage of examining the series 
of 122 specimens taken by Slevin near Boquete, Panama (Slevin, 
1942). 1 

Having seen this series I am impressed by the similarities of 
the various populations. There is indeed considerable variation 
within A. hiporcatus, as I understand it. Thus, the number of 
scales between the semicircles, the number of scales between 
interparietal and semicircles, the number of loreal rows, the 
amount of keeling- of the head scales, the contact or separation 
of suboculars and supralabials, and the length of the hind limbs 
all appear to be quite variable from individual to individual. 
Yet the species has a strong habitus character and is quite recog- 
nizable in spite of very striking variation. 

A very few characters appear to sort out geographically. The 
supraocular scales and those surrounding the interparietal 
are extraordinarily small in Ecuadorian and southern Colombian 
specimens. In these same populations the ear opening is circular 
and very small in contrast to the rather oval opening of northern 
specimens. 

Since there seem also to be some color differences of both body 
and dewlap in these same populations — which are preciselj^ those 
sympatric with fraseri — it seems permissible to regard the 
Ecuadorian and southern Colombian animals as representing a 
distinct subspecies. Some discussion of available names is there- 
fore required. 

Anolis solifer Ruthven, type locality La Concepcion, Santa 
Marta Mountains of northern Colombia, is the only name thus 
far proposed for a South American population of this species. 
That solifer does belong in hiporcatus and may provisionally be 
synonymized with the nominate race seems clear from the com- 
parison of the type with Panamanian and more northern material. 
The type of solifer can be matched by Panamanian material 
(except for the exceptionally short hind limbs) and Panamanian 
material is not separable by any constant characters from more 
northern specimens. The character of the very short hind limbs 
may define a population in northeast Colombia and Venezuela. 
One of the two available Venezuelan specimens has limbs as 

1 1 am indebted to Dr. AUan Leviton of the California Academy of Sciences for 
this opportunity. 



1966 



ANOLIS BIPORCATUS AND ANOLIS FRASERI 



7 



short as those of the solifer type but the other does not. In any 
event, the name solifer is clearly not available for the distinctive 
Ecuadorian and south Colombian populations. In the critical 
characters solifer resembles northern hiporcafus. 

The other names referable to the species hiporcatus are equally 
unavailable. None of them refer to South American material. 
The nominate race hiporcatus Wiegmann 1834 has the type 
locality "Mexico," now restricted to Piedra Parada, Chiapas. 
A. copei Bocourt 1873 has the type locality Santa Rosa de Pansas, 
Guatemala. Anolis hrevirostris Peters 1873, next in date, has the 
type locality Chiriqui Prov., Panama. A7wlis hrevipes Boettger 
1893 came from Cairo Plantation, La Junta near Limon, Costa 
Rica. Every one of these seems at present best grouped under 
the name hiporcatus. 

The southern race is thus without a name. In reference to the 
small ear it may be called : 



Anolis biporcatus parvauritus new subspecies 

Holotype: MCZ 78935, an adult i , "banana plantation, woods 
and penal colony camp, northern Gorgona Island. Cauca, 
Colombia, 5-45 meters altitude,'' collected by F. Medem and L. 
Salazar G., 1 to 23 February 1961. 

Paratypcs: Colombia. Cauca: MCZ 78933-34, 78936-41, same 
data as type. Narino: MCZ 79142, Rio Mataje. Ecuador. Es- 
meraldas: USNM 157105, Cachavi ; B:M 1901.6.27.1, Carondolet; 




Fig. 3. Anolis hiporcatus biporcatus, MCZ 56243, from Los Diamantes, 
Costa Rica. Dorsal view of head. 



8 BREVIORA No. 239 

AMNH ()967, Rio Capavas ; BM 1907.7.29.10-11, ZMB 18213, 
VM 12755, Rio Sapayo; BM 1901.8.3.1, Salidero; BM 1901.6.27.1, 
San Javier; Orces 3898, San Lorenzo. Imhahura: B^.I 98.4.28.13, 
Paramba. Pichincha: Orces 4596, Santo Dominj^o de los 
Colorados. "West Ecuador": BM 1946.8.13.21 (/Va.srr? syntype). 
"Ecuador": USNM 20610, ANSP 7910. 

Diagnosis. A subspecies of Anolis hiporcatus distinguished by 
the small size of the scales surrounding the interparietal which 
are smaller than or only equal to middorsal scales (instead of 
distinctly larger than middorsal scales), by the small round ear 
rather ventrally placed (instead of a large oval ear extending 
well dorsall}'), and hy a black-edged dewlap. 

Comment. None of the conventional numerical counts in 
Anolis (e.g. scales across snout, scales between semicircles, scales 
between interparietal and semicircles, etc.) reveal the distinct- 
ness of parvanritiis, though it is visually apparent immediately 
on inspection of the specimens (Figs. 2 and 3). The number of 
scales between interparietal and semicircles might l)e expected 
to correspond neatly to tlie very striking difference in the size 
of scales surrounding the interparietal in the nominate race and 
in parvauriius. lu fact, however, there is wide overlap, and no 
discrimination could be based on the numerical character. How- 
ever, comparison of the scales around the interparietal and the 
scales on the midline of midbody of the same animal gives instant 
conviction of the difference between the two races. 

The character of the small size of supraocular scales is equally 
apparent on inspection. Here there is in parvaurifns an absence 
of clear definition of any supraocular disk. This, however, is 
true also in many specimens of the nominate race. Here, again, 
comparison with middorsal scales (or direct confrontation with 
representatives of the northern race) convinces one of the exist- 
ence of a diff'erence. It is, however, a difference which is some- 
what less striking than in the case of the scales surrounding the 
interparietal. 

The size of the ear is again a character easily api^reciated 
visually (Fig. 4), but it does not reveal itself by comparison with 
the size of the interparietal, which itself varies and may be very 
small. Comparison with the temporal area is more useful but the 
size and shape of the ear are sometimes distorted by preservation. 
Certain specimens seem intermediate : MCZ 79656-57 from Villa 
Arteaga, Antioquia, Colombia, have the ear round but rathei- 
large. In other regards (scales around interparic^tal, supraocular 
scales) these are northern in type. MCZ 79842 from Sincelejo, 



19()(; 



AXOLIS BIPORCATUS AND ANOLIS PRASERI 





Fig. 4. Ear size in A. biporeatus hiporcatiis (top) and A. hiporcatii^t 
parvniiritus (bottom). Hpecinieiis are of the same snout-vent leugtli. 



Bolivar, on the other hand, has the ear -small as in the southern 
race and the supraocular scales definitely large as in the northern 
race. A juvenile from Pizarro, Choco, Colombia (CXHM 43778). 
appears, in ag-reement with expectations from its geographic 
position, to be intermediate in size of ear and size of scales 
around interparietal and, like the nominate race, in size of 
supraocular scales. From this small sample one may perhaps 
infer that the ear character and the other characters that dis- 
tinguish the races vary erratically and not in any correlated 
fashion in northern Colomljia. 



10 BRE\aoRA No. 239 

The color of the southern race seems different in preserved 
specimens from that of any northern animals including the 
northern Colombian intermediates, but this is very difficult to 
define in animals with strong powers of color change. In the 
Gorgona Island material, which is the largest series and the 
most recently collected, there is strong reticulation enclosing 
light areas which may, in animals in the dark phase, give an 
appearance of light spotting. This reticulation tends to extend 
to the belly, but the throat is not spotted with darker as it typi- 
cally is in the northern race. The north Colombian intermediates 
also lack throat spotting but otherwise seem like other northern 
populations in color. 

The small dewlap is conspicuously black in preserved specimens 
of the southern race. Examined under the microscope, it is seen 
that the scales are intensely black while the skin may have a trace 
of red. According to F. Medem's notes on the Gorgona Island 
series, the dewlap in life is orange and red at the center, black 
at the edge. 

Slevin's 1942 notes on the nominate race at Boquete, Chiriqui, 
Panama, contrast with this : ' ' The rather large dewlap of the 
males had a ground color of orange with rows of bluish scales." 

Duellman (1963) has described the dewlap of a specimen from 
20 km NNW Chimapa, Guatemala, in very similar terms, as 
having "the outer part pale orange, the median part purplish 
blue." J. M. Savage has sent me a diagram of dewlap coloration 
in a male from the Tilaran area in northwest Costa Rica. From 
this it appears that the free edge of the dewlap is red-orange 
while a small anterior part of the base is white, a larger posterior 
portion powder blue.^ 

These three descriptions, all in essential agreement, are from 
specimens which bracket much of the range of the typical race 
and indicate a dewlap color consistently and sharply different 
from that of the southern race. Unfortunately, the dewlap color 
in Venezuelan specimens, for which "solifcr" is an available 
name, is unknown. 

Behavioral notes. F. Medem (field notes) remarks on the 
Gorgona Island topotypic series of parvauritus that the gular 
sac is "extended laterally" and that this species is here found 



1 The description by Taylor (19.56, p. 1.36) of the dewlap of Costa Rican speci- 
mens as "white or slightly sreenish" appears on careful reading to result from a 
complex confusion of notes for A. capita with the text for A. hiporcaUis, and ap- 
parent resulting omission of some of the pertinent data for biporcatiix. See the 
reference to A. capita, top of page 137. 



19()(j ANOLIS BIPORCATUS AND ANOLIS FRASERI 11 

on trees up to 5 meters or more in height^ rarely on the ground. 
It is seen most often early in the morning (7:30-9:30 am). 

A. biporcatus is everywhere known as a strongly arboreal 
species. Slevin (1942) reports on it at Boquete, Panama, as 
follows : "It was generally found in the larger and higher trees, 
but occasionally on fence posts and once or twice on the ground. 
While most frequently seen on the tree trunks, it was often dis- 
covered on fairly high limbs, crouching down as if to avoid 
detection." This compares very well with Medem's notes on 
parvauritus, and the two races are probably very similar 
ecologically. 

INTRASPECIES VARIATION: A. FRASERI 

A. fraseri, with a much smaller geographic range and a smaller 
available sample, seems rather consistent in scale characters.^ 
There is no hint of racial differentiation. In color, however, there 
are two striking variants: strong transverse banding extending 
onto flanks (see fig. 3 in Williams, 1963), or the restriction of 
alternate light and dark to the vertebral zone. This statement, 
of course, applies to preserved specimens ; there is unfortunately 
no information on color in life and these two patterns may very 
well be part of the repertoire of one individual. 

THE DISTRIBUTION OF THE TWO SPECIES 
IN SOUTH AMERICA 

The number of specimens of A. fraseri and of South American 
A. biporcatus that I have been able to examine is large only by 
the standards peculiar to the generality of South American 
anoles: for A. fraseri, 14 specimens; for A. biporcatus, 32. A. 
biporcatus, as may often be true for an invader from Central 
America, extends not only down through the Chocoan area of 
Colombia into Ecuador west of the Andes but also along the 
northern edge of Colombia in the Santa ]\Iarta Mountains into 
Zulia and Taehira in Venezuela. A. fraseri — a South American 
endemic — is apparently restricted to Colombia and Ecuador 
west of the Andes, and is mainlv Ecuadorian. I list ])elow the 



^ Anolis devillei Boulenger 1880 (type examined) is an unequivocal synonym of 
A. fraseri Giinther 1851). and Boulenger (188.")) himself synonymized it. The 
juvenile specimen referred to deviUei by Boulenger (1880) is very poorly preserved 
and indeterminable ; Boulenger utilized it with great hesitation. 



12 BREVIORA No. 239 

South American A. hiporcatiis examined (except for the i^ara- 
types of parvauritus, which have been listed pi-eviously) and also 
all specimens of A. fraseri. 

Anolis hiporcatiis. Colombia. Antioquia. Villa Arteaga, MCZ 
79656-57. Magdalena. La Concepcion, Santa Marta Mtns., MCZ 
6549 (type of solifer Ruthven). Bolivar. Sincelejo, MCZ 78942. 
Turbaco (?), Leiden 2807. Norte de Santayider. Near Rio Zulia, 
82 km N Cucuta, ILS 19. Choco. Pizarro, CNHM 43778. 
Venezuela. Tdchira. La Fria, UMMZ 55994. Zulia. La Kasmera, 
Parija 300 m, UCV 8023. ? BrasU. "Silva, See Saraca," V:\l 
5904.1 

Anolis fraseri. Colombia. Caiica. Buenaventura, CNHM 
43771; Ei Tambo, ANSP 25563; Jamundi, CNHM 43772. 
Ecuador. Esmeraldas. San Mateo, CNHM 27681. Imhahura. 
Apuela, JAP 4331. Pichincha. Mindo, UMMZ 55525; Naneg'al, 
BM 83.2.23.11, JAP 925; Quito, BM 72.2.26.16. Guayas. Rio Pes- 
cado, nr. Naranjal, AMNH 23432 ; Bucay, AMNH 23030. El Oro. 
7 km SE Buenavista, JAP 2485. "West Ecuador" : BM 60.6.16.36 
(1946.8.8.47) (lectotype of fraseri). "Equateur": IRSN 200(; 
(type of dcvillei Boulenger) . 

REFERENCES CITED 

Baeisoi'r, T. 

19.34. The anoles. II. The mainland species from ^Mexic-o southward. 
Bull. Mus. Comp. Zool., 77: 121-1.5.5. 

BOETTGER, O. 

1893. Kalaloj; der Keptilien-Sammlung iiii Museum der Senekenber- 

gischen uaturforschenden Gesellschaft in Frankfurt am Main. 

I Teil. (Elivneophalen, Sohildkrotcn, Krokodile, Eidechsen, 

Chamiileons). x + 14(1 p]). Fiankfurt-a-lM. 
BOCOURT, F. 

187.J. Etudes sur les reptiles. ^lission scientitique au Mexique et dans 

I'Amerique Centrale. Pt. 3. Sauriens. Livr. 2: 33-112. Paris. 

BOTLEKGER, G. A. 

1880. Reptiles et batraciens reeueillis par M. Emile DeVille dans les 
Andes de I'Equateur. Bull. Soe. Zool. France, 1880: -11-48. 

1885. Catalogue of the lizards in the British Museum (Xatural His- 
tory), 2: ix + 497 pp. London. 
Burt, C. E. and M. D. Burt 

1931. South American lizards in the collection of tlie .American Museum 
of Natural History. Bull. Amer. Mus. Nat. Hist., 61: 227-395. 



1 I assume that this is Lajro SaracA, Aniazunas, but tliis locality is very remote 
troni any otlier reconl. Tliere are no other specimens of biporcattia from Brasil. 
I,ocalit\"r( cords in tlie Vienna Museum are so often erroneous that I cannot lucepi 
this without confirmation. 



1966 AXOLIS BIPORCATUS AND AXOLIS FRASERI 13 

DfELLMAX, W. E. 

1963. Amphibians and reptiles of the rain forests of southern EI Peten, 
Guatemala. Univ. Kansas Publ. Mus. Nat. Hist., 15: 205-249. 
Dunn, E. R. 

1937. The giant mainland anoles. Proc. Xcw England Zool. Club, 16: 
5-9. 
Etheridge, R. 

1960. The relationships of the anoles (Reptilia: Sauria : Iguanidae) : 
an interpretation based on skeletal morphology. University Micro- 
films, Inc., Ann Arbor, Michigan. 
GtJNTHER, A. 

1859. Second list of cold-blooded vertebrates collected by Mr. Eraser in 
the Andes of western Ecuador. Proc. Zool. Soc. London, 1859: 
402-420. 
Peters, W. 

1874. Uber neue Saurier {Spoeriodacfylus, Anolis, Phrynosovia, 
Tropidolepisvia, Lygosoma, Ophio seine us) aus Centralamerica. 
Mexico, und Australien. Monatsber. Akad. Wiss. Berlin, 17 
November 1873: 738-747. 
Ri"th\t;n, A. 

1916. Three new species of Anolis from the Santa Marta Mountains, 
Colombia. Occ. Pap. Mus. Zool. Univ. Michigan, 32: 1-8. 
Slevin, J. R. 

1942. Notes on a collection of reptiles from Bocjuete, Panama, with the 
description of a new species of Hi/dromorphus. Proc. California 
Acad. Sci., Ser. 4, 23: 463-480. 
Smith, H. and E. H. Taylor 

1950. An annotated checklist and key to the reptiles of Mexico, ex- 
clusive of the snakes. Bull. U. S. Nat. Mus., 199: 1-253. 
Stuart, L. C. 

1948. The amphibians and reptiles of Alta Verapaz, Guatemala. Misc. 
Publ. Mus. Zool. Univ. Michigan, 69: 1-109. 

1955. A brief review of the Guatemalan lizards of the genus Anolis. 
Misc. Publ. Mus. Zool. Univ. Michigan, 91 : 1-31. 

Taylor, E. H. 

1956. A review of the lizards of Costa Rica. Univ. Kansas Sci. Bull., 
38: 1-322. 

WiEGMANN, A. F. A. 

1834. Herpetologia Mexicana. Berlin, vi + 54 pp. 
WlLLL\MS, E. E. 

1963. Studies on South American anoles. Description of Anolis minis, 
new species, from Rio San Juan, Colombia, with comment on 
digital dilation and dewlap as generic and specific characters in 
the anoles. BulL Mus. Comp. Zool., 129: 463-480. 

(Received 15 September 1965.) 



14 



BREVIORA 



No. 239 




:}c Anolis fraseri 

• A. bi porcQtus parvauritus 



200 mi. 



Fig. 5. Map of distributions of A. fraseri and A. b. parvauritus. Inter- 
mediates and representatives of the typical race of hiporcatus in northern 
Colombia and Venezuela omitted. 



BREVIORA 

seiam of CoiniipsiFative Zoology 



Cambridge, Mass. February 25, 1966 Nimber 240 



GYMNOTHOBAX GALETAE, A NEW MORAY EEL 
FROM THE ATLANTIC COAST OF PANAMA 

By 

Ira Rubinoff 

Museum of Comparative Zoology, Harvard University 

The collection of fishes which included this new species was 
taken by the author in 1962 at Galeta Island on the Atlantic 
Coast of Panama. The type locality is identical to that from 
which Rubinoff and Rubinoff (1962) reported several new Pana- 
manian records of apodes. 

The fish here described as new is immature and unique. It is 
nevertheless described because of the proposed construction of a 
sea-level canal across the Middle-American Isthmus. The removal 
of tiiis land barrier to interoceanie dispersal places an obligation 
upon biologists for a thorough pre-canal recording of the popula- 
tions and distributions of marine fishes in this still relatively 
poorly studied region. 

Family MURAENIDAE 

Genus GymXOTHOEAX Bloch 1795 

Gymnothorax galetae, new species 

Figure 1 

Type locality: Rubinoff Station No. 79, Galeta Island, 9°24'20"N 
79°52'18"W Canal Zone, Atlantic Coast of Panama. Collected at 
a depth of less than 6 inches from the flat of reef, with "Pro- 
Noxfish'' that was introduced to the seaward edge of the reef 
and which was rapidly dispersed over the reef by the flooding 
tide, 1300 hrs., 25 May 1962. 

Measurements of the holotype (in mm) : MCZ 44035, an im- 
mature fish. Total length 128, head and trunk 59.0, tail 69, 
head 16.5, snout 2.7, eye 1.7, depth at gill opening 6.1, depth at 
anus 5.1, cleft of mouth 6.6, length of gill opening 1.1, fleshy 
interorbital 2.0, snout to origin of dorsal fin 11.8, diameter of 
typical body spot 3. 



2 BREVIORA No. 240 

Description: Body elongate, somewhat laterally compressed, 
this compression somewhat more pronounced posteriorly. Greatest 
body depth about 21 in total length. Head and trunk shorter than 
tail by about six-tenths of the length of the head. Head 7.75 in 
total length. Eye 9.7 in head, 1.6 in snout. Snout 6 in head. 
Cleft of mouth 2.5 in head, mouth closing completely, teeth 
all enclosed by lips. Fleshy interorbital 8.25 in head. Dorsal 
fin inserted well in advance of gill opening, distance from snout 
to dorsal insertion about 1.4 in head. Anal fin beginning im- 
mediately behind anus. Dorsal fin confluent with anal around 
tip of tail. No paired fins present. The anterior nostrils are 
simple tubes. The posterior nostrils have slightly raised rims 
and are above and slightly in advance of the margins of the 
eyes. There are two pores just anterior to and slightly above the 
gill slits. There are four pores along the margin of the upper 
jaw and five along the lower jaw. The total number of vertebrae, 
as determined by X-ray, is 158. 

The terminology of the following description of the dentition 
corresponds to that of Ginsburg (1951). Upper "jaw teeth" 
include 4 anterior and 14 posterior teeth on the left side, with 9 
anterior and 10 posterior teeth on the right side. The "pre- 
maxillary" has 3 enlarged canines, and there are 7 small conical 
"palatal teeth." The dentary has an outer row of 19 teeth on 
the left side, and 21 on the right. Anteriorly, 4 enlarged canines 
are present on each side medial to the outer dentary series. 

Color description: The following color observations are made 
from Kodachrome II photographs which were taken immediately 
after the capture of the specimen. Head light orange-brown, 
without spots. Snout maroon, darker than rest of head (con- 
siderably darker than indicated in the Figure). Anteriormost 
spot above, and just posterior to origin of dorsal fin. Laterally, 
maroon-brown spots begin at the level of the gill slits and con- 
tinue posteriorly with approximately equal density and size 
to the margin of the tail where the confluence of the dorsal 
and anal fin is pigmented light orange, a hue similar to that of 
the head. Between the pale orange of the head and tail, the back- 
ground is a pale reddish brown. Thin striae of maroon pigment 
radiate from the dense circular lateral maroon spots. These 
lateral spots are generally quite discrete, with the margins of 
two adjacent spots only occasionally overlapping. The spots do 
not look as if they might merge to form a reticulated pattern 
with growth. Upon preservation the body spots have become 
brown and the orange color of the head and tip of the tail has 
changed to a pale reddish brown matching the rest of the body 
ground color, which has remained relatively unchanged. 



1966 NEW PANAMANIAN MORAY EEL 3 

Remarks: Gijmnothorax galetae does not resemble the adult or 
immature stages of any other Atlantic species of GymnotJwrax. 
It is readily distinguished from all other Atlantic species by 
its large maroon spots on a paler ground color. Immature speci- 
mens of G. moringa and G. vicinns are common inhabitants of 
reef interstices at Galeta Island. The possibility that the present 
species might be a color variant of these species has been con- 
sidered but eliminated by a comparison of their total number of 
vertebrae : ca. 135 for G. vicinus, ca. 141 for G. moringa, and 
158 in the type of G. galetae. 

Due to the proximity of the type locality to the Panama Canal, 
Eastern Pacific members of the genus Gymnothorax were also 
examined for possible evidence of affinities to the new species. 
There are, however, no records of members of this genus entering 
brackish and fresh waters of the Panama Canal System (Hilde- 
brand, 1938, and personal observation at draining of Gatun 
Locks in 1961). The color patterns of several Indo-Pacific species 
such as G. isingtccna and G. stcllatus superficially resemble the 
pattern of G. galetae. However, these can be readily distin- 
guished by the continuation of their body spotting onto the 
head. The head of G. galetae is not spotted. 

The relationships among the species of Gymnothorax are in 
general too poorly understood to further attempt to discuss 
the affinities of the new species, particularly in view of the lim- 
ited material available. 

ACKNOWLEDGMENTS 

I wish to acknowledge the aid which the Evolutionary Biology 
Committee of Harvard University and the National Science 
Foundation (through GP 3450) have extended to the author. 

LITERATURE CITED 

GiNSBURG, I. 

19.51. The eels of the northern Gulf Coast of the United States and 
some related species. Texas Journal of Science, 3 (3) : 431-485. 

HiLDEBBAND, S. F. 

1938. The Panama Canal as a passageway for fishes, with lists and 
remarks on the fishes and invertebrates observed. Zoologiea 
(N. Y.),24 (3): 15-45. 

RUBINOFF, I., AND R. W. RUBINOFF 

1962. New records of fishes from the Atlantic Coast of Panama. 
Breviora, No. 169: 1-7. 

(Received 20 September 1965.) 



BREVIORA 



No. 240 








BREVIORA 

Mmseium of Comparative Zoology 

Cambridge, Mass. February 25, 1966 Number 241 



AVOCETTINOPS YANOI, A NEW NEMICHTHYID EEL 
FROM THE SOUTHERN INDIAN OCEAN 

By Giles W. Mead and Ira Rubinoff 

Museum of Comparative Zoology, Harvard University 

The oceanic eels of the genus Avocettinops lack the prolonged 
jaws or snout typical of other snipe eels, although they do have 
fleshy anterior appendages ; and they can be distinguished from 
all other apodal fishes by the complete absence of teeth, if not 
from their bizarre physiognomies alone. The number of known 
specimens is small: the type of A. schmidti (Roule and Berlin, 
1924) ; a second specimen taken off Zanzibar during the "John 
Murray" expedition (Norman, 1939; the specimen rather casu- 
ally christened Avocettinops iiormani by Bertin in 1947) ; a speci- 
men collected from the Arcturus by William Beebe, New York 
Zoological Society, off New York and reported as A. schmidti by 
Bohlke and Cliff (1956) (SU 47758); and the fish described 
below. Bohlke and Cliff (1956) also discussed in some detail the 
nomenclatorial and taxonomic entanglements generated by earlier 
authors, and reviewed the relationship of the genus to other 
snipe eels. 

Avocettinops yanoi, new species 
Figure 1 

Holotype. — A specimen 620 mm long collected by R/V Anton 
Bruun during the International Indian Ocean Expedition, Cruise 
VI, Sta. 350 B, APB label 7314; 27 June 1964; 28°05' S, 64°58' 
E to 28°28' S, 65°04' E; depth of bottom 2200-4000 m; 10-ft. 
Isaacs Kidd Midwater Trawl w4th catch dividing device nomi- 
nally set to operate at 350 m ; deep fraction of catch the probable 
depth of capture between the maximum depth reached. 1750 m 
and 125 m. MCZ Catalog No. 44404. 



2 BREVIORA Xo. 241 

Distinctive characters. — Avocettinops yanoi differs from its 
congjeners in the position of the dorsal fin, which originates well 
in advance of the gill slit, and by the more anterior anal origin, 
the preanal distance being about 1.3 times the length of the head 
(c/. 2.0 or 2.1 in specimens hitherto described). The species has 
a correspondingly low number of preanal vertebrae (13 cf. 20 
or more), and of lateral line pores between the temporal pore 
and that above the origin of the anal fin (13 cf. 18 or more). 

Description. — Meristic data and measurements, which are ex- 
pressed in per cent of head length to facilitate comparison with 
prior catches, are provided in Table 1. 

Body about two-thirds as broad as deep anteriorly, becoming 
more compressed posteriorly. Head 12 in total length. Jaws 
coterminal and bearing fleshy protuberances anteriorly both of 
which were damaged during capture. Angle of gape under pos- 
terior edge of eye. Mouth without teeth but with minute denticles 
on skin overlying jaws and roof of mouth. Anterior nostril be- 
fore center of orbit, the tube anteriorly directed. Posterior 
nostrils large deep pits, the posterior edges of which lie on a 
tangent with anteriormost points of orbits. Gill slits short (13 
in length of head), ventrally directed, and placed below bases of 
pectoral fins. Eye circular, its diameter 7 in length of head. 
Acustico-lateralis system on head well developed (Fig. 1), the 
pores large. The system includes series of lappets, which are 
presumably sensory, such as the vertical row behind and the 
horizontal series above and behind the eye. Similar lappets are 
interspersed at irregular intervals between pairs of pores in 
the lateral line along the flank. These lappets are not bilaterally 
symmetrical. The lateral line is continuous and complete. 

Dorsal and anal fins originating far forward and nearly 
continuous around tip of tail. Predorsal distance 17 in total 
length, preanal 10 in length. Pectoral fin broad, short, and 
lying in a horizontal plane when expanded. Anus and urogenital 
openings immediately anterior to origin of anal fin. The fish is 
completely black externally. Internally, the peritoneum, linings 
of mouth, and pharyngeal cavities are white. The coelom extends 
posteriorly far beyond the anus to about the midpoint of the 
total length. ^lost of this space is filled by the swollen, con- 
voluted and apparently mature pair of gonads. These appear to 
be ovaries, although they are too decomposed for close study. 
The muscular stomach, which is placed anterior to the anus, is 
empty. 



1966 NEW NEMICHTHYID EEL 3 

The success of any trawling expedition is dependent in no 
small measure on those responsible for the maintenance and op- 
eration of the nets and associated equipment. Throughout the 
trawling activities which produced the collections of which this 
specimen is a part, Mr. Shigeru Yano and his associate, Mr. C. P. 
Lee, devoted themselves to this equipment with ability, under- 
standing, and scrupulous care; without these master fishermen 
these cruises would have fallen short of their goals. With respect 
and professional admiration we take pleasure in naming this 
new eel in honor of Shigeru Yano, friend and fellow^ fisherman. 

Remarks. — The identity of the nominal species of Avocet- 
tinops remains in doubt. The type of A. schmidti has been cleared 
in potassium hydroxide and stained with alizarine. Norman's 
"John Murray" specimen from off Zanzibar lacks the tail, and 
has a fragmentary head which has also been cleared and stained, 
while the Atlantic individual discussed by Bohlke and Cliff is 
also fragnuentary. Hence meaningful morphological comparison 
is impossible. A qualitative study of the latter specimen and the 
published accounts of the two others, in comparison with A. 
yanoi, suggests that the Zanzibar specimen is probably identical 
with A. schmidti, while the western North Atlantic specimen of 
Bohlke and Cliff probably represents a distinct and unnamed 
species. 

This specimen was taken during the American Program in 
Biology, International Indian Ocean Expedition, a program 
financed by the National Science Foundation and under the gen- 
eral scientific direction of Dr. John H. Ryther of the Woods 
Hole Oceanographic Institution. To the National Science Foun- 
dation, which has also financed the research of which this is a 
part through GF 147 with Harvard University, and to Dr. 
Ryther the authors express here their sincere appreciation. 



BREVIORA No. 241 



Table 1 



Proportional dimensions, expressed as per cent of length of head, 
of Avoceftinops schmidti and A. yanoi. Data for the type of A. 
schmidti taken from the descriptions and figures of Roule and 
Bertin (1924, 1929) ; those of the Stanford University specimen 
from Bohlke and Cliff (1956), and from the specimen. 



Total length (mm) 
Length of head (mm) 

Depth of body at anus 

(%ofh.) ' 
Greatest depth of body 
Width of body at anus 
Greatest width of l)ody 
Greatest depth of head 
Greatest width of head 
Length of snout 
Diameter of orbit 
Postorbital length of head 
Length of upper jaw 
Length of lower jaw 
Length of gill slit 
Interorbital width 
Predorsal length 
Preanal distance 
Width of base of pectoral tin 
Length of pectoral fin 

Dorsal fin rays 

Anal fin rays 

Pectoral fin rays 

Total number of vertel)rae 

Preanal vertebrae (±1) 

Total number pores in lateral 

line 188 — 185 

Lateral-line pores, temporal 

pore to anus 18 ? 23 13 



Type of 
A. schmidti 


Western North 
Atlantic 

(SU 47758) 


Type of 
A. yanoi 


510.0 


— 


620.0 


32.0 


32.0 


50.0 


21.9 




24.4 


43.8 


28.4 


35.8 


9.4 


ca. 7.8 


15.4 


10.9 


— 


16.4 


25.6 


ca. 23.4 


26.4 


18.8 


17.5 


19.8 


25.0 


25.0 


24.0 


15.6 


12.5? 


13.4 


59.4 


55.9 


62.4 


39.1 


33.1 


34.8 


32.8 


24.7 


32.8 


13.1 


17.2 


7.6 


14.1 


— 


16.6 


90.6 


108.8 


73.8 


203.1 


207.8 


127.4 


14.1? 


9.7 


9.0 


28.1 


24.7 


29.0 


340 


— 


285 


315 


— • 


266 


16-17 


15-15 


12-1;: 


194 


— 


184 


20 


24 


13 



1966 



NEW NEMICHTHYID EEL 




6 BREVIORA No. 241 

LITEEATURE CITED 

Bertin, Leon 

1947. Notules iehthyologiques. 1. — Complements .sur le genre Avocet- 
tiriops. Bull. Soc. Zool. France, 72:54-55. 
BoHLKE, James E., and Frank S. Cliff 

1956. A discussion of the deep-sea eel genus Avocettinops, with notes 
on a newly diseovered specimen. Copeia, 1956:95-99. 
Norman, J. R. 

1939. Fishes. Sci. Rep. "John Murray" Expedition, 7(1) : 116 pp. 
EouLE, Louis, and Leon Bertin 

1924. Notice preliminaire sur la collection des nemichthydes recueillie 
par 1 'expedition du "Dana," (1921-1922), suivie de considera- 
tions sur la classification de cette section des poissons apodes. 
Bull. Mus. Nat. Hist. Natur., Paris, 30:61-67. 
1929. Les poissons apodes appartenant au sous-ordre des Nemichtliydi- 
formes. Rep. "Daaia" Exp., 1920-1922, No. 4, 113 pp., 19 pis. 

(Received 9 November 1965.) 



BREVIORA 

Miisenam of Comparsitive Zoology 

Cambridge, Mass. February 25, 1966 Number 242 

THE SUPPOSED "SPONGE SPICULES" OF 

MERRILL, 1895. FROM THE LOWER CRETACEOUS 

(ALBIAN) OF TEXAS 

By William A. S. Sarjeant 

Geology Department 
University of Xottingham, Nottingham, England 

INTRODUCTION 

In 1895, a i)aper was published by J. A. Merrill giving' the 
results of a study of chert nodules from Texas. In this he de- 
scribed and figured a number of microscopic structures which 
were interpreted as sponge spicules and referred, hesitantly, to 
six known genera; eight new species were proposed, all of Avhich 
were doubtfully attributed to the living genus Geodia. 

This work escaped attention until 1945, when Glaessner noted 
that the supposed spicules of Geodia were in fact hystricho- 
spherids (p. 20). As such, they constitute the first Lower Cre- 
taceous hystrichosphere assemblage and hence the first record 
of Mesozoic dinoflagellate cysts from the Ignited States. Subse- 
quently, the present author proposed the provisional reallocation 
of Merrill's species, on the basis of study of his figures and de- 
scriptions, to various dinoflagellate cyst genera (Sarjeant, 1964, 
p. 175). 

Through the courtesy of Dr. H. B. Whittiiigton, the author 
has been permitted to examine Merrill's t^^De material, from the 
collections of the Museum of Comparative Zoology, Harvard. 
The results of this study are presented here. 

STRATIGRAPHY AND MATERIAL 

The specimens studied by Merrill were nodules of chert col- 
lected by Mr. Edward E. Cauthorne from "a quarry near 
Austin, Texas." They were obtained from the Caprina Chalk 
(now known as the Edwards Limestone), a member within the 



2 BREVIORA No. 242 

Frederieksburg' Formation, Comaiiehean Stage. This formation 
is considered to be equivalent to the middle and the lower part 
of the Upper Albian of Europe (Murray, 1961, p. 349). 

The material in the Harvard collections consisted of three 
slides and seventeen partial nodules or slices of chert (MCZ 
185A). Two of the three slides measure 2" by 1" and bear 
sections of flint cut to a thickness of between two and three 
times that of a normal petrological section (0.03 mm ) ; these 
were unlabelled and are here referred to as slides I and II. The 
third slide consisted of a square of glass measuring 3" by 2" by 
1/4^' thick, bearing a wedge-shaped chert slice (varying in thick- 
ness from approximately eight times to approximately twice the 
thickness of a normal petrological section) which was surrounded 
by Canada balsam in an irregular ring. The shape of the l)alsam 
ring indicated that the chert slice had originally been fully twice 
its present size, but that one half had become detached and lost 
at some period. The thickness of the glass mount was too great 
to permit satisfactory examination ; the surviving portion of 
chert was therefore remounted for study (slide III). 

Merrill states that "several .slides" were made from eacli 
nodule ; it is therefore possible, in absence of any clear labelling, 
that the three surviving slides do not contain the holotype mate- 
rial of his species. As will be detailed later, a number of speci- 
mens showed a general correspondence to one or other of Mer- 
rill's figures, but the figures are unaccompanied by photographs 
and are not drawn with sufficient accuracy to permit any certain 
linking of specimen to figure. There remains a residue of figures 
which do not correspond even in a general fashion to any speci- 
men located ; it must be presumed in these cases, either that the 
holotypes were contained in other slides now lost, or that they 
were contained in the missing half of flint section III. 

With Dr. Whittington's courteous permission, sections were 
made from the remaining chert specimens. Nine further mounts 
were made and examined (slides IV-XII) ; they were cut to a 
thickness of between three times and twice that of a normal 
petrological section. It was hoped that the lithological character 
would prove sufficiently variable to enable correlation of the 
new with the old slides and thus to link the latter to their source 
nodules ; unfortunately, this did not prove possible, the sections 
showing insufficient distinctive features. 

Merrill also mentions (p. 10) having made a comparative 
study of English cherts ; three sections were made by Merrill, 
but these proved virtually barren of microorganisms and thus 



1966 CRETACEOUS CHERT FOSSILS 3 

cannot correspond to the surviving slides. Similarly, since Eng- 
lish cherts are not mentioned on the specimen label, it must be 
assumed that the surviving chert specimens are entirely from 
Texas. 

GENERAL DISCUSSION OF MERRILL'S RESULTS 

The earliest studies of microfossils in chert flakes were made by 
C G. Ehrenberg, in 1836 ; he assumed wrongly that the liystri- 
chospheres were silicified and placed them in the freshwater 
desmid genus Xanthidiiun. The name "xanthidia" was used 
also in descriptions of similar fossils by a group of English 
microscopists, but G. A. Mantell conclusively demonstrated their 
organic rather than siliceous composition and thus precluded 
placing them iu Xanthidium (18-15). 

Merrill was clearly unaware of these earlier studies and re- 
peated Ehrenberg 's error of assuming his microfossils to be 
silicified. He states (p. 8) : ". . . we must suppose either that the 
spicules have been replaced by amorphous silica, or that they 
are still in the hyaline or colloidal state as found, and coloured 
yellow by some organic agent perhaps." He noted the presence 
in the cherts of foraminifera (rotaliids and textularids), either 
as ' ' ghosts ' ' entirely replaced by silica, or with the organic shell 
linings persisting ; the latter he described as " . . . replaced by 
amorphous silica, the outlines remaining in a dark substance 
which has the appearance of an organic residue." This is one 
of several indications that Merrill was unaware of, or discounted, 
the possibility of structures formed of organic substances incor- 
porated into the cherts and persisting almost unchanged. He 
effectively assumed the whole microfossil content of the chert 
sections to be originally siliceous or secondarily silicified. This 
attitude clearly orientated his thinking during his stud}', for, as 
will be shown in the ensuing section, the microfossils he described 
as sponge spicules include spores, pollen, hy.strichospheres. wood 
and plant fragments, and even carbonate crystals. 

Merrill noted the relatively low concentration of microfossils in 
the chert (p. 6) : ". . . the numoer of organic remains is few, and 
the massive silica greatly in excess." This comment is wholly 
endorsed by the present author ; the number of microfossils other 
than P'oraminifera encountered in the nine new slides (IV-XII) 
averaged less than two. Foraminifera were comparatively abun- 
dant, averaging more than a dozen per slide. 



4 BREVIORA No. 242 

DISCUSSION OF THE FORMS FIGURED BY MERRILL 

In the presentation of his results, Merrill described the forms 
sequentially according' to their arrangement in his plate (here 
reproduced as Text-fig. A). This procedure is again followed 
here: the numbers, and names given by Merrill (in quotes), are 
first stated, and a reinterpretation of the specimens follows. 

1. and 2. "Monactinellid : Axinella I sp." The figured spec- 
imens were not relocated ; however, they appear to consist of 
sections through shell fragments which have l)een entirely re- 
placed by silica. Merrill noted (p. 7) the finding of supposed 
shell fragments replaced by silica, but described them as a 
"bright transparent yellow" and thus was probably misinter- 
preting organic tissue fragments. 

3. and 4. "Monactinellid: Ecnicra ? sj)." Also probably 
oblique sections through shell fragments replaced by silica. The 
figured specimens were not positively relocated, but a number 
of similar fragments were noted. 

5. "Monactinellid; aeuate spicule." Neither description nor 
figure is sufficiently informative to permit either certain recog- 
nition of the figured specimen or precise determination of its true 
nature. However, it is most probably a fragment of organic tissue 
( ?plant) ; such fragments are quite common in the cherts. 

6. "Monactinellid." The figure and description support the 
presumption that this is a wood fragment ; such fragments occur 
infrequently in the cherts. 

7. "Monactinellid." Described by Merrill (p. 13) as "the most 
abundant of the sponge spicules found in the flint"; micro- 
organisms of this type, varying considerably in the detail of size 
and shape, were encountered in all twelve cherts examined ; the 
figure is not adequate to permit certain recognition of the holo- 
type. The presence of septae crossing the cone at right angles to 
its long axis makes it probable that these are the shell linings of 
unilocular foraminifera (PI. 1, fig. 1). 

8. "Monactinellid." Also most probably a wood fragment; 
not relocated. 

9. "Monactinellid: E'spen^es ?sp." Although stated by Mer- 
rill to be "a very common form" (p. 13), this figure was not 
correlated with certainty with any microstructure seen in the 
slides. It may possibly represent a i-eplaced shell fragment or a 
fragment of organic tissue. 



1966 CRETACEOUS CHERT FOSSILS 5 

10. '^MoivAciineUid: Renter a ? sp." Whilst neither figure nor 
description permit certainty, it is most probable that this repre- 
sents a fragment of organic tissue ( ?plant). Plant fragments of 
similar appearance, some arcuate, were encountered with fair 
frequency. 

11. "Tetractinellid : GeocHa ? aiistini n. sp." This species was 
tentatively transferred to the genus SysteiiiafopJiora in an earlier 
paper (Sarjeant, 1964, p. ITo). A specimen closely resembling 
the holotype is present in slide I (specimen lA: PI. 1, fig. 10). 
From examination of both specimen and figure, there can be no 
question that this species is a junior synonym of Hystricho- 
sphaera ramosa (Ehrenberg, 1838) O. Wetzel 1933, emend. Davey 
and Williams 1965a. The species name austini must therefore be 
i-ejected. under Article 63 of the "International Code of Botan- 
ical Xomenclature." 

12. "Tetractinellid: (reodia ? erefacea n. sp." This species 
was transferred to the genus Bnltisphaerielium in an earlier 
paper (Sarjeant, 1964). .Several representatives of this species 
have been encountered in slide III. The holotype cannot be dif- 
ferentiated with complete certainty ; the specimen most similar 
to the figure is specimen III J (PL 1, fig. 6). Hystrichospheres of 
this type are frequent in the Upper Cretaceous. Ehrenberg 
(1838) was the first to record such forms, applying the name 
Xanthidiitnt liirsi(tnm, which is that of a living desmid. Lejeune- 
Carpentier ( 1941 ) redescribed Ehrenberg 's fossil material, under 
the name Hijsfrichospliacridiiun hirHutum, but Deflandre (1946) 
correctly pointed out that the trivial name hirsutuni could not be 
legitimately transferred to another genus and applied to the fossil 
forms. Deflandre had earlier (1937) proposed a new species 
//. striolatnm for similar Upper Cretaceous forms having numer- 
ous slender, simple or branching spines, and a striate shell sur- 
face ; there can be little doubt that Merrill's species erciaceum is 
a senior synonym of sfriolatum, although confirmation of the 
striate nature of the shell surface did not prove possible. 

Under strict application of the rules of priority (Article 63 of 
the "International Code of Botanical Nomenclature") the name 
cretaceum is senior and should be retained; the name striolatum 
is junior and should be rejected. However, the sjiecies siriolaium 
was soundly based, fully described and adequately figured ; the 
species o-etaceuni was. in contrast, profoundly misinteri)reted 
and inadequately figured and described. ^Moreover, conq^lete cer- 
tainty in the recognition of the holotype is not possible ; and full 



BREVIORA No. 242 



Text-figure A 

A reproilurtion of Merrill's plate (1895). The specimens are here labelled 
according to the interjiretation given in the present paper. Fuller discussions 
of the assignations are given in the text. 

1-4. Shell fragments sectioned at varying angles. 

5. A fragment of organic material. 

6. 8. Wood fragments. 

7. Shell lining of a unilocular foi-aminifer (f). 
9. CSiaracter not clear. 

10. Fragment of organic material. 

11. Hystrichosphaera ramosa (Ehrenberg, 1838). 

12. Exoc^hosphaeridium crctaceum (Merrill, 1895) = 
E.striolatnm (Deflandre, 1937). 

13. Eystrlchodinium pulchrtini Detlandre (?). 

14. Oligo.sphaeridium complex (White, 1842). 

15. Hystricli(>sp]iaira raniusa (Ehrenberg, 1838). 

16. Spore. 

17. Character not clear. 

18. Baltisphacridium texanitm (Merrill, 1895). 

19. ChJamydophorella ii\^. (?). 

20. Hystrichosphatra rantosa (Ehrenberg, 1838). 

21. Eystrichosphacridium iuhifcrum (H. H. White, 1842). 

22. Detached spines of Hystrichosphaera ramosa (Ehrenberg, 
1838). 

23-26. Character not clear. 

27. " Brown bodies" of Bryozoa (?). 

28. Carlionate crystals. 

29. Dinoflagellate cyst, prol)ably Microdinium sp. (?). 

30. 31. Character not clear. 
32-34. Proljalile sponge spicules. 

35. Fragment of TTiistriehosphnrra ramosa (Ehrenberg, 1838) 

en. 

36. Fragment of organic material. 



1966 



CRETACEOUS CHERT FOSSILS 




8 BREVIORA No. 242 

and accurate redescriptioii of the holotype, even if certainly re- 
located, would not be possible because of the difficulty in studying 
chert-enclosed specimens at high magnification. The name strio- 
lafuiit is in widespread use; the name cretaceum has not been 
reused since its original pul^lication, other than in taxonomic lists. 
Were a generic name concerned, an adequate case for conserva- 
tion could be made under Article 14; unfortunately, this Article 
does not apply to species names. Nonetheless, it is here suggested 
that striolatum should continue to be used, in preference to the 
name crcfacciim, in view of its more adequate description. The 
correct generic assignation is currently to Exochospliacridiuni 
Davey, Downie, Sarjeant and AYilliams, 1966. 

13. " Tetractinellicl : Gcodia f spi)ii-ciiri'ata n. sp. " This species 
was transferred to tlie genus BaltispJtacridium in an earlier 
paper (Sarjeant, 1964, p. 175). No specimen comparable to the 
holotype or attributable, on grounds of general morphology, to 
this species, was encountered. The figure shows a fonn with 
relatively few, simple spines and having a precingular archae- 
opyle ; it is probably referable to the genus Hystrichodinium De- 
flandre 1935 emend. Sarjeant 1965, perhaps to H. pulchruni De- 
fiandre 1935, a species very frequent in English cherts. How- 
ever, since the holotype is lost and since neitlier description nor 
figure permit certain statement of its characteristics, the species 
name spini'Cnrvatnm cannot continue in use. 

14. "Tetractinellid : Geodia ? irregularis n. sp." This species 
was transferred to the genus Husirichosphacridinm in an earlier 
paper (Sarjeant, 1964, p. 175). No specimen comparable to the 
holotype or attributable, on grounds of general morphology, to 
this species, was encountered. However, despite Merrill's state- 
ment (p. 15) that "Nothing similar to this has been found 
figured," there can be little douI)t that this species is a junior 
synonym of Oluiosphctcridium complex (H. H. White, 1842), 
Davey and Williams 1966a, a species frequent in the Cenomanian 
a]id known to range down well into the Lower Cretaceous. 

15. '"Tetractinellid: Geodia ? tripunctafa n. sp." Rejection 
of this species on the grounds that it is a junior synonym of 
Hifstrichosphaera )-aniosa (Ehi-enberg, 1838) O. Wetzel 1933, 
emend. Davey and Williams l!)66a, has already been proposed 
(Sarjeant, 1964, p. 175). A number of representatives of the 
species //. ramosa were encountered, including perhaps Merrill's 
holotype (specimen IIIC : PL 1, fig. 4) ; the earlier judgement is 
fully su])i)orted. 



1966 CRETACEOUS CHERT FOSSILS 9 

16. "Tetractiiiellid : Hyino-aphia f sp." The figured specimen 
was not found; liowever, Merrill's figure shows a trilete mark, in- 
dicating that his specimen was in all probability a spore. .Spores 
and pollen occur infrequently in the chert sections; two some- 
what similar forms are specimens IIIF (which resembles the 
figured specimens, but lacks a comparable clear ti-ilete mark) and 
VIA (PI. 1, fig. 5). 

17. "Tetractinellid: Chondrilla ? sp." The figure and descrip- 
tion of this form are so indefinite as to render impossible deter- 
mination of its true character. One specimen encountered (IIC: 
PI. 1, figs. 2, 3), a dinoflagellate cyst (questionably referable to 
the genus Gonyaulacysta and seen in oblique antapical view) may 
perhaps be Merrill's specimen, but this is incapable of confirm- 
ation. 

18. "Tetractinellid: Gcodiaf tcxana n. sp." This species was 
transferred to the genus Baltisphaeridium in an earlier paper 
(Sarjeant, 1964. p. 175). Neither the holotype nor any com- 
parable specimen was located; the morphology, as illustrated, is 
insufficiently characteristic to permit fuller comment on its prob- 
able affinities. Since the holotype is lost, a fuller analysis of the 
characteristics of the species is not possible ; the name texanum 
cannot, therefore, continue in use. 

19. "Tetractinellid: Hymeraphia ? sp." The figure is not 
especially informative, but shape, spine cover, and a faint indi- 
cation of a cingulum combine to suggest a dinoflagellate cyst. The 
combination of characters strongly suggests a species of the genus 
ChlamydophorcUa ; unfortunately, this probable assignation 
cannot be confirmed, since the figured specimen was not relocated. 

20. "Tetractinellid: Gcodiaf spinipcnisata n. sp." This 
species was transierred to the genus Hystrichosphaeridiiou in an 
earlier paper (Sarjeant, 1964, p. 275). It is probable that a speci- 
men encountered in slide III (specimen III B : PI. 1, fig. 7) is the 
holotype. There can be no question that this is yet another speci- 
men of Hystrichosphaera ramosa (Ehrenberg, 1838) 0. Wetzel 
1933. emend. Davey and Williams 1966a; this species shows some 
range of variation in morphology (cf. Davey and Williams, 
1966a), and also varies in appearance according to orientation. 
The spines are not tubular, so that assignation to HystricJio- 
sphaeridium is incorrect. The species spimpnnmtum nnist be re- 
jected, as a junior synonym of ramosa. 



10 BKEVIORA Xo. 242 

21. "Tetractiiicllid : Gcodiaf liilU ii. s])." This species was 
transferred to the genus Hystrichosphacridium (Sarjeant, 19{)4). 
No specimen comparable to the holotype or attributable, on 
grounds of general morphology, to this species, was encountered. 
However, despite Merrill's repetition of his statement, earlier 
applied to No. 14, "Nothing similar has been found figured," the 
specimen he figures is without question attributable to the species 
HifsfricJwsphacrkUiDii f}(])lfrn()v (Ehrenberg, 1838) 0. Wetzel 
1933, emend. Davey and Williams 1966a, a species common 
throughout the Cenomanian and likely to be present in the Al- 
bian. The species hilli must, therefore, be rejected as a junior 
synonym of tubifcnoii. 

22. "Tetractinellid : Geodia iripuncfaia ? n. sp. Fragments 
resulting from solution." Unquestionably these are detached 
spines of Hystrichosph(u ra ranioui. A number of such detached 
spines were seen, none corresi)ondiiig precisely to the grouping 
.shown in the figure, which might in any ease be idealised. 

23. "Lithistid ? Flesh spicule." This is described as "yellow- 
ish" and must be presumed to be a plant or wood fragment. The 
figure, by it generalised nature, precludes certain recognition ; 
similar fragments were by no means uncommon. 

24. 25, 26. " Tetractinellids ; Hymerapliia ? sp." (_)ver-simpli- 
fied figures and descriptions again render certain recognition 
difficult. Pollen grains of compai'able morphology were noted, 
and also (iii slide X) a species of dinoflagellate cyst questional)ly 
attributable to the genus ProUxusphaeridium Davey, Downie, 
Sarjeant and AVilliams 1966, with a sinnlarly elongate shai)e and 
cover of simple spines ( PI. 1, fig. 8 ). 

27. "Teti-actinellid : Geodia ? sp." Groups of circular, disc- 
like structures were encountered in several cherts (e.g. VIl B; 
PL 1, fig. 11). Similar structures occur widely in jialynological 
material from the Upper Mesozoic ; Merrill's descrijition (p. 17) 
is quite accurate. Their interpretation is doubtful: Otto Wetzel 
(1961, pi. 1, figs. 8-10) has figured similar forms as "brown 
bodies" of Bryozoa ; it is also possible that th(>y represent remains 
of colonial algae, possibly Chlorophyta. 

28. "Tetractinellid: dermal spicules .'" These ar(> abundant 
in most of the cherts examined and are undoubtedly rhomb-shaped 
carbonate ( .' dolomite) crystals. Carbonate crystals are fre- 
quently encountered in cherts and are somc^times considered to 



1966 CRETACEOUS CHERT FOSSILS 11 

indicate that the ehert was formed by replacement of a pre-exist- 
ing limestone. ^lerrill's misinterpretation of them is surprising 
(PI. l,fig. 12). 

29. "Tetractiiicllid : (itodia sj)." The figure may correspond 
to a sjiecimen in slide III, which resembles it in general features 
(specimen II III). Although full study is difficult because of its 
situation deep in the chert, this is certainly a dinotlagellate cyst, 
and most probably a species of the genus Microdinium, species of 
which are known to occur in the Cenomanian. 

.'30. "Tetractinellid : globo-stellate of dermal layer." This 
structure was not relocated and is of dubious reference, possibh' 
])ollen. 

81. "Uiiodia .'" The figure and description are inade(|uate to 
permit recognition of this form. 

32. ^'Gcodia ? Pyramidal or zone spicule." Not relocated; 
most ju'obably correctly interpreted as sponge spicules. 

33,34. "Hexactinellid. SiaurdciiueJUa .' sp." Not relocated : 
most probably correctly interpreted as sponge spicules. 

3.1. "Frame work of a Hexactinellid.'" This appears to be a 
fragment of a HystricJiospJiacra furatia, although the figure is 
extremely generalised and precise interpretation is difficult. A 
similar fragment was encountered in slide III (specimen IIIE). 

36. "Monactinellid: Reniera ? sp." The sausage-shaped ob- 
ject in Merrill's figure is most prol)ably to be interpreted as a 
fragment of organic tissue ( ? plant ) ; many similar fragments 
were seen. 

COXCLrSIONS 

A re-examination of Merrill's text and figures, in conjunction 
with study of the three surviving chert sections (one broken) and 
of new sections made from surviving fragments of cherts, makes 
it clear that, of the .'36 microscopic structures he figures and de- 
scribes, only three (none confirmed) can be regarded as represent- 
ing sponge spicules. Nine of his figures certainly, and four others 
possibly, represent dinotlagellate cysts (mainly hystricho- 
spheres) ; these include his eight proposed new species, of whi"h 
five are junior synonyms of previously described species. The 
three remaining species retain technical validity, but the holo- 
types of two are lost, the descriptions and figures being insuffi- 
cient to enable their accurate characterisation, and the liolotype 
of the third species does not permit full study. 



12 BREVIORA No. 242 

The reinainino' 20 mieroseopic .structures tig'ured and described 
by Merrill are of very diverse character. Four are considered 
to represent shell fragments sectioned in varying directions; one 
is thought to be a f oraminiferal shell lining ; one is a spore ; three 
possibly pollen ; one represents an association of disc-like struc- 
tures, possibly "brown bodies" of Bryozoa ; one comprises carbon- 
ate crj^stals ; two represent wood fragments ; four represent frag- 
ments of organic ( ^ plant ) tissue of varying sha]:)e ; and three 
are of indeterminate character. 

The currently valid species of dinoflagellate cysts recognised 
in Merrill's figures and material are Hystricliosphaera furcata, 
Hystrichosphaeridiioii ti(hif(riniK Exochospliacridium striolatum, 
OligosphocridiuDi complex, Rystricliodinium pulchrum (doubt- 
ful), and undetermined species of the genera (TO)iyaulocijsta, Pro- 
lixosphaeridiioit, Hiistrichodiniiim, Chlamydophorella and Micro- 
dinium. Although the number of individuals encountered is small, 
and a full picture of the dinoflagellate cyst assemblage cannot be 
said to have l)een obtained, the species represented are all ones 
which would be likely to occur in Middle to Tapper Albian prep- 
arations. 

EEFERENCES 

Davey, E. J., C. DowNiE, W. A. S. Sarjeant and G. L. Williams 

1966. Fossil dinoflagellate cysts attributable to Baltlspraeridium. In 

Studies of Mesozoic and Cainozoic dinoflagellate cysts. Bull. 

Brit. Mus. (Nat. Hist.) Geol., Suppl. 3 (in press). 
Davey, R. J. and G. L. Williams 

1966a. The genera Rystricliosphaera and Achomosphaera. In Studies of 

^Slesozoie and Cainozoic dinoflagellate cysts. Bull. Brit. Mus. 

(Nat. Hist.) Geol., Suppl. 3 (in press). 
19661). The genus Hystrichosphaeridium and its allies. In Studies of 

Mesozoic and Cainozoic dinoflagellate cysts. Bull. Brit. Mus. 

(Nat. Hist.) Geol., Suppl. 3 (in press). 
Deflandre, G. 

1935. Considerations biologiques sur les micro-organismes d 'origine 

I)lanctonique conserves dans les silex de la craie. Bull. Biol. Fr. 

Belg., vol. 69, pp. 213-44, pis. 5-9. 
1937. Mierofossiles des silex cretaces II. Ann. Paleont., vol. 26, pp. 

51-103, pis. 8-18. 
1946. Remarques sur la systematique des hystriehosphaerides. C. R. 

Somm. Soc. Geol. Fr., No. 7, pp. 100-111. 



1966 CRETACEOUS CHERT FOSSILS 13 

Ehrenberg, C. G. 

1838. Uber das Massenverhaltnis der jetzt labenden Kieselinfusorien 
und iiber ein neues Infusorien-Conglomerat als Polierschiefer von 
Jastraba in Ungarn. Abh. K. Akad. Wiss. Berlin (1836), vol. 1, 
pp. 109-35, pi. 1. 
Glaessner, M. F. 

1945. Principles ot" micropalaeontology. Melbourne: University Press, 
296 pp., 1-4 pis. 
Lejeune-Carpentier, M. 

1941. L 'etude microscopique des silex (9 ieme note). Sur Hystricho- 
sphaeridium hirsutum (Ehr.) et quelques formes voisines. Ann. 
Soe. Geol. Belg., vol. 63, no. 3, pp. 71-92. 
Mantell, G. a. 

1845. Notes of a microscopical examination of the Chalk and Flint of 
southeast England. Ann. Mag. Nat. Hist., vol. 16, no. 103, pp. 
73-88. 
Merrill, J. A. 

1895. Fossil sponges of the flint nodules in the Lower Cretaceous of 
Texas. Bull. Mus. Comp. Zool., Harvard. (Geology Ser. Ill), 
vol. 28, no. 1, pp. 1-26, 1 pi. 
Murray, G. E. 

1961. Geology of the Atlantic and Gulf Coastal Province of North 
America. New York : Harper, 692 pp. 
Sarjeant, W. a. S. 

1964. Taxonomic notes on hystrichospheres and acritarehs. J. Paleont., 

vol. 38, pp. 173-7. 
1966. Dinotlagellate cysts with a Gonytiulax type tabulation. In Studies 
of Mesozoic and Cainozoic dinoflagellate cysts. Bull. Brit. Mus. 
(Nat. Hist.) Geol., Suppl. 3. (in press). 
Wetzel, O. 

1933. Die in organischer Substanz erhaltenen Mikrofossilien des Bal- 
tisehen Kreide-Feuersteins. Palaeontographica, vol. 77, pp. 141- 
88; vol. 78, pp. 1-110, pis. 1-7. 
1961. New microfossils from Baltic Cretaceous flintstones. Micro- 
palaeontology, vol. 7, no. 3, pp. 337-50, pis. 1-3. 
White, H. H. 

1842. On fossil Xanthidia. Microse. J., vol. 11, pp. 35-40, pi. 4. 

(Received 25 October 1965) 



14 BREVIORA No. 242 



Plate 1 

Fig. 1. Presumed shell lining of a unilocular Foraminifer (Fig. 7 
" Alonactinellid " of Merrill). X 500. 

Figs. 2, 3. Gonyaulacysta sp. indet., seen in oblique apicaJ view (Fig. 2) 
and, by transparency, in oblique antapical view (Fig. 3) — the antapex is 
at left. Figure 2 closely resembles ' ' Tetractinellid ; Chondrilla ? sp. ' ' of 
Merrill. X 500. 

Fig. 4. Eystric'liosphaera ramoaa (Ehrenberg). The presumed holotype 
of Merrill's invalid species Geodia .' iripunciata. X 500. 

Fig. 5. H ystrichosphaera ramosa (Ehrenberg). Possibly the holotype of 
Merrill's invalid species Geodia ? spini-pansafa. X 500. 

Fig. 6. Exochosphaeridium crefaeeum (Merrill). The specimen may be 
the holotype. X 500. 

Fig. 7. Hystrichospliaera nnno.sa (Ehrenlierg). A specimen in terminal 
view, illustrating the high degree of variation in appearance given by 
orientation. X 500. 

Fig. 8. Prolixospltaeridium sp. indet. (Possibly corresponding to Merrill's 
Fig. 26b "Tetractinellid; Eymcraphia ? sp. "). X eSOO. 

Fig. 9. Spore, gen. et sp. indet. (Similar to Merrill's Fig. 16 "Tetracti- 
nellid; Hymeraphia ? sp. "; but lacking a clear trilete mark.) X c-300. 

Fig. 10. Hystrichosphaera ramosa (Ehrenberg). Possilily the holotype of 
Merrill's invalid species Geodia ! austini. X 500. 

Fig. 11. Group of disc-like organisms, possibly "brown bodies" of 
Bryozoa (equivalent to Merrill's Fig. 27 "Tetractinellid; Geodia ? sp. "). 
X 500. 

Fig. 12. A carbonate ( ? dolomite) crystal. Similar crystals are described 
by Merrill as "Tetractinellid: dermal spicules?" X 500. 



1966 



CRETACEOUS CHERT FOSSILS 



15 




BREVIORA 

Miiiseiuiini of Comparative Zoology 

Cambridge, ^Iass. April 29, 1900 Number 243 

QUATERNARY FISH FOSSILS FROM WEST OF LAKE 
RUDOLF, KENYA 

By Keith Stewart Thomson 

Department of Zooloj^y, University College London. i 

INTRODUCTION 

The fossils to be described in this paper were collected during 
the summer of 1963 by a party from the Museum of Comparative 
Zoology, consisting of Professor Bryan Patterson, Messrs. A. D. 
Lewis, C. T. Williams, and the author. It is a pleasure to ac- 
knowledge the assistance given to us by Dr. L. S. B. Leakey, and 
by Mr. John Walsh of the Mines and Geological Department, 
Kenya, who called our attention to the deposits discussed in this 
paper. 

The late Pleistocene and Quaternary deposits west of Lake 
Rudolf, North Kenya (Turkana District), were visited by us 
during a survey of the Miocene formations in the region. Ac- 
counts of earlier visits to the region are given by Arambourg 
(e.g. 1943), and Fuchs (1939). A general view of the area con- 
cerned is shown in the accompanying map. A range of hills (in- 
cluding Losidok) west of the lakeshore is Tertiary in age, consist- 
ing of grits and volcanics. Along the base of this range there are 
exposed large and extensive beds of Pleistocene age — largely 
unfossiliferous lake beds, including conglomerates of small clay 
pebbles with, in places, small rocks derived from the volcanics. 
Away from the bases of the hills — and here we explored only 
eastwards — are flat plains covered Avith gravel and windblown 
sands ; close to the lake large sand dunes occur. These plains 
extend from the hills down to the lakeshore. Nearer the lake, as 
Fuchs has noted (1939), there is evidence of at least two former 
beaches, some 220 and 90 feet above the current level of the lake. 
Along these beaches fossils may be found on the surface, although 
their derivation is difficult to determine, and "reefs" of the 



1 Present aiklress : Dept. of Kiulogy and Pcalioily MiKseuni of Natural History, 
Yale University, Xew Haven, Conn. 



2 BREVIORA No. 243 

freshwater oyster, Aetheria elliptica, also occur. We were able to 
follow the beaches for several miles and found fossils even down 
to the lakeshore. In several places drainage from the plains has 
cut into the deposits and we found a number of large gullies or 
washes along the sides of which the bedding of the deposits — 
apparently with a slight eastward dip — was visible, and fossils 
could be collected in situ. The principal place from which fossils 
were collected was an extensive exposure at Kangatotha (see 
map), where the TurkAvell River has made a large cut through 
some hundred feet of the lake deposits. Patterson and Williams 
measured the exposed section as follows : 

white sands 

black clay 

white coarse sands with gravel lenses 

grey clay 

micaceous sand 

black clay 

coarse sand and gravel 

black clay 

white sand 

lilack clay 

interbcdded black clays and grey 
sands 

black clays 

fine grey sand 

fine grey sand highly indurated with 
iron 
" 3 5 feet fine wliite sands, black micaceous 

flakes 
" 2 4 feet medium yellow sands, black micaceous 

flakes 
" 1 1-% feet coarse grey sands with large clay 

pebbles in upper unit (base con- 
cealed) 

"Upper unit" and "lower unit" are informal, non-committal 
terms ; formal stratigraphic naming should follow, not precede, a 
general survey of the Lake Rudolf Quaternary. The two units 
are readily distinguishable, the upper being predominantly dark 
in colour, the lower light. Slightly indurated fillings of plant 
roots are abundant in the lower unit. 

Partly due to the swollen state of the river and partly due to 
lack of time, a similar cliff on the south bank of the watercourse, 
here about half a mile wide, was not visited. The tip of northern 
cliffs corresponds to the 220 foot level of Fuchs. 



Upper unit, Ijed 10 


Surface 


( ( 


9 


4 feet 


( ( 


8 


16 feet 


i i 


7 


21/2 feet 


i I 


6 


10 feet 


I i 


5 


4 feet 


( I 


4 


1/2 to 5 feet 
5 feet 
% foot 
21/2 feet 


i I 


.> 


9 feet 


^< 


1 


12 feet 


Lower unit, bed 


5 


7 feet 


" 


4 


31/2 feet 



1966 



QUATERNARY FISHES FROM KENYA 



Mollusc, fish, reptilian and mammalian remains are common in 
the upper unit of the exposures, mollusc and fish in the lower. A 
human skull, a partial mandible, and various skeletal fragments 
were found isolated in bed 8 of the upper unit, and poorly pre- 
served parts of a human skeleton were found in the overlying 
black clay of bed 9. Artifacts and pottery fragments are common 
on the surface and both were encountered in situ in bed 7 of the 
upper unit. 




Miles 20 15 



Sketch map of Lake Eudolf and the Lodwar region (Turkana District, 
Kenya ) showing position of the Kangatotha locality. 



4 BREVIORA No. 243 

FOSSIL FISHES 

All the fish remains collected consisted of isolated bones, never 
associated, and very often fragmentary. Few showed great signs 
of weathering or of having been ' ' rolled. ' ' The bones were highly 
mineralized, often showing considerable iron content, despite 
their relatively recent age (see below). 

From lower to upper strata, the fish were distributed as fol- 
lows: 

Lower unit, base of bed 5, seven feet below junction with upper 
unit 
Order Ostariophysi 
Suborder Siluroidea 
Family Mochokidae 
Synodontis sp. 

Seven pectoral spine fragments 

Family Clariidae 
Clarias sp. 

One articular (posterior portion), one quadrate 
(articular portion) 

Suborder Cyprinoidea 
Family Cyprinidae 
cf . Barhus 

Two dorsal pterygiophore fragments 

Order Percomorphi 
Suborder Percoidea 
Family Centropomidae 

Lates cf. Lates niloticus Cuvier and Valenciennes 
One incomplete parasphenoid (length 150 mm), 
three vertebrae (diam. 25, 33, 46 mm), four fin 
spines (38, 52, 55, 62 mm) 

Family Cichlidae 
Tilapia sp. 

Two fragments of pterygiophores, six fin spines (2 
nearly complete, 55 and 47 mm), two branched 
fin rays, and one vertebra 



1966 QUATERNARY FISHES FROM KENYA 5 

Lower unit, bed 5 (general) 
Order Ostariophysi 
Suborder Cyprinoidea 
Family Cyprinidae 
cf . Barhus 

Tentativel}-^ referred to this genus are sections of 
two fin spines. 

Order Pereomorphi 
Suborder Percoidea 
Family Centropomidae 

Latcs cf. Lates niloticus Cuvier and Valenciennes 
Four dorsal fin spines (28, 42, 45, 51 mm) 

Family Cichlidae 
Tilapia sp. 

One pterygiophore — first interliaemal (70 mm), 
two dorsal fin spines (incomplete), four ver- 
tebrae: two abdominal (diam. 10, 11 mm), 
one caudal (diam. 15 mm), one terminal 
(diam. 8 mm, length 26 mm) 

Upper unit, bed 5 
Order Ostariophysi 
Suborder Siluroidea 
Family Bagridae 

Bagrus cf. Bagriis hayad (Geoffroy) 

Posterodorsal portion of a single occiput (estimated 
height 28-30 mm) 

Family Mochokidae 

Sy7iodontis cf. Synodontis scliall Cuvier 

Humeral process of left cleithrum (49 mm long), 
one posttemporal bone, and fragments of four 
pectoral spines 

Family Clariidae 

Clarias cf. Clarias lazera Cuvier and Valenciennes 
Posterior portion of dermethmoid bone (breadth 25 
mm), median portion of supraoccipital (max. 
breadth 35 mm) and eleven vertebrae (diam. 8, 
12, 13, 13, 13, 15, 15, 16, 17, 17, 17 mm) 



BREVIORA No. 243 

Suborder Cyprinoidea 
Family Cyprinidae 

Barhus cf . Barhus hynni Cuvier and Valenciennes 
Three dorsal pterygiophores (incomplete), one 
pharyngeal (length approx. 37 mm), one ver- 
tebra (diam. 6 mm) 

Order Percomorphi 
Suborder Percoidea 
Family Centropomidae 

Lates ef . Lates niloticus Cuvier and Valenciennes 
The most abundant fish in this unit ; twenty-three 
vertebrae (diam. 12, 13, 16, 19, 20, 20, 24, 25, 
26, 26, 26, 27, 27, 30, 34, 35, 35, 37, 42, 42, 50, 
51, 53 mm), the articular regions of four quad- 
rates, one incomplete preoperculum, four in- 
complete fin spines, three fragments of hyo- 
mandibulars, one dermethmoid, one partial 
parasphenoid, two left premaxillae (proximal 
ends), one maxilla (distal end), and two iso- 
lated neural arches (probably from anterior 
vertebrae) 

Family Cichlidae 
Tilapia sp. 

One dorsal pterygiophore, two spines, nine verte- 
brae (diam. 8, 7, 6, 6, 6, 6, 5, 5, 5 mm) 

Upper unit, bed 8 
Order Ostariophysi 
Suborder Siluroidea 
Family Mochokidae 

Synodontis cf . Synodontis schall Cuvier. 

The humeral process of one cleithrum (length 55 
mm), one dorsal fin spine, three pectoral fin 
spine fragments 

cf. Synodo7itis 

Fragments of three spines 



1966 QUATERNARY FISHES FROM KENYA 7 

Family Clariidae 

Clarias cf. Clarias lazera Cuvier and Valenciennes 
Three dermethmoids (2 complete, lengths 34, 45 
mm; one incomplete, anterior width 50 mm), 
one posttemporal, two sections of supraoceipital 
(one anterior, one posterior), one pectoral fin 
spine fragment, two vertebrae (diam. 13, 19 
mm) 

Suborder Siluroidea indet. 

One vertebra (diam. 23 mm), two dorsal fin spines 
(80, 48 mm) and several spine fragments 

Suborder Cyprinoidea 
Family Cyprinidae 

Barhus cf. Barhus hynni Cuvier and Valenciennes 
Four pharyngeal bones (approx. length 30, 30, 40, 
45 mm), two ( fdorsal) pterygiophore frag- 
ments 

cf . Barhus 

Two vertebrae (diam. 14, 15 mm) 

Order Percomorphi 
Suborder Percoidea 
Family Centropomidae 

Lates cf. Lates niloticus Cuvier and Valenciennes 
Six articular bones (posterior sections), five incom- 
plete quadrates (articular sections), four in- 
complete preoperculars, the proximal ends of 
five premaxillae, the anterior part of one right 
palatine, one dermethmoid, five dorsal fin 
spines, twenty-two vertebrae (1 second verte- 
bra — diam. 25 mm ; 3 third vertebrae — diam. 
20, 23, 27 mm ; 18 abdominal vertebrae — diam. 
17, 20, 20, 20, 22, 24, 24, 28, 28, 28, 29, 29, 32, 
33, 34, 40, 48, 49 mm), plus a number of shat- 
tered and unidentifiable fragments of large 
cranial elements 



8 BREVIORA No. 243 

Upper unit, bed uncertain 

These fishes were collected at one of the smaller exposures 
near the lake shore, about two miles north of the mouth of the 
Turkwell. 

Order Ostariophysi 
Suborder Siluroidea 
Family Bagridae 

Bagrus cf. Bagrus hay ad (Geoffroy) 

One occiput, incomplete (max. depth 69 mm) 

fFamily Clariidae 

Dermethmoid portion of skull, incomplete 

Order Percomorphi 
Suborder Percoidea 
Family Centropomidae 

Lates cf. Lates niloticiis Cuvier and Valenciennes 
Posterior portion of a skull (depth of occipital re- 
gion 80 mm), one premaxilla (length 127 mm), 
three vertebrae (diam. 29, 30, 88 mm). Note 
the extreme size of these specimens. 

Collected on lake shore, horizon not known, but probably 
upper unit 
Order Percomorphi 

Family Centropomidae 

Laics cf . Lates niloticus Cuvier and Valenciennes 
Eight vertebrae (diam. 14, 18, 20, 24, 25, 26, 27, 30 
mm) and three dorsal fin spines (40, 44, 46 
mm) 

Family Cichlidae 
Tilapia sp. 

One incomplete fin spine 



DISCUSSION 

Of the living species to which the fossils are most closely com- 
parable — Bagrus hayad, Synodontis schall, Clarias lazera, Lates 
niloticus, Barhus tynni, and Tilapia nilotica — only Bagrus 
hayad, according to Worthington and Riccardo (1936), does not 



1966 QUATERNARY FISHES FROM KENYA 9 

frequent sheltered or shallow waters. Synodontis schall is dis- 
tributed universally in Lake Rudolph. With the possible excep- 
tion of the bigger Tilapia and Lates, the others are confined to 
the sheltered and shallower areas of the lake, such as, for ex- 
ample, Ferguson's Gulf. A tentative conclusion from examina- 
tion of the fossils would therefore be that bed 5 of the upper unit 
was laid down in more open and less shallow waters than bed 8 
of this unit or the upper bed of the lower unit. 

An interesting feature of the fish fauna is the very great size 
of the largest vertebra of Lates from the fifth locality (lake- 
shore, horizon uncertain) ; this is far larger than is known from 
present-day Lates. Daget (1959) and White (1926) have de- 
scribed similar giant Quaternary Lates, and it is interesting to 
discover that such forms were still extant in such relatively very 
recent times, as shown in these Rudolf materials. 

The nature of the fish fauna does not afford any direct evi- 
dence as to the age of the Kangatotha beds, since the Nilotic na- 
ture of the Rudolf fish fauna has been stable at least since the 
Lower Pleistocene disconnection of the lake basin from the Nile 
drainage (see Fuchs, 1939, for general chronology of the area). 
The mammalian fauna, so far as studied, is of Recent aspect. 

The Kangatotha beds belong to the very latest stage in the his- 
tory of the Rudolf basin. Their level and position would suggest 
that they were laid down while the lake was at its largest extent 
after the Gamblian Pluvial, presumably during the Makalian or 
Nakuran "wet phases" (see Cooke, 1957, 1963). A radio-carbon 
date of 4,800 ± 100 years was obtained from a sample of Etheria 
elUptica from the "beaches" on the lake shore, noted above. 
This attests to very recent age of the deposits and implies an ex- 
tremely rapid fossilization. It is of great interest to note that the 
giant forms of Lates, described by Daget, Wliite, and in this 
paper, survived to a very recent date. Finally, with respect to 
the time scale, a note may be added here about one of the arti- 
facts collected on the surface of bed 7 of the upper unit. This is 
a uniserially barbed bone harpoon (now in the Peabody Mu- 
seum, Harvard University). De Heinzelin (1957) lists the dis- 
tribution of similar harpoons and gives a scheme of the evolution 
of harpoon design with an estimate of the equivalent culture. 
(To de Heinzelin 's list should be added a specimen from the late 
Upper Kenya Capsian of Gamble's Cave described by Oakley, 
1961). At first view, the Kangatotha harpoon seems to resemble 
the penultimate stages of design as shown by specimens from the 
"S.F.M." and "G.Y." levels at Ishango (de Heinzelin, 1957), 



10 BREVIORA No. 243 

whereas the "N. tuff." specimens from Ishango, the specimen 
found by Arambourg (1943) at Nanoropus near the end of Lake 
Rudolf, and the Gamble's Cave specimen are of a more advanced 
style. 

ACKNOWLEDGMENTS 

The expedition was financed by NSF Grant No. GP 1188. In 
addition to the acknowledgment made in the introduction, it is a 
pleasure to thank Dr. P. H. Greenwood for his enthusiastic as- 
sistance in the study of the material and Professor Bryan Patter- 
son for valuable comments on the manuscript and for providing 
the sketch map (Fig. 1). 

LITERATURE CITED 

Arambourg, C. 

1943. Mission Scientifique de I'Omo 1932-1933. Tome 1. Geologie- 
Anthropologie. Fasc. II. Mus. Nat. Hist. Natur. (Paris). Pp. 
1-74. 
Cooke, H. B. S. 

1957. Observations relating to Quaternary environments in east and 
southern Africa. Trans. Geol. Soc. S. Africa, 60 (Annex.), 73 pp. 
1963. Pleistocene mammal faunas of Africa, with particular reference 
to southern Africa. Pp. 65-116 in F. C. Howell and F. Bourliere, 
eds., African Ecology and Human Evolution. Aldine Co., Chi- 
cago. 
Daget, J. 

1959. Restes de poissons du Quaternaire saharien. Bull. Inst. Franc. 
Afr. Noire, 23: 182-191. 
Dh Heinzelin, J. 

1957. Les feuilles d 'Ishango. Exploration du Pare Albert, 1957: 46- 
61. 
FucHS, V. E. 

1939. The geological history of the Lake Rudolf basin, Kenya Colony. 
Phil. Trans. Roy. Soc. London, (B) 229: 219-274. 
Oakley, K. P. 

1961. Bone harpoon from Gamble's Cave, Kenya. Antiq. J., 41: 86-87. 
White, E. I. 

1926. The geology and paleontology of the Kaiso bone-beds. Part II. 
Palaeontology. Fossil Pisces. Occas. Paper Geol. Surv. Uganda, 
2: 45-51. 
WORTHINGTON, E. B. AND C. K. RiCARDO 

1936. Scientific results of the Camliridge expedition to the East Afri- 
can lakes, 1930-1, no. 15. The fish of Lake Rudolph and Lake 
Baringo. J. Linn. Soc. London, Zool., 39: 353-389. 

(Received July 26, 1965.) 



BREVIORA 

Miasenam of Comparative Zoology 

Cambridge, ^Mas:-. April 29, 1966 Xttmber 244 

A NEW SPECIES OF ASHMUXELLA FROM WEST TEXAS 
(MOLLUSCA: PULMONATA) 

By W. J. Clexch axd W. B. Miller^ 



The first specimens of this new species were collected by our 
colleag'ue, Dr. J. C. Beqiiaert, in the Davis Mountains, West 
Texas, in May and September 1961. All of the specimens were 
dead shells. During June 1965 live specimens of this species were 
collected by Dr. J. C. Bequaert and W. B. Miller from whicli 
the anatomical drawings were made by the junior author. 

ASHMUNELLA BEQUAERTI Uew SpCCieS 

Plate 1 ; text figures 1, 2 

Holotypi : MCZ 260274, taken from a rockslide about I4 mile 
up a tributary canyon (locally known as Goat Cave Canyon) to 
Little Aguja Canyon near Buffalo Trail Boy Scout Camp at the 
base of the northeastern slope of Black Mountain in the Davis 
Mountains, Jeff Davis County, Texas. Elevation 4,900 ft. (J. C. 
Bequaert and W. B. ^Miller, 5 June 1965). 

Igneous rockslides line both sides of the canyon, below verti- 
cal cliffs; AsJununclla heqiiaerti was found on both sides of the 
canyon, at elevations from 4800 ft. to 4900 ft., along with 
Humboldt ian a palmcri Clench and Rehder, in association with 
Quercus texana, Q. hypoleucoidcs, Acer grandidentata and Rhus 
trilohata. 



1 I'liivtM-sir.v of Arizona, l)t'i)r. of Zo(iloy:y. 



HKKVIOHA Xo. 244 

Paratypcs: AK'Z 2(j()27") I'roni the same locality. 





(ilT.-ltcl- 


r>osser 




[eiglit 


1 )i;i iiictiT 


Diameter 




null 


111111 


111111 




3.9 


12.2 


n..") 


llolotype 


3.8 


13.0 


11.7 


Paratype 


3.7 


]2.4 


11.0 




3.8 


12.2 


11.1 


i i 


3.5 


11.7 


10.5 


i I 


3.4 


11.3 


10.1 


I i 


3.2 


10.7 


9.5 


C i 


3.0 


10.5 


9.3 


I i 



D( script ion : Shell leiitieular, slightly convex above, moder- 
ately convex below, acutely carinate, widely umbilicate, thin, pale 
brown. The surface, above and below and into the umbilicus, is 
sculptured with fine growth wrinkle-striae from which project 
numerous cuticular scales giving a pilose appearance to fresh 
clean shells ; the scales also extend well into the umbilicus. In 
live and recently dead animals, the scales usually hold dirt and 
debris, giving a dusty, dirty appearance to the surface. In older 
specimens where the scales have worn off, there remain raised 
hyphen-shaped papillae, parallel to the growth striae, giving a 
granular ai)pearance to the surface. There are six whorls, the 
first half whorl of the embryonic shell glossy with only a few 
incipient radial striae, the remainder with a silken appearance, 
the fine growth wrinkle-striae and the papillae and scales. The 
first two and a half whorls are convex ; subsequent whorls flat- 
tened. Last whorl descends slightly to the aperture and is 
deeply guttered close behind the outer and basal margins of the 
lip. The under surface of the body whorl is marked additionally 
by very fine, microscopic, spiral striae. The aperture is small and 
very oblique ; peristome white to light brown, reflected except 
near the upper insertion of the lip, the terminations connected 
by a slightly raised callus. Parietal wall with two teeth, the 
larger one situated basally, sinuous, diverging posteriorly toward 
the smaller upper one, thicker anteriorly ; the smaller one close 
to the upper insertion of the lip, raised posteriorly. Outer mar- 
gin thickened, with a flat-topped rectangular tooth set trans- 
versely across the aperture, spanning and overlapping the gap 
between the parietal teeth. On the basal margin, two longitudi- 
nally compressed teeth are connected by a raised ridge along the 
peristome. Interdental intervals are about equal. The umbilicus, 



1!)66 XE\V SI'ECIKS OF ASHMUXKLLA 3 

measured from lower suture of body wliorl just behiud peristome 
to opposite side of body whorl in the pit, is contained about four 
times in the greater diameter of the shell. 

The mantle over the lung is clear except for rare, very small 
•rroups of liiiht »ii-ey pi^i'ment spots. 

Jaw with ten ribs. 

Genitalia. Penis with lower sac short and wide, upper sac 
longer and narrower. In the upper sac there are three longitudi- 
nal ridges, in addition to a small nodule which is attached to the 
wall of the penis at its junction with the epiphallus. In whole 
mounts, this nodule gives the appearance of a short verge when 
seen by transparency ; dissection of the penis, however, reveals it 
as a growth on the side of the penial wall. It is possible that this 
nodule, together with the tip of the muscular walls of the epiphal- 
lus, acts as a papilla when the penis is completely everted. The 
lower sac has two thickened processes w'hich come together near 
the lower end and form a constriction within the penial cavity. 
There is a short penial retractor on the epiphallus which is in- 
serted in the floor of the lung cavity ; loose strands of connective 
tissue connect the penis with base of epiphallus. Talon of the 
multilobar type. Length of penis 5.0 mm, epiphallus 24.0 mm, 
flagellum 1.5 mm, penial retractor 1.3 mm, spermatheca and duct 
31.0 nun, vagina 4.0 nnn, free oviduct 2.5 mm, atrium 1.0 mm. 

Remarks: Ashmu7ieUa hequaerti is the most easterly known 
species of Ashmunella in the U. S. It is most closely related to 
the mearnsi group, in shell characteristics as well as geographi- 
cally. In paratypes, the spire varies from moderately raised 
conical to completely flat ; the diameter varies from 10.5 mm to 
13.0 mm. Its nearest relative appears to be A. hchardi Pilsbry 
and Vanatta, with which it agrees in the lenticular shape of the 
shell, the sharply carinate periphery, and the general shape of 
the teeth, including the two parietal teeth. In .4. hequaerti, 
however, the shell is generally more flattened, the edge of the 
parietal callus is appressed and very weak, not raised into a low, 
free ridge as in hehardi, and the upper insertion of the peristome 
is only weakly descending, nearly straight, not strongly descend- 
ing as in hehardi. Anatomically, the internal nodule at the 
upper end of the upper penial sac has not been seen or reported 
in any other Ashmunella. It remains to be seen from additional 
dissections, however, whether this is a consistent characteristic. 

The new species is named after Dr. Joseph C. Bequaert, life- 
long malacologist and entomologist, who first discovered this 



BREVIORA 



No. 244 




Fig. 1. A.slnii undid heqtuu-rti Cleiii-h and Miller. Dissection of reproduc- 
tive system with illustrations made from stained whole mount, a. Entire 
system, b, Enlargement of the distal end of the epiphallus at its junction 
with the vas deferens where the two are bound togetlier. This may be ealled 
the flagellum portion of the epiphallus. There is no free flagellum. 

1, Genital oritice; 2, Atrium; 3, Vagina; 4, Oviduct; 5, Prostate; 6, 
Uterus; 7, Albumen gland; 8, Talon; 9, Hermaphroditic duct; 10, Sperma- 
theeal duct; 11, Lower sac of penis; 12, Upper sac of penis; 13, Penial 
retractor; 14, Epiphallus; 15, Yas deferens; 16, Connective tissue; 17, 
Flagellum bound with vas ileferens; 18, Penis papilla. 



19(36 



XEW SPECIES OF ASIIMUXELLA 



snail on 2 May 1961, obtaining quantities of dead shells hut no 
live animals. On 5 June 1965, he and the junior author returned 
to the locality and were successful in obtaining 4 live adults and 
3 live immatures. Dead shells were very numerous. One of the 
live adults was designated the holotype. Two others were dis- 
sected to corroborate anatomical findings. The fourth live adult 
and the three live immatures are being kept alive in a terrarium 
at the University of Arizona, in the hope of obtaining additional 
studies on the adult anatomy for comparison. 




5 mm. 

Fig. 2. Ashniiinella hequucrti CleiR-h and Miller. Lower portion of repro- 
dnctive system enlarged. (Xunil)ering as in Figni-e 1.) 



EEFEKEXCES 

PiLSBRY, II. A. 

19-1:0. Land Mollnsca of North America. Academy of Natural Sciences, 
Philadelphia, Monograph No. 3, vol. 1, pt. 2, pp. 912-977. 

19-18. The genera Hiimboldtiana, SonoreUa, Orcohelix and Ashmunella. 
Proc. Acad. Nat. Sci., Philadelphia, vol. 100, pp. 199-203. 

(Received 24 January, 1965.) 



BREVIORA 



No. 244 




Figs. 1-2. AsltmuneUa bequaerti Cleiu-h iind ^filler, from Goat Cave 
Canyon, Black Mt., Davis :Mts., Texas. Fig. 1. I'aratype, MCZ 260275 (4 
X). Fig. 2. IldlotyiK'. MCZ 2(i(i274 (5.4 X). 



BREVIORA 

Mmseiuim of Comparative Zoology 

Cambridge, Mass. April 29, 196() Number '24!\ 

NOTES AND DESCRIPTIONS OF NEW UROCOPTIDAE 

FROM CUBA AND HISPANIOLA (MOLLUSCA: 

PULMONATA) 

By William J. Clench 



The followino- notes and descriptions of new species are based 
upon material in the Museum of Comparative Zoology which has 
been collected over the past several years. All species are mem- 
bers of the family Urocoptidae, one of the most dominant fami- 
lies in the West Indies and in a lesser way in Central America 
and in the southwestern United States. This family occurs also 
in northern South America, but species are few and present only 
a very minor element in the molluscan fauna there. 

UROCOPTIDAE 

Archegocoptis Pilsbry 

Archegocoptis Pilsbry, 1903, Manual of Conchology (2) 15:301 (type 
species, CylindreUa crenata Weiiiland and Martens, original designation). 

So far as known, this genus is confined to the southwestern or 
Tiburon Peninsula of Haiti. Specific localities for the few spe- 
cies are all from the Departement du Sud. 

According to Pilsbry, Archegocoptis is nearest in relationship 
to Eucalodium Crosse and Fischer, a genus limited in distribu- 
tion to southern Mexico and Guatemala. 

The known species are as follows : 

Archegocoptis barbot'ri Clench 

Archegocoptis barbouri Clench, 1935, Proc. Boston Soc. Nat. Hist. 41:5, 
pi. 1, tigs. B, F (Tardieu, Mt. LaHotte, Haiti, ± 3000 feet). [Holotype, 
MCZ 108593.1 



2 BREVIORA No. 245 

Archegocoptis crenata (Weinland and Martens) 

CylindreUa crenaia Weinland and Martens, 1859, Malakozoologische Blat- 
ter 6:54 (Jeremie, Haiti); Pfeiffer, 1869, Xovitates Conchologicae 3:440, 
pi. 97, figs. 33, 34 (Haiti). 

Archegocoptis darlingtoni Clench 

Arcliegooo'ptis darlingtoni Clench, 1935, Proc. Boston See. Nat. Hist. 41:6, 
pi. 1, figs. C, G (Tardieu, Mt. LaHotte, Haiti, ± 3000 feet). [Holotype, 
MCZ 108592.] 

Archegocoptis decapitata (Roding) 

Cerion decapifatum Eoding, 1798, ^luseum Boltenianum, p. 90 (no locality 
given [Haiti]). [See below.] 

Archegocoptis eximia (Pfeiffer) 

CylindreUa eximia Pfeiffer, 1857, Malakozoologische Blatter 4:232 (lo- 
cality unknown) ; Pfeiffer, 1869, Novitates Conchologicae 3:439, pi. 97, figs. 
30-32 (Haiti). 

Archegocoptis tiburonica Clench 

Archegocoptis tihuronlca Clench, 1935, Proc. Boston Soc. Nat. Hist. 41:6, 
pi. 1, fig. I. (Tardieu, Mt. LaHotte, Haiti, between 3-4000 feet). [Holo- 
type, MCZ 108591.] 

Archegocoptis haitiexsis new species 
Plate 1, fio-nres 1, 2 

Holotype. MCZ 252056, from the top of Morne Rochelois, 25 
km WSW of Miragoane, Dept. dn Snd, Haiti, about 3000 feet, 
collected by W. J. Eyerdam, July 1927. 

Paratypes. A series of 5 paratypes from the same locality, 
MCZ 252057. 



Height 


Width 




mm 


mm 




41 


10 


Holotype (see Remarks) 


27 


10 


Paratype 


26.5 


10 


i ( 


26.5 


10 


i I 



Description. Shell medium in size, reaehino- 42+ n\m in 
length (decollated), minutely umbilicate and coarsely sculptured. 



1966 NEW SPECIES OF UROCOPTIDAE 3 

Color a more or less uniform choeolate-brown and shining on the 
coarse, axial riblets and dull in between. AVhorls 15-|- and flat 
sided. Suture well defined. Spire extended, the upper portion 
slightly concave. Aperture holostomatous, subcircular, slightly 
flaring and reflected. Axis simple. Sculpture consisting of a 
series of somewhat irregular axial costae which are diagonal and 
fine, irregular axial threads in between the costae. There is a well 
defined basal ridge. 

Remarks. In relationship, this species is nearest to ArcJiego- 
coptis iihnronica Clench, differing, however, by being much lar- 
ger, lacking the whorl shoulder and in having a much larger 
basal ridge. The sculpture is similar in both species, though 
coarser in A. tihuronico. 

Many species in the family Urocoptidae lose many of the 
upper whorls when approaching the adult stage or during their 
adult life. They produce a plug in one of the mid or earlier 
whorls and later the whorls above the plug are lost. The fracture 
is a mechanical one and not automatic, as quite frequently a 
few adult specimens in a single population may still retain their 
early whorls. It is of considerable interest to note that A. haitien- 
sis apparently does this twice, as three of the five paratypes 
have plugs seven whorls above the aperture ; the remaining two 
paratypes are young specimens and have lost the early three or 
four whorls. Onh' the holotype has retained the mid-whorls. 

Archegocoptis decapitata (Roding) 

Cerion decapitatum Roding, 1798, Museum Boltenianuni, p. 90 (refers to 
Chemnitz, 1786, Conchylien-Cabinet (1)9: pi. 136, figs. 12.56-1257 [fig. 1255, 
in error] ). 

Cyclostoma fasciatn Lamarck, 1816, Eneyclopedie Metliodique, Atlas, p. 
12, pi. 461, fig. 7. 

Helix truncafa Dillwyn, 1817, Descriptive Catalogue of Recent Shells, 
London 2:948 (Santo Domingo). 

Helix fasciata Lamarck: Ferussac, 1822, Tableaux Systematiques, p. 61, 
no. 503 (The Antilles). 

Cylindrella fasciata 'Chemnitz' Pfeiifer, 1862, Malakozoologische Blatter 
9:199 (Corail [Haiti]). 

Cylindrella fasciata 'Chemnitz' Crosse, 1891, Jour, de Conch. 39:134 (Co- 
rail, near Jeremie, SW Haiti). 

Urocoptis tntncata (Dillwyn) : Pilsbry, 1903, Manual of Conchology 
(2)15:154, pi. 39, figs. 27-28 (Corail, near Jeremie, Haiti). 

Cyclostoma fasciata Lamarck: Mermod, 1952, Revue Suisse de Zoologie 
59:46, fig. 111. 



4 BREVIORA No. 245 

Remarks. This species has had a long and varied history. The 
only specific locality previously known was that of Weinland 
who had collected a single worn specimen in a small water course 
at Corail, Haiti (Crosse, 1891). From this, Pilsbry had assumed 
that it had been carried down this stream from the mountains 
above. In 1927, W. J. Eyerdam collected several dead specimens 
under stones at the base of a cliff in a forested area on Grande 
Cayemite, a small island about 61^ miles NE of Corail. It is 
possible that the specimen found by AVeinland may have drifted 
from Grande Cayemite. The specimens collected by Eyerdam 
are certainly in the genus Archegocoptis and appear to be A. 
decapitata Eoding, differing only in that the color band of dull 
red is on a whorl lower than those figured by both Chemnitz and 
Lamarck. All specimens so far as known were collected dead. 

Urocoptis (Gongylostoma) wiiittumi new species 
Plate 2, figure 1 

Holotype. MCZ 59580, from Guabairo, near Soledad, Cien- 
fuegos. Las Villas, Cuba, collected by Clench, Crozier and Xavez, 
October 1928. 

Paratijpcs. MCZ 59284, 86289, 105148, all from the above lo- 
cality, and collected by various students and visitors to the 
Harvard Tropical Gardens. 



Height 


Width 




mm 


mm 




11.5 


2.7 


Paratype 


11.6 


2.5 


1 1 


14.8 


3 


i i 


16 


3 


( ( 


17.3 


3.2 


Holotype 


17.5 


3 


Paratype 


18.5 


3 


" 


20.5 


3.5 


" 



Description. Shell somewhat fusiform, usually decollated, nar- 
row, widest point a little below the middle in the decollated speci- 
mens. Color very pale reddish brown, darker on the earlier 
whorls. Whorls 18 to 22 (not always decollated at the same 
whorl). Whorls rather flat, evenly tapering towards the spire 
tip. Last whorl free for one-quarter to one-third of its length. 
Spire acute, first two whorls a little larger than the third and 
fourth. Aperture holostomatous, circular, white and reflected. 



1!)6G NEW SPECIES OF UROCOPTIDAE 5 

Columellar axis stout, encircled by a single, wide lamella which 
bears numerous deep, spine-like crenulations. Sculptured by nu- 
merous strong, almost straight riblets, about 17-18 on the second 
whorl. 

Remarks. This species belongs to the group of U. canteroiana, 
but differs in having fewer and stronger riblets, different shape 
and proportions, and by its internal axis, which is stouter; it is 
also encircled by a larger lamella, the margin of which possesses 
a greater number of tooth-like processes. 

This shell is very variable in size, but very constant in its rela- 
tive proportions. The riblets are approximately the same in 
number on the different sizes. 

Named for Mr. Walter Whittum, now of Springfield, Massa- 
chusetts, whose kindness and courtesy made many trips possible 
in the rich collecting ground about Guabairo, at the time he 
was in charge of a colonia for the Soledad Sugar Company in 
1928. 

Urocoptis (Goxgylostoma) exquisita new species 
Plate 2, figure 2 

Holotype. MCZ 59286, from one-half mile E of Guabairo, 
Soledad, Cienfuegos, Cuba, W. J. Clench collector, December 
1927. 

Paratypes. MCZ 59285, 59579, from the same locality. 



ire specimens 




Decollated specimens 


Height 




Height 


mm 




mm 


14.4 


Holotype 


12.7 


14.4 




12.5 


12.9 




11.9 



Description. Shell fusiform, slender, generally entire ; when 
decollated, limited to only a very few of the early whorls. Widest 
point about midway in decollated shells. Color light horn, not 
uniform, but generally somewhat marbled with patches of lighter 
and darker areas, occasionally albinistic ; surface somewhat glis- 
tening, often translucent in certain areas. Whorls convex, evenly 
tapering towards both the spire tip and the aperture. In complete 
shells whorls number 21-23; in decollated specimens 15-18 whorls. 
Spire acute, first tw^o whorls slightly larger than the third and 
fourth. Last whorl free, developed with a long, slightly curved 
neck. Aperture holostomatous, circular, white and well reflected. 



6 BREVIORA No. 245 

Axis of columella thin and sinuous with a single spiral thread. 
Suture somewhat impressed. Sculpture consisting of very fine 
oblique axial riblets except on the free whorl which has many 
relatively coarse riblets, irregularly spaced and nearly encircling 
the whorl except on the face or aperture side. 

Remarks. This species belongs in the suligenus Gongylostoma 
Albers, near Urocoptis harhouri Torre and Clench. It differs 
from U. harhouri, however, in being smaller, and with a larger 
number of whorls in the decollated specimens, and is relatively 
more slender and more fusiform. The color is very similar, 
though the shell is a little more translucent in cxquisita. The 
free aperture whorl is also much longer than in TJ. harhouri. 

The habitat of this species is quite peculiar for the genus in 
this region, as specimens were found in more or less heavy woods, 
on the rocky walls of cave entrances, or in other large rocky 
fissures. The other urocoptids occupied more open areas, espe- 
cially U. Jivida harhouri and V . livida aikinsi Torre and Clench. 

Urocoptis (Gongylostoma) canteroiana (Arango) 
Plate 2, figure 4 

CyllndrcUa canieroiana 'Gundlac-h' Arango, 187.") [1876], Aiiales Real 
Academia Ciencias Medicas, Fisieas y Naturales, Hal^aiia 12:284 (La Vigia, 
Trinidad [Cuba]); Arango, 1878, Fauna Malacologica Cubana, Haliana, p. 
117 (environs of Vigia, Trinidad). 

Urocoptis canteroiana 'Gundlacli' Arango: PilsV)ry, 1903, Manual of 



Conehology (2)15:2^4. 




Height 


Widtli 


mm 


umi 


12.5 


2.5 



Lectotype 

Lectotype. Here selected, MCZ 189252, from La Vigia, Trini- 
dad, Cuba, collected by Gundlach and sent to MCZ by Kafael 
Arango. 

ReiHarks. This species has not been figured previously. It is 
comparatively rare as only a few specimens have been collected. 
We have specimens from La Vigia, Finca Cantero, and Pinca La 
Pastora, all in the vicinity of Trinidad, Las Villas, Cuba. 

Urocoptis (Gongylostoma) diagonalis new species 
Plate 2, figure 3 

Holotypc. MCZ 59292, from Mina Carlota, Sierra de San 
Juan, 8 miles S of Cumanayagua, Las Villas, Cuba, W. J. Clench 
and Calvin Goodrich collectors, November 30. 1927. 



1966 NEW SPECIES OF UROC'OPTIDAE 7 

Paratypc. A single i)aratyi)e from the same locality is iii the 
Museum of Zooloo'v, University of Michigan. 



eight 


Width 




mm 


mm 




16 


3.3 


Holotype 


15 


3 


Paratype 



Description. Shell reaching 16 nnn in height (decollated), 
fusiform, imperforate and sculptured. Color a dull gray. Whorls 
16 (remaining), very slightly convex, the last free for about one- 
fifth of a whorl. Suture indented. Spire decollated, a loss of 
12-14 early whorls. Aperture holostomatous, white and flaring. 
Sculpture consisting of numerous fine, axial riblets which are 
slightly diagonal and are flatly sigmoid in shape. On the free 
portion of the last whorl these riblets become rings and are 
closer together. 

Remarks. This species is related to U. canteroiana (Arango), 
but differs in several of its characters. U. diagonalis is larger, 
has finer sculpture of diagonal riblets which are straight in can- 
teroiana and flattened sigmoid in shape in diagonalis. 

Urocoptis (Urocoptola) cayemitensis new sj^ecies 
Plate 1, figure ^ 

Holotype. ]\IC'Z 254666. from XE Grande Cayemite, a small 
island about 614 miles XE of Corail, Departement du Sud, Haiti, 
collected by W. J. Eyerdam, July 1927. 

Paratxjpes. MCz' 254667, population 1; and MCZ 254668, 
population 2. From the same locality as the holotype. 

Height Width 

mm mm 

Holotyi^e, population no. 1 ^ 
Paratype ' ' 



population no. 2 ^ 



Description. Shell oblong, glossy, having the greatest diameter 
above the middle, imperforate, sculptured and rather thin in 
structure. Color a pale brownish pink. AVhorls very slightly 

1 Both populations from the same general area. 



17.2 


6.5 


18.5 


7 


17.3 


5.8 


15.8 


6 


20.5 


7.1 


20.4 


7 


20.1 


7 



8 BREVIORA No. 24o 

convex and tapering above the mid-whorls to the truncated sum- 
mit. Last whorl with a distinct basal ridge. Spire extended. 
Aperture nearly circular, the parietal area adnate to the whorl 
above, lip flaring. Suture moderately impressed. Sculpture con- 
sisting of numerous, slightly diagonal, axial costae, which do not 
produce crenulations at the suture. Axis with a single twist. 

Remarks. This species is a member of the U. sericea (Pfeiffer) 
complex which is widely distributed along coastal Haiti within 
the Gulf of Gonave. Urocoptis caycmitensis diifers from F. seri- 
cea by being much smaller, imperforate, and in having a uniform 
sculpture throughout. The spire is truncated between the sev- 
enth and eighth whorls from the aperture. 

Urocoptis (Urocoptola) ekmani new species 
Plate 1, figure 5 

Holotype. MCZ 260871, from La Source, NW Gonave Island, 
Haiti, collected by W. J. Eyerdam, August 1927. 

Paratypcs. MCZ 260872, from the locality of the holotype. 

Height Width 



25 


11 


Holotype 


26 


11 


Paratype 


22.5 


10.2 


i i 



Description. Shell large, elliptical in shape, remaining whorls 
reaching 26 mm in height, imperforate, rather solid and sculp- 
tured. Whorls 8 (remaining) and moderately convex. Flesh 
colored with a few, small, irregular, dark brownish red flecks on 
all of the whorls. Spire extended and tapering rapidl.y from the 
mid-area and produced at an angle of about 55°. Aperture sub- 
circular, holostomatovis, one specimen out of eight being adnate 
above. There is a slight and closed rimation at the umbilical 
area. Suture slightly indented. Sculpture consists of diagonal 
and straight, fine axial riblets on the upper whorls and diagonal 
and arcuate axial riblets on the lower whorls. Protoconch un- 
known. 

Remarks. This species is related to Urocoptis hencomoi Clench, 
but is smaller, proportionally narrower, and has the lip holosto- 
matous. 

Named for Dr. Erik L. Ekman, the Swedish botanist who was 
with W. J. Eyerdam during his trip to Gonave Island. 



1966 NEW SPECIES OP UROCOPTIDAE 9 

Urocoptis (Urocoptola) caribbaea new species 
Plate 1, figure 3 

Holotype. MCZ 260873, from 1 mile E of Pointe a Raquette, 
south central Gonave Island, Haiti, collected by W. J. Eyerdam, 
July 1927. 

Poratypes. MCZ 260874, from the same locality. 

Height Width 

mm mm 

24.8 12 Holotype 

22 11.6 Paratype 

22.5 11 

22.5 11.4 

Description. Shell large, reaching about 25 mm in height, im- 
perforate, rather solid and sculptured. Whorls 7 (remaining) 
and moderately convex. Flesh colored with a few small, irregu- 
lar, dark brownish red flecks on all of the whorls. Spire extended 
and tapering to the apex and the base from the mid-area and 
produced at an angle of about 60°. Aperture subeircular and 
adnate on the parietal area. Suture slightly indented. Sculpture 
consists of diagonal and straight, fine axial riblets and somewhat 
arcuate on the last whorl. Riblets on the two remaining early 
whorls more widely spaced. Protoconch unknown. 

Remarks. This species is closely related to U. ekmani, differing 
in being shorter, with different proportions, having all specimens 
adnate on the parietal area and in having the axial riblets more 
widely spaced on the remaining early whorls. 

Urocoptis (Idiostemma) perplicata (Beck) 

Helix (CocJdodina) perplicata Ferussac, 1821 [1822], Tableaux Sys- 
lematiques Animaux Mollusques, p. 61 or 65, no. .^06 (The Antilles) [no7nen 
nudum]. 

BrachypodeUa perplicata Ferussac: Beck, 1837, Index Molluseorum, p. 
89 [refers to Ferussac 's pi. 163, fig. 9]. 

Clausilia perplicata Ferussac: Deshayes [in] Lamarck, 1838, Animaux 
sans Vertelires (2)8:216. 

CyUndrella perplicata Ferussac: Philippi, 18-17, Abbildungen Conchylien 
2:217, pi. I (Achatina), fig. 9 (The Antilles). 

CyUndrella perplicata Ferussac, 1851, Histoire Naturelle Generale et 
Particuliere 2:229, pi. 163, fig. 9 (The Antilles); non CyUndrella perplicata 
Ferussac. Pt'eiflfer 1840 [in] Weigniann, Archiv fiir Xaturgeschicte 1:41 
(Fundador, Matanzas, Cuba). 



10 BREVIORA No. 245 

CyUndrella fastigiata 'Gundlach' Pfeift'er, 1860, Malakozoologische Blat- 
ter 7:20 (Baracoa, Mata, and Yunque, Cuba); Pfciffer, 1865, Novitates 
Conchologicae 2:263, pi. 65, figs. 23-25 [Syntypes, MCZ 26639, 86621]. 

Urocopti.s fasiigmia Pfeiffer: Pilsbry, 1903, Manual of Conchology 
(2)15:171, pi. 45, figs. 36-41 (Baracoa, El Yunque, Mata, Oriente, Cuba). 

BrachypodeUa perplicata Ferussae : Pilsbry, 1903, Manual of Conchology 
(2) 16:83, pi. 7, figs. 23-24 (Antilles). 

Remarks. Considerable confusion has existed concerning the 
proper name for this species. Pilsbry retained both perplicata 
and fastigiata in two different genera but noted under perplicata 
that "the ribs follow one another from whorl to whorl, as in 
Urocoptis fastigiata,''^ but he gave no indication that the two 
names applied to the same species. 

Since Pilsbry 's Manual of Conchology was published in 1903 
far more material in this family is available for study, not only 
from Cuba but from the entire West Indies as well. 

The credit for the name of this species must go to Beck rather 
than to Ferussae, as Ferussae did not publish this name until 
1851. The plates were issued much earlier but there were no 
names attached to them. Through some source, other than publi- 
cation, these names were available, and Beck was the first to cite a 
plate and figure for this species. 

Specimens examined. CUBA : Oriente, Farallones de Ba- 
rigua; Yunque de Baracoa; Silla de Baez, W of Baracoa; Sa- 
bana, Cabo Maisi ; Zona de la Caleta, Baracoa ; Arriba Mandinga, 
Baracoa; Arriba Guandao, Mandinga; Mata, Punta Maisi; Boca 
de Taco, Nibujon, Baracoa. 

Brachypodella (Brevipedella) imitatrix Pilsbry 

BrachypodeUa {Brevipedella) imitatri.r Pilsliry, 1903, Manual of Conchol- 
ogy (2)16:47, pi. 8, figs. 54-55 (Port-au-Princo, Sans-Souci, St. Marc, and 
La Perriere, Haiti). 

Remarks. This species has a much wider distribution than 
most species in the Urocoptidae, extending as it does from Haiti 
to the Samana peninsula in eastern Santo Domingo. 

A closely related species, B. anguUfcra (Gundlach), occurs in 
eastern Cuba. 

Specimens examined. Haiti: Bizonton, Lasbaines; Port-au- 
Prince and Diquini. Santo Domingo : Mt. Isabel de Torres, 
Puerto Plata, at 1200 to 1600 feet; Sanchez; Pefion de Basiles 
and Peiion de Maria Luisa, both Santa Barbara de Samana, and 
Punta Lirio, 2 miles E of Santa Barbara de Samana. 



l!)(i(i NEW SPECIES OF UKOCOPTIDAE 11 

Brachypodella (CtYRAxis) samana new sj)ecies 
Plate 2, figure 5 

Holofype. MCZ 57214, from Pefioii de Maria Luisa, Santa 
Barbara de Samana, Republica Dominicana, collected by W. J. 
Clench, H. D. Russell and R. A. McLean, August 1937. 

Paratypes. MCZ 57218, from the same locality ; and MCZ 
57213 from Peiion de Basiles in the same general area. 



h'ight 


Width 




mill 


inui 




14.5 


2.5 


Holotype 


12.5 


2.4 


Paratype 


14.5 


2.5 


( i 


12.8 


2.4 


i i 



Description. Shell slender, imperforate, generally entire, 
widest at the mid-section of the shell and finely sculptured. 
Color marbled in yellowish and light brown with some areas 
white, the brownish areas translucent, the white and yellowish 
areas somewhat opaque. AYliorls 13 to 14 in entire specimens 
and moderately convex witli the last whorl free. Suture well 
defined. Sjnre acute, aperture holostomatous, sul)circular with 
the lip fiaring slightly. Axis gyrate. Sculpture consisting of 
numerous, fine, axial, slightly diagonal and slightly curved 
costae ; the first two whorls finely and axially costate. On the 
early whorls certain of these costae in small groups impinge 
slightly on the suture. 

Remarks. This species is closely related to Brachypodella 
{Gyraxis) scricata Pilsbry. It differs by being larger, having 
slightly coarser thread-like costae and having the marbled colora- 
tion, scricata being a dull white. 

Brachypodella scricata Pilsbry was collected originally by 
William M. Cabb during 1869-71 while on a geological survey of 
Santo Domingo. Unfortunately, the recent mollusks collected by 
him seldom had specific locality data. We collected this species 
in some numbers at San Lorenzo Bay, Bahia de Samana. This 
locality can be considered the type locality as Gabb surveyed this 
area about the Bahia de Samana. 

(Received December 1, 1965.) 



12 BREVioRA No. 245 



Plate 1 

Figs. 1-2. Archegocoptis haitiensis n. sp. Monie Eochelois, 25 km WSW 
of Miragoaiie, Dept. du Sud, Haiti, at 1000 meters. Fig. 1, Paratype, MCZ 
252057; fig. 2, Holotype, MCZ 252056 (both 3X). 

Fig. 3. Urocoptis (Urocoptola) caribhaea n. sp. 1 mile E of Pte. a Ea- 
quette, S central Gonave Island, Haiti. Holotype, MCZ 260873 (3X). 

Fig. 4. Urocoptis {Urocoptola) cay emit ens is ii. sp. NE Grande Cayemite, 
Dept. du Sud, Haiti. Holotype, MCZ 254666 (4X). 

Fig. 5. Urocoptis (Urocoptola) ekmani n. sp. La Source, XW Gonave 
Island, Haiti. Holotype, MCZ 260871 (3X). 



1!»G6 



NEW SPECIES OF UROCOI'TIDAE 



13 




^f 4 






X 



.#'^'i^. 




I 2 

J 





Plate 1 



14 



BREVIORA 



No. 245 




Plate 



Fig. 1. Urocoptis {Gongy]os1om(t ) irJiiftiinu ii. sp. (hinliaini, Soledad, Las 
Villas, Cuba. Holotype, MCZ .59580 (5.5 X). 

Fig. 2. Urocoptis (Gongylostoma ) ('.rquisita, n. sp. % mile E of Guabairo, 
Soledad, Las Villas, Cuba. Holotype, MCZ 59286 (5.5 X). 

Fig. 3. TJrocoptis (Gongylostoma) diagoruilis n. sp. Miua Cailota, Siena 
de San Juan, 8 miles S of Cumanayagua, Las Vill.-is, Cul>,i. Holotype, M<'Z 
59292 (5.5 X). 

Fig. 4. Urocoptis (Gongylostoma) canteroiana (Aiango). (La Vigia, 
Trinidad [Las Villas, C\iba]). Lectotype, MCZ 189252 (5.5 X). 

Fig. 5. Brachypodella (Gyraxis) samann n. sp. T'eiion de Maria Luisa, 
Santa Barbara de Samana, Repulilica Doniiuii ana. Holotype, MCZ 5721-t 
(5.5 X). 



BREVIORA 

Miuseiiiim of Coinnp,arative Zoology 

Cambridge, Mass. AIay 3, 1966 Number 246 

PSEUDANTHE88IUS PR0CURREN8 N.SP., A CYCLOPOID 

COPEPOD ASSOCIATED WITH A CIDARID ECHINOID 

IN MADAGASCAR 

By Arthur G. Humes 

Boston University, Boston, Massachusetts 

and 

Associate in Marine Invertebrates, Museum of Comparative Zoology 

INTRODUCTION 

During an extensive search in 1963-64 for copepods associated 
with marine invertebrates at Nosy Be, in northwestern Mada- 
gascar, 109 adults and 9 copepodids of the new lichomolgid cope- 
pod described below were recovered from the sediment obtained 
after washing 30 large pencil urchins, Phyllacanthus imperialis 
(Lamarck), in weakly alcoholized sea water. (The host echinoid 
is widespread in the Indo-Pacific region, where it occurs, for ex- 
ample, in Australia, the Marshall Islands, the Philippine Islands, 
Ceylon, the Red Sea, and Zanzibar) . This new form brings the 
total number of species known in the genus Pseudanthessius to 
24 (including tlie 22 species listed by Stock, Humes, and Gooding, 
1963, and a new species from a polychaete annelid in Madagascar 
M^hose description by Humes and Ho is in press). 

ACKNOWLEDGMENTS 

The copepods were collected by the author while participating 
in the 1963-64 activities of the U.S. Program in Biology of the 
International Indian Ocean Expedition. 

The study of the specimens has been aided by a grant (GB- 
1809) from the National Science Foundation of the United 
States. 

I wish to thank Dr. H. Barraclough Fell, Professor of In- 
vertebrate Zoology at the Museum of Comparative Zoology, for 
the identification of the echinoid host, and to acknowledge the 
assistance to the field work given by the staff of the Centre 
d'Oceanographie et des Peches at Nosy Be. 



2 BREVIORA No. 246 

DESCRIPTION 

Family LICHOMOLGIDAB Kossmann, 1877 
Genus Pseudanthessius Glaus, 1889 

PSEUDANTHESSIUS PROCURRENS ^ n.Sp. 

Figures 1-29 

Type material. — 16 females, 9 males, and 1 copepodid from 
washings of 3 pencil urchins, Phyllacanthus imperialis (La- 
marck), in 1 meter depth among dead coral (Acropora) at Pte. 
Ambarionaomby, Nosy Komba, near Nosy Be, Madagascar. Col- 
lected November 28, 1963. Holotype female, allotype, and 19 
paratypes (13 females and 6 males) deposited in the U. S. Na- 
tional Museum, Washington, and the remaining paratypic adults 
(dissected) together with the copepodid in the collection of the 
author. 

Other specimens (all from Phyllacanthus imperialis collected 
in 1963 at the type locality). — 2 females from 1 host, July 3; 
10 females and 7 males from 3 hosts, July 18 ; 14 females and 5 
males from 6 hosts, August 23 ; 13 females, 6 males, and 4 cope- 
podids from 7 hosts, October 30 ; and 16 females, 11 males, and 
4 copepodids from 10 hosts, December 14. This last collection is 
deposited in the Museum of Comparative Zoology. 

Female. — The body (Figs. 1 and 2) has a broadened pro- 
some. The length (excluding the setae on the caudal rami) is 
0.95 mm (0.90-1.01 mm) and the greatest width is 0.44 mm 
(0.42-0.46 mm), based on 10 specimens. The ratio of the length 
to the width of the prosome is 1.28 :1. The segment bearing leg 1 
is almost completely fused with the head, the only indication of 
separation being a short weak crease on each side. The lateral 
areas of the metasomal segments are rounded. 

The segment of leg 5 (Fig. 3) is expanded laterally, being 50 \>. 
in length and 133 [x in width. Between the segment of leg 5 and 
the genital segment there is a ventral intersegmental sclerite (see 
Fig. 2). The genital segment (Fig. 3) is 127 ii long. Anteriorly 
its lateral margins form 2 rounded, strongly sclerotized lobes 
(the width of the segment at this level being 115 \).). The width 
at the level of the dorsolateral areas of attachment of the egg sacs 
is 107 [i. Behind each attachment area the segment is slightly 



1 The specific name proctirrcns, from Latin procurrcre, meaning to bulge out or 
project, alludes to the outer expansion on the coxa of leg 1 and to the 2 rounded 
lobes on the anterior part of the genital segment in the female. 



1966 NEW COPEPOD FROM AN ECHINOID 3 

constricted with nearly parallel margins (the width in this re- 
gion being 86 [x). Each egg sac attachment area (Fig. 4) bears 
anteriorly a slender, slightly haired seta (21 i;. long) and jnst 
posterior to it a short naked seta (6.5 {x long) composed of an ex- 
panded sclerotized basal portion and a slender hyaline distal 
part. Medial to the latter seta there are 2 small spinelike proc- 
esses. The three postgenital segments are 44 x 81, 39 x 75, 
and 65 x 72 [j. from anterior to posterior. The anal segment bears 
on each side on its distal margin a dorsal and ventral row of 
spinules. 

The caudal ramus (Fig. 5) is elongated, with a terminal 
ventral expansion whose margin bears a row of spinules. The 
length along the inner side of the ramus to the end of the ex- 
pansion is 114 pL, along the outer side 104 \i, and the width at the 
level of the outer seta is 24 [f.. The ratio of length to width is 
about 4.5 :1. The outer seta, inserted 70 [i from the base of the 
ramus, is naked and 56 [>. in length. The pedicellate dorsal seta is 
33 [I and slightly haired. The outermost terminal seta (100 \l) 
and the innermost terminal seta (12 [).) are haired. The 2 long 
median terminal setae are 177 and 250 [x in length respectively 
and haired. A minute lateral setule is borne on the outer basal 
margin of the ramus. The dorsal and ventral surfaces of the 
ramus bear a few refractile points. 

The dorsal surface of the prosome and the dorsal and ventral 
surfaces of the urosome bear minute setules and refractile points. 
In addition, the outer ventral areas of the head carry a sub- 
marginal row of refractile points (Fig. 6). The ratio of the 
length of the prosome to that of the urosome is 1.53 :1. 

The egg sacs (Fig. 1) are moderately elongated, often rather 
pointed posteriorly, and contain numerous eggs. In one female 
the egg sacs measured 385 x 220 [jl, with each egg about 57-60 (jl 
in diameter. 

The rostral area (Fig. 7) is moderately well developed. Be- 
tween this area and the front of the labral region there is a 
slight protrusion on the ventral surface of the head. 

The first antenna (Fig. 8) is 7-segmented, with the lengths of 
the segments (measured along their posterior non-setiferous mar- 
gins) 24 (39 [J. along the anterior margin), 103, 21, 39, 33, 20, 
and 17 [x respectively. The formula for the armature is 4, 13, 6, 
3, 4 -f 1 aesthete, 2+1 aesthete, and 7+1 aesthete. All the 
setae are naked. 

The second antenna (Fig. 9) is 4-segmented, with the last seg- 
ment elongated (69 [jl along the shorter ventral margin, 93 [x along 



4 BREVIORA No. 246 

the dorsal margin, and 19 [x in width). Each of the first two seg- 
ments bears a small ventral seta, the third segment bears 4 
slender setae (one of them very small), and the last segment 
bears 2 unequal slender recurved claws (47 and 25 [i respectively 
along their axes) and 5 setae, one of them very long (99 \).). The 
extremity of the last segment is swollen, so that the 2 claws 
insert at one side rather than directly on the tip of the segment. 
All the setae are naked. 

The labrum (Fig. 10) consists of 2 diverging, pointed lobes 
with their medial edges straight and finely dentate, both arising 
from a large, conspicuous, sclerotized area which projects (Fig. 
2) from the ventral surface of the head. On the posterior wall of 
the labrum, in front of the mouth area, there is a pair of small 
sclerotized lobes. The surface of the labrum lacks fine ornamenta- 
tion. 

The mandible (Fig. 11) has on the concave side of the blade 
an oblique row of spinules followed distally by a spinelike process 
lying parallel to the blade and evidently not articulated with it. 
The convex side of the blade bears a fringe of graduated spinuli- 
form structures without definite articulations. The paragnath 
(Fig. 12), lying medial to the base of the first maxilla (as in 
Fig. 16), is a rounded lobe bearing a small sclerotized outer 
process, a small postero-inner knob, and a posterior group of 
hairs. The first maxilla (Fig. 13) is a single segment bearing 4 
naked elements, comprising terminally 2 obtuse subequal spines 
(10 and 8 [jl long) and a shorter pointed spine {5 [).) and sub- 
terminally a naked hyaline seta (11 \k). The second maxilla 
(Fig. 14) is 2-segmented. The first segment is unarmed but has a 
small sclerotized protuberance on its expanded margin. The sec- 
ond segment is produced to form a long bilaterally spinulose 
lash ; the dorsal surface bears a proximal seta 29 \j. in length 
(bearing lateral spinules along one edge) and a distal row of 3-5 
spinules. The maxilliped (Fig. 15) is 3-segmented. There are 2 
naked setae on the second segment. The terminal segment bears 
proximally a spine with unilateral spinules, a hyaline naked 
setule, and near the base of the latter a minute setule ; the seg- 
ment is produced to form an attenuated spinelike structure with 
a row of long spinules along one side and a row of minute 
spinules near the opposite margin. 

The postoral area (Fig. 16) shows between the paragnaths a 
shield-shaped area which projects ventrally to form a low median 
process. Posterior to the sclerotization which almost joins the 



protopod 


0-1; 


1-0 


exp 


I-O; 


I-l; 


111,1,4 








end 


0-1; 


0-1; 


1,2,3 


protopod 


0-1; 


1-0 


exp 


I-O; 


I-l; 


111,1,5 








end 


0-1; 


0-2; 


1,11,3 


protopod 


0-1; 


1-0 


exp 


I-O; 


I-l; 


111,1,5 








end 


0-1; 


0-2; 


1,11,2 


protopod 


0-1; 


1-0 


exp 
end 


I-O; 
II 


I-l; 


11,1,5 



1966 NEW COPEPOD FROM AN ECHINOID 5 

bases of the maxillipeds, the ventral surface of the cephalosome 
protrudes slightly ( best seen in a lateral view, as in Fig. 2 ) . 

Legs 1-4 (Figs. 17, 18, 19, and 20) have trimerous rami except 
for the endopod of leg 4 which is unimerous. The armature of 
the legs is as follows (the Roman numerals indicating spines, the 
Arabic numerals setae) : 

PI 
P2 
P3 
P4 



The inner seta on the coxa of legs 1-3 is long and plumose, but 
in leg 4 this seta is minute (7 [jl long) and naked. In the first 3 
legs the inner margin of the basis bears a short row of hairs, but 
in leg 4 these hairs are lacking. The outer coxal margin of leg 1 
is expanded to form a prominent lobe (Fig. 17) ; in legs 2 and 3 
this expansion is much less prominent and in leg 4 it is ap- 
parently absent. The tips of the outer spines on the exojjods are 
slightly recurved posteriorly and the more proximal ones have 
minute terminal flagella. The 3 spines on the last segment of 
the endopod of leg 2 are 14, 10, and 13 ;J^ in length from proximal 
to distal, with the middle one having a short terminal flagellum. 
In leg 4 the exopod is longer than in any of the preceding legs. 
The endopod measures 53 x 16 pi. and has nearly parallel margins 
without a constriction or notch. It bears a row of hairs on its 
outer proximal third and an anterior row of small spinules near 
the insertions of the 2 divergent terminal spines (the outer 18 [jl 
long with a minute flagellum, the inner 37 [x long with a more 
strongly spinulose flange along the outer side than along the 
inner side). 

Leg 5 (Fig. 21) consists of a strong spine (33 y. long) and an 
adjacent seta (21 \i), together with a dorsal seta (22 [x), orna- 
mented as in the figure. External to the spine and seta there is a 
row of minute blunt spinules at the apex of the segment. Al- 
though, as in other species of Pseudanthessius, there is no free 
segment of leg 5, it is likely that the spine and its adjacent seta 
correspond to the terminal armature in other lichomolgids. 

Leg 6 is probably represented by the 2 setae near the attach- 
ment of each egg sac (see Fig. 4). 



6 BREVIOEA No. 246 

The color in life in transmitted light is translucid, the eye red, 
the intestine black, the ovary gray, the egg sacs opaque gray. 
(Although in specimens preserved in 70 per cent ethyl alcohol 
the color is an opaque grayish brown, the color changes quickly 
to a bright red when the copepods are placed in lactic acid.) 

Male. — The body (Figs. 22 and 23) has a much narrower 
prosome than in the female, but otherwise resembles that sex in 
general form. The length (not including the setae on the caudal 
rami) is 0.76 mm (0.73-0.78 mm) and the greatest width is 0.27 
mm (0.25-0.28 mm), based on 10 specimens (including the allo- 
type, 8 paratypes, and 1 specimen from Pte. Ambarionaomby on 
October 30). The ratio of length to width of the prosome is 
1.57:1. 

The segment of leg 5 is similar to that of the female, and meas- 
ures 31 X 94 \j.. The genital segment (Pig. 24) is nearly as long 
as wide, 104 x 110 y., and in dorsal view has a subspherical out- 
line. In lateral view (Fig. 25) the anteroventral part of the 
segment projects noticeably. There is no intersegmental sclerite 
between the segment of leg 5 and the genital segment. The 4 
postgenital segments are 39 x 52, 32 x 51, 30 x 50, and 43 x 53 [;. 
from anterior to posterior. 

The caudal ramus resembles that of the female, but is a little 
shorter, the inner length being 78 [x, the outer length 73 \)., the 
width 23 [J., and the ratio of length to width 3.26 :1. 

The surfaces of the prosome and urosome bear minute setules 
as in the female. The ratio of the length of the prosome to that 
of the urosome is 1.31 :1. 

The rostral area, first antenna, second antenna, labrum, man- 
dible, i)aragnath, and first maxilla are like those in the female. 
The second maxilla also resembles that of the female, but lacks 
the small sclerotized protuberance on the expanded margin of 
the basal segment. The maxilliped (Fig. 26) is much elongated, 
slender, and 4-segmented (assuming that the fourth segment is 
represented by the proximal part of the claw). Its entire length 
including the claw is about 300 [jl. The first segment is unarmed. 
The second bears 2 unequal inner setae and 2 rows of hairs, one 
along the inner margin distal to the setae and another starting on 
the proximal inner margin and passing obliquely to the distal 
posterior surface of the segment. The very short third segment is 
unarmed. The terminal recurved claw, 135 ;;. in length along its 
axis, bears a conspicuous terminal lamella. The slightlj^ crenated 
fringe along its concave margin is interrupted about midway. 
Near the base of the claw on its posterior surface there is a seta 



1966 NEW COPEPOD FROM AN ECHINOID 7 

42 ijL long with minute lateral spinules, and on its anterior sur- 
face there are 2 small naked setules, one 10 [i, the other 4 [i. long. 

The postoral area is like that of the female. 

Legs 1-4 resemble those of the female except that the last seg- 
ment of the endopod of leg 1 is more elongated (Fig. 27), and 
the terminal segment of the endopod of leg 2 is more elongated, 
and its 3 spines are longer (24, 22, and 25 \}. from proximal to 
distal), as seen in Figure 28, 

Leg 5 is similar to that of the female. 

Leg 6 (Fig. 29) consists of a posteroventral flap on the genital 
segment. Beyond the rim of the segment the leg projects con- 
spicuously (see Fig. 24) as a large, pointed, ventral sclerotized 
process dorsal to which there is a shorter rounded process bearing 
2 naked setae 22 and 24 jx in length. 

The spermatophore, seen only inside the body of the male 
(Fig. 24), is oval, about 72 x 45 [x, with a short neck. 

The color in life in transmitted light resembles that of the 
female. 

RELATIONSHIP TO OTHER SPECIES IN THE GENUS 

Twelve species of Pseudanthessius may be readily distin- 
guished from P. procurrens in that they lack the prominent 
outer expansion on the coxa of leg 1 and do not bear the two 
rounded sclerotized lobes on the anterior part of the genital 
segment in the female. These species are : P. aestheticus Stock, 
Humes, and Gooding, 1963, P. concinnus Thompson and A. Scott, 
1903, P. deficiens Stock, Humes, and Gooding, 1963, P. duhius 
G. 0. Sars, 1918, P. graciloides Sewell, 1949, P. luculentus Humes 
and Cressey, 1961, P. mucronatus Gurney, 1927, P. nemertophi- 
lus Gallien, 1935, P. notabilis Humes and Cressey, 1961, P. pec- 
tinifer Stock, Humes, and Gooding, 1963, P. tortuosus Stock, 
Humes, and Gooding, 1963, and a new species (Humes and Ho, 
in press) from a polychaete annelid in Madagascar. 

Eight other species may be separated from P. procurrens in 
that they lack the two lobes on the anterior part of the genital 
segment in the female and have a different armature on the last 
segment of the second antenna. (Unfortunately, in the original 
descriptions of these species, definite information on the con- 
dition of the outer coxal margin of leg 1 was not given.) These 
species are : P. gracilis Claus, 1889, P. latus Illg, 1950, P. oh- 
scurus A. Scott, 1909, P. saiivagei Canu, 1892, P. spinifer Lind- 
berg 1945, P. tenuis Nicholls, 1944, P. thorelli (Brady, 1880), 
and P. weheri A. Scott, 1909. 



8 BREVIORA No. 246 

Three species remain to be compared with P. procurrens : P. 
liber sensu Sewell, 1949, P. liber (Brady, 1880), and P. assimilis 
G. 0. Sars, 1917. The species referred to as P. liber (Brady and 
Robertson) by Sewell (1949) is probably a new and unnamed 
species of the genus (see Humes and Cressey, 1961, pp. 80-81). 
It differs from P. procurrens chiefly in the presence of two long- 
elements (aesthetes ?) on the basal segment of the first antenna, 
in the inwardly curving terminal spine on the last segment of 
the exopod of legs 2-4, and in the notch on the outer margin of 
the endopod of leg 4. No information was given for this form 
regarding the nature of the genital segment in the female, and 
the condition of the outer coxal margin of leg 1 is not clearly 
shown in Sewell 's text figure 32 D. 

P. liber (Brady, 1880)- lacks the two lobes on the anterior 
l)art of the genital segment of the female, has a caudal ramus 
about twice as long as wide, and has a different armature on the 
last segment of the second antenna. Brady did not mention an 
outer coxal expansion on leg 1, but Sars (1917) both mentioned 
and figured such an expansion in specimens taken in Norway. 
The expansion illustrated on Sars' plate XCIV is, however, less 
pronounced than in P. procurrens. The armature for the last 
segment of the endopod of leg 3 in the female is, according to 
Sars, 1,11,3, instead of 1,11,2 as in the Madagascan species. 

P. assimilis G. 0. Sars, 1917, is said by Sars to be closely allied 
to P. liber (Brady, 1880). Like the latter species it lacks the two 
lobes on the genital segment of the female and has an armature 
on the last segment of the second antenna unlike that of P. pro- 
currens. Sars stated that the legs are "almost exactly as in P. 
liber," implying that there is a similar outer coxal expansion 
on leg 1. 

Judging from the available information, P. procurrens appears 
to be near both P. liber (Brady, 1880) and P. assimilis G. 0. 
Sars, 1917, resembling them in having an outer coxal expansion 
on leg 1, but differing in having two lobes on the anterior part 
of the genital segment in the female and in having a different 
armature on the last segment of the second antenna. 

SUMMARY 

The new species Pscuelanthcssius procurrens is associated with 
the cidarid echinoid Phyllacanthus imperialis (Lamarck) in the 



2 Not IJi-ady and Uobertsou, 1S70. See StmU. Hiinies. and Goodius, I'.Mi:}, p. 10, 
footnote. 



1966 NEW COPEPOD P^ROM AN ECHINOID 9 

region of Nosy Be, northwestern Madagascar. Within the genus 
the eopepod may be recognized by the conspicuous expansion on 
the outer coxal area of leg 1 and by the two rounded lateral lobes 
on the anterior part of the genital segment in the female. The 
species nearest to the new form appear to be P. Z('&er (Brady, 
1880) and F. assimilis G. 0. Sars, 1917. Although the host 
echinoderm is widely distributed in the Indo-Pacific area, the 
new eopepod associated with it is known at present only from the 
type locality in Madagascar. 

REPEEENCES CITED 

Brady, G. S. 

1880. A monograph of the free and semi-parasitic Copepoda of the 
British Islands. Ray Society, London, 3:1-83. 
Brady, G. S. and D. Robertson 

1876. Report on dredging off the coast of Durham and North York- 
shire in 1874. Rept. British Ass. Adv. Sei. (Bristol), 45: 185- 
199. 
Canu, E. 

189:2. Les Copepodes du Boulouuais, morphologic, embryologie, taxono- 
mie. Trav. Lab. Zool. mar. Wimereux-Ambleteuse (Pas-de- 
Calais), 6:1-354. 
ChAL-s, C. 

1889. Uber neue oder wenig bekaniite halbparasitische Copepoden, 
insliesondere der Liehomolgiden- und Aseomyzontiden-Gruppe. 
Arl). Zool. Inst. Univ. Wien, 8(3) :327-370. 
Gali-iex, L. 

1935. Ps(ii(lant]ic.stiiu.s nenurtopJiilus nov. sp., copepode commensal de 
JAiieiis lo7igi.ssiinufi Sowerby. Bull. Soc. Zool. France, 60:451-459. 
GlR.NEY, R. 

1927. Zoological results of the Cambridge expedition to the Suez Canal, 
1924, XXXIII. Report on the Crustacea: — Copepoda (littoral 
and semi-parasitic). Trans. Zool. Soc. London, 22(4) :451-477. 
Humes, A. G. and R. F. Cressey 

1961. Deux nouvelles especes de PseudantJicssius (Copepoda, Cj'clo- 
poida) parasites des oursins a Madagascar. 2ilem. Inst. Sci. 
Madagascar, ser. F, 1959, 3:67-82. 
Humes, A. G. and J.-S. Ho 
In pre s. Xew cyclopoid copepods associated witli polyt-liacte annelids in 
Madagascar. Bull. Mus. Comp. Zool. 
Il.LG, P. 

1950. A new eopepod, Pseitdanthessiiis latus (Cyclopoida :Lichoniolgi- 
dae) commensal with a marine flatworm. Jour. Washington 
Acad. Sci., 40(4) : 129-133. 



10 BEEVIORA No. 246 

LiNDBERG, K. 

1945. Un iiouveau copepode poecilostome cle I'Inde de la famille des 
Lichomolgidae ; Pseudanthessms spinifer, n. sp. Bull. Soc. Zool. 
France, 70:81-84. 
NiCHOLLS, A. G. 

1944. Littoral copepods from South Australia (II). Calanoida, Cyclo- 
poida, Notodelphyoida, Monstrilloida and Caligoida. Eec. So. 
Austr. Mus., 8(l):l-62. 
Sars, G. O. 

1917. An account of the Crustacea of Norway with short descriptions 
and figures of all the species. Vol. 6, Copepoda, Cyclopoida, pts. 
11 and 12, Clausidiidae, Lichomolgidae (part), pp. 141-172. 
Bergen Museum, Bergen. 

1918. An account of the Crustacea of Norway with short descriptions 
and figures of all the species. Vol. 6, Copepoda, Cyclopoida, pts. 
13 and 14, Lichomolgidae (concluded), Oncaeidae, Corycaeidae, 
Ergasilidae, Clausiidae, Eunicicolidae, Supplement, pp. 173-225. 
Bergen Museum, Bergen. 

Scott, A. 

1909. The Copepoda of the Siboga Expedition. Part I. Free-swim- 
ming, littoral and semi-parasitic Copepoda. Siboga Exped., 
29a: 1-323. 
Sewell, R. B. S. 

1949. The littoral and semi-parasitic Cyclopoida, the Monstrilloida and 
Notodelphyoida. John Murray Exped. 1933-34, Sci. Eepts., 
9(2):17-199. 
Stock, J. H., A. G. Humes, and R. U. Gooding 

1963. Copepoda associated with West Indian invertebrates IV. The 
genera Octopicola, Pseudanthessius and Meomicola (Cyclopoida, 
Lichomolgidae). Studies Fauna Curasao and other Caribbean 
Is., 18(77) :l-74. 
Thompson, I. C. and A. Scott 

1903. Report on the Copepoda collected by Professor Herdman at Cey- 
lon in 1902. Rept. Govt. Ceylon Pearl Oyster Fish. Gulf of 
Manaar, pt. I, Suppl. Rept. No. 7:227-307. 

(Received December 10, 1965.) 



EXPLANATION OF THE FIGURES 

All the figures have been drawn with the aid of a camera lucida. The let- 
ter after the explanation of eacli figure refers to the scale at which the figure 
was drawn. 

Ablneviations: ai := first antenna, md = mandible, p = paragnath, mxi = 
first maxilla, mx2 = second maxilla, mxpd = maxilliped, pi = leg 1. 



1966 



NEW COPEl'OD FKOM AN ECHINOID 



11 




Figures 1-7. Pseudanthessius procurrens n.sp., female. 1, dorsal (A) ; 2, 
lateral (A) ; 3, uiosome, dorsal (B) ; 4, area of attaclmient of egg sac, dor- 
sal (C) ; .J, caudal ramus, ventral (D) ; 6, edge of cephalosome, ventral 
(E) ; 7, rostral area, ventral (F). 



12 



BREVIORA 



No. 246 




Figures 8-16. Pseudanthessius procurrens ii.sp., female (continued). 8, 
first antenna, anterodorsal (F) ; 9, second antenna, inner (F) ; 10, labrum, 
ventral (D) ; 11, mandible, inner (D) ; 12, paragnath, ventral (D) ; 13, first 
maxilla, outer (G) ; 14, second maxilla, dorsal (D); 15, maxilliped, inner 
(D) ; 16, postoral area, ventral (H). 



1966 



NEW COPEPOD FROM AN ECHINOID 



13 





Figures 17-21. Pseudanthes-sius procurrens n.sp., female (continued). 17, 
leg 1, anterior (F) ; 18, leg 2, anterior (F) ; 19, leg 3, posterior (F) ; 20, 
leg 4, anterior (F) ; 21, leg 5, dorsal (D). 



14 



BREVIORA 



No. 246 




Figures 22-29. Pseudanthessius procurrcns n.sp., male. 22, dorsal (A); 
23, lateral (A) ; 24, urosome, dorsal (B) ; 25, genital segment, lateral (H) ; 
26, maxilliped, anterior (F) ; 27, endopod of leg 1, anterior (F) ; 28, endo- 
pod of leg 2, anterior (F) ; 29, leg 6, ventral (D). 



BREVIORA 

Museiuiiiiti of Compairative Zoology 

Cambridge, Mass. May 3, 1966 Number 247 

THE CHAXAKE8 (ARGENTINA) TKIASSIC REPTILE 

FAUNA 

I. INTRODUCTION 
By Alfred Sherwood Romer 

THE 1964-65 LA PLATA-HARVARD EXPEDITION 

This is the first of a proposed series of papers giving the scien- 
tific results of a fossil-collecting expedition to western Argentina 
made during the summer of 1964-65 under the joint auspices of 
the Universidad de La Plata and Harvard University. The proj- 
ect grew out of the realization, during an earlier expedition 
of this sort, that further work in this area might result in im- 
portant additions to the early tetrapod fauna of South America. 

In 1958 a group of Harvard paleontologists under the direction 
of Prof. Bryan Patterson and myself, in cooperation with the 
Museo Argentina de Ciencias Naturales "Bernardino Riva- 
davia" de Buenos Aires, spent some months in exploring for 
vertebrate fossils in western Argentina. In part. Tertiary mam- 
mals were collected, and in part, fishes and brachyopid labyrin- 
thodonts from the Triassic Cacheuta series of Mendoza Province 
(cf. Rusconi, 1951, etc., and Romer, 1960). Most especially, how- 
ever, we explored and developed the Triassic beds of the Ischi- 
gualasto region in northeastern San Juan Province (cf. Romer, 
1962) ; these proved exceedingly productive. Work on them has 
been continued by the Instituto Lillo of Tucuman, at first under 
Dr. Osvaldo Reig, currently under Sr. Jose F. Bonaparte. 

The Harvard members of the 1958 party included, in addition 
to Prof. Patterson and myself, Arnold D. Lewis, chief preparator 
of the Museum of Comparative Zoology, James A. Jensen, then 
of the Harvard staff' but now in charge of geological and paleon- 
tological collections at Brigham Young University (Provo, 
Utah), and Mrs. Ruth H. Romer. Prof. Patterson's interests have 
now shifted to another field. The others of the group, however, 



2 BREVIOBA No. 247 

became deeply interested in the problems of the early history of 
tetrapod life in South America, and felt that, if the opportunity 
should be presented, a second trip of exploration in western 
Argentina might quite possibly result in a major addition to 
this story. 

Such an opportunity arose in 1964 through a grant from the 
National Science Foundation, from which could be met most of 
the costs of a field trip of the sort proposed. Accordingly, an 
agreement to this end was drawn up between the Harvard Mu- 
seum of Comparative Zoology and the Museo de Universidad 
Naeional de La Plata. Under Argentinian law 9080 of 1911, all 
fossils are declared the property of the national government, un- 
der the Ministry of Education. A commission of three directors 
of leading museums is empowered by the Ministry to regulate 
the collection and disposition of fossil materials. Our agreement 
with the Museo de La Plata was approved by this commission. 
As provided by law, type specimens and other representative 
materials, or casts thereof, are to be deposited in the Museo de 
La Plata; other specimens remain the property of the Harvard 
Museum. Further, we are to be allowed to have for two years 
exclusive right to the study and description of specimens from 
any new fossil deposits discovered during the course of the work. 

Our objective was to find fossil localities for reptiles and am- 
phibians in horizons in the Permian or in Triassic beds earlier 
than that of Ischigualasto. Although the position of the Ischi- 
gualasto fauna is somewhat debatable, it is essentially middle 
Triassic in character, predominantly one of rhynchosaurs and 
cynodonts of gomphodont type ; there are known a few advanced 
thecodonts, but only the bare beginnings of the dinosaur fauna 
which dominates the scene in the late Triassic. Faunas com- 
parable to that of Ischigualasto are known in Brasil (Santa 
Maria) and in Africa (Manda beds) ; but while antecedent Per- 
mian and Lower Triassic faunas are abundant in Africa, almost 
nothing was known of earlier tetrapods in South America. On 
that continent no tetrapod has ever been reported from the 
Permian of Argentina, and elsewhere nothing is known except 
for the aberrant aquatic Mesosaurns of southern Brasil (and 
South Africa) and a single labyrinthodont from the Brasilian 
Permian (Price, 1948). The earlier Triassic was, before our re- 
cent exploration, almost equally unknown as regards reiitilian 
faunas. It is possible that, despite general beliefs as to its late 
Triassic position, the Cacheuta series may be of early Triassic 
age (cf. Reig, 1961), but the evidence is far from conclusive, and 



1966 CHANARES REPTILES 3 

little reptilian material is known from this series. Recently, two 
forms, a dicynodont and a gomphodont, were discovered in south- 
ern Mendoza Province and are now being studied by Sr. Bona- 
parte; these are probably early Triassic in age. 

This short list is a complete roster of all tetrapod vertebrates 
known from any part of South America in deposits older than 
the relatively late Triassic of the Ischigualasto Formation and 
the essentially comparable Santa Maria beds of Brasil. A great 
deal of stratigraphic work has been done in western Argentina 
by the geologists of the Instituto Nacioual de Geologia y Mineria, 
the Yacimientos Petroliferos Fiscales and the Comision Nacional 
de Energia Atomica, and some thousands of square miles of con- 
tinental deposits have been mapped as presumably Permian or 
Triassic in age. But apart from a few finds of fishes, not a single 
vertebrate fossil was discovered in any of these seemingly barren 
beds. It was thus not impossible that our trip might have been 
a complete failure, and that the entire four months planned for 
field work might have ended without our finding a single speci- 
men. 

Geologists, however, intent on stratigraphic work, are no- 
toriously liable to overlook fossil materials for which the eye of 
the paleontologist is trained. Our obvious plan of campaign was 
to visit, one after another, "sample" areas in which Permian or 
Triassic rocks had been mapped, and to hope that, ultimately, one 
or more of these widespread regions might yield a worthwhile 
fauna. We found the geologists of the various Argentinian serv- 
ices mentioned exceedingly generous and helpful in giving us 
their advice and free access to unpublished maps, manuscripts, 
and field notes to aid in our laying out a plan of campaign and 
in discussing matters of stratigraphy and faunal succession. We 
wish to thank these friends for their help. They include, amongst 
others : Director General Felix Gonzalez Bonorino, Dr. Roberto 
V. Tezon, Vocal del Directorio del Instituto, Dr. Juan C. Turner, 
Dr. Eduardo Holmberg, Dr. Francisco Fidalgo, Dr. Guillermo 
Furque, Dr. Emilio Gonzalez Diaz, and Sr. Carlos Turcogreco, of 
the staff of the Instituto Nacional de Geologia y Mineria ; Dr. 
Eduardo Rolleri, Dr. Miguel Flores, Dr. Alberto Mingramm, Dr. 
Pedro Criado Roque, Dr. Hector F. de la Mota, of the Yacimien- 
tos Petroliferos Fiscales ; Profs. Sergio Archangelsky, Angel V. 
Borrello, and Alfredo Cuerda, of the Universidad de La Plata; 
Prof. Arturo Amos of the Universidad de Buenos Aires; Dr. 
Pedro N. Stipanicic, director of geological work for the Comision 



4 BREVIORA No. 247 

Nacional de Energia Atomica ; and Sr. Jose P. Bonaparte of the 
Instituto Miguel Lillo de Tucuman. 

The major part of the funds which made the expedition pos- 
sible was furnished by grant GB-2454 of the National Science 
Foundation. The cost of the motor vehicles and a fraction of the 
travel expense were supplied from private sources. We wish to 
thank Brigham Young University for allowing Mr. Jensen to ac- 
company us, and the Museum of Comparative Zoology for mak- 
ing it possible for Mr. Lewis to take part. Publication of this 
paper was aided by NSP grant GB-4615. 

Our collaboration with the Museo de La Plata proved exceed- 
ingly advantageous. Professor Rosendo Pascual aided us in 
every way before, during, and after our months in the field, and 
for a large fraction of the time was an active member of the field 
party. We are very grateful to Dr. and Senora Pascual for their 
hospitality in La Plata before and at the close of the field season. 
Dr. Mario Teruggi, Dean of Natural Sciences and Director of the 
La Plata Museum, aided greatly in solving our difficulties with 
the provincial government. Very welcome was the hospitality 
given us by the University at Samay Huasi, "the house of re- 
pose." This had been the country estate of Dr. Joaquin B. Gon- 
zalez, famous Argentinian author, statesman, and educator, and 
is located near the city of Chilecito in La Rioja Province. Willed 
by him to the University at his death, it is now used as a resort 
for artists, writers, and faculty members. An irrigated desert 
oasis of some 40 acres or so, with splendid groves of trees, vine- 
yards, and gardens, it contains a spacious and well-equipped 
house, with a permanent staff. Most of our work proved to lie in 
La Rioja Province ; Chilecito was the best available town to visit 
periodically for supplies, and we greatly enjoyed on these occa- 
sions the amenities afforded by Samay Huasi through Sr. and 
Sra. Amoretti and their staff*, and Prof. Bongiorno of the Univer- 
sity who has charge, in addition to other duties, of a museum at 
Samay Huasi. 

Our equipment for the trip was carefully selected by Mr. 
Lewis, on the basis of his experience on the previous expedition, 
to the end of attempting to make our work under the somewhat 
trying conditions of summer in the arid regions of western Ar- 
gentina as efficient and as comfortable as possible. For rolling 
stock we took with us a 4-wheel-drive Jeep and a i/o ton stake- 
bodied truck, likewise with 4-wheel traction, and equipped with a 
75-gallon water tank and spare gasoline tanks. Most useful, as it 
proved, was a ' ' tote-gote " — a motorcyclette of a type recently 



1966 CIIANARES REPTILES 5 

developed in the western United States for traversing rough 
country and steep grades, and able to carry a modest load of ma- 
terials in addition to the rider. Without this, our work would 
have been severely handicapped, for part of our best collecting 
areas proved to be one or two miles from the closest point we 
could reach with a jeep ; the laborious business of attempting to 
carry in on our backs many hundreds of pounds of plaster and 
hundreds of gallons of water, and return with some thousands 
of pounds of plaster blocks, would have greatly reduced our 
results. 

Our party arrived in Argentina the latter part of October, and 
set out for the field after a short stay in La Plata and Buenos 
Aires to complete our equipment and obtain all possible data 
and advice. We then began our planned series of visits to typical 
areas of sediments mapped as presumed Permian or Triassic in 
western Argentina. As the discussion above would suggest, our 
expectations of success at any given locality were none too san- 
guine, and for the first two months of the trip our gloomiest 
forebodings appeared to be realized ; during this time we failed to 
find the slightest scrap of tetrapod bone in any new area. 

The Santa Clara Formation occupies a considerable area in the 
precordillera between Mendoza and San Juan. It has sometimes 
been thought to be Triassic in age, but is believed by Dr. H. J. 
Harrington (1941, 1956) to be Permian. A few fishes (Bordas, 
1944) and a single footprint (Rusconi, 19ola, 1956, and Peabody, 
1955) had been described from the Santa Clara, but no tetrapod 
fossils. We first camped near the puesto of Santa Clara, near 
the north end of the area covered by the formation, at the east 
margin of the precordillera in southern San Juan Province, and 
explored this area. Here the Santa Clara shales and sandstones, 
some thousands of feet thick, form a series of mountains sepa- 
rated b}^ deep valleys. We found further remains of fishes in the 
area of original discovery, and a further fish locality in shales 
north of the Quebrada de la Montaiia a short distance east of 
the boundary between the Santa Clara beds and the Villavicencio 
group. In addition, a series of trackways of rather poor quality 
was found and cast ; these were situated a short distance up a 
senda leading south from the Quebrada de la Montaiia to Puesto 
Alojamiento. But as far as our sampling of these vast exposures 
weiit, they appeared to be entirely barren of tetrapod osseous 
remains. 

A second camp was made in the southern area of the Santa 
Clara Formation, in the canyon of the Rio de las Peiias, between 



6 BREVIORA No. 247 

Cerro Pefia and Cerro Palado de las Higueras. Footprints had 
been previously reported from this area. We learned later that 
Sr. Bonaparte had made a visit to this area previous to ours. 
Like him, we found the region to be exceedingly rich in reptilian 
trackways, in a fashion comparable to areas of the famous Con- 
necticut Valley Triassic. The trackways, hoM'ever, are in the 
main obviously those of dinosaurs or advanced thecodonts, and 
the beds are presumably middle or late Triassic in age, despite 
the presence of '' ChirotJurium "-like footprints. As in the north- 
ern part of the Santa Clara Formation, we failed to find any 
bones; not even fish specimens were discoverable. 

Bodenbender (1911, etc.) had described at length as "Paganzo 
II" a very thick series of red sandstones, presumably Permian 
in age, from La Rioja Province. These are present in the region 
of the Sierra de Saiiogasta, its southern continuation, the Sierra 
de Vilgo, and the country south of the mountains, between the 
village of Paganzo and the railway junction of Patquia to the 
east. We spent many weary days walking these sandstones from 
the region of Patquia northward towards Chilecito, and in the 
region of Los Colorados railway station. We found not the slight- 
est trace of animal life in these abundant exposures of ' ' Paganzo 
II"; not a bone, not even a fish scale or a footprint. Following 
this discouraging period, we took time out for a brief visit to 
Tucuman, where we were hospitably received at the Instituto 
Lillo by Director Willink, and profited l)y discussion with Sr. 
Bonaparte. 

Permo-Triassic beds had been studied in the Sierra de las Qui- 
jadas in northwestern San Luis Province by Sr. Jose Roman 
Guinazti of San Luis, and footprints from this region were de- 
scribed by Lull (1942). The group spent some time there. The 
best exposures lay in a deep amphitheatre, difficult of access, 
west of Puesto Hualtaran. From the nearest approach by car 
there was necessary a long, tortuous walk, followed by a horse- 
trail descent of several hundred meters into the base of the 
amphitheatre ; the return at the end of a day involved the weari- 
some ascent of the canyon walls. Here, as in other areas pre- 
viously explored, not a bone was found, although a number of 
footprints of middle to late Triassic type were seen in the deepest 
and lowest beds of the area. 

Permian sediments had been reported in the Sierra de los 
Llanos of southern La Rioja (Braeaccini, 1946). Here we found 
massive red conglomerates and grey plant-bearing shales. Nei- 
ther type of sediment appeared promising as a possible source of 



1966 CHANARES REPTILES 7 

tetrapod remains, and only a very short time was spent in this 
area. 

Having thus prospected for the better part of two months in 
some five areas with completely negative results, we had become 
somewhat disheartened. As some slight alleviation, we decided to 
visit for a short time our old "stamping ground" of Ischigua- 
lasto, for the purpose of collecting fresh material of rhyncho- 
saurs, which by agreement with Sr. Bonaparte, I am studying. 
We had, in our discouragement, half come to believe that perhaps 
we had lost the ability to see fossil bones even if present. Ischi- 
gualasto disabused us of that apprehension ; a good selection of 
rhynchosaur specimens was obtained. 

The Christmas season had now arrived, and we stopped work 
to spend a few days at Samay Huasi. But despite the beauty and 
amenities of that place, the holidays were far from joyful for us. 
Half of our field time had been consumed, and we had thus far 
met only with utter failure. 

But a complete reversal of fortune now occurred. Among the 
places remaining for possible investigation, one which appeared 
particularly attractive was an area in western La Rioja Province, 
west and south of Chilecito, west of the Sierra de Sanogasta, and 
about three hours distant from Chilecito by car. Here, at the east 
margin of the barren Campo de Talampaya, along the rivers Ta- 
lampaya, Chaiiares, and Gualo, geological reconnaissance by Bo- 
denbender (1911, etc.) and, in recent years, by de la Mota and 
Fidalgo (unpublished) had shown the presence of a varied series 
of sediments of Permian and Triassic age. No fossils had been 
seen, but in view of the considerable range in time occupied, 
and the probability of a variety of sediments in the succession of 
formations which appeared to be present, one might not unrea- 
sonably hope that vertebrates might somewhere be found. 

This hope was abundantly justified. We first pitched camp in 
the southern part of the area, near the elevation known as the 
Mogote del Gualo and near the only spring of any importance in 
this arid region. On our first day of prospecting, fragmentary ver- 
tebrate remains were discovered ; the next day several good speci- 
mens of dicynodonts came to light ; from that day on until we 
stopped work some seven weeks later, not a day passed without 
the discovery of worthwhile reptilian material. From our first 
camp we presently moved some miles to the northwest to camp 
near the point where the Rio Gualo emerges from the hills into 
the flat Campo de Talampaya. Finding that still richer fossil de- 
posits were present farther to the north, in the valley of the Rio 



8 BHKVIOKA No. 247 

Chanares, we camped for the remainder of oin- stay on the banks 
of that stream. 

In the following article in this series there will be given a 
resume of the geology of the area. The region is onlj^ a short 
distance from the Ischigualasto Valley on the opposite, western, 
side of the Campo de Talampaya, and several of the later Trias- 
sic formations in the Chaiiares-Gualo region could be readily 
identified with the Los Colorados, Ischigualasto, and Los Rastros 
formations with which we had become familiar on our earlier 
trip. No fossils were found by us in the limited exposures of the 
first two ; in the carbon-bearing Los Rastros a small amount of 
fish material was seen, and more could probably have been col- 
lected had we not found other, more profitable work. In rather 
lower beds, perhaps of Permian age, which we are terming the 
Tarjados Formation, fragmentary reptile remains were found in 
two areas, and not improbably intensive work Avould result in 
further finds. Nearly all our finds, however, were derived from 
a series of volcanic tuffs underlying the Los Rastros beds, 
which we are terming the Chanares Formation. These in- 
clude some 75 meters of distinctive blue-white beds, easily trace- 
able across the region despite the presence of numerous faults. 
Almost all the fossil remains, we presently discovered, occurred 
in the lowest 10 meters or so of these beds, and in most areas a 
careful following of the base of the Chanares resulted in the dis- 
covery of fossil specimens at frequent intervals. Some of the 
specimens were found simply buried in the tuffs. In a large pro- 
portion of cases, however, they were enclosed in grey-brown 
colored nodules of coarser tuffs. A fraction of such nodules 
proved to be barren ; in some areas, however, almost every nodule 
would contain a skull or skeleton, and in some cases, two or three 
individuals would be represented in a single nodule. Never had 
any of us encountered a fossil-collecting area of equal richness. 
To add to our pleasure, it was obvious in the field that prepara- 
tion of the specimens (unlike much of the material collected at 
Ischigualasto on our earlier expedition) could be readily ac- 
complished. 

Apart from a large dicynodont, almost all the specimens were 
forms of small size ; the skulls averaged but a few inches in 
length. More than 150 good skulls were collected and, in addi- 
tion, large amounts of postcranial materials. At the time this is 
written, little of the material has been prepared, but apart from 



1966 CIIANARES REPTILES 9 

a relatively small fraction of " problematical' the general sys- 
tematic position of most of the specimens was evident when col- 
lected. There appears to be but a single type of dicynodont, ap- 
parently of the general Kanncmeyeria type. A considerable frac- 
tion of the finds were those of small thecodonts, some, at least, 
aj)i)earing to be forms on the pattern of Euparkcria of the Cynoy- 
nathus zone of South Africa (recently redescribed by Ewer, 
396-')). The greater part of the collection, however, consists of 
small cynodonts; part of them are carnivorous, but the greater 
number are "gomphodonts, " in which the cheek teeth form a 
grinding battery. 

THE TRTASSIC FAUNAE SEQUENCE 

The position of the Chaiiares fauna in the Triassic sequence 
will, I trust, become apparent as its components are described in 
future numbers of this series. Meantime, a brief discussion of our 
current knowledge of the sequence of Triassic continental verte- 
brate faunas seems merited. The Triassic is commonly divided 
into Lower, Middle, and Upper portions stratigraphically — or in 
chronologic terms, early, middle, and late — in agreement with the 
three-fold division of the European Triassic (as indicated by the 
period name) into Bunter, Muschelkalk. and Keuper. On the 
basis of marine faunas, a series of stages — Scythian, Anisian, 
Ladinian, Carnian, Norian, and (as a transition to the Jurassic) 
Rhaetic — have been erected. It is frequently attempted to fit 
the vertebrate faunas into this sequence of stages. For the most 
l)art, however, this cannot be done with any degree of accuracy, 
because of the rarity of actual continuity of beds containing con- 
tinental vertebrate faunas with marine strata which can be as- 
signed a definite place in the stage sequence ; most of the stage 
names are derived from Alpine beds far removed from continen- 
tal areas. The only major exceptions have to do with the 
"Eotriassic" Scythian zone. As both Watson (1942) and Lehman 
and others (1959) have shown, the Cynognathus and Lystrosau- 
)-}(s zones of South Africa are definitely to be regarded as 
"Eotriassic'' and Scythian in age' mainly on the basis of laby- 
rinthodonts ; the Moenkopi of the early North American Triassic 
and tlie Bunter of Europe are equally well assignable to the 



1 Huene (1956) has at times tabulated the Ci/nognathiis zone as middle Triassic. 
hut has cited no reason for this. 



10 BREVIORA No. 247 

Scythian on the basis of their content of labyrinthodonts — laby- 
rinthodonts of types not too far removed from those of the 
"Botriassic" of Greenland and Madagascar. At present it is 
laerhaps best to avoid commitment of specific faunas to specific 
marine stages in attempting to define a continental vertebrate 
sequence. 

In the early years of this century our knowledge of a Triassic 
vertebrate sequence was very limited. For the early Triassic, we 
knew little of continental forms (apart from amphibians) ex- 
cept in the Upper Beaufort {Lystrosanrus and Cynognathus 
zones) of South Africa. Here, therapsids, most notably a variety 
of cynodonts, constituted the bulk of the fauna ; archosaurs, in 
the form of primitive or aberrant thecodonts, were present in 
modest numbers ; other reptiles were small in numbers and va- 
riety. At the other end of the Triassic sequence, a late reptilian 
assemblage of very different nature, consisting almost entirely of 
dinosaurs and advanced thecodonts, was known from a variety of 
redbeds formations, such as the Keuper of Europe, the Newark, 
Chinle, etc. of North America, and the redbeds of the Upper 
Stormberg of South Africa. Between, continental faunas of pos- 
sible middle Triassic age were practically unknown, for the Euro- 
pean Mnschelkalk is marine, other middle Triassic beds of the 
northern continents are also marine, and the Molteno of South 
Africa had yielded no fossils except plants. 

With the predominance of therapsids in the early Triassic, 
and of thecodonts and dinosaurs toward the end of the period, it 
was generally assumed that middle Triassic reptile faunas, if and 
when discovered, would exhibit a simple transition between these 
two extremes, with therapsids on the wane and with thecodonts 
advancing and showing the beginning of the dinosaur radiation. 
We are now in process, however, of gaining from Africa and 
South America a seemingly representative series of middle Trias- 
sic faunas — from the Santa Maria beds of Brasil, the Manda 
beds of East Africa and, most recently, the Ischigualasto beds of 
Argentina. To some degree these faunas show expected inter- 
mediate characters, for they contain advanced thecodonts and the 
beginnings of the dinosaurs. But the faunas show striking posi- 
tive characters. Therapsids are rather restricted in variety, but 
show a surprising abundance of "gomphodont" cynodonts. 
Equally surprising and striking in these faunas is the overwhelm- 
ing abundance of rhynchosaurs, almost unknown in earlier or 
later beds. 



1966 CHANARES REPTILES 11 

While it is premature, I think, to tie them down to specific 
marine stages, or to characterize them as "early," "middle" or 
"late" in any restrictive way, I believe it is clear that there are 
three successive stages in the faunal history of continental Trias- 
sic vertebrates : 

(A) An early assemblage, of which the faunas of the Lystro- 
snurus and Cynognathiis zones of South Africa, and the Wuh- 
siang beds of Shansi (China) are typical. There are abundant 
therapsids, principally cynoclonts — mainly carnivorous, but 
with a few "gomphodonts" with molar-like cheek teeth. Theco- 
donts include generalized primitive forms such as Euparkeria 
and aberrant archaic types such as Erythrosuchus and Chasmato- 
saurus. Primitive rhynchosaurs are present but rare. 

(B) The type of assemblage found in the Maleri, Santa Maria, 
and Ischigualasto beds. Carnivorous cynodonts persist, but are 
greatly overshadowed by abundant gomphodonts. Typical rhyn- 
chosaurs constitute a large fraction of the fauna. Thecodonts of 
advanced type are present, and a few early dinosaurs. 

(C) The typical assemblages of the late Triassic, such as the 
Upper Keuper, South African redbeds, Lufeng series of Yunnan, 
Newark, Chinle, and Dockum of North America, and Los Colo- 
rados of Argentina. Therapsids are close to extinction. There 
are various advanced thecodonts, such as aetosaurids and (in the 
northern continents) phytosaurs, and numerous and varied dino- 
saurs, but rhynchosaurs are generally extinct. 

One would, of course, expect assemblages transitional between 
these stages. It is probable, for example, that the Chanares fauna 
will prove to be intermediate l)etween (A) and (B), and the same 
may be true of the Ntawere fauna of Zambia (Brink, 1963 ; Kitch- 
ing, 1963). Of the faunas which we have listed under (B), that 
of Ischigualasto is perha])s relatively late, since several forms 
have already crossed the thecodont-saurischian boundary. 

The typical (C) faunas are characterized in general by an 
abundance and variety of dinosaurs, the presence, in the North- 
ern Hemisphere, of metoposaurs and phytosaurs, and the absence 
of rhynchosaurs and practical absence of therapsids. Certain 
beds usually placed in this category appear, however, either to be 
of earlier age than that of the typical (C) faunas, or else indicate 
regional late survival of certain of the forms characteristic of the 
(B) faunas. For example, the Maleri beds of India (Jain, Robin- 
son, and Chowdhury, 1964) are generally assumed to be of rela- 
tively late Triassic age because of the presence of metoposaurs 
and phytosaurs. However, there is, in the restricted fauna 



12 BREVIORA No. 247 

known, only one possible dinosaur, and rhyncliosaurs persist. We 
may have here a (B)-(C) intermediate. 

Baird and Take (1959) have recently discovered a reptilian 
fauna in the AVolfville sandstone of Nova Scotia, generally re- 
garded as a basal member of the Upper Triassic Newark group. 
But quite in contrast with typical Newark faunas, Dr. Baird in- 
forms me that not only rhynchosaur remains, but a gomphodont, 
as well, are present. Quite surely we are here dealing with a 
fauna transitional between (B) and (C). 

Rhynchosaurs are present in the Triassic fauna of the region 
of Elgin, Scotland, currently being restudied by Walker. Be- 
cause of the close resemblance between Stagonolepis of Elgin and 
Aetosaurus of the German Keuper, Walker (1961) believes the 
Elgin fauna to be late Triassic in age. There is, however, no rea- 
son to believe that the aetosaurid type of thecodont was particu- 
larly short-lived ; it was already present in the Ischigualasto 
fauna (Casamiquela, 1960). Possibly the Elgin fauna is tran- 
sitional between (B) and (C) ; but the absence of dinosaur re- 
mains, except for Ornithosiichns (Walker, 1964) — a form so 
primitive that it has generally been regarded as a thecodont — 
strongly raises the question as to whether the Elgin fauna may 
not really belong in the (B) category and be of middle Triassic 
age. Rhynchosaurs are known from several English localities 
which are classified as ' ' Keuper. ' ' But it will be noted that there 
is no Englisl] equivalent of the Muschelkalk of the continent ; the 
early Triassic "Bunter" equivalents are immediately followed 
by redheds of "Keuper" type, and it is quite possible that part 
or all of the English rhynchosaurs are of middle Triassic age. 

In sum, it appears to be a reasonable working hypothesis that 
the Triassic included a sequence of three successive continental 
vertebrate faunas characterized by a dominance of (A) therap- 
sids, (B) gomphodonts and rhynchosaurs, and (C) dinosaurs. 
We cannot, of course, expect completely clean-cut distinctions 
between the three ; but a clear pattern seems to be emerging. 

LITERATURE CITED 

Baird, D., and W. F. Take 

1959. Triassic reptiles from Nova Scotia. Bull. Geol. Soc. Amer., 
70: 15(55-1566. 
Bodenbi;nder, G. 

1911. Coiistitucion geologica de la parte meridional de La Eioja y 
regiones limitrofes (Repiiblica Argentina). Bol. Acad. Nac. 
Cienc. Cordoba, 19: 1-1220. 



1966 CHANARES REPTILES 13 

BORDAS, A. F. 

1944. Feces triasicos de la Quebrada de Santa Clara (Mendoza j San 
Juan). Physis, 19: 453-460. 
Bracaccini, O. 

1946. Los estratos de Paganzo y sus niveles plantiferos en la Sierra de 
Los Llanos (Provincia de La Eioja). Eev. Soc. Geol. Argent., 
1: 19-61. 
Brink, A. S. 

1963. Two cynodonts from the Ntawere Formation in the Luangwa 
Valley of Northern Rhodesia. Palaeont. Afr., 8: 77-96. 

Casamiquela, E. M. 

1960. Noticia preliminar sobre dos nuevos estagonolepoideos argen- 
tinos. Ameghiniana, 2 : 3-9. 
Ewer, E. F. 

1965. The anatomy of the thecodont reptile Euparkeria cai)ensis 
Broom. Phil. Trans. Eoy. Soc. London, B 248: 379-435. 
FLXRRINGTON, H. J. 

1941. Investigaciones geologicas en las Sierras de Villavieencio y Mai 
Pais, Provincia de Mendoza. Bol. Direccion Minas y Geologia, 
Rep. Argent., 49: 1-31. 

1956. Argentina. In: Handbook of South American Geology. Mem. 
Geol. Soc. Amer., 65: 131-186. 

HUEXE, F. 

1956. PaUiontologie und Phylogenie der niederen Tetrapoden. Jena, 
716 pp. 
Jain, S. L., P. L. Eobinson, and T. K. E. Chowdhury 

1964. A new verteljrate fauna from the Triassic of the Deccan, India. 
Quart. Jour. Geol. Soc. London, 120: 11.5-124. 

KiTCHING, J. W. 

1963. The fossil localities and mammal-like reptiles of the Upper 
Luangwa Valley, Northern Ehodesia. South Afr. Jour. Sci., 
59: 259-264. 
Lehman, J. -P., C. Chateau, M. Laurain, and M. Nauche 

1959. Paleontologie de Madagascar. XXVIII. Les poissons de la 
Sakamcna moyenne. Ann. Paleont., 45: 177-219. 
Lull, E. S. 

1942. Triassic footprints from Argentina. Amer. Jour. Sci., 240: 421- 
425. 

Peabody, F. E. 

1955. Occurrence of Chirotherium in South America. Bull. Geol. Soc. 
Amer., 86: 289-240. 
Price, L. I. 

1948. Um anfibio labirintodonte da forma?ao Pedra de Fogo, Estado 
do Maranhao. Ministerio da Agricultura. Bol. Serv. Geol. Min. 
Brasil, 124: 1-32. 



14 BREVIORA No. 247 

Eeig, O. a. 

1961. Acerca de la posicion sistematica de la familia Eauisueliidae y 
del genero Saurosudhus (Eeptilia, Thecodontia). Publ. Mus. 
Ciene. Nat. Mar del Plata, 1 : 73-114. 

KOMEK., A. S. 

1960. Vertebrate-bearing continental Triassic strata in Mendoza re- 
gion, Argentina. Bull. Geol. Soc. Amer., 71 : 1279-1294. 

1962. The fossiliferous Triassic deposits of Ischigualasto, Argentina. 
Breviora, Mus. Comp. Zool., No. 156, 1-7. 

EuscoNi, C. 

1951a. Laberintodontes triasicos y perniicos de Mendoza. Eev. Mus. 

Hist. Nat. Mendoza, 5: 33-158. 
1951b. Eastros de patas de reptiles permicos de Mendoza. Eev. Soc. 

Hist. Geog. Cuyo, 3 : 1-14. 
1956. Mares y organismos extinguidos de Mendoza. Eev. Mus. Hist. 
Nat. Mendoza, 9: 2-88. 
Walker, A. D. 

1961. Triassic reptiles from the Elgin area; Stagonolepis, Basygna- 
thus and their allies. Phil. Trans. Eoy. Soc. London, B 244 : 103- 
204. 

1964. Triassic reptiles from the Elgin area: Ornithosuchus and the 
origin of carnosaurs. Phil. Trans. Eoy. Soc. London, B 248 : 53- 
134. 
Watson, D. M. S. 

1942. On Permian and Triassic tetrapods. Geol. Mag., 79: 81-116. 

(Eeceived January 6, 1966.) 



BREVIORA 

Museum of Comparative Zoology 

Cambridge, Mass. 29 July, 1966 Number 248 



A TRIASSIC AMMONITE FROM THE HINDUBAGH 
REGION, BALUCHISTAN, WE8T PAKISTAN 

By Bernhard Kummel 

Ammonites are extremely rare in the thick development of 
Triassic strata between Quetta and Hindnbag-h in West Pakistan. 
Only five specimens, mostly very poorly preserved, have been re- 
ported to date. Because of their rarity the documentation of one 
additional s])ecimen of rather good preservation is M'arranted. 

The first record of Triassic ammonites from this region of 
Baluchistan was by Moj.sisovics (1896, p. 611; 1899, p. 44) who 
described a single specimen, collected by C. L. Griesbach from a 
loose block, 7 miles south of Hindubagh. This specimen was 
described as Didijmiiis afghanicus Mojsisovies (1896; 1899, pi. 
20, fig. 9) and assigned a Norian age. Yreclenburg (1904), on 
the basis of surveys carried out in the summer of 1901, con- 
tributed some r(^gional, stratigraphic, and paleontological data 
on these Triassic formations. This author reported the presence 
of Monotis cf. salinaria in fair abundance at a few horizons and 
localities and illustrated one specimen. In addition, he illustrated 
an anummite-Halorites sp. The specimen was not found in place 
but from local conditions Vredenburg concluded that it had not 
been transported very far. These specimens plus others collected 
by Vredenburg were submitted to C. Diener who published his 
results in 1906. In this small collection Diener (1906) was able 
to recognize the following forms : 

Monotis salinaria Schlotheim 
Halorites sp. ind. aff. suhcaienaio Mojsisovies 
CeJfites sp. ind. (group of '? acufepUcafi) 
Paratihefites sp. ind. ex aff. tornq)(isti Mojsisovies 
Dittmariies or Disfichites ? sp. ind. 
RhacophyJIitcs vrcdrnhurgi Diener 



2 BREVIORA No. 248 

The generally poor state of preservation of the specimens avail- 
able to Diener is well reflected in the uncertainty in the identi- 
fications. Of these specimens only Monotis salinaria and Halorites 
sp. ind. aff. suhcatenaio were obtained from the spot where they 
probably weathered out. All the rest of the specimens were 
picked up amongst transported boulders from the beds of rivers 
within the Triassic outcrop area. On the basis of this scanty 
data, both Vredenburg and Diener concluded that the strata en- 
closing these fossils are of Norian age. Additional geological 
observations on the Triassic formations of the Quetta-Hindubagh 
area have been made by Gee (in Heron, 1939, p. 26-27), by 
Williams (1959), and by the Colombo Plan Project (1961), but 
in these reports there is little additional paleontological data. 

I have had the opportunity to spend a few days in the region 
between Quetta and Hindubagh examining these Triassic forma- 
tions but was not successful in finding any fossils. Mr. S. A. 
Bilgrami entrusted to the writer a specimen found in a stream 
bed near a mine of the Pakistan Chrome Ltd., two miles east of 
Gwal Railway Station. This specimen is fairly well preserved 
and can be assigned with confidence to Arietoceltites arietitoides 
Diener, a species first described from the Tropites limestone ex- 
posed at Kalapani, Byans, in the Himalayas. This new specimen 
thus provides an additional link in the correlation of these Trias- 
sic formations of Baluchistan with the much better documented 
Himalayan sequence. The Tropites limestone of Byans has 
yielded 168 species of invertebrates of which 155 are annnonites. 
The fauna as a whole shows very strong atfinities to the Upper 
Triassic faunas of the Mediterranean region. All of the fossil 
species were collected from a three foot bed of limestone; of 
special interest is the mixture in this bed of typical Carnian 
and Norian ammonite species. Since Arietoceltites arietitoides 
was a unique species previously known only in Byans, and its 
presence in a mixed fauna at that, it does not help in refining 
the age determination of the Baluchistan Triassic formations. 
Diener 's (1912, p. 150) final words on the age of these Triassic 
rocks were that thev were Norian in age. 



1966 TRIASSIC AMMONITE 3 

SYSTEMATIC DESCRIPTION 

Family TROPICELTITIDAE Spatli, 1951 

Genus Arietoceltites Diener, 1916 

Arietoceltites arietitoides (Diener) 

Fio-ures la, b 

Tropiceltites arietitoides Diener, 1906, p. 156, pi. 3, fig. 12; Diener, 1912, 
p. 125; Diener, 1916, p. 101; Spath, 1951, p. 94; Kummel, in Arkell, 
1957, p. L171, fig. 201, 4. 

This species is represented by a single specimen of 56.5 mm in 
diameter, 13.1 mm for the height of the adoral whorl, 13.8 mm 
for the width, and 35 nun for the diameter of the umbilicus. In 
degree of involution and general proportion, this specimen is 
very much like the type specimen. There are slight differences 
in ornamentation : in the Baluchistan specimen the ribs are 
rusiradiate in the adoral 1.5 volutions, whereas in the Byans 
specimen the ribs are more radial. In both specimens the ribs 
turn strongly forward on approaching the marginal furrows 
aligning the median keel. I do not believe that these differences 
in ornamentation are of any taxonomic significance. The Byans 
material available to Diener (1906) consisted of the type speci- 
men plus a few fragments. ^Vith this one additional specimen 
there is insufficient data available to evaluate the range of vari- 
ability in these features. However, on the basis of experience 
with populations of comparable forms, one should expect a range 
of variability that would include that expressed by these two 
specimens. The suture was only partially exposed on Diener 's 
type specimen and is not preserved in the specimen recorded here. 

Occurrence. — From stream bed near mine of Pakistan Chrome 
Mines Ltd., two miles east of railway station at Gwal. presumably 
from Alozai grou]). l)etween Quetta and Ilindubagh, West Pak- 
istan. 

Repository. — Geological Survey of Pakistan, Quetta, West 
Pakistan. 

Acknoivhdgcmenis. — I wish to thank Mr. S. A. Bilgrami for 
the loan of the specimen and for his hospitality during a visit to 
Hindubagh. ^W visit to Pakistan was made possible by the aid 
of National Science Foundation Grant No. G19066. 



BREVIORA 
EEFEEENCES 



No. 248 



Arkkll, W. J. ei al. 

1957. Treatise ou invertebrate paleontology, Part L, Mollusca 4, Ceph- 
alopoda, Ammonoidea. Geol. See. Am. and Univ. Kansas Press, 
490 pp. 
Colombo Plan Project 

1961. Ret-onnaissanee geology- of jiart of West Pakistan. Toionto, 
550 pp. 
DiENEK, Carl 

190(i. Himalayan fossils. Fauna of the Tropilr.s Limestone of Byans. 

India Geol. Surv., Palaeont. Indii-a, (15) 1: 1-1201, pis. 1-17. 
1906. Notes on an Upper Triassie fauna from the Pishin District, 
Baluchistan, collected ))y E. VreiUnlmrg in the year 1901. Rec. 
(ieol. Surv. India, 34: P2-21, jils. H, 4. 
191:J. The Trias of the Himahiyas. Mem. Geol. Surv. India, 36: 1-159. 
19 Ki. Kinige Bemerkungen zur Nomenklatur der Trias-cephalopoden. 
Centralhl. Mineral, (ieol. Paliiont., No. 5: 97-l(»5. 
Heron, A. M. 

1939. General report of the geological survey of Ind'.a foi- the year 
1938. Eec. Geol. Surv. India, 74: 1-132. 
Mojsisovics, E. V. 

1896. Beitriige zur Kenntniss der oliertriadis(dien Cephalopoden- 
Faunen des Himalayas. Denkschr. Akad. Wiss. Wien, 63: 575- 
701, pis. 1-22. 
1899. Upper Triassic Ceplial()])(id;i faunae of the Himalay;i. India 
Geol. Surv., Palaeont. Indica, (15) 3: 1-157, pis. 1-22. 
Spatii, L. F. 

1951. Catalogue of the fossil Cei)halopoda in the British Museum 
(Natural History). Part 5, The Ammonoidea of the Trias. 
London, 228 i)p. 
Vredenburg, E. 

1904. On tlie occurrence of a species of IJalorilrs in tlie Trias of 
Baluchistan. Rec. Geol. Surv. India, 31: 162166, pi. 17. 
Willl\ms, M. Dean 

1959. Stratigraphy of the Lower Indus Basin, West Pakistan. Fifth 
World Petroleum Congress, See. 1, Paper 19: 1-15. 



1966 



TRIASSIC AMMONITE 





Figure 1. Side and ventral view of Avieioceltlies arietitoides (Diener) 
from Triassic formation near Hindubagh, West Pakistan. X 1- 



BREVIORA 

Mniseiimi of Comparative Zoology 

Cambridge, Mass. 29 July, 1966 Number 249 

ADDITIONAL NOTES ON THE AMPHISBAENIDS OF 
GREATER PUERTO RICO 

By Richard Thomas^ 

The overall faunal similarity of Puerto Rico and the Virgin 
Islands has led to the use of the term ' ' Greater Puerto Rico ' ' for 
their combined areas (Schmidt, 1928, used the term for a sup- 
posed continuous land mass of the past). The amphibaenids of 
this region have been dealt with thoroughly by Gans and Alex- 
ander (1962), in their comprehensive work on the Antillean 
members of the family. More recently, Gans (1964) has de- 
scribed a new species, Amphishaena schmidti, from Puerto Rico. 
Thorough as it has been, the w^ork of these authors has been 
hampered by a dependence upon specimens with insufficient or 
doubtful locality data. 

Herpetological and ornithological collections made under the 
sponsorship of Dr. Albert Schwartz in the Virgin Islands (sum- 
mer, 1964) and Puerto Rico (January -March and July 1965, and 
previous visits) have added new data to the knowledge of the 
distribution, taxonomy and ecology of the Greater Puerto Rican 
amphisbaenids with many new specimens and one new species. 

PHYSIOGRAPHIC FEATURES OF PUERTO RICO 

Before proceeding with the discussion of the species, a brief 
outline of the major physiographic features of the island of 
Puerto Rico is pertinent. Puerto Rico may be broken into three 
general physiographic regions, each of which is, in a broad sense, 
ecologically uniform. 

1. The northern limestone region extends west from roughly 
the city of San Juan to the western extremity of the island at 
Aguadilla. This region at its extreme width is one-third that of 
the island. Most of it is occupied by an extremely eroded to- 
pography of close set, conical nwgotes (to above 1500 feet), often 
called the Haystack (or Pepino) Hills; but some areas of it are 

1 10,000 SW 84th St., Miami, Florirla .''..S143. 



BREVIORA 



No. 249 




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1966 AMPHISBAENIDS OF PUERTO RICO 3 

rolling' or flat, especially along the coastal margin. The ter- 
mination of this region to the south is usually abrupt and fre- 
quently evident as an escarpment. This limestone region varies 
from xeric to mesic ; parts are moist enough for the growing of 
coffee. 

2. The central highlands, primarily igneous in lithology. com- 
prise the main mass of the island and are composed of a very 
complexly subdivided system of mountains in which few major 
units or trends are evident. Heights range to over 4000 feet in 
places, but generally the elevations are moderate (1000-2000 
feet). Except for the eastern and western coastal margins it is 
predominantly mesic and at one time was doubtless almost com- 
pletely forested. 

3. The southern coastal plain is directly continuous with the 
central highlands but is in contrast ecologically in its aridity. 
Outstanding faunistically in this region is the area to the south 
of the Valle de Lajas in the extreme southwest ; it is comprised 
in general of a discontinuous range of low coastal hills, which at 
one time probably constituted a chain of offshore keys or a penin- 
sula joining the main island in the vicinity of Guayanilla. This 
southern coastal region, particularly the portion just mentioned, 
is probably the strongest area of endemism in Puerto Rico, 
where such forms as Phyllodactylus wirshingi, Sphaerodactylus 
roosevelti, Ameiva wetmorei, Anolis poncensis, Anolis cooki (= A. 
cristatellus cooki Grant), and the very distinctive Typhlops 
granti have their primary distributions. 

SPECIES DISCUSSION 
Amphisbaena bakeri Stejneger 

Specimens examined: new localities: PUERTO RICO: 4.2 km 
S Mora, 500' (150 m), ASFS V5335 ; 7 km W Maricao, ca. 300 
meters, ASFS V6424. Previously recorded localities: "Las 
Marias," MCZ 66512 (= UPR 1?); "Maricao,"' MCZ 66513 
(= UPR 2?) ; "Mayagiiez," UMMZ 55824. 

Amphisbaena hakeri is known only from a few specimens (12 
including the new records cited herein) from a cluster of locali- 
ties in the north-central portion of extreme western Puerto Rico 
(Fig. 1). The previous localities for this species were all merely 
town names. From some of the localities were also recorded 
specimens of caeca, but due to the imprecise nature of the locali- 
ties, it could not be determined whether the two forms were 
truly sympatric. A. caeca was unknown from the northwestern 



BREVIORA 



No. 249 



corner of the island, and the only localities from which it was 
known "with" hakeri were those which were possibly peripheral 
to the range of hakeri (cf. Gans and Alexander, 1962, fig. 4). 

A. hakeri differs from the wide-ranging A. caeca principally 
in being larger in size, in having a higher number of body 
annuli, longer internasal suture, absence of a third row of chin 
segments, and lack of caudal autotomy (Gans and Alexander, 
1962). Only the second (although possibly the third) has been 
recognized as an absolute difference from caeca. The specific dis- 
tinctness of hakeri has therefore not been established with cer- 
tainty. 

One of the recent specimens (ASFS V5335) represents only a 
minor range extension but constitutes the first record from the 
northern limestone region ; ASPS V6424 is from within the pre- 
viously known range. There is no evident explanation for the 
small, very restricted, apparent range of hakeri. The fact that 
a number of specimens of caeca (and no hakeri) were collected 
in the vicinity of Utuado indicates that hakeri probably does not 
occur much farther east than it has been previously reported. 

The coloration in life of these two specimens of A. hakeri was 
noted as being "pale pinkish gray"; ASPS V6424 Avas noted as 
being "pink below" (ventrally). The right hemipenis of ASFS 
V6424 is everted and appears to be very similar to the hemipenis 
of A. caeca (Pig. 2). 





B 



Fig 2. Hemipenes of A, Amphisbaena schmidti (ASFS V5871), and B, 
AmpMsbaena caeca (ASFS X4111) ; view ceplialad with organs normal to 
body axis. 



1966 AMPHISBAENIDS OF PUERTO RICO 5 

ASFS V5335 from the northern limestone region was col- 
lected in a small grassy pasture, which formed the bottom of a 
doline in a mesic portion of the Haystack Hills. It was found 
beneath a rock where it was seen disappearing into a vertical 
burrow in moist clayey soil from which it had to be dug. ASFS 
V6424 was taken in a typical coffee producing area of the cen- 
tral highlands. It was found in a rotten log along the edge of a 
clearing overgrown with short, weedy vegetation. The nature 
of the ecological preferences of hakeri cannot, of course, be 
judged from just two encounters. The locality for the first 
specimen (4.2 km S Mora) is clearly within the range of caeca, 
the nearest locality for that species being less than three kilo- 
meters to the south. A. schmidii also occurs in this part of the 
limestone hills; the nearest locality for that species was 0.2 
kilometers to the north of the locality for the first specimen of 
hakeri. Thus, in this area, the three known Puerto Rican species 
occur sympatrically, whatever their ecological preferences (see 
below for caeca and schmidti) . 

Amphisbaena caeca Cuvier 

Specimens examined: new localities: PUERTO RICO : 13.3 km 
E Utuado, ASFS V4443-48 ; 17.7 km NE Utuado (= approx. 8 
km straight line distance), 1100' (335 m), ASFS V4482, V4499, 
V4500-01, V4623, V4449, V5860; 8 mi. (13.5 km) NW Utuado, 
1500' (407 m), ASFS V4548 ; 9 mi. (15 km) E Lares, ASFS 
V4656; 7.0 km S Mora, 800' (244 m), ASFS V5152, V5329. 
V5332; 5 mi. (8 km) NE Lares, 1200' (366 m), ASFS V5267 ; 
Ramey Air Force Base, Rifle Range Beach, ASFS V5430-31 ; 
2.3 km E Juana Diaz, 200' (61 m), ASFS V5755 ; 3 km NE 
San German, ASFS V6407 ; 4.6 km W, thence 4.6 km NW 
Juana Diaz, ASFS V6620-28, V6631-35; 3.6 km W, thence 9.7 
km N Juana Diaz, ASFS V6658-59 ; 3.6 km W, thence 8 km N 
Juana Diaz, ASFS V6660-61 ; 10.2 km E Dos Bocas, ASFS 
V5861-62; 18.8 km SW Arecibo, 800' (244 m), ASFS V5908- 
09; 11.2 km NW Utuado, 1300' (400 m), ASFS V5922 ; 
1 mi. (1.6 km) NW Dorado, ASFS V5939 ; 4.1 km NE Villa 
Perez, 2200' (670 m), ASFS V5973 ; 2 km NE Barranquitas. 
2100' (640 m), ASFS V6027 ; 1.8 km S Adjuntas, ASFS V6207 ; 
12.3 km SE Patillas, ASFS V4798 ; 10 km SE Patillas, ASFS 
V4802; 12.9 km SW Fajardo, 800' (244 m), ASFS V5081. 
VIEQUES : Cayo de Afuera, ASFS V6168. Previously recorded 
localities: PUERTO RICO: Isla Verde, ASFS X937-43, X4104- 



6 BREVIORA No. 249 

25, X7381-98; 2.2 mi. SW Sabana, ASFS X7433-34; Mayagiiez, 
UPR 84c, 87, 88, 90, 92, 140. 

Large numbers of A. caeca were examined by Gans and Alex- 
ander, but they had large series from the eastern part of the 
island only and relatively few specimens from the west. The 
species was not then known from the northwestern corner, an 
area bounded roughly by Mayagiiez, Lares and Arecibo. The 
present collection also has an eastern bias due to the large series 
from Isla Verde, but a more thorough coverage has been made 
in the west, including the formerly blank northwest portion. 

With the new material I have not undertaken a detailed 
analysis of trends such as that done by Gans and Alexander 
(fig. 29). However, I have compared trends in number of body 
annuli of selected eastern (Isla Verde and east), central (Juana 
Diaz region), west central (Utuado-Ad juntas region), and west- 
ern (Ramey Air Force Base to south of Mora) "samples" 
(these do not include all specimens examined) with those shown 
by Gans and Alexander for this character (in which the strong- 
est trends were evident). An agreement in direction of trends 
was found. Both pre- and postcloacals average fewer in eastern 
specimens ; this is also in agreement with the observations of 
Gans and Alexander. Additionally, internasal suture length 
(expressed as a fraction of interprefrontal suture length) aver- 
ages longer in the eastern specimens. 

The largest specimen (ASFS V6207) is a male that measures 
291 mm total length (tail 21 mm), which appears to be the 
largest caeca recorded (the largest seen by Gans and Alexander 
was 269 mm total length). A body annulus count of 214 (ASFS 
V4120) is lower than the low value of 217 recorded by Gans 
and Alexander. 

The Vieques specimen is, in a way, the first record for that 
island. The specimen was not taken on the main island but on 
an offshore key, Cayo de x\fuera. However, the species may be 
assumed to occur on Vieques proper. It has long been a matter 
of speculation as to which species of Amphishaena, would be 
found on Vieques and Culebra, caeca or fenestra fa. Some have 
tacitly assumed the former. Of course, the finding of one spe- 
cies does not preclude the possibility that the other might also 
occur on the island. The Cayo de Afuera specimen has a high 
number of ventral midbody segments ; it is also near the upper 
extreme of caeca in body annuli. 



1966 AMPHISBAENIDS OF PUERTO RICO 7 

A. caeca seems to prefer a mesic, usually shady, habitat (see 
below for comparisons with other species) ; it occurs from mon- 
tane cafetales at elevations of at least 2200 feet (ASFS V5973, 
apparently the highest elevation record for caeca) to the coast. 
It still appears to be absent from the extremely arid southern 
regions; the record of Fowler (1918) for Guanica probably does 
not pertain to this form (vide infra). One specimen (ASFS 
V4798) was collected approximately 40 feet from the ocean's 
edge and shed skins were found in the vicinity. Three specimens 
were collected in rotting wood, two in logs that were not even in 
firm contact with the ground. The most xeric habitat in which I 
have found caeca was on Cayo de Afuera ; even there it was in a 
wooded and somewhat shady, although not moist, situation. 

Amphisbaena xera new species 

Holotype: ^ICZ 81019, an adult male from 7 km E of Guanica, 
Puerto Rico, elevation 600' (183 m), one of a series collected on 
6 March 1965 by Albert Schwartz and Richard Thomas. 

Parafypes: PUERTO RICO: UIMNII 56910, ASFS V5646, 
USNM 152588, 7.3 km E Guanica, 2 March 1965, R. Thomas; 
ASFS V5659, 7.1 km E Guanica, 3 March 1965, R. Thomas; 
ASFS V5662, 7.3 km E Guanica, 3 March 1965, R. Thomas; 
ASFS V5722-23, AMNH 94170, same data as type (foregoing 
localities are at the same elevation as that of the type) ; UMMZ 
73844, Sabana Grande: ASFS Y5800, 6.2 km E Juana Diaz, 
10 March 1965, R. Thomas; ASFS V6408, 5 mi. (8 km) ESE 
San German, south slope Cerro Algarrobo, 9 July 1965. R. 
Thomas; ASFS Y6148, 5 mi. (8 km) ESE San German, south 
slope Cerro Algarrobo, 10 July 1965, R. Thomas; ASFS V6423, 

1.5 km AA'NW San German, 11 July 1965, R. Thomas; ASFS 
V6427, 3 km NE San German, 12 July 1965, R. Thomas; ASFS 
V6587, 2.3 km E Juana Diaz, 26 July 1965, R. Thomas ; ASFS 
V6616-18, 4.6 km W. thence 1.1 km NW Juana Diaz, 27 July 
1965, R. Thomas: ASFS V6619, 4.6 km W. thence 4.6 km NAV 
Juana Diaz, 27 July 1965, R. Thomas; ASFS V6636-40, 28 July 
1965, R. Thomas; ASFS V6646-57, 4.6 km W, 1.9 km NW 
Juana Diaz, 28 July 1965, R. Thomas; CM 40577-78, RT 1333-34, 

4.6 km W, thence 1.9 km NW Joaua Diaz. 29 July 1965, R. 
Thomas; UM:MZ 73846, 3 mi. (4.8 km) E Juana Diaz, 27 Novem- 
ber 1931, Chapman Grant; MCZ 36301, 10 mi. (16 km) E Juana 
Diaz, 27 November 1931, C. Grant ; UPR 84a-b, Mayagiiez, 10 



BREVIORA 



No. 249 



December 1957, J. A. Ramos; UPR 86, Mayagiiez, 8 September 
1949, Nestor Nazario ; UPR 91, no data. 

Diagnosis: A form of AmpMshaena lacking fusion of head 
scales, of closest affinities to the species hakeri and caeca, difiPer- 
ing from the former in having a low number of body annuli 
(225-234), the presence of a third row of chin segments (most 
specimens), and much smaller size. From caeca it differs in pos- 
sessing a relatively longer internasal suture (32 to 53 per cent 
of the length of the interprefrontal suture), lack of a postmalar 
row of chin segments, fewer average midbody segments, caudal 
autotomy absent or ill-defined, and smaller size. 






Fig. 3. Dorsal, lateral and ventral views of the head of the type of A. 
xera (MCZ 81019). Line represents 1 mm to scale. 



Description of type (Fig. 3) : Meristic and mensural data for 
type and paratypes are found in Table 1. The type is a mature 
adult male with the hemipenes partially everted. The first two 
body annuli correspond to three dorsal half -annuli ; the median 



1966 AMPHISBAENIDS OF PUERTO RICO 9 

segments of the second dorsal half-annulus are enlarged to form 
a pair of parietals in broad contact at the midline; the median 
segments of the third dorsal half-annulus are not enlarged to 
form a second pair of parietals. In the chin region two of the 
three segments of the first postgenial row are fused with two in 
the second row so that the second row is represented by only 
one segment (Fig. 3) ; the mental is partly fused with the genial. 
The postmalar^ row contains four scales and is bounded laterally 
by the malars which abut on the first body annulus. The dorsal 
groove is weakly indicated by an alignment and some medial 
rounding of the posterior paravertebral segments. 

Variation: In size, only five specimens exceed the type; 42 
(90 per cent) are smaller. The head scalation described for the 
type is characteristic ; a second pair of parietals is indicated in 
only four specimens. The internasal sutures range from 32 to 53 
per cent of the length of the interprefrontal sutures. In the chin 
region the most variability is encountered in the second row of 
postgenials. Its scales are often variously fused with the first 
and third rows, or it may be represented only by one or two small 
and irregular scales crowded between the first and second rows. 
Eighteen specimens have only two postgenial rows and thereby 
have the chin configuration of hakeri. It is as if the second 
postgenial row were being lost. The lack of a postmalar row is 
characteristic; one specimen (ASFS V6637) appears to have 
this row but the malar is abnormally divided. Autotomy con- 
strictions are evident in a few specimens at the level of the fifth 
to seventh caudal annulus ; slightly heavier pigmentation of a 
caudal annulus may mark the autotomy level in others. Other 
specimens either definitely do not have an autotomy constric- 
tion or the presence of one is indeterminate. Cloacal pores are 
four, with the exception of one specimen which lacks two on one 
side. Hemipenes are verv similar to those of hal'eri and caeca 
(Fig. 2)._ 

Coloration: The dorsal coloration in life of this form is typi- 
cally a pale tan, very reminiscent of the two living specimens of 
hakeri I have seen. The coloration lightens gradually towards 
the ventral surface and pigmentation is almost absent on the 
four ventralmost rows. Dropping out of pigmented ventral seg- 
ments takes place in some specimens but is not prominent be- 
cause of the fading out of all pigmentation ventrally. Dorsally 



II have objected elsewhere (Thomas, 196.5) to the use of the term postmalar; 
however, this was clue to my misunderstanding of the terminology as it applied 
to Amphisbaena innocens. 



10 BREVIORA No. 249 

and ventrally the coloration is darker (browner) anteriorly. In 
life the venter is pink due to the transiucence resulting from 
lack of pigmentation. The temporal region is deep reddish 
purple, probably another product of superficial vascularity and 
pigmentation. The snout (including prefrontals) is light (buffy) 
as is the posterior half of the edge of the lower jaw. 

Comparisons and discussion: The most pertinent comparisons 
of xera are with caeca and haker^i (Fig. 4). Seven specimens, 
having no postmalar row of chin segments and regarded by 
Gans and Alexander as A. caeca, are here regarded as A. xera. 
Aside from one specimen I have not seen (SU 7775, examined 
by Gans and Alexander) which may also be referable to xera, 
and a possible exception noted above, the absence of a postmalar 



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INTERPREFRONTAL 

Fig. 4. Scatter diagram of internasal versus interprefrontal suture 
lengths for A. caeca (stippled squares), A. haheri (solid triangles), and 
A. xera (hollow circles). Numbers in the xera symbols indicate the number 
of specimens possessing that value. Values were not plotted for all speci- 
mens of caeca examined, only enough to show variation including extremes. 
The symbols at the extreme right for xera and caeca represent the values 
for the largest specimen of each of these two forms. 



1966 AMPHISBAENIDS OF PUERTO RICO 11 

row distinguishes xera from caeca in all cases. Internasal suture 
length varies from 32 to 53 per cent of the interprefrontal suture 
in xera; in caeca the longest interprefrontal sutures (29 per cent 
of the interprefrontal length) are found in four specimens from 
eastern Puerto Rico, where the internasal suture length averages 
higher than in material from the rest of the island (and interest- 
ingly an area where sympatry with the two other long internasal 
suture forms is absent). A. xera is characterized by a low num- 
ber of segments around midbody; one specimen (from Maya- 
giiez) has 32 midbody segments, otherwise the range is 26 to 30; 
a count of 30 or below is found in less than 20 of over 300 speci- 
mens of caeca examined by Gans and Alexander and by me. In 
the low average number of cloacal scales xera is more like eastern 
caeca than like those from other parts of the island. In coloration 
xera is paler than most specimens of caeca which are typically 
brown or gray -brown rather than pale (or even reddish) tan 
(although this color does occur in caeca). Specimens of xera 
are usually identifiable on first sight by their paler color and 
more slender proportions {caeca is characteristically stouter, 
doubtless a correlative of the greater number of midbody seg- 
ments). The tails are tapered towards the tip and are not so 
cylindrical and rounded terminally as is typical of caeca; this 
is a distinguishing character in living specimens. The pale 
snout of xera is also not found in caeca (or is not so extensive). 

A. caeca and A. xera are allopatric for the most part; they 
have been taken together at four localities (Mayagiiez, 3 km NE 
San German, 4.6 km W, thence 4.6 km NW Juana Diaz, 2.3 km 
E Juana Diaz), all peripheral to the range of xera and appar- 
ently to that of caeca. At these localities there is no evidence of 
intergradation. Relatively short internasal sutures are found in 
the four Mayagiiez xera and in the one from Sabana Grande (30 
to 33 per cent of the length of the interprefrontal suture), but 
these do not overlap with internasal suture lengths for caeca 
from any part of the island. One Mayagiiez specimen (UPR 
84a) has 32 segments at midbody (16/16), the only xera with 
counts higher than 30. 

A. xera is strikingly like hakeri in several respects (internasal 
suture, lack of caudal autotomy, lack of a postmalar row, pale 
coloration), although in others it is significantly different (fewer 
body annuli, size, gradual fading of color on venter). In size, 9 
of the 12 known specimens of hakeri are larger than 42 (90 per 
cent) of the specimens of xera. The configuration of the chin 
segments may well be thought of as intermediate between the 



12 BREVIORA No. 249 

conditions in hakeri and caeca but tending more towards the 
former. The 18 .vera with but two postgenial rows are even more 
suggestive of a trend towards hakeri. 

Schmidt (1928) commented on three specimens from Maya- 
giiez that appeared to be intermediate between hakeri and caeca. 
Two of these (noted as having an internasal length of about one- 
half the interprefrontals) almost certainly pertained to A. xera 
(these specimens do not appear to be extant). The other speci- 
men (CNHM 12473) was identified by Gans and Alexander as 
A. hakeri. Two specimens of hakeri are known from Mayagiiez ; 
one (CNHM 12473) has a body annnlus count of 240, which is 
near the extreme low for that form ; another has a count of 255 
which is at the upper extreme. A specimen (UPR 7, Gans and 
Alexander), which has no locality data other than Puerto Rico, 
is probably from the Mayagiiez region ; it has a body annulus 
count of 239. It should also be recalled that three of the Maya- 
giiez (and the one from. Sabana Grande) specimens are at the 
upper extreme in size for .vera. More specimens with precise 
data from the Mayagiiez vicinity may show that hakeri and 
xera are subspecifically related. I am reluctant to so allocate 
these forms now because hakeri is so little known. 

A. schmiclti is distinguished from xera by its lower number 
of body annuli (192-207 versus 225-234), higher number of cau- 
dal annuli (18-22 versus 10-14), the presence of caudal autot- 
omy, larger size, different proportions, and overall darker color- 
ation. The two are additionally distinguished by differences in 
relative size of some of the head scales and by different chin 
segment configurations. 

Range: Known from the central to western parts of the south- 
ern xeric coastal plain of Puerto Rico (Fig. 1) and as far north 
as Mayagiiez, San German, Sabana Grande, and to the northwest 
of Juana Diaz. In all probability it occurs on Caja de Muertos/ 
whence there are shed skins yielding midbody segment counts 
of 14/16 (1) or 14/14 (2) and caudal annuli of 12 (2), 13 (1) 
or 14 (1). The specimen recorded by Fowler (1918) from 
Guanica may well have been of this new form. 

Hahitat: The topotypical paratypes and type (7.0-7.3 km E 
Guanica) were collected in the xeric woods which form the 
more luxuriant covering of the limestone hills of this region. 



iHeatwole, Torrps and Pleatwole (Stahlia, 1965. no. 4. p. 1) recorded taking a 
specimen of AviphiRhnrnn on Caja de Muertos. I have not examined tliis speci- 
men, but almost certainly it pertains to A. xera. as it was noted to laciv tiie post- 
malar row of chin segments and also agreed with xera in other counts given. 



{ 



1966 AMPIIISBAENIDS OF PUERTO RICO 13 

The soil varied from sandy with moderate organic (plant) con- 
tent to loamy; there was a uniform cover of leaf litter. At the 
type locality, A. xera occurs syntopically with Typhlops richardi 
and Typhlops granti. 

The specimen from 6.2 km E Juana Diaz was collected in a 
region of xeric foothills ; it was found at night under a rock 
which was not at all imbedded in the soil. The area was exposed 
and had very sparse vegetation. In the heat of the day it was 
doubtless a most inhospitable place, and the chance of finding 
an Amphishaena near the surface would have been negligible. 

At the following three localities I was able to personally col- 
lect and observe caeca and xera together. 

3 km NE San German. This locality is in the foothills to the 
north of the San German valley ; it is transitional between the 
arid region to the south and the mesic highlands, but closer to 
the latter. The single specimen of caeca was collected on a 
boulder-strewn and somewhat wooded hillside surrounded by 
pasture ; the specimen of xera was collected near a small stream 
at the base of the hillside. No ecological differences were dis- 
cernible. 

4.6 km W, thence 4.6 km NW Juana Diaz. This locality is in 
the valley of the Rio Inabon ; it too is transitional between the 
xeric south and the mesic interior. On the first visit, nine speci- 
mens of caeca and one of xera were found in a patch of sandy, 
mesic, river-border woods. The surrounding regions (where nat- 
ural growth was evident) were covered by semi-arid scrubby 
woods; tlie spiny palm (Acroconiia), a typically semi-xeric form, 
was seen on some of the nearby hillsides. On the second visit an 
attempt was made to collect in a more open, scrubby, overgrown 
field nearby. Five specimens of xera and five of caeca were ob- 
tained. Not quite three kilometers to the south, another area of 
shady (but distinctly more xeric) riverine woods was visited. 
Eighteen specimens of xera (two of which have not been exam- 
ined for this paper) were collected and no specimens of caeca. 
Less than a kilometer more to the south in still arid country, 
three more specimens of xera but no caeca were taken. 

3.3 km E of Juana Diaz. This locality is a mesic, Avooded ra- 
vine surrounded by typically xeric to semi-xeric regions. One 
specimen each of both species was found in the ravine. 



14 BREVIORA No. 249 

Amphisbaena schmidti Gans 

Specimens examined: new localities: PUERTO RICO: 5.5 mi. 
(9 km) N Lares, 1100' (346 m), ASFS V5197-99, V5258-60; 
5 mi. (8 km) NE Lares, 1200' (366 m), ASFS V5268 ; 4 km S 
Mora, 500' (153 m), ASFS V5334; Ramey Air Force Base, Rifle 
Range Beach, ASFS V5432-37 ; 3 mi. (4.8 km) W Las Llanadas, 
600' (183 m), ASFS V5871 ; 18.8 km SW Arecibo, 800' (244 m), 
ASFS V5910; 11.2 km NW Utuado, 1300' (400 m), ASFS 
V5923-24; 1 mi. (1.6 km) NW Dorado, ASFS V5940; 3.5 mi. 
(5.4 km) W Playa de Vega Baja, ASFS V5959 ; 2.5 km SW 
Florida, ASFS V5999 ; 5.6 km NE Morovis, 400' (122 m), ASFS 
V6028-29. 

Amphisl)ae7ia schmidti was described (Gans, 1964) on the 
basis of six specimens from four more or less coastal localities, 
three in the northwest section of the island and one ( ' ' Salinas ' ' ) 
presumably on the south coast. Professor Manuel J. Velez, col- 
lector of the type and one paratype of schmidti, has informed 
me {in litt.) that the type locality of A. schmidti should cor- 
rectly be "Isabela, orilla (cuneta) Carretera Caiio." The lo- 
cality is in the district of Isabela about six kilometers southeast 
of the town of Isabela hy the side of Carretera Caiio (= road 
number 113). 

The distribution of schmidti was given as "coastal Puerto 
Rico." The new localities cited here are restricted to the north- 
ern limestone section (up to elevations of 1300 feet), and the 
species is in all likelihood endemic to this region (Fig. 1). No 
specimens were found elsewhere on the island. 

Of the numbers of amphisbaenids collected in the southern part 
of Puerto Rico (A. caeca and A. xera) and of the shed skins 
that were often found there, none pertained to A. schmidti. 
The apparent absence of schmidti from the southern mar- 
ginal region plus its absence from the coastal regions east of 
San Juan (in spite of the large series of caeca from there) indi- 
cate that its distribution is not coastal Puerto Rico and, at the 
same time, throw considerable doubt on the Salinas record. Con- 
ceivably it refers to another Salinas (a not uncommon coastal 
name in Latin America), possibly Punta Salinas on the north 
coast to the west of San Juan, which is probably in the range of 
schmidti. 

The new specimens of schmidti agreed in coloration (in life) 
very closely with the coloration recorded by Gans ; they were 
usually noted as being purplish brown dorsally with darker 



1966 



AMPHISBAENIDS OF PUERTO RICO 



15 



brown heads and tails. The venters are heavily pigmented but 
sometimes pinkish. In general proportions and appearance this 
species is very distinct; the blunted and almost triangular head 
is characteristic and is accentuated by the deeply constricted 
neck. Indeed, this species, in proportions, color and scutellation, 
is possibly the most "different" of the species of the Greater 
Puerto Rican region. The enlarged parietals and enlarged scales 
at the end of the third row of chin segments are very character- 
istic. 

Completely everted hemipenes (Fig. 2) are jjresent in two 
specimens. The organs are relatively slender and deeply bilobed ; 
the sulcus spermaticus forks in the crotch, and the apices are 
flattened, a condition that has been noted for innocens and not 
for caeca which has rounded apices (Thomas, 1965). Autotomy 
level varies from the sixth to eighth caudal annulus. 

Different ecological preferences were observed for schfnidti 
and caeca in the jointly occupied parts of their ranges. A. 
schmidti was found in more open and exposed areas, the edges of 
pastures, the cleared sides of gradually sloping dolines — areas 
where the tree cover is lacking, or high and sparse. A. caeca, 
although wider in its ecological range, prefers shady areas such 
as coffee groves or wooded ravine bottoms. The following table 
indicates briefly the type of situation in which either species 
was collected at particular localities within their joint range ; 
the number of specimens at each locality is indicated in paren- 
theses. 



schmidti 
acacia scrub (6) 

moderate to well-exposed, scrubby 
ravine sides (6) 

moderatelj' exposed hillside (1) 



exposed hillside pasture (2) 

high, open ravine woods (1) 

moderately exposed edge of 
coffee grove (1) 



caeca 
coastal almond (Tenninalia) 
woods (2) 



coffee grove (3) 

coffee grove (1) 

niesic woods adjacent to 
pasture (1) 

exposed ravine side (1), shady 
ravine woods (1) 

interior of coffee grove (1) 



16 BREVIOBA No. 249 



schmidti 



gutiva scrub at rim of 
doline pasture (1) 



moist, open pasture (2) 
cleared slope of doline (1) 



in coffee grove (2) 



moderately exposed roadside shady coastal woods (1) 

ditch (1) 

open lake margin (1) 



Amphisbaena fenestrata Cope 

Specimens examined: new localities: VIRGIN ISLANDS. 
St. John: Lameshur Bay, VINP 1478; Annaberg Road near 
Frederickdal, VINP 1479 ; Caiieel Hill Trail, VINP 1485 ; Coral 
Bay, VINP 1490 ; Windberg ruins, ASFS V7504-05 ; Frederick- 
dal ruins, ASFS V7564, V8066. Tortola: Baugher's Bay, ASFS 
V7921; Jackass Gut, ASFS V7940, V7984-85, RT 977. Virgin 
Gorda: SW slope of Gorda Peak, ca. 500' (150 m), ASFS V3805- 
06. Great Camanoe: between Lee Bay and Cam Bay (shed skin). 
Previously recorded localities: St. John: VINP 1496-97. 

In the report on A. fenestrata by Gans and Alexander, "Doro- 
thea" and "West End of Great St. James" are listed in "local- 
ity records" under St. John. Dorothea is on St. Thomas and 
Great St. James is a small satellite island of St. Thomas. 

When the data from the new specimens are placed with the old 
data from the various islands, the trends noted by Gans and 
Alexander are almost completely obliterated. The following 
ranges summarize the new data (data from Gans and Alexander 
are in parentheses) : 

body caudal ventral 

Island annuli annuli segments posteloaeals 

St. Thomas (242-247) . . (12-14) ... (14-16) . . (8-10) 

St. John 241-251 (247-249) 12-13 (12-14) 14-17 (14-15) 8-11 (9-10) 

Tortola 243-248 (236-242) 13-14 (13-14) 14-16 (14-16) 8-11 (9-12) 

Virgin Gorda 240-245 12-13 . . 16-17 . 9 

The shed skin from Great Camanoe is too fragmentary to 
yield counts. The Virgin Gorda and Great Camanoe records 
are the first for those islands. The two specimens from Virgin 
Gorda have higher dorsal midbody counts than do those of the 
other islands; the total midbody counts are higher than all but 
one of the specimens from the other islands. 



1966 AMPHISBAENIDS OF PUERTO RICO 17 

Gans and Alexander have said (1962:95) that the dorsal 
groove in West Indian Amphisbaena "may be suggested by an 
alignment alone." However, in well-injected specimens of A. 
fenestrata the dorsal groove is structurally evident on approxi- 
mately the posterior two-thirds to three-fourths of the body. 
The paramedian segments are rounded medially and the result- 
ing space is occupied by "fold granules." The appearance of 
this groove at its fullest development is much the same as that 
of the lateral grooves. It becomes increasingly indistinct ante- 
riorly and ends at the approximate level of the thirty-sixth to 
sixty-seventh postoral annulus. Xo other specimens of West 
Indian species seen by me {caeca, schmidti, hakeri, cubana, inno- 
cens, gonavensis, manni) have so prominent a dorsal groove, 
although one specimen of hakeri has a moderately well indicated 
dorsal groove. 

Amphisbaena fe7iestrata inhabits islands that are preponder- 
antly xeric; however, it prefers (or at least is most easily col- 
lected in) the most mesic areas that can be found. When we 
visited Virgin Gorda the weather was unseasonably dry. Shed 
skins of these animals were found regularly under rocks in the 
more shady parts of the low, xeric woods that cover much of the 
island. After almost a week of persistent rock turning, two 
specimens were obtained on a heavily wooded hillside after a 
heavy morning shower. Typhlops richardi, which was more 
abundantly collected in all parts of the island, was also found 
in moist piles of coconut husks near the coast, where Amphis- 
baena fenestrata was not. This preference for the more mesic 
areas was noticed also on St. John and Tortola. The presence 
of a soil of high organic content seems to be an important fac- 
tor. A brief visit to the island of Anegada yielded no specimens 
or signs of amphisbaenids, although six specimens of Typhlops 
were collected. 

This species is now known from the islands of St. Thomas, 
Great St. James, St. John, Tortola, Great Camanoe, and Virgin 
Gorda. Its true range is probably all of the Virgin Islands large 
enough to bear a suitable habitat, St. Croix excepted. Anegada 
is geologically distinct and well set ofiF from the rest of the Vir- 
gins, but its fauna is not radically different ; fenestrata may be 
found there. 

ACKNOWLEDGEMENTS 

The collecting that preceded this study and the study itself 
were supported by Dr. Albert Schwartz, to whom I am most in- 



18 BREVIORA No. 249 

debted. Sr. Felix Inigo of the Department of Agriculture and 
Commerce of the Commonwealth of Puerto Rico has been prompt 
and courteous in his attention to requests for collecting permits ; 
Mr. Jesse E. Williams was of great assistance to our collecting 
on Ramey Air Force Base, Puerto Rico. Dr. Carl Gans and Mr. 
A. Allan Alexander have helpfully read and criticized the manu- 
script. I wish to express appreciation to Mr. Donald W. Buden, 
Mr. Gerald D. Gagnon, Mr. David C. Leber, and Mr. Ronald F. 
Klinikowski for their most able assistance in the field. 

Much appreciation is due to the following people who loaned 
material for study: Mr. Hugh B. Muller, Virgin Islands Na- 
tional Park (VINP) ; Dr. Charles F. Walker and Mr. George 
R. Zug, University of Michigan Museum of Zoology (UMMZ) ; 
Dr. Ernest E. Williams, Museum of Comparative Zoology at 
Harvard (MCZ) ; Dr. Juan A. Rivero and Mr. Horaeio Mayorga, 
University of Puerto Rico at Mayagiiez (UPR). The following 
institutions have received paratypes of Amphishaena .vera: the 
U.S. National Museum (USNM), the University of Illinois Mu- 
seum of Natural History (UIMNH), and the Carnegie Museum 
(CM). RT indicates the Richard Thomas private collection. 

LITEEATURE CITED 

Gans, Carl 

1964. Amphisbaerm schmidti, a third species of the genus from Puerto 
Rico (Amphisbaenia, Reptilia). Breviora, Mus. Comp. Zool., No. 
198, pp. 1-11. 

Gans, Carl and A. Allan Alexander 

1962. Studies on amphisbaenids (Amphisbaenia, Reptilia). On the 
amphisbaenids of the Antilles. Bull. Mus. Comp. Zool., Vol. 128, 
no. 1, pp. 65-158. 
FoAVLER, Henry W. 

1918. Some amphibians and reptiles from Porto Rico and the Virgin 
Islands. Pap. Dept. Marine Biol. Carnegie Inst., vol. 12, pp. 
1-15. 
Schmidt, Karl Patterson 

1928. Amphibians and land reptiles of Porto Rico, with a list of those 
reported from the Virgin Islands. Sci. Surv. Porto Rico and 
Virgin Islands, vol. 10, part 1, pp. 1-160, 4 pis. 
Thomas, Richard 

1965. Two new subspecies of Amphishaena (Amphisbaenia, Reptilia) 

from the Barahona Peninsula of Hispaniola. Breviora, Mus. 
Comp. Zool., No. 215, pp. 1-14. 

(Received 2 November, 1965) 



1966 AMPHISBAENIDS OF PUERTO RICO 19 

KEY TO GREATER PUERTO RICAN SPECIES OF AMPEISBAENA 

1. Nasals separated by posterior extension of rostral which contacts pre- 
frontals; body annuli 236-249 fenestrata 

Nasals in contact with one another 2 

2. Number of body annuli low (192-207); high number of caudal annuli 

(18-22) schmidti 

Number of body annuli higher (more than 214) ; caudal annuli lower 
(17 or less) 3 

3. Internasal suture short (less than one-third interprefrontal suture) ; 

body annuli 214-237 caeca 

Internasal suture long (one-third or more the length of interprefrontal 
suture) 4 

4. Number of body annuli higher (239-255) ; size large haJceri 

Number of body annuli lower (225-237), size small xera 



TABLE 1 

Meristic and mensural data for specimens of Amphisbaena examined. 
Tail and total length measurements are in millimeters. Internasal and in- 
terprefrontal suture lengths are in ocular micrometer units; one unit equals 
0.125 mm. Asterisks indicate abnormality or autotomized tails. 



20 



BREVIORA 



No. 249 



SPECIMEN NUMBER 



ANNULI SEGMENTS 



tj <i 



W o Eh 

US M to 
o m o 



Eh M 



M HI E-i 



A. BAKEHI 






















ASFS V5335 
ASFS VSkZk 
MCZ 66512 
MCZ 66513 
UMMZ 55824 


251+ 
255+: 
253+ 
245 + 


4 +12 
3/4+14 
4 +13 
4 +13 


16/16 
14/16 
16/17 
16/18 


3 
3 
3 

3 


4 
4 
4 
5 


- - 


4+7+11 
4+6+10 
4+6+11 
4+8+11 


9 

9 

11 

11 

9 


20 
20 
21 
27 
17 


243 12 
236 15 
269 17 
322 20 


A. CAECA 






















ASFS V543O 
ASFS 75431 
UPR 84c 
UPH 87 
UPR 88 


229 + 
226+ 
227+ 
229 + 
223+ 


3 +14 
3 +14 
3 +15 
3 +16 
3 +15 


16/16 
16/16 
16/16 
14/16 
17/18 


3 
3 
3 
3 
3 


4 
4 
3 
4 
4 


- 9 

- 9 
5 9 

- 9+10* 

- 9 


4+6+14 
4+6+12 
4+8+13 
4+8+13 
4+7+11 


3 
4 
4 
5 
4 


22 
22 

24 
22 

28 


231 15 
196 13 
193 13 
215 18 
198 14 


UPR 90 
UPR 92 
UPR 140 
ASFS V5I52 
ASFS V5329 


227 + 
225+ 

228 + 
234 + 
233 + 


3 +15 
3 +6* 
3 +15 
3 +14 
3 +15 


16/17 
16/17 
16/16 
16/18 
16/18 


3 
3 
3 
3 
3 


4 
4 
4 
4 
2 


- 8 

- 9 

- 9 

- 8 

- 8 


4+6+13 
4+8+12 
4+6+12 
4+8+13 
4+8+12 


4 
5 
5 
3 
6 


26 
23 

21 
23 

24 


196 14 

170 13 
206 14 
202 11 


ASFS V5332 
ASFS V53O5 
ASFS V5267 
ASFS V6407 
ASFS V5908 


231 + 
232+- 
225+ 
216 + 

227+ 


3 +15 
3/4+14 
3 +5* 
3 +16 
3 +5» 


16/18 
16/16 
16/18 
17/16 
16/18 


3 
3 
3 
3 
3 


4 
4 
4 
4 
4 


- 3 

- 9 

- 9 

- 9 

- 9 


4+8+12 
4+8+12 
4+8+12 
4+8+12 
4+7+12 


6 
4 
4 
4 
3 


24 
21 
24 
19 
25 


225 15 
186 13 
206 7* 
172 14 
218 7* 


ASFS V5909 
ASFS V4656 
ASFS V4548 
ASFS V5939 
ASFS V5973 


221 + 
221 + 
229 + 
227 + 
232 + 


3 +15 

3 +15 

4 +15 
3 +14 
3 +5* 


16/18 
16/16 
16/17 
15/17 
15/18 


3 
3 
3 
3 
3 


4 
4 
4 
4 
1* 


- 8 

- 9 

- 9 

- 9 

- 6» 


4+8+12 
4+6+12 
4+8+13 
4+8+11 
4+6+12 


3 
2 

3 

4 
4 


19 
26 
25 

24 
20 


160 13 
217 17 
225 16 
215 16 
148 5* 


ASFS V6207 
ASFS V4443 
ASFS V4444 
ASFS V4445 
ASFS V4446 


231+ 3 +14 
232+2/3+14 
233+ 3 +15 
234+2/3+15 
232+ 3 +14 


16/17 
14/16 
16/17 
16/16 
17/17 


3 
3 
3 
3 
3 


3 

5 
4 
4 
6 


- 8 

- 10 

- 9 

- 9 

- 9 


4+6+13 
4+6+12 
4+7+13 
4+7+12 
4+8+13 


5 
3 

4 
3 
4 


34 
18 
27 
26 
27 


291 21 
241 16 
231 16 
210 14 
224 16 


ASFS V4447 
ASFS V4448 
ASFS V4449 
ASFS V4482 
ASFS V4499 


235 + 
230 + 
232 + 

230 + 

231 + 


3 +15 

3 +14 

4 +15 
3 +15 
3 +6* 


16/18 
14/16 
14/16 
16/17 
14/16 


3 
3 
3 
3 
3 


4 
5 
3 
4 
4 


- 10 

- 8 

- 7 

- 9 

- 8 


4+8+13 
4+6+13 
4+7+12 
4+7+14 
4+7+13 


4 
4 
4 
4 
4 


24 
21 

24 
21 
27 


195 13 
174 12 
210 16 
165 11 
226 7< 


ASFS V4500 
ASFS V4501 
ASFS V4623 
ASFS V586O 
ASFS V5861 


230 + 
229+; 
2 31+; 
234+ 
235+ 


3 +15 

2/3+14 

3/3+15 

3 +15 

3 +14 


14/16 
15/16 
15/16 
16/18 
14/16 


3 
3 
3 
3 
3 


4 
4 
4 
4 
4 


- 9 

- 8 
3* 9 

- 8 

- 9 


4+8+12 
4+8+14 
4+8+13 
4+8+12 
4+8+13 


4 
4 
3 
3 
4 


26 
21 
24 
26 
20 


230 14 
171 11 
198 15 
209 15 
160 10 



1966 



AMPHISBAENIDS OF PUERTO RICO 



21 



ASPS V5862 
ASF3 V6620 
ASFS V6621 
ASFS V6622 
ASFS V6623 

ASFS V6624 
ASFS V6625 
ASFS V6626 
ASFS V6627 
ASFS V6628 

ASFS V663I 
ASFS V6632 
ASFS V6633 
ASFS 7663^+ 
ASFS V6635 

ASFS V6658 
ASFS V6659 
ASFS V666O 
ASFS V666I 
ASFS V5755 

ASFS V5939 
ASFS V6O27 
ASFS X937 
ASFS X938 
ASFS X939 

ASFS X940 
ASFS X9'il 
ASFS X9'+2 
ASFS X9'+3 
ASFS X4104- 

ASFS X4105 
ASFS X4106 
ASFS Xi+107 
ASFS X4108 
ASFS X4109 

ASFS X4110 
ASFS X4111 
ASFS X4112 
ASFS X4113 
ASFS X4114 

ASFS X4115 

ASFS Xi+lie 

ASFS Xkll7 

ASFS X4118 

ASFS X'+119 

ASFS XiH20 
ASFS X'+121 
ASFS X4122 
ASFS X'+123 
ASFS X4124 

ASFS Xi+125 

ASFS X73BI 

ASFS X7382 

ASFS X7383 

ASFS X7384 

ASFS X7385 
ASFS X7386 
ASFS X7387 
ASFS X7388 
ASFS X7389 



226+2/3+15 
223+2/3+1'+ 
223+ 3 +1'+ 
219+ 3 +1^* 
220+ 3 +1^ 

226+ 3 +12 
219+2/3+13 
221+ 3 +1^+ 
221+ 3 +13 
22U+ 3 +13 

220+ 3 +1^ 
222+ 3 +14 
222+ 3 +1^+ 
219+ 3 +1'* 
223+2/3+1^+ 



220+ 3 
220+ 2 
223+ 3 
222+ 2 
225+ 4 



+13 

+1^* 
+1'+ 
+lil 
+1U 



14/16 
16/16 
16/16 
16/16 
16/17 

17/16 
18/18 
16/16 
16/16 
16/16 

16/16 
16/16 
16/16 
16/16 
16/16 

16/18 
16/18 
16/16 
16/18 
17/18 



223+ 4 +14 14/16 

226+3/4+12* 18/18 

224+ 4 +12 16/18 

222+ 4 +12 16/18 

226+ 3 +13 16/18 



223+3/'*+13 
229+ 3 +14 
220+ 4 +13 
224+2/3+14 
222+ 3 +14 

225+ 4 +13 
228+ 3 +14 
229+ 4 +15 
220+ 3 +13 
223+ 3 +14 

218+ 3 +14 
230+3/4+13 
223+3/'*+13 
218+ 3 +14 
228+3/4+14 

228+ 4 +13 
221+ 4 +14 
221+ 3 +11 
228+ 4 +14 
222+3/4+13 

214+ 4 +11 
225+ 4 +14 
226+ 4 +14 
222+ 3 +13 
222+ 3 +13 

226+ 3 +14 
225+ 4 +14 
227+3/4+12 
221+ 3 +14 
229+ 3 +12 



17/18 
16/16 
16/18 
15/16 
16/17 

16/18 
16/18 
16/16 
14/16 
16/18 

16/18 
16/16 
16/16 
16/18 
16/18 

16/18 
15/16 
16/16 
16/18 
16/17 

16/17 
16/18 
15/16 
15/17 
16/16 

16/18 
16/18 
16/18 
16/18 
15/16 



224+ 3 +13 16/18 

224+ 3 +14 16/18 

220+3/4+13* 16/18 

227+ 3 +14 15/18 

224+ 4 +12 16/16 



- 9 

- 10 

- 10 

- 9 

- 10 

- 10 

- 7 

- 8 

- 9 

- 9 

- 9 

- 9 

- 9 

- 9 

- 9 

- 9 

- 6 

- 9 

- 9 

- 9 

- 9 



- 9 

- 7 

- 9 

- 9 

- 9 

- 8 

- 7 

- 7 



4+8+12 
4+7+14 
4+6+12 
4+8+12 
5+8+14 

4+8+12 
4+7+12 
4+8+12 
4+6+12 
4+7+13 

4+6+13 
4+7+13 
4+7+12 
4+8+12 
4+8+12 

4+8+13 
4+8+13 
4+6+13 
4+7+12 
4+8+12 

4+7+13 
4+8+12 
4+6+12 
4+6+12 
4+7+13 

4+6+13 
4+6+12 
4+6+12 
4+8+11 
4+6+10 

4+7+10 
4+6+12 
4+6+13 
4+6+12 
4+6+10 

4+6+13 
4+6+11 
4+6+12 
4+6+11 
4+6+13 

4+6+12 
4+6+11 
4+6+13 
3+6+12 
4+6+12 

4+8+10 
4+6+11 
4+6+11 
4+6+11 
4+6+12 

4+6+12 
4+6+11 
4+6+10 
4+8+10 
4+6+13 

4+6+11 
4+6+10 
4+6+13 
4+7+10 
4+8+12 



19 
27 
27 
23 
19 

25 
25 
22 
24 
23 

27 
25 
23 
24 
23 

18 
23 
25 
23 
23 

23 

24 
28 
24 
23 



5 25 

5 24 

4 20 

4 18 

4 22 



23 
23 

25 
24 
23 

23 
24 
24 
23 
22 

24 
20 
22 
23 
25 

22 
26 
21 
19 
19 

18 
24 
23 
25 
23 

22 
23 
25 
21 
21 



148 9 
227 17 
218 15 
205 15 
205 14 

197 13 

196 14 
145 14 

193 12 
185 13 

218 16 
202 14 

202 15 
177 13 
191 14 

227 16 
185 13 
215 15 
210 15 
199 13 

208 14 

220 17 
245 15 

221 15 
208 15 

214 15 
207 14 
177 12 
109 7 

214 16 

222 16 
220 15 

226 17 
205 14 
199 I't 

220 16 

227 16 
201 13 

203 15 

197 1^+ 

229 16 
188 14 

194 14 

215 16 



181 12 
227 16 
187 14 
131 7 
126 8 

113 8 
224 15 
211 14 
222 16 

209 15 

215 13 

210 14 
214 13* 
195 !>* 
183 12 



22 



BREVIORA 



No. 249 



ASF3 X7390 
ASF3 X7391 
ASFS X7392 
A3FS X7393 
ASFS X739'+ 

ASFS X7395 
ASFS X7396 
ASFS X7397 
ASFS X7398 
ASFS V4798 

ASFS V4802 
ASFS V5081 
ASFS X7433 
ASFS X7'+34 
ASFS V6168 

A. XERA 

UPH eUa 
UPR 84b 
UPH 86 
UPH 91 
ASFS 76408 

ASFS V6418 
ASFS V6423 
ASFS V6427 
UMMZ 73844 
UIMNH 56910 

ASFS V5646 
U3NM 152588 
ASFS V5659 
ASFS V5661 
MCZ 81019 

ASFS V5722 
ASFS V5723 
AMNH 94170 
ASFS V6616 
ASFS V6617 

ASFS V66I8 
ASFS V6619 
ASFS V6636 
ASFS V6637 
ASFS V6638 

ASFS V6639 
ASFS V6640 
ASFS V6646 
ASFS V6647 
ASFS V6648 

ASFS V6649 
ASFS V6650 
ASFS V665I 
ASFS V6652 
ASFS V6653 

ASFS V6654 
ASFS V6655 
ASFS V6656 
ASFS V6657 
CM 40577 

CM 40578 
RT 1333 
HT 1334 
ASFS V6587 
UMMZ 73846 
ASFS V58OO 
MCZ 36301 



226+ 3 +13 
227+3A+13 
227+ 4 +14 
225+ 4 +13 
222+ 3 +14 

229+ 3 +13 
227+ 3 +14 
224+ 4 +14 
226+4/3+14 
219+ 3 +15 

232+4/2+15 
228+ 3 +13 
233+ 3 +14 
229+ 3+15 
236+ 3 +15 



224+ 4+13 
221+2/3+14 
224+2/3+13 
234+ 3 +16 
221+ 3 +13 

220+ 3 +13 
220+ 2 +13 
233+ 2 +15 
226+ 3 +14 
225+3/4+10 

234+ 2 +13 
231+ 2 +14 
225+ 3+13 
232+ 3 +13 
232+ 2 +13 

226+ 3 +13 
231+ 2 +13 
229+ 3 +14 
233+ 2 +13 
224+ 3 +14 

228+ 3 +13 
227+ 2 +13 
231+ 2 +14 
237+ 3 +14 
234+ 2 +13 

232+ 2 +14 
229+2/3+13 
230+ 2 +14 
228+3/2+14 
228+3/4+12 

228+2/3+15 
229+ 3 +13 
234+ 3 +14 
223+ 3 +13 
225+ 2 +14 

227+ 3 +13 
229+ 3 +13 
225+ 2 +14 
230+ 2 +15 
228+ 3 +14 

225+ 2 +5* 
227+ 3 +12 
228+ 2 +14 
231+ 3 +14 
235+ 3 +15 
231+ 3 +14 
230+ 4+13 



15/18 3 3 

16/19 3 4 

16/18 3 4 

17/18 3 4 

16/18 3 4 

16/18 3 4 

16/17 3 4 

16/18 3 4 

16/16 3 4 

16/16 3 4 



16/18 3 

17/19 3 

16/18 3 

16/16 3 

14/18 3 



16/16 
14/16 
14/16 
14/14 
14/14 

14/16 
14/16 
14/16 
13/15 
14/16 

14/16 
14/16 
14/16 
14/15 
14/16 

14/16 
14/14 
14/16 
14/15 
13/15 

14/16 
14/15 
14/14 
14/14 
12/14 

14/14 
14/15 
13/14 
14/14 
14/16 

14/16 
14/14 
14/15 
14/14 
13/14 

14/16 
14/14 
14/16 
14/14 
14/14 

14/15 
14/14 
13/14 
14/15 
14/16 
14/16 
14/14 



2 5 

1 4 
4 5 
4 - 

2 4 



- 9 

- 9 

- 9 

- 9 

- 8 

- 9 

- 9 



5 - 
4 - 



5 - 

4 - 

4 - 

5 - 



5 - 



6 - 



4+6+12 
4+6+12 
4+6+12 
4+6+12 
4+6+11 

4+6+12 
4+8+13 
4+6+11 
4+6+11 
4+9+13 

4+8+13 
4+8+12 
4+6+11 
4+8+11 
4+8+12 



4+6+10 
4+8+10 
4+8+11 



4+8+12 
4+6+13 
4+7+11 
4+6+12 
4+6+10 

4+6+8 

4+6+10 

4+7+12 

4+6+10 

4+6+12 

4+6+10 
4+6+10 
4+6+10 
4+7+10 
4+8+12 

4+8+11 
4+8+12 
4+8+11 
4+8+13 
4+7+11 

4+6+11 
4+8+10 
4+8+11 
4+7+13 
4+8+11 

4+7+10 
4+8+11 
4+7+10 
4+8+10 
4+8+10 

4+8+10 
4+8+10 
4+8+11 
4+8+11 
4+8+9 

2+8+10 
4+7+11 
4+8+10 
4+6+12 
4+6+11 
4+8+12 
4+7+12 



23 
22 
21 
22 
22 

22 

17 
18 
16 
16 

24 
25 
26 
27 
19 



8 25 

10 32 

8 24 

7 22 

4+6+12 7 17 



16 
18 
19 
22 
18 

17 
16 
17 
17 
19 

18 
18 
16 
17 
17 

17 
17 
17 
16 
17 

17 
15 

17 
18 
17 



7 16 

8 16 

8 16 

9 17 
8 16 



17 
17 
17 
15 
17 



7 17 

8 16 
8 17 
8 18 

7 16 

8 16 
7 18 



194 13 

189 14 
194 14 

182 13 

190 14 

190 13 
106 6 

102 7 
238 19 

230 16 
227 15 
226 14 
240 16 

183 13 



236 16 
181 14 
245 18 
241 18 
168 10 

140 9 
162 11 

202 14 

237 20 

193 11 

186 12 
170 11 

188 13 

191 12 
210 13 

190 14 

195 12 
158 12 

194 13 
175 12 

185 12 

177 12 

186 12 

187 14 

178 11 

170 11 
162 11 

203 14 

192 13 
200 13 

193 13 

191 13 
187 12 
187 13 
183 12 

186 13 
181 12 
181 12 
139 9 

196 14 

180 4« 

181 11 
191 14 
221 15 
147 11 
191 13 

189 14 



1966 



AMPHISBAENIDS OF PUERTO RICO 



23 



A. SCHMIDTI 



ASFS V5432 
ASFS 75433 
ASFS V5434 
ASFS V5435 
ASFS V5436 


200+ 3 
201+ 3 
199+ 3 
200+ 3 
200+ 3 


+19 
+20 

+18 
+20 
+20 


14/16 
13/16 
14/16 
13/16 
13/16 


2 
2 
2 
2 
2 


3 - 

4 - 
3 - 
3 - 
3 - 


7 
7 

6 
7 
6 


4+6+12 

4+5+8 

4+6+13 

4+8+10 

4+6+12 


192 19 

182 17 
189 19 
176 17 
159 17 


ASFS V5437 
ASFS V5334 
ASFS V5I97 
ASFS V5198 
ASFS V5I99 


196+ 3 
199+ 3 
200+ 3 
196+ 3 
201+ 3 


+8* 
+21 
+22 
+20 
+20 


14/16 
14/16 
14/16 
14/16 
14/16 


2 
3 
2 
2 
2 


3 - 

4 - 
4 - 

3 - 

4 - 


7 
7 
7 
6 
6 


4+^+11 
4+6+11 
4+8+10 
4+6+11 
4+8+10 


138 7» 
119 12 
224 26 
201 23 
180 20 


ASFS V5258 
ASFS V5259 
ASFS V526O 
ASFS V5268 
ASFS V5910 


202+ 3 +21 
202+ 3 +21 
204+ 3 +21 
200+3/4+20 
197+ 2 +8* 


14/16 
14/16 
14/16 
16/17 
14/16 


2 
2 
2 
3 
2 


3 - 

4 - 
2 - 

4 - 
4 - 


6 
7 
6 
6 
7 


4+6+11 

4+8+8 

.4+6+11 

4+6+11 

4+6+11 


221 23 
208 24 
105 11 
123 13 
195 11* 


ASFS V5923 
ASFS V5924 
ASFS V587I 
ASFS V5999 
ASFS V6O28 
ASFS V6029 
ASFS V5959 
ASFS V5940 


205+3/2+21 
207+ 3 +21 
201+ 4 +21 
200+ 3 +21 
201+ 3 +21 
192+ 3 +20 
196+3/2+20 
195+ 3 +7* 


14/16 
14/16 
14/16 
14/18 
14/17 
14/16 
14/17 
14/16 


2 
2 
2 
2 
2 
2 
2 
2 


4 - 

4 - 

5 - 

2 - 
4 - 
4 - 

3 - 
1 - 


7 
6 
7 
6 
6 
7 
6 
6 


4+8+9 

4+8+10 

4+8+14 

4+6+10 

4+7+9 

4+6+12 

4+6+11 

5+6+12 


194 21 
183 19 
240 26 
212 24 
220 24 
200 22 
137 14 
200 11* 


A. FEt.'SSTHATA 



















ASFS V7504 
ASFS V7505 
ASFS V7564 
ASFS V8O66 
VI NP 1478 

VINP 1479 
VINP 1485 
VINP 1490 
VINP 1496 
VINP 1497 

ASFS V7921 
ASFS V7940 
ASFS V7984 
ASFS V7985 
HT 977 
ASFS V3805 
ASFS V3806 



243+ 
250 + 
243+ 
248+ 
247+ 



+13 
+13 
+12 
+12 
+13 



243+3/2+13 
246+ 2 +13 
246+3/2+13 
251+ 2 +12 
241+2/3+12 



+14 
+13 



247+ 2 
248+ 2 .^^ 
243+ 3 +14 
248+ 2 +14 
242+ 2 
240+ 2 



+14 
+12 



245+ 3 +13 



14/16 
14/16 
14/16 
14/15 
14/16 

14/16 
13/14 
14/16 
14/14 
14/17 

14/14 
14/14 
14/15 
14/16 
14/15 
15/17 
15/16 



3 4 

3 4 

3 4 

3 ^ 



10 

9 

10 

10 



4+6+10 

4+6+10 

4+6+8 

4+6+9 

4+6+9 

4+6+10 
4+6+11 
4+6+10' 
4+6+11 
4+6+9 

4+7+8 

4+7+10 

4+6+9 

4+6+10 

4+6+11 

4+6+9 

4+6+9 



238 15 
166 11 
208 11 
252 14 

250 17 

251 18 
237 15 
232 16 
262 16 
232 16 

148 10 
230 13 
228 13 
226 14 
155 11 
257 15 
225 14 



BREVIORA 

Museum of Comparative Zoology 

C'ambridgf,, Mass. X()vp:mber 4, IQBd Number 250 

THE HKAIX OF THE EMT 
DROMAEUS NOVAEHOLLANDIAE 

II. Anatomy of the Principal Nerve Cell Ganglia and 

Tracts! 

By Stan^ley Cobb 

INTRODUCTION 

This contribution is to supplement a jiaper published recently 
(Cobb and Edinger, 1962) on the gross anatomy of the brain 
and pineal body of the emu. Microscopic serial sections^ ^vere 
made of the whole of one of these brains (4^104), embedded 
in celloidin, and cut at 25 microns thickness. The sections were 
stained with cresyl violet for cell bodies and w4th Loyez modi- 
fication and Weigert's stain for myelinated nerve fibers. Pieces 
of the other brain (i^So) were cut for Golgi and Bielschowski 
stains, for axons and glia. The report here presented, however, 
is far from being a thorough description of the histology of 
the emu's brain; it is a description of the large neuronal divi- 
sions, the gray masses of nerve cells, and the conspicuous fiber 
tracts. This is to enable the reader to obtain a general picture 
of the distribution of the gray matter in ganglia, nuclei, and 
cortex, the relation of the main ganglia to each other, and their 
main connections by tracts. Such a picture may serve as a 
basis for comparing the general neuronal arrangement of the 
emu's brain with other avian brains. An attempt has been made 
to estimate the size of different ganglia, but because of shrinkage 
in fixation and embedding, the absolute volumes are difficult 
to measure. A comparison of the size of one ganglion to another 
in the same brain, however, is possible Avith reasonable accuracy. 
Such measurenients have significance in relation to function 
(Cobb, 1964). 

1 Fi-din tbe Museum of Comparative Zoology, and the Department of Neurologv 
and I's.vchiatry, Harvard University, and the Laboratory for Psychiatric Research. 
-Massachusetts General IU)spital. 

This investigation was aided by a grant from the National Institute of Neuro- 
logical Disease and Blindness. Grant #NB-0342!t-(ir>. 

I wish to thanl< Dr. Paul I. Yakovlev for his help in preparing the serial sec- 
tions and in photographing and naming the various structures shown. 



2 BREVIORA No. 250 

THE CERElUiAL VEXTRTCEES 

In order to understand the brain of any bird, one ninst get 
an nnderstanding- of how the ventricular spaces are arranged, 
because they have a special relation to the cortex and corticoid 
areas. The ventricles of the avian brain are unlike those of 
other vertebrates, and among birds vary in shape and size. The 
ventricular system is filled with fluid and extends from the 
olfactory bulb to the spinal cord. In much of its course it is 
narrow; in the cerebral hemispheres it is often only a potential 
space, moistening two layers of brain which lie one upon the 
other (see Fig. 9). 

In the olfactory bulbs the size of the ventricle varies greatly, 
from a tiny ependymal slit in the single fused bulb of small 
passerine birds, such as Passer (house sparrow), to a large 
open chamber in Diomeclea (albatross) which has big separate 
bulbs. In the emu the ventricle is distinct but not large (Figs. 
2, 3). The ventricle of the olfactory l)ulb is directly connected 
with the lateral ventricle of the cerebral hemisphere. This lateral 
ventricle lies between septum and striatum with its long dimen- 
sion vertical (Figs. 4, 7). As it spreads backward it becomes 
larger, and as the mid-sections of the brain are reached the 
ventricle is seen to extend more and more laterally over the 
striatum (Figs. 8, 9). Here it lies under a sheet of tissue com- 
posed anteriorly of dorsal hippocampus and parahippocampus 
(Fig. 7), and farther back of corticoid tissue of three kinds: 
parahippocampal, dorsolateral and periamygdalar (Figs. 8, 10). 
Thus, over approximately the caudal third of the emu's cerebral 
hemispheres, the A'entricle is a narrow, potential space separat- 
ing a sheet of supraventricular tissue from the main mass of 
the underlying ganglia. The relation to the nucleus diffusus, in- 
wulst (Fig. 4) is important. In the emu the wulst is frontally 
placed so the ventricles remain largely medial and vertical 
initil the wulst becomes smaller caudally and the vallecula flat- 
tens out (Figs. 4, 5). Farther back, the ventricle spreads widely 
to each side and around to the base, making a complete covering 
of the occipital pole of the hemisphere (Figs. 9, 10). 

The two lateral ventricles join at the liase of the hemisphere, 
forming the third ventricle. This is continuous with the single 
ventricular space of the midbrain (aqueduct) which is large 
with lateral out-pocketings into the big optic lobes (Fig. 11). 
Much of these lateral ventricles of the midbrain appears to be 



1!)66 THE BRAIN OF THE EiSIU 3 

merely potential spaces, but at tlieir mesial portions, where 
they join the aqueduct, the lumen is distinctly patent. Below 
the level of the midbrain the fourth A-entricle is seen, wide be- 
neath the cerebellum and narrowinp: down to liecome covered 
with nervous tissue in the medulla near the s]iinal cord. 

ANATOMY OF THE FOREBRATN 

In Itii'ds, the main mass of the forebrain is composed of lar^re 
o'ray fi'an<>-lia (the "striatum") with a relatively small corticoid 
part, and variable, but usually small olfactory bulbs. Strictly 
speaking-, the striatum of higher mammals is composed of the 
more dorsal cerebral gray ganglia traversed by bands of Avhite 
fibers. Many of these run to and from the neocortex, making the 
striped appearance that gave the area its name. Such a combina- 
tion of gray and white matter develops of necessity wherever 
there are gray ganglia forming axonal connections with each 
other. The naming of these ganglia in mammals and birds, as 
if they were homologous structures, has led to no little mis- 
understanding (Table 1). Rose (1914) was wise enough to sim- 
ply name his ganglia A, B, C, etc., and Kuhlenbeck (1938), 
relying on embryological evidence, developed a nomenclature 
for the cerebral ganglia of birds that largely disregarded mam- 
amalian analogies. His work has been substantiated and effec- 
tively used by Jones and Levi-Montalcini (1958). Kuhlenbeck 's 
investigations show that the ganglia (he calls them "nuclei") 
of the avian cerebral hemisphere arise from four embryonic 
zones, dorsomedial and A^entromedial (the larger olfactory part) 
and dorsolateral and ventrolateral (the striatal nuclei) ; the lat- 
ter two form the main masses of the cerel)ral hemisphere. A 
conspicuous boundary, the lamima medullaris dorsalis (Fig. 6) 
separates the striatal nuclei into two divisions, the epibasal 
nuclei and the basal. This seems to be a basis for a more rational 
nomenclature than the one built up by Kappers ft al. (1936) 
on the comparative neurology of living reptiles and birds. This 
nomenclature is. therefore, used in this paper in describing 
the forebrain. Karten and Hodos (1966) in their recent atlas 
of the pigeon 's l)rain use a modified Kappers nomenclature with 
abbreviations, as shown in my Table I. In mentioning different 
parts of the striatum, 1 use Kuhlenbeck 's names and put 
Karten and Hodos' abbreviations in parentheses; for example: 
nucleus epibasalis caudalis (A) is the archistriatum. (See Table 
I and Fig. 1, and the list of abbreviations.) 



BREVIORA 



No. 250 






Ml M 

Q CS O 

H oQ 1;^ 

M W ^ 

^ ^ ^ 

00 ^ 



w 

M S > 
^ rH H 



^ tK r-^ 



<1 

o 

> 

O 





cv, 


'^ 


CI 


^ 


c. 


03 


T-H 


W 


^ 


H 


PQ 


m 


ac 


P 


O 


w 


O 



_— ^ (^ M r-^ 



o t* 



>^ K 



o 

K 






C a:' 



J^ K 



o 

o 






1066 



THE BRAIN OP THE EMU 



Figvn-e 1 is drawn from a reconstruction of tlie foreln-ain 
of the emu, the main ganglia or nnclei being shown in seven 
sections at the levels indicated l)y the vertical lines drawn on 
the hiteral view of the whole brain (insert). The wulst or 
nucleus diffusus (HxV, HD) is seen in five sections; beginning 
with the most anterior it is the largest; in the second and third 
it is large and then tapers off in the fourth and fifth. The largest 
nucleus of all, the nucleus epibasalis centralis (N and E), ap- 
pears as a small part of the third section and enlarges caudally 
until it is the only ganglion present in the most caudal section. 
()ther main nuclei can be similarly traced to show their antero- 
posterior r('lati()ushij)s. Tlic relation of one nucleus to another 
in any one section can be seen in the photographic reproduction 



LAMINA FRONTALIS SUPREMA- 
LAMINA FRONTALIS SUPERIOR- 



LAMINA 
HYPERSTRIAT 




PARAHIPPOCAMPAL 
CORTICOID 
AREAS 



VENTRICLE 



Fig. lA. A diagrammatic representation of the dorsoventral relations of 
the principal forebrain ganglia, but no one section could show all of these. 



BREVIOBA 



No. 250 



of the sections that represent various levels (Figs. 2 to 11). The 
nucleus iutercalatus is found in most birds. In the emu it is a 
rather inconspicuous part of the wulst lying between the fibers 
of the lamina frontalis (Fig. 4). It is composed of small nerve 
cells and is more compact than the rest of the nucleus diffusus 
of which it is a part (if the nucleus diffusus is defined, as in 
this paper, as that part of the gray mass lying within or above 
the lamina frontalis). 

The division between wulst (nucleus ditfusus) (HA, IID) 
and nucleus epibasalis dorsalis (HV) is quite clear in the emu, 
the lamina frontalis being well developed, but in my specimen 
it is not split into two parts, superior and suprema. Therefore, 
one cannot make out a distinct nucleus lying between the parts 




Fig. IB. A simplified diagram of the principal forebrain ganglia showing 
perspective, to give some idea of the anteroposterior relationships. The 
serial sections from which the diagrams were draw'n are (from front to back) 
#161, #361, #501, #701, #901, #1201, #1281. Their position on the 
whole brain is shown by vertical lines on the small inset. 



1966 THE BRAIN OF THE EMU 7 

of the lamina as described in some birds' brains. Rather, one 
sees a branching" of the lamina frontalis with man,v cells lying- in 
the interstices. Some of these are small and might be said to 
form a group which could be called nucleus intercalatus (Figs. 
6, 7). This is not clear in sections farther forward (Fig. 4). 
In fact, no l)Oundaries would seem to be definite enough to 
justify describing a distinct nucleus; in short, I do not find in 
emu a nucleus epibasalis dorsalis, pars superior (HD). Below 
the lamina frontalis the nucleus epibasalis dorsalis (HY) is well 
defined and bounded ventrally by the lamina hyperstriatica 
(Figs. 4, 6). 

CORTEX AND CORTICOID AREAS 

The question of the avian cerebral cortex (pallium) is com- 
plex and controversial. Craigie meticulously described the corti- 
coid areas of the emu (1935 A, B). In the next j'ear Kappers, 
Huber and Crosby (1936) modified Craigie 's findings to some 
extent. Bures, Fifkova and Marsala (1960) have made a good 
summary of the "cortex question," but, unfortunately, their 
nomenclature differs from others. I am using the names of 
corticoid areas and hippocampus as used by Kappers, Huber 
and Crosby (1936) because they seem simpler and freer from 
implied homologies to mammalian brains. 

In the emu there is a well developed archicortex, the hippo- 
campus, with three or four obvious layers. These become less 
distinct in the dorsal hippocampus (Fig. 7), and lamination 
cannot be seen clearly at all in the parahippocampal corticoid 
areas which have a thin single or sometimes double layer of 
cells lying over the surface of the wulst. Further laterally, the 
dorsolateral corticoid area is even thinner and less distinct (Fig. 
8). As one goes towards the posteroventral part of the hemis- 
phere, a definite corticoid area with two layers of cells is seen 
over the nucleus epibasalis eaudalis (A). This is called the 
periamygdaloid corticoid area (Fig. 8). Also ventral but far 
forward is seen another conspicuous corticoid area, the praepyri- 
form. It lies about the root of the olfactory bulb where it joins 
the frontal pole of the hemisphere (Fig. 3). 

The relationship of these corticoid areas to the cerebral cortex 
of mammals is not well understood. The fiber connections of the 
parahippocampal areas suggest a function analogous to that of 
neocortex in mammals, but these areas are surelv not homologous 



8 BREVIORA No. 250 

witli mammalian neocortex; their homology to the "general cor- 
tex" of reptiles is more clear. The area described by Goldby 
and Gamble (1957) as "dorsal cortex" seems to correspond witli 
what is called here "parahip}iocam])al cortieoid," and called 
"general cortex" by Kappers, Hnl)er and Crosby (1986). A 
thickening found in this "dorsal cortex," which overlies the 
anterior dorsal part of the striatum, occupies a position in the 
pallium from which, in flic ciiihryo, cortex is seen to develop. 
In such a topographical sense only can this cortex be said to be 
homologous with mammalian neocortex. 

There is better evidence that the periamygdalar cortieoid areas 
are homologous with the temporal mesocortex of mammals, and 
that the praepyriform cortieoid areas are homologous with the 
entorhinal areas of mannnals. One tiling seems sure, and that 
is that birds (including the emu) have no neocortex. For a 
scholarly and thorough discussion of this vexing question, the 
reader is referred to Kuhlenbeck's recent book (196()). 

COMPARATIVE :MEASUREMENTS OP GANGLIA IN THE 

FORE BRAIN 

Believing that increase of any function in an animal is posi- 
tively correlated Avith increase in size of the parts of the brain 
significantly involved, it is of interest to compare the sizes of 
the different i)arts of the brains of birds of various families and 
varied types of behavior. This has been done in relation to the 
auditory nuclei (Winter, 1963), the olfactory bulbs (Cobb, 1960). 
auditory and visual centers of the midbrain (Cobb, 1964), and 
ganglia of the forebrain (Fritz, 1949). Since we know so little 
about the function of the diff'erent ganglia of the forebrain, it 
is difficult to choose which ganglia to measure. Fritz showed 
differences in size of the ganglia he measured, ])ut no correlations 
with any functions. He chose to measure the volume of "hyper- 
striatum" (n. ditt'usus plus n. epibasalis dorsalis), "neostria- 
tum" (n. epibasalis centralis), "archistriatum" (n. epibasalis 
caudalis), and "paleostriatum" (n. basalis plus n. entopeduncu- 
laris). The boundary between the neostriatum and archistriatum 
is not always clear, as pointed out by Fritz. 

I decided to measure: 1) n. diffusus, both ])arts ])lus ii. 
intercalatus (i.e. all the ganglionic mass above the lamina 
frontalis superior) ; 2) the n. epibasalis dorsalis (HV) ; 3) the 
n. epibasalis centralis (N, E). Tlie first was chosen because 
it seemed, grossly, to vary so much from species to s])ecies, the 



I 



1 ;)()() THE BRAIX OF THE EMU 9 

other two becuuse they are large forebrain g-anolia and are 
possibly associative in function and hence related to "liigher 
integrations'" and "intelligent" behavior (Table II). 

In the brainstem tlie n. isthnii, n. mesencephalicus, and tectum 
opticnm "were chosen because of their relation to liearing and 
vision, as described in a previous paper (Col)b, 1964), and l)y 
Karten (19()6). The method of estinuiting the volume of each 
of these ganglia and of the whole hemisphere, with which they 
are compared, has been described previously (Gobi), 1964). 
Briefly, it is to project and draw the structures to scale, and, 
by knowing tlie thickness of the sections and their distances 
apart and number, to compute the volume. Absolute values for 
volumes of different structures are not reliable 1)ecause of dif- 
ferences in shriidvage from one brain to another, but relative 
measurements in any one series of slides made from one brain 
are significant. 

All attcmi)t was made to measure the areas of the hippocampus 
and tlic corticoid structures, but because of their long and nar- 
row sliapes, it was difficult to use tlie i^lanimeter to measure 
their areas. Photographs were tlierefore made to scale and the 
desired areas were cut out of each print and weighed. By this 
method (Fritz, 1949), it was estimated that the n. diff'usus 
(HA and HD) had a vohnne 28 per cent as large as the hemis- 
phere, and that the hippocampus plus the corticoid areas had a 
volume 1:5 ])er cent of the size of the hemisi)here. By the projec- 
tion ])lanime1er method (Cobb, 1964) tli(> n. diffusus (HA and 
111)) was also found to be 28 i)er cent of tlie hemisphere. For 
these (juantilative data on the (^nu brain see Table II. 

Table II 
^'oLI'^iES OF Certain Parts of Emi' Brain 

Voliinie of one ceiebial hemisphere (before fixation) approximately 6.3 cc 

percentage of liemisphere 
Nucleus Diffusus (HA & HD) 28 

X. ?]pil)asalis Dorsalis (HV) 11 

X. Epiljasalis Centralis (X & E)^ 32 

A'olume of one optic lobe (before fixation ) approximately 0.5 cc 

percentage of optic lohe 
X. ]\lesencephalicus Lateralis .j.-i 

X"^. Istlimi Magnocellularis 1.9 

X. Isthmi Parvocellularis 2.5 

Tectum Opticum 81.0 

1 As (Ictiiii'd ill sccrioii mi alibrcviations ami iiiiiiiciiclaturc and Fig. 1. 



ID BREVIORA No. 250 

As to the compactness of the emu's brain, it is oln'ions by 
microscopic inspection that there are fewer nerve cell bodies 
(perikarya) in a given area than in the brains of smaller birds. 
Comparison was made by counting the number of cells in 0.0143 
of a square millimeter in microscopic slides of brain tissue of 
the emu and of a hummingbird {ScIaspJwrus platycercns) . 
Knowing the thickness of the section, the number of cells per 
cubic millimeter was computed for stratum E of the optic tec- 
tum of the midbrain, and for the midposterior part of the n. 
epil)asalis centralis (N). In each of these cases the emu was 
found to have less than half as many cells per cubic millimeter 
as the hummingbird (see Table III). Purkinje cells in the 
cerebellum were also counted and measured in these tAvo birds, 
and the ratio of 1 to 2 was also approximated. Because of the 
linear arrangement of Purkinje cells along the edge of a stratum, 
cells per volume could not be counted, but cells per linear micron 
of the layer were easily tigured. One should not lay too much 
claim to accuracy in these determinations of cells per given 
volume, because in the process of fixing and embedding in 
celloidin, there is much shrinkage of the brain tissue, and the 
amount of shrinkage in two separately prejiared series of brain 
sections cannot l)e accurately estimated. In my series of brains 
it seems to be about 20 or 23 per cent of the volume of the fresh 
brain. The figures are given in tabular form, with emphasis on 
the fact that they are only approximate (Table III). 

Table 111 

Estimates of Compactness of Emt Brain 

Compared to Hummingbird 

Bromaeiu^. ^plasphorus 

novaehoUandiar platycercns 

Number of cells per mm^ of n. epi- 42,800 125,000 

basalis eeiitralis (N). 

Number of cells per mm3 of stratum 40,000 100,000 

E of tectum 

Purkinje cells per 100 linear niicra 2.2 4.8 

Average diameter of Purkinje cells 26.0 micra 12.0 micra 



106C THE BRAIN OF THE EMU 11 

GKAY NUCLEI AND LAMINAE OF MIDBRAIN AND 
HINDBRAIN 

Figures 8 to 12 show the main gray masses of the midl)rain 
and hindbrain; these masses are eitlier nuclei, large and small, 
or laminated structures such as the nerve cell layers of the tec- 
tum opticum and cerebellum. This laminate pattern has neurons 
oriented in three planes, the axons usually parallel and the 
dendrites and collaterals at right angles, making a criss-cross 
arrangement in which layers of cells are more or less clearly 
separated by layers of fibers. Such an arrangement of neurons 
probably is more plastic in function than the arrangement in 
nuclear masses, allowing more rapid and efficient adaptations to 
environmental change (Yakovlev, 1952). The acme of such de- 
velopment is seen in the mammalian cerebral neocortex. In birds, 
where vision is of such great importance, it is remarkable that a 
visual cortex has developed in the tectum of the midbrain, l)ut 
none in the forebrain. 

The cerebellum, in the hindbrain, is an example of an organ 
with laminated cortex, subserving rapid coordination in wide- 
spread neuronal integrations. The three-layered arrangement of 
neurons is uniform throughout the wide extent of this cortex 
(Fig. 12). 

Sections were made of the pineal body, l)ut were stained only 
witli hematoxalin and eosin, so no refinements of histology could 
be identified. However, one could see the structures described by 
Quay (1965) : a sac-like formation, tubules and follicles, and a 
mass of cellular parenchyma. In two areas there were con- 
spicuous lymphoid nodules. In general, the histology of the 
pineal body is more like that of the Galliformes than the 
Passeriformes. 

SUMMARY 

The emu, like the ostrich, is an enormous running bird and 
cannot fly. This specimen weighed 31 kilograms. The brain of 
the emu is one of the largest avian brains, and less compact than 
the brains of smaller birds. The forebrain is well developed 
with medium sized hippocampus and parahippocami^al corticoid 
areas. The size of these areas is probably related to the medium 
size of the olfactory bul])s. The gray nuclei that make up the fore- 
brain are much like those of Gallus in arrangement and relative 
size, except for the nucleus diffusus (wulst or hyperstriatum 



12 BREVIORA No. 250 

aecessorium) Avliieh is relatively larger and is bounded below by 
a loosely arranged lamina frontalis. This eannot be divided into 
two parts, superior and suprenia. The nucleus interealatus is 
represented l)y a group of small eells in the lateral part of the 
area crossed ])y the sparse upper fibers of the lamina frontalis. 
Little is known about the function of the wulst. its connections 
with the optic tectum of the midbrain indicate that it has some- 
thing to do with vision, and the large size of the enni's eye sug- 
gests a reason for the big wulst. 

The midbrain is remarkable for its large optic tectum with 
seven definite layers, but this visual cortex is conspicuous in 
all birds, the emu rating above the average of most birds in 
relative size of tectum, but not as high as the Falconidae. In the 
hindbrain there is nothing remarkable ; the cerebellum is well 
developed but not relatively as large as in many bii'ds with skill 
in flying. 

REFERENCES 

BuRES, J., E. FiFKOVA and J. ^Iarsala 

1960. Leao 's spread in depression in pigeons. J. Conip. Neurol., 114:1. 

COBH, S. 

ll»(3ll. A note on tlie size of the avian olfactory bnlb. Epilepsia, l:H;i4- 

W2. 
I!)(i4. A comparison of the size of an auditory nucleus ( n. nieseuce- 
phalicus lateralis, ]iars dorsalis) with the size of the optic lobe 
in 27 species of l)irds. J. Comp. Neurol., 122: 271-280. 
COBB, S. and T. Edinger 

]9fi2. The brain of the emu (Droniaciis itovachoUcuKUac, Lath) 1. 
Gross anatomj' of the Inain and pineal Imdy. Breviora, Mus. 
Comp. Zool. No. 170: 1 IS. 
Craig IE, E. H. 

IDS-la. The hippoeampal and imrahippocampal cortex of the emu. J. 

C'omp. Neurol., 61: -IGo. 
I!t3-")b. The cerebral hemispheres of the kiwi and emu. J. Anat. 69: 
SSO-SO.S. 
Fritz, \\^ 

]!>4!». Yergleichende Studien iiber den Antiel von Striatumteilen am 
Hemisphiirenvolumen des Vogelliirns. Rev. Suisse Zool., 56: -tlil- 
4<tl. 
GoLDBY, F. and H. J. Gamble 

19.'i7. The reptilian cerebral hemispheres. Biol. Rev., 32: .'583-420. 
Huber, C. G. and E. C. Crosby 

1!)2!). The avian diencephalon. J. Comp. Neurol., 48: 1-177. 



1966 THE BRAIN OF THE EMU 13 

Jones, A. E. and E. Levi-Montalcini 

1958. Patterns of differentiation of the nerve centers and fiber tracts 
in the avian cerebral hemispheres. Arch. Ital. Biol., 96: 231-281. 
Kappers, C. U. a., G. C. Huber and E. C. Crosby 

1936. The comparative anatomy of the nervous system of vertebrates, 
including man. New York, MacMillan, 2 vols. 
Kaktex, H. J. and W. Hodos 

1966. Stereotaxic atlas of the pigeon 's brain. (In press). 

KUHLENBECK, H. 

1938. The ontogenetic development and phylogenetic significance of 
the cortex telencephali in the chick. J. Comp. Xeurol., 69: 273- 
295. 
1966. Comparative anatomy of the vertebrate brain. (In press). 
Quay, W. B. 

1965. Histological structure and cytology of the pineal organ in birds 
and mammals. Progress in Brain Eesearch, 10: 49-86. 
Rose, M. 

1914. Ueber die cytoarchitektonische Gliederung des Yorderhirns der 
Yogel. J. Psychol. Neurol., 21: 278-352. 
Smith, G. E. 

1919. A preliminary note on the morphology of the corpus striatum 
and the origin of the neopallium. J. Anat., 53: 271-291. 
Winter, P. 

1963. Yergleichende qualitative und quantitative Uutersuchungen an 
der Horbahn von Yogeln. Z. Morphol. Okol. Tiere, 52: 365-400. 
Yakovlev, p. I. 

1952. Patterns of neuronal assemblies and the anatomical substratum 
of seizures. Epilepsia, (Ser. 3)1: 51-68. 



ABBREYIATIONS AND NOMENCLATURE 

USED IN THE FIGURES 

(See also Table I) 

A. SEPTAL. Area septalis 

ANT. COM. Anterior commissure, interconnects n. epi. cauda- 

les of the two hemispheres 
ANT. MED. YEL. Anterior medullary velum merges anteriorly with 

tectal commissure, which is the dorsal part of 

post. com. 
CBL Cerebellum. Its cortex has 4 layers: 

1. molecular 

2. Purkiuje cells 

3. granular 

4. medullary 

CENTR. GRAY Central gray matter around ventricle 



14 



BREVIORA 



No. 250 



CHORIOID 

DORS. LAT. CORTI- 

COID OR DORSOLAT 

CORTICOID 
FASC. LONG. POST. 
HABENULA 
HEM. 

HIPPOCAMP. 
INNER GRAN. 
LAM. FRONT. 

LAM. HYPBRSTRIAT. 
LAM. MEDUL. DORS. 

OR LAM. MED. DORS. 
LAT. F. B. BUNDLE 

LAT. LEM. 

LAT. LEM., D.P. 

LAT. LEM., V.P. 

LAT. VENT. 

LAT. VENT. M.B. 

N. BASALIS 



N. CEREBEL. LAT. 
N. DIFF. DORS. 

N. DORSOLAT. 
N. DORSOMED. 
N. ENTOPED. 

N. EPI. CAUDALIS 

N. EPI. CENTRALIS 

N. EPI. DORSALIS 



N. GENIC. 

N. HYPOTHAL. 



Plexus choiioideus of the third ventricle 
Dorsolateral superficial corticoid area, anteriorly 
on surface of wulst, posteriorly outside the ven- 
tricle 

Fasciculus longitudinalis posterior 
An olfactory way station of eijitlialanins 
Cerebral hemisphere 
Hii3i)ocampus or cortex ammonalis 
Inner granular layer of cells. 

Lamina frontalis. Sometimes separated into su- 
perioris and suprema 
Lamina hyperstriatica 
Lamina meduUaris dorsalis 

Lateral forebrain bundle, contains tracts to thala- 
mus, midbrain, and hindbrain 

Lateral lemniscus, lies in fibrous capsule of n. 
mesen. lat. 
Dorsal peduncle 
Ventral peduncle 

Lateral ventricle of the cerebral hemisphere 
Lateral out-pocketing of the midl)rain ventricle 
beneath the tectum 
Medulla oblongata 

Paleostriatum augmentatum, the nucleus separa- 
ted from neostriatum al>ove and laterally Ijy the 
lamina medullaris dorsalis; PA (Karten) ; Field 
H (Rose) 

Nucleus cerebellaris lateralis 

Nucleus diffusus dorsalis or hyperstriatum acces- 
sorium, HA (Karten), Field B (Rose) 
Dorsolateral nucleus of the thalamus 
Dorsomedial nucleus of the thalamus 
Nucleus entopeduncularis, or paleostriatum primi- 
tivum; PP (Karten) ; Field J (Rose) 
Nucleus epibasalis centralis or archistriatum 
(amygdala) ; A (Karten) ; Field K (Rose) 
Nucleus epibasalis centralis or neostriatum, the 
ganglion just below lamina hyperstriatica, N and 
E (Karten) ; Field G (Rose) 

Nucleus epibasalis dorsalis pars inferior or hy- 
perstriatum ventrale, the ganglion lying just Ije- 
low the lamina frontalis; IIV (Karten) ; Field D 
(Rose) 

Nucleus geniculatus lateralis 
Nucleus hypothalamicus 



1966 



THE BRAIX OF THE EMU 



15 



N. IXTEECAL. 



X. IXTERPED. 
X. ISTHMI 

X. :\r]-:8EX. lat. 

X. :\rESEX. PROF. 

X. XERV. ABDUC. 

X. OCULOMOT. 

X. OLIV. SUP. 

X. OVOID. 

X. PARAYEXT. 

X. PRETECT. 

X. ROTUXDUS 

X. RUBER 

X. SPIRIFORM. 

X. 8UBPRETECT. 

X. YESTIB. LAT. 

X. VESTIB. VENTEO- 

LAT. 
OLE. 

OLE. A^EXT. 
OP. 

op. chiasm 
oi^ter grax. 
parahip. corticoii) 
peria:\iyg. corti- 

COID 

POXT. GRAY. 
POST. COM. OR P. 
COM. 

PREPYR. CORTICOII) 
STRAT. FIBR. 
SUPRAOPTIC. 
TECTUM OP. 



TR. FROXTO- 
ARCHISTRIAT. 



Nucleus iuterealatus hyperstriaticus ; HIS (Kar- 
tell) ; Field A (Rose), of nucleus diffusus dorso- 
lateralis 

Xucleus interpeduiicularis 
Nucleus isthmi, pars magnocellularis 
Xucleus meseueephalicus lateralis, pars dorsalis, 
also called torus semicircularis 
Nucleus mesencephalicus profundus or tegniento- 
pedunculopontinus 
Nucleus uervus abducentis 
Nucleus oculomotorius 
Nucleus olivarius sux)erior 
X'ncleus ovoidalis 

Nucleus paraventricularis magnocellularis 
X'ucleus pretectalis 
Nucleus rotundus 
Nucleus ruber or red nucleus 
Xucleus spirif oruiis 
Xucleus sul)i)retectalis 

X'ucleus vestilnilaris lateralis, Deiter's nucleus 
X'ucleus vestibularis ventrolateralis 

Olfactory bulb 
Ventricle of olfactory l)nll) 
Optic lobe 
Optic chiasm 

Outer granular layer of cells 
Parahippocampal corticoid areas 
Periamygdalar corticoid area, contiguous Avitli 
the parahippocampal areas; not outside the ven- 
tricle 

Pontine gray nuclei 

Posterior commissure, from tectum to opposite 
tectum and also connects spiriform :\ud dorsal 
mesencephalic nuclei 
Praepyriform corticoid area 
Stratum fibrosum 
Dorsal supraoptic decussation 

Tectum opticum of the midbrain. Divided into 
six layers or strata (Huber & Crosby 1924) 

A. Stratum opticum 

B. Stratum fibrosum et griseum superficiale 

C. Stratum album centrale 

D. Stratum griseum centrale 

E. Stratum griseum periventricnhirc 

F. Stratum fibrosum periventricuhire 
Tractus fronto-archistriaticus 



16 BREVIORA No. 250 

TB. OCCIP.-MESEX. Tractus occipito-mesencephalicus 

TR. OP. MARG. Tractus opticus marginalis, lamina A of the op 

tic tectum. Input from optic nerve to tectum 

TR. SEPTOMESEN. Tractus septomesencephalicus 

TR. THAL-RTRIAT. Tractus thalamostriatalis 

V. OR VAL. Vallecula 

VENT. M.B. Midbrain ventricle or aqueduct of Sylvius 

W. Wulst 



1966 



TPIE BRAIN OF THE EMU 



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CEREBELLUM 



MOLECULAR LAYER 
PURKINJE CELLS 
GRANULAR LAYER 

MEDULLARY LAYER 



N. CEREBEL. LAT 



N VESTIB LAT 



FASC. LONG. POST 
N. OLIV. SUP 
N. NERV ABDUC. 



PONTINE GRAY 
Fig. VI. Serial section #Hi'Sl. Ciesyl Violet stain, magnification X 5.0. 



BREVIORA 

Musemim of Comparsitive Zoology 

Cambridge, Mass. Xovembek 4, 1966 Number 251 

CHRONOLOGICAL SURVEY OF THE 
TETRAPOD-BEARING TRIASSIC OF ARGENTINA 

By J. F. Bonaparte^ 

SUMMARY OF PREVIOUS WORK 

Ever since 1958, Argentinian paleontologists have been work- 
ing intensively on continental Triassic outcrops bearing tetrapod 
remains, inspired by A. S. Romer's successful expedition to 
Mendoza and San Juan provinces in that year. The resultant 
collections are from both new and previously known localities ; 
they come mainly from the Ischigualasto Formation, worked on 
by Romer, but also include material from a number of other 
formations. Exploring and collecting trips to La Rioja, San 
Juan, San Luis, Mendoza, and Santa Cruz provinces were made 
possible by many grants from the Consejo Nacional de Investi- 
gaciones Cientificas y Tecnieas. This support has enabled us not 
only to make good collections from previously known fossil-bear- 
ing localities, but also to discover new fossiliferous areas, which 
have broadened our view of Argentinian Triassic faunas. Ma- 
terials collected in recent years include fossils from three locali- 
ties in the Ischigualasto Formation, from the Los Colorados 
Formation of La Rioja and San Juan provinces, from the Ca- 
cheuta Formation in northern ^Mendoza Province, and, in addi- 
tion, from a neAv locality in the Puesto Viejo Formation south 
of San Rafael, Mendoza Province, discovered by the former 
Direccion Nacional de Geologia y ^lineria. A great proportion 
of the tetrapod material collected has been studied, and it is 
possible to identify the remaining material with satisfactory 
approximation ; these studies enable us to attempt an interpreta- 
tion of the chronology of some of the Argentinian Triassic 
formations. 



1 Fundacion Miguel Lillo, Univevsidad Nacional de Tucuman; and Consejo Nacional 
de Investigaciones Cientificas y Tecnieas, Buenos Aires, Republica Argentina. 



2 BREVIORA No. 251 

The chronology of these Triassic series has, in the past, been 
interpreted largely on the basis of their paleobotany, which was 
well summarized by Bonetti (1963). These interpretations, un- 
fortunately, are not in complete agreement with those ol)tained 
from a study of tetrapod remains ; the paleobotanic data indicate 
little differentiation from one formation to another, while the 
tetrapod data suggest clear differences, and allow^ a better 
analysis of chronology. 

Bonetti 's work is the most recent on the subject. She accepts 
the opinion of Groeber and Stipanicic, 1953, and Stipanicic, 
1957, that the known Argentinian Triassic series with Dicroidium 
flora are (when present) in discordance over rocks synchronic 
with those known as Choiyoilitense from southern Mendoza and 
northern Neuquen provinces. The correlation proposed by these 
authors, between the Argentinian Choiyoilitense and the Porfidic- 
queratofiric series of the Chilean coast south of Los Vilos, is also 
accepted by Bonetti. She concludes that the Argentinian Trias- 
sic series with Dicroidium flora are essentially of Keuper age 
(Carnian at the lower levels, Norian at the upper). This conclu- 
sion is based on the presence of an ammonite and pelecypod fauna 
in sediments underlying the effusive rocks near Los Vilos, as well 
as on the affinities of a flora found in a lutitic level intercalated in 
the upper part of the Chilean effusive complex with the Argen- 
tinian Triassic flora from the Potrerillos, Cacheuta, Barreal, Cor- 
taderita, etc., formations. In her clear and brief analysis, how- 
ever, Bonetti points out the need for re-examination of the 
determinations of most of the marine fauna below the Los Vilos 
eft'usives. Quite apparently, the chronology of the Chilean effu- 
sive series is far from definitely clear. 

In recent years some attempts at Triassic chronology have been 
based on tetrapod remains. Owing to the presence of Chirothe- 
rium footprints studied by Rusconi (1951b) and reconsidered by 
Peabody (1955), Stipanicic (1957) considered that the Sierra 
de las Higueras Formation in northern Mendoza was the oldest 
Triassic formation of Argentina, and referred it to the Neo- 
seythian — Eoanisian. Homer (1960, 1962b) believes that the age 
of the Ischigualasto Formation is upper Middle Triassic : "surely 
pre-Norian and not improbably pre-Carnian." Reig (1961) con- 
sidered briefly the age of four Triassic formations. He believes 
that the Cacheuta Formation, with both brachyopids and a 
proterosuchian, is more reasonably assigned to the "Eotriasico 
Superior" (Scythian) than to any later time. As to the age of 
the Sierra de las Higueras Formation as given by Stipanicic, 



1966 



TETRAPOD-BEARIXG TRIASSIC OF ARGENTINA 



Reig points out the limited significance of the footprint data. 
As regards the Los Rastros and Ischigualasto formations, after 
some analysis and comparisons, he assigns an upper Ladinian 
age to the Ischigualasto fauna. 





Stipanicic 
1957 


Bonetti 
1963 


Reig 
1961 , 1963 


Romer 
I960 , 1962 


NORIAN 


Los Colorados 


Los Colorados 
Ischigualasto 






Ischigualasto 
Los Rastros 


CARNIAN 


Ischichuca 


Los Rastros 
Ischichuca 






LADINIAN 






Ischigualasto 


Ischigualasto 
? 







Table 1. Recent chronological interpretations of some Triassic forma- 
tions of Argentina. 



ANALYSIS OF VERTEBRATE FAUNAS 

Puesto Vie jo Formation (Mendoza Frovince). — The fauna of 
these red sandstones comprises, so far as is now known, only 
therapsids of the infraorders Cynodontia and Dicynodontia. 
The cynodont Fascualgnathns polanshii Bonaparte (in press. b), 
known from very complete remains, is a diademodontid. It has 
affinities of great significance with Trii'achodon, and of lesser sig- 
nificance with Diademodon. Among these points of similarity 
are important characters of the skull and jaws, the tooth row, 
the secondary jDalate, and so forth ; in the axial region of the 
postcranial skeleton we also find good characters for the family 
assignation. Pascualgnathus is closer to African genera than 
to other cynodonts from South America or from other conti- 
nents; it represents, therefore, an assemblage different from 
that common in South America, and provides new evidence of 
some type of vinculum between Africa and South America not 
detected in the case of other tetrapods from the South American 
Triassic. 

Recent work on Diademodon rhodesiensis Brink (1963) and 
on a Trirachodon-like genus (Brink, 1963; Kitching, 1963) 
demonstrates that these genera survived beyond the end of the 



4 BREVIORA No. 251 

Lower Triassic. They have been found in the Ntawere Forma- 
tion, Avhich immediately follows the Cynog^iothus Zone and is 
thus of probable Anisian age. As Pascualgnathus is but little 
advanced anatomically beyond Trirachodon and Diademodon, 
I have considered this genus to be also Anisian (Bonaparte, 
in press.b,c). 

The dicynodont of this formation has been referred to the 
African genus Kannemeyerw., as K. argentinensis Bonaparte 
(in press. b), because of the great similarity between its skull and 
that of Kannemeyeria sp. (Watson, 1948, fig. 17) from the 
Cynognathus Zone. {Kannemeyeria argentinensis is a smaller 
species than the African representatives of the genus, so that 
postcranial similarities are less noticeable.) There are clear 
generic differences between K. argentinensis and other kanne- 
meyerid genera. Until recent years, Kannemeyeria was unknown 
outside of the Cynognathus Zone, but a form referable to Kanne- 
meycrisi is cited by Kitching (1963) from the Ntawere Forma- 
tion, and a Kannemeyeria from the Manda Formation of the 
Ruhuhu Valley in Tanganyika is reported by Cox (1965) and 
Brink (1963). It is thus apparent that Kannemeyeria survived 
into the Anisian ; we can, consequently, consider K. argentinen- 
sis to be either Scythian or Anisian in age. The presence of 
Pascualgnathus, however, leads us to consider the therapsids of 
the Puesto Viejo Formation as Anisian. 

PotrerilJos Formation (Mendoza. Province). ■ — -Fragmentary 
remains of a cynodont jaw constitute the only available tetrapod 
material from this formation. ]\Iinoprio (1954) studied these 
remains, and erected a new genus, Colhertosaurus, which he 
placed in the Ictidosauria. However, the principal anatomical 
feature used by Minoprio in his identification of Colhertosaurus 
as an ictidosaurian — the placement of the posterior part of the 
tooth row — is a common feature among the gomphodont cyno- 
donts from the African ]\Iiddle Triassic and the South American 
Middle or Upper Triassic ; Colhertosaurus is thus more reason- 
ably considered a gomphodont cynodont than an ictidosaurian 
(see Bonaparte, 1962). Colhertosaurus can best be compared 
with Pascualgnathus, from the Puesto Viejo Formation. The 
two have many features in common : the type of postcanine im- 
plantation, the number and size of the incisors, the large section 
of canines, the larger size of the postcanines in the middle of 
the tooth row, and the type of jaw constriction behind the 
canines. Pascualgnathus has eleven postcanines and Colherto- 
saurus nine. 



I 



1966 TETRAPOD-BEARING TRIASSIC OF ARGENTINA 5 

It is evident that there is a significant, although limited, re- 
semblance between the tAvo genera. The Potrerillos Formation 
may therefore be considered, at least on the basis of present 
knowledge, as very close in age to the Anisian Puesto Viejo. 

Cachciita Formation (Mcndoza Province). — From this for- 
mation we know two Ijrachyopid labyrinthodonts, apparently 
representing two separate genera — Pelorocephalus Cabrera 
(1944) and Chigutisaurus Rusconi (1948). We also have the 
postcranial skeleton of a proterosuchian, Cuijosuclius Reig (1961), 
which was described by Rusconi in 1951 as a brachyopid. 

Watson (1956) agreed that Pelorocephalics is a brachyopid, 
and accepted as valid the affinities claimed by Cabrera between 
Pelorocephalus and Batrachosuchus from the Cynognathus 
Zone. As Watson pointed out, brachyopid labyrinthodonts are 
known only from the Permian and Lower Triassic throughout 
the world. 

These labyrinthodonts from the Cacheuta Formation require 
further study ; it is possible that in the palatine features of the 
two genera there are similarities with Batrachosuchus, but the 
difference in skull height in Chigutisaurus, as well as its larger 
size, may indicate that the Argentinian brachyopids, perhaps 
because of peculiar ecological conditions, outlasted the Scythian. 
It would, of course, be out of the question to call them Keuper ; 
tentatively w^e may consider them to be of Anisian age. Cuyo- 
suchiis, as mentioned above, is considered by Reig (1961) to be 
a proterosuchian; Hughes (1963) and Tatarinov (1961) agree 
with this designation. At present, all the more or less well 
known proterosuchians are of Scythian age, so that there is 
chronological agreement between them and the brachyopids. 
Thus, taking into consideration the possible special ecological 
conditions mentioned above, as well as some features of the 
Chigutisaurus skull, the fauna of the Cacheuta Formation is 
probably Anisian. 

Sierra de las Higueras Formation (Mendoza Proinnce). — The 
only fossil remains from this formation are a group of footprints 
representing five or six different forms, most of them observed 
and collected in 1963. Rusconi (1951b) studied a Chirotherium 
footprint showing an incomplete track of a hind limb and a 
probable track of the forelimb. Rusconi called them C. higueren- 
sis; Peabody, in 1955, restudied these footprints, and assigned 
them to C. harthi from the Moenkopi Formation of the United 



6 BREVIORA No. 251 

States. Keig (1961) and Casamiqiiela (1964) have demonstrated 
the weakness of such identifications, particularly as they are 
based on footprints which are far from clear. As mentioned 
above, Stipanicic (1957) considered the Sierra de las Higueras 
Formation to be Neoscythian-Eoanisian. 

Among the footprints collected and observed in 1963 near 
Puesto de las Higueras, there is one similar to that of Rigalites 
Huene (1931) from the Los Rastros Formation. The others are 
referable to a lacertiform and to therapsids. It seems useful to 
examine the chronological position of C. higiierensis and the 
Rigalites-\ike genus together, rather than to examine C. higiieren- 
sis alone. A first point is that the structure of the foot of 
Aetosauroides from the Ischigualasto Formation, which is known 
more or less in detail, is in agreement with the footprint of Q. 
higiierensis. A second is that the footprints assigned to Rigalites 
from Las Higueras are probably not older than the Rigalites 
from the Los Rastros Formation. My interpretation is, of 
course, open to question, but it appears highly unlikely that 
the upper section of the Sierra de las Higueras Formation would 
be older than the Los Rastros. In consequence, I assume a 
Ladinian age for this formation. 

Los Rastros Formation (San Juan Province). — The only 
known remains from this formation are good footprints of an 
archosaurian, Rigalites ischigualastianus Huene (1931). Huene 
considered them to be footprints of an ornithischian, but this is 
far from well established. Reig (1961), on the other hand, after 
study of the limbs assigned to Saurosuchus from the Ischigualasto 
Formation, believes that these footprints may well have been 
made by a pseudosuchian of great size, comparable to Sauro- 
suchus. This idea appears to be more acceptable. Despite the 
fact that these footprints are very clear, their age is not easy 
to ascertain. Some inferences may be made from the fact that 
the Los Rastros Formation is overlain coneordantly by the Ischi- 
gualasto Formation, the age of which is fairly well defined. On 
this basis, we conclude that the Los Rastros would not be younger 
than Upper Ladinian. 

Ischigualasto Formation (San Juan and La Rioja provinces). 
— Cabrera (1944) first studied remains from the Ischigualasto 
Formation, which had been collected by Frenguelli. In recent 
years, a goodly number of papers on this fauna have been pub- 
lished, greatly increasing our knowledge of this, the richest 
Triassic tetrapod fauna of Argentina. These include papers by 
Reig (1959, 1961, 1963), Casamiquela (1960, 1962), Cox (1962, 



1966 TETRAPOD-BEARING TRIASSIC OF ARGENTINA 7 

1965), Romer (1962a), and Bonaparte (1962, 1963a-e, in press.a, 
b). The fauna of the Ischiofualasto Formation is composed of six 
different gTOups; these will be discussed individually. 

(1) Labyrinthodontia. Promastodonsaurus hellmanni Bona- 
parte (1963a) is represented by incomplete remains of the skull, 
jaws, and pectoral girdle. It has been assigned to the Capito- 
sauridae, and closely resembles Mastodonsaurus from the Lower 
and Upper Triassic of Europe. As an indication of the chronol- 
ogy of the formation, it is, thus, of little value. 

(2) Cynodontia. The cynodonts are very numerous in this 
formation, and are found in nearly all its levels. The car- 
nivorous cynodonts are represented by a few specimens referred 
to Chiniquodon from the Santa Maria Formation of Brasil 
(Bonaparte, in press.a). The gomphodont cynodonts are Exaereto- 
don Cabrera (1944), Proexaeretodon Bonaparte (1963d), and 
Ischignathits Bonaparte (1963c) ; all three genera are found at 
approximately the same levels. Exaeretodon and the apparently 
somewhat less specialized Proexaeretodon are comparable with 
the Brasilian Traversodon and with Scalenodontoides from the 
Middle Triassic sensu lato of Africa (Crompton and Ellenber- 
ger, 1957). Ischignathus shows clear advances over the Brasilian 
and African Middle Triassic cynodonts, particularly in its pala- 
tine structure, which resembles that of the tritylodonts. Present 
knowledge indicates that the cynodonts from Ischigualasto are a 
little younger than those from Brasil ; they are obviously older 
than the tritylodonts from the redbeds of Africa. 

(3) Dicynodontia. The only known genus of this group is 
IscJiigualastia (Cox, 1962, 1965). Cox believes that Ischi- 
gualastia gives us no decisive chronological data, as it could be 
considered either Ladinian or Carnian. 

(4) Pseudosuchia. The genera of Pseudosuchia from this for- 
mation are Saurosuchus and Proterochampsa (Reig, 1959, 1961) 
and Aetosaiiroidcs and Argcntinosuchus (Casamiquela, 1960, 
1962). Reig considers that the rauisuchid ornithosuchian Sauro- 
suchus is not only the largest genus of this family, but has a 
longer ischium and ilium, and a weaker femur than do Presto- 
stichiis from Brasil or StagonosHchiis from Africa. AVe can as- 
sume that these characters represent anatomical advances over 
the Brasilian and African rauisuchids, comparable to the posi- 
tion noted in the ejaiodonts. Proterochampsa is considered by 
Reig as a true crocodilian. New materials of this genus, how- 
ever, indicate the need for a reconsideration of its position, par- 
ticularly because of the primitive palatine structure. Aetosau- 
roides, well analyzed by Casamiquela (1962), shows important 



8 BREVIORA No. 251 

similarities to Aetosaurus (from the German Keuper) in charac- 
ters of the skull, vertebral column, pelvic and pectoral girdles, 
limbs, and armor. Some differences may be noted in the teeth, 
which indicate that Aetosaiiroides may be slightly more primi- 
tive than Aetosaurus (Casamiquela, pers. com.). Aetosaurus 
and the related genus Stagonolcpis {fide Walker, 1961) are from 
the European Norian. In consequence, it is clear that Aetosau- 
roides, if not actually Norian, cannot be too far from this age. 
Argentinosuchus is known only from a humerus, some fragments 
of the radius and ulna, and some dermal scutes, and is, tenta- 
tively assigned to the Stagonolepidae. As the principal diagnos- 
tic parts of this genus are missing, however, its chronology is 
dubious. 

(5) Saurischia. Three genera of saurischian dinosaurs are 
known (Reig, 1963) ; a carnosaurian, Herrerasaurus ; a coeluro- 
saurian, Triassolestes; and Ischisaurus, of uncertain affinities. 
Herrerasaurus, known from good material of the postcranial 
skeleton, and from fragments of jaws which have been referred 
to it, is considered by Reig as more advanced, in some features 
at least, than other Triassic carnosaurians. Triassolestes is based 
on an incomplete skull and jaws, and some postcranial material. 
Reig reports some affinities with CoeJophysk from the Upper 
Triassic, the only genus included in his comparison, and con- 
siders as possible the inclusion of Triassolestes in the Podoke- 
sauridae. The affinities of Ischisaurus, which is based on some 
postcranial remains and fragments of the skull and jaws, are 
uncertain ; there are possible resemblances in the limb bones to 
Triassic pachypodosaurians. 

Little can be inferred regarding chronological position from 
these incompletely known saurischians. Since all of the compari- 
sons have been made with Upper Triassic forms, we have no valid 
arguments at present to consider them older than the Keuper ; 
we may, then, assign them to the Carnian. 

(6) Rhynchosauridae. Complete remains of these curious 
forms have been found at different levels of the Ischigualasto 
Formation. Until the j^resent, unfortunately, no study has been 
made of them. Some general features of the skull and jaws of 
these rhynchosaurs resemble those of Cephalonia (or Scaphonyx) 
from Brasil. No chronological data can be drawn from these 
unstudied forms. 

In several genera of the fauna of the Ischigualasto Formation, 
then, we find some advances over related Brasilian genera. It is 
also evident that the aetosaurid Aetosauroides, the traversodon- 
tid IscJiignathus, and the three saurischians appear as forms 



1966 TETRAI'OD-BEARING TRIASSIC OF ARGENTINA 9 

more related to Upper than to Middle Triassic faunas. On the 
other hand, the numerous cynodonts indicate a Middle Triassic 
age for this formation ; their abundance, however, could be ex- 
plained by sui)posing- that ecological conditions permitted a later 
expansion of these forms. The advanced features of the cyno- 
dont IscliUjnatlius, which has departed far from the known 
structure of the African Middle Triassic cynodonts, would sup- 
port this view. The presence of both rhynchosaurs and cynodonts 
has been considered by Romer (lf)60, 1962b) as strong evidence 
that this is an upper Middle Triassic fauna — as noted above, 
"surely pre-Xorian and not improbably pre-Carnian.' ' But the 
fact that rhynchosaurs from the Norian or Carnian of India 
have been found associated with a phytosaur, with a form com- 
parable to a coelurosaurian, and with prosauropod fragments 
(Jain, Robinson, and Chowdhury, 1964), makes his hypothesis 
of limited validity, in addition, rhynchosaurs have been re- 
ported from England, found in association wdth the supposedlj^ 
Norian Stagonolepis. In consequence, we may consider the 
fauna of the Ischigualasto Formation as of Carnian age. 

Los Colorados Formation (San Juan and La Rioja prov- 
inces). — In the past three years, much good tetrapod material 
has been found in this redbeds formation. This material is pres- 
ently under study, but some general considerations are possible 
at this time. The Los Colorados Formation lies concordantly 
over the Ischigualasto Formation, and is about 800 m thick. 
At its base has been found a skull and jaws comparable to Lschi- 
(jualastia, indicating the possibility of the survival of some 
genera connnon to the Ischigualasto Formation. But from the 
middle and upper sections of the Los Colorados we have found a 
completely different assemblage of tetrapods. At least three or 
four genera of prosauropods, some of great size, have been 
found. In addition, almost complete remains have been discov- 
ered of an aetosaurid closely related to Aetosauroides, but with 
more highly specialized teeth and cranial features ; apparently 
this is a Norian fauna. 

Our attempts to interpret the chronology of these Argentinian 
Triassic formations have been based on the currently accepted 
diagnoses of the Upper Beaufort and Stormberg Series of Af- 
rica, the Upper Scythian for the Cynognathus Zone, and so on. 
These attempts have been based solely on tetrapod remains from 
which excellent data have been obtained, particularly from the 
numerous forms comprising the fauna of the Ischigualasto For- 
mation and from the two genera of the Puesto Viejo, which are 



10 



BREVIORA 



No. 251 



SO closely comparable to African genera. This work raises the 
question of the age generally accepted for the Choiyoilitense and 
its correlations. Apparently this effusive series must be placed 
in the lower Middle Triassic, or else its correlations with Famati- 
nense and Paganzo III are in need of re-examination. 



San Rafael, 



San Juan 









Mendoza 




La Rioja 




Mendoza 




Mendoza 




o 

I/O 

< 

on 

1— 

Qi 

LlJ 

Q. 
Q. 

Z3 


< 

on 
o 




Los Colorados 




2 
< 

on 
<c 

o 


Ischlgualasto 








Los Rastros 




Sierra 

de las 

Higueras 




O 

on 
1/1 
< 

on 

(— 

LiJ 

o 

i 


■z. 
< 

o 

< 


Ischichuca 
















■z. 
<. 

oo 

z. 










Cacheuta 






Puesto 
Viejo 




Famatinense 

Dananvn 1 1 1 




Potrerillos 
















' "S""<-" 1 ■ ■ 


Las Cabras 





Table 2. A new attempt at chronology of some Tiiassic formations in 
Argentina. 



LITERATURE CITED 

Bonaparte, J. F. 

1962. Descripcion del craneo y mandibula de Exaeretodon frenguelUi 

Cabrera y su comparacion. . . . Publ. Mus. Cienc. Nat., Mar 

del Plata, 1: 135-202. 
19G3a. Promastodonsaurus bellmanni n.g. et n.sp., capitosaurido del 

Triasico Medio de Argentina. Ameghiniana, 3: 67-78. 
1963b. Descripcion del esqueleto postcraneano de Exaeretodon. Acta 

Geol. Lilloana, 4: 5-52. 
1963c. Descripcion de Ischignathus sudamericanus n.gen. et n.sp., 

nuevo cinodonte gonfodonte del Triasico Medio superior de San 

Juan, Argentina. Acta Geol. Lilloana, 4: 111-128. 



1966 TETRAPOD-BEARING TRIASSIC OF ARGENTINA 11 

lJ)()3d. Uii imevo ciuodoiite gonfodonte del Triasico Medio superior de 
San Juan, Proexaeretodori vincei. Acta Geol. Lilloana, 4: 129- 
133. 

19G3e. La familia Traversodontidae. Acta Geol. Lilloana, 4: 163-194. 

(In press. a) Chiniqtiodon Huene, en el Triasico de Ischigualasto, Argen- 
tina. Consideraciones sobre su asignacioii familiar. Acta Geol. 
Lilloana, 8: 

(In press. b) Sobre nuevos terapsidos Triasicos hallados en el centre de 
la Provincia de Mendoza, Argentina. Acta Geol. Lilloana, 8: 

(In press.c) Una nueva "fauna" Triasica de Argentina. Consideraciones 
filogeneticas y paleobiogeografieas. Aineghiniana. 
BONETTI, M. I. R. 

1963. Contribucion al conocimiento de la flora fosil de Barreal, Dpto. 
de Calingasta (San Juan). Thesis, University of Buenos Aires. 
Brink, A. S. 

1963. Two cynodonts from tlie Ntawere Formation in the Luangwa 
Valley of Northern Rhodesia. Palaeont. Afr., 8: 77-96. 

Cabrera, A. 

1943. El primer hallazgo de terapsidos en la Argentina. Notas Mus. 
La Plata (Paleout. No. 55), 8: 317-331. 

1944. Sobre un estegocefalo de la Provincia de Mendoza. Notas Mus. 
La Plata (Paleont. No. 69), 9: 69, 421-429. 

Casamiquela, R. M. 

1960. Noticia preliminar sobre dos nuevos estagonolepoideos argen- 

tinos. Anieghiniana, 2: 3-9. 
1962. Dos nuevos estagonolepoideos argentinos. Rev. Asoc. Geol. 
Argentina, 16 (3-4) : 143-203. 

1964. Estudios ienologicos. Gobierno Prov. Rio Negro. Min. Asuntos 
Sociales. Buenos Aires, 229 pp. 

Cox, C. B. 

1962. Preliminary diagnosis of Ischigualastia, a new genus of dicyno- 
dont from Argentina. Breviora, Mus. Comp. Zool., No. 158: 8-9. 

1965. New Triassic dicynodonts from South America, their origins 
and relationships. Phil. Trans. Roy. Soc. London, B 248: 457- 
516. 

Crompton, a. W. and F. Ellenberger 

1957. On a new cynodont from the Molteno beds and the origin of 
the tritylodouts. Ann. S. Afr. Mus., 44: 1-14. 
Groeber, p. F. C. and p. N. Stipanicic 

1953. Triasico. In: Geografia de la Republica Argentina. Gaea, 
2 (1): 7-141. 
Huene, F. v. 

1931. Die fossilen Fahrten im Rhat von Ischigualasto in Nordwest- 

Argentinien. Paleobiol., 4: 99-112. 
1935-42. Die fossilen Reptilien des siidamerikanischen Gondwanalandes. 
Munich, 332 pp. 



12 BREVIORA No. 251 

Hughes, B. 

1963. The earliest archosaurian reptiles. S. Afr. J. Sci., 59: 221-241. 
Jain, S. L., P. L. Eobinson, and T. K. E. Chowdhury 

1964. A new vertebrate fauna from the Triassic of the Deccan, India. 
Quart. J. Geol. See. London, 120: 115-124. 

KiTCHING, J. W. 

1963. The fossil localities and mammal-like reptiles of the Upper 
Luangwa Valley, Northern Rhodesia. S. Afr. J. Sci., 59: 259- 
264. 

MiNOPRIO, J. L. 

1954. Theriodonte en el Triasico de Mendoza. An. Soc. Cien. Argen- 
tina, 157: 31-37. 

Peabody, r. L. 

1955. Occurrence of Cltirothrriiim in South America. Bull. Geol. Soc. 
Amer., 66: 239-240. 

Reig, O. a. 

1959. Primeros datos descriptivos sobre nuevos reptiles arcosaurios del 
Triasico de Ischigualasto (San Juan, Argentina). Rev. Asoe. 
Geol. Argentina, 13: 257-270. 

]961. Acerca de la posicion sistematica de la familia Rauisuchidae y 
del genero Saurosuchus (Reptilia, Thecodontia). Publ. Mus. 
Cien. Nat. Mar del Plata, 1: 73-114. 

1963. La presencia de dinosaurios saurisquios en los " Estratos de 
Ischigualasto" (Mesotriasico superior) de las provincias de 
San Juan y La Rioja (Republica Argentina). Ameghiniana, 
3: 3-20. 

ROMER, A. S. 

1960. Vertebrate-bearing continental Triassic strata in Mendoza Re- 
gion, Argentina. Bull. Geol. Soc. Amer., 71: 1279-1294. 

1962a. La evolucion explosiva de los rhynchosaurios del Triasico. Rev. 

Mus. Arg. Cien. Nat., Cien. Zool., 8 (1): 1-14. 
1962b. The fossiliferous Triassic deposits of Ischigualasto, Argentina. 

Breviora, Mus. Comp. Zool., No. 156: 1-7. 
RUSCONI, C. 

1948. Nuevos laberintodontes del Triasico de Mendoza. Rev. Mus. 

Hist. Nat. Mendoza, 2: 225-229. 
1951a. Laberintodontes Triasicos y Permicos de Mendoza. Rev. Mus. 

Hist. Nat. Mendoza, 5: 33-158. 
19511). Rastros de patas de reptiles Permicos de Mendoza. Rev. Soc. 

Hist. Geogr. Cuyo, 3: 1-14. 
Stipanicic, p. N. 

1957. El sistema Triasico en la Argentina. Internat. Geol. Congr., 

20th, Mexico, Tr. Sec. 2: 73-111. 
Tatarinov, L. p. 

1961. Materiales de pseudosuehios de la LTRSS. Paleont. Zhurn. 
URSS, No. 1: 117-132. [In Russian.] 



1966 TETRAPOD-BEARING TRIASSIC OF ARGENTINA 13 

Walker, A. D. 

1961. Tiiassic reptiles from the Elgin area: Stagonolepis, Dasygna- 
thns and their allies. Phil. Trans. Eoy. Soc. London, B 244: 103- 
204. 
Watson, D. M. S. 

1948. Dicynodon and its allies. Proc. Zool. Soc. London, 118: 823-877. 
1956. The brachyopid labvrinthodonts. Bull. Brit. Mus. (Nat. Hist.), 
Geol., 2: 315-392. 

(Received 6 May, 1966.) 



BREVIORA 

Mmseiiiinti of Cojmparsitive Zoology 

Cambridge, Mass. November 4, 1966 Number 252 



THE CHANARES (ARGENTINA) TRIASSIC REPTILE 

FAUNA 

II. SKETCH OF THE GEOLOGY OF THE RIO 
CHANARES-RIO GUALO REGION 

By Alfred Sherwood Romer and James A. Jensen 

TALAMPAYA BASIN 

111 the preceding article in this series an account was given of 
the joint La Plata-Harvard expedition of 1964-65 and the dis- 
covery during its course of an excellent fauna of Triassic reptiles 
in the region of the Chanares and Gualo rivers in La Rioja 
Province. The present paper is intended to give a rough outline 
of the geology of the collecting area so that the fossil horizon 
can be placed in its proper setting. Since our expedition was 
aimed solely at fossil collecting, proper instruments for geologic 
Avork were not part of our equipment, and mapping of the area 
was done by one of us (J.A.J.) with the aid of such instru- 
ments as could be improvised. Our map is thus provisional 
in nature; it is our hope that Argentinian geologists may make 
more definitive studies in this portion of an extremely interesting 
sedimentary basin. 

Our area of concern is part of a basin of late Paleozoic and 
Mesozoie continental sediments located mainly in western La 
Rioja Province south of Villa Union. No name has been applied 
to this region ; we shall here term it the Talampaya Basin, be- 
cause of the central position in it of the Campo de Talampaya.^ 
Its boundaries, as regards presumed Triassic elements, at least, 
are outlined by Frenguelli in figure 1 of his 1948 discussion of 
the Argentinian "Rhaetic." To the east, the basin is bordered 
by the north-south range of the Sierra de Saiiogasta, a southern 



1 Frenguelli (1948) speaks of the "Cueiica de Ischigualasto-Ischichuca," but 
this properly applies only to the southwestern portion of the basin. 



2 BREVIORA No. 252 

continuation of the Nevada de Famatina, and by this sierra's 
southern terminations, such as the Sierra de Vilgo and the Sierra 
de Cerro Blanco ; to the west, its boundary is the Rio Bermejo, 
east of which the beds of the basin are sharply down-tilted, and 
to the north, its tributary, the Eio Guandacol (or de la Troya). 
The sediments of the basin extend northward to the Villa Union 
region. To the south, the basin narrows in the region of the little 
settlement of Los Baldecitos, where the southwestern-trending 
Sierra de Cerro Blanco approaches the Sierra de Valle Fertil, 
but (although the situation is obscured by a covering of Recent 
deposits) the sediments of the basin appear to be more or less in 
continuity with other late Paleozoic and Triassic beds extending 
eastward toward Paganzo and Patquia and possibly southward 
toward Valle Fertil in San Juan Province. Very probably, be- 
fore the diastrophic movements which resulted in the rising of 
the Sierra de Saiiogasta and the down-thrusting east of the 
Bermejo, the basin may have been of much greater extent, run- 
ning from the Patquia district on the east to the Iluaco region 
in San Juan Province to the west. The central portion of the 
basin is a nearly flat plain covered by Pleistocene sediments and 
Recent sands. The northern portion of this plain is drained by 
the Rio Vinchina which at the south breaks through the hills at 
the west of the basin and joins the Rio de la Troya (or Rio 
Guandacol) to form the Rio Bermejo. The barren Campo de 
Talampaya occupies most of the southern part of this plain. Its 
main drainage is the northward-flowing Rio del Alto (or Arroyo 
Manero), with main tributaries from the east, in north-south 
sequence, including the Pagancillo, Medanos, Talampaya, 
Chanares, and Gualo rivers. Along the western margin of the 
basin extends a broken range of north-south hills which are 
essentially a sedimentary northward continuation of the Sierra 
de Valle Fertil, including the long Sierra ilorada, Cerro Rajado, 
Cerro Bola, and Cerro Villa Union; contained in the first of 
these is the depression of the Cuenca de Ischigualasto. On the 
east, the northern part of the plain is separated by well-devel- 
oped foothills from the main Sierra de Saiiogasta. East of the 
village of Pagancillo is the Loma de los Loros, composed of 
Tertiary deposits (de Alba, 1954, pp. 54-56), farther south the 
striking Sierra de los Tar j ados, formed by Permo-Triassic sedi- 
ments. South of the region of the Rio Talampaya, the west flank 
of the Sierra de Safiogasta descends abruptly (due, presumably, 
to major faulting [Fidalgo, 19G3]) to a sand-covered plain, 
averaging 10 to 15 kilometers in breadth, which slopes gently 



1966 



RIO CHANARES-RIO GUALO GEOLOGY 



westward toward the Campo de Talampaya. But for the last few 
kilometers before the campo is reached, erosion in the valleys 
of the Chaiiares and Gualo rivers, which drain this area, has in 
great measure removed the alluvial covering and developed a 
north-south series of foothills in which a considerable series of 
Permian and Triassic sediments is exposed. 

It is this last mentioned area in which our fossil finds were 
made and with which the present paper is concerned. Figure 1 
is a sketch of a portion of La Kioja and San Juan provinces to 
indicate the position of the small area covered by our geologic 
map. 

We have in the first number of this series expressed our thanks 
to the numerous Argentinian geologists who aided our expedi- 
tion by furnishing us with geological information and advice. 




©Voile Fertll 



67' 



Fig. 1. Sketch map of parts of La Eioja and San Juan provinces, to 
show the position of the area studied. Elvers and principal roads indi- 
cated; provincial boundary in broken line. Scale approximately 1/1,200,000. 



4 BREVIORA No. 252 

During the preparation of the present paper we have further 
corresponded, to our advantage, with Dr. Pedro N. Stipanieic of 
the Comision Nacional de Energia Atomica, Dr. Hector F. de la 
Mota of the Yacimientos Petroliferos Fiscales, and Dr. Francisco 
Fidalgo of the Instituto Nacional de Geologia y Mineria. 

In the entire area of the sedimentary basin described above, 
the population is small, due to great aridity (the average rainfall 
is about 2 inches [5 cm] per annum) ; railroads are entirely lack- 
ing, and roads few. Irrigation from the Rio A'inchina has per- 
mitted the growth of population to about 3500 in the Villa 
Union region ; the water of the Rio Pagancillo has allowed the 
development of a village of that name. But from Pagancillo 
southward some 90 kilometers to the little settlement of Los 
Baldecitos — an area of perhaps 5000 square kilometers — not a 
single inhabitant, not even a goat-herder, is to be found. At 
the north, Villa Union is connected by national roads with Guan- 
dacol to the west, and, to the east, with Chilecito via Puerto 
Alegre and the Cuesta de Miranda ; Pagancillo has road connec- 
tions with both Villa Union and Puerto Alegre. Until recently, 
almost no roads of any sort were present in the remainder of the 
basin ; travel and exploration were exceedingly difficult, not 
merely because of the absence of highways, but also because of 
the nearly complete absence of fodder and water for the mules 
which formed the main means of transport.^ A few years ago, 
however, a well-graded road, aimed to become eventually an in- 
ternational highway to Chile, was built from the railway junction 
of Patquia westward to Baldecitos, and thence north along the 
Campo de Talampaya to Pagancillo. From this highway (at 
kilometer post 100) a track has been worked out westward by 
Dr. Ramon de la Vega, via the Ischigualasto Valley, to the Los 
Rastros coal mine; at kilometer post 116 a track, traveled by 
occasional miners, runs eastward and northward along the plain 
at the foot of the Sierra de Saiiogasta. In general, the sand}^ 
"river" channels, dry except after rare rainy-season showers, 
can be readily traveled by motors with four-wheel traction. 

As said above, the presence here of a great basin of late 
Paleozoic and Mesozoic sediments was early recognized. Some 
early observations were made by Stelzner (1885), who traversed 



1 Note Bodenhemler's emijhatic (ami italicizpd) warning (1911, p. 96): 
"NOTA — A los futitros explorades hngo prescnte que no se puedc contar — 
rrcepto nnos (Ic miicha lluria — con sufflciente pasto para los animales," etc. 
There is. however, a mule track, of considerahle antiquity and still used, from 
.Jacluil to Patquia, which crosses the Rio Bermejo at Paso Ferreyro, traverses the 
Ischigualasto Valley, and continues on to the southeast. 



1966 RIO CHANARES-Rio GUALO GEOLOGY 5 

the area in 1873, and later by Boclenbender, as noted in his pub- 
lications of 1896, 1897, and 1902. The first published work of 
importance, however, was that of Bodenbender in 1911, in which 
he gave a general survey of the geology of southern La Rioja 
Province and adjacent regions of San Juan. It is to Boden- 
bender that we owe the invention of the term "Paganzo," com- 
monly used })y later writers in variable and often confusing 
fashion. Groeber in 1940 summarized what was then known of 
this region. De la Mota has done much valuable work, unfortu- 
nately unpublished, in various portions of the Talampaya Basin ; 
Rigal, Harrington, and Ramaccioni have also worked here, but 
have not published ; Fidalgo is currently studying the eastern 
portion as part of a survey of the geological Hoja Catizaco 
(which includes as well the Sierra de Sanogasta and territory to 
the east of it). The sediments of the most northerly part of the 
basin are described in de Alba's bulletin (1954) on the Villa 
Union sheet of the geological map of Argentina. Heim published 
on coal deposits in this area in 1946 and 1947, and in his 1949 
studies of the Los Rastros coal in the western part of the basin, 
contributed valuable observations on that area from his own 
observations, and to some degree from those of Ramaccioni. 
Most important are Frenguelli's studies of the area (1944a, b, c, 
1945, and especially 1948), in which he appears to have relied 
greatly on the work of de la Mota and Ramaccioni. An excellent 
resume of this work, as regards the Triassic beds, is given by 
Groeber and Stipanicic (1953). Recently Dr. Apolo Ortiz of the 
Y.P.F. has made important investigations, as yet unpublished, 
in the southwestern part of the basin. The greater part of pre- 
vious studies has dealt with the western portions of the basin, 
west of the Campo de Talampaya ; little study has been made of 
the eastern section, with which we are here concerned. 

The general topographic features of the region of present in- 
terest are shown on the accompanying geological map. This is 
an area running for about 25 kilometers along the eastern border 
of the campo from the mouth of the Rio Talampaya on the 
north to a point somewhat south of the Rio Gualo. It extends 
eastward from the campo to the point, averaging about 10 
kilometers, where the Permo-Trias beds become covered by the 
sui^eriicial deposits of a plain which slopes up eastward to the 
foot of the high mountains. At the northern end of the region 
studied, the Rio Talampaya, which drains a considerable area of 
the sierra and its foothills, emerges south of the Sierra de los 
Tarjados onto the campo through a spectacular high-walled 



BREVIORA 



No. 252 




^^^ Sand Dunes 
'/'/// Undifferentiated 
'^^■4■ Tertiary 

-^nv Los Colorados 
Ichigualasto 
Los Rastros 
Chonares 
Tarjodos 
Tolampoya 






PROVISIONAL GEOLOGIC MAPv^^?-<^ 



TALAMPAYA- CHANARES 
GUALO REGION, 



LA RIOJA, 

ARGENTINA 



flppro«imale Scole 




Fig. 2. 



canyon, the Puerta de Talampaya. For some kilometers to the 
south of the river there is an elevated barren plain, with con- 
spicuous exposures of red sandstones and with a series of parallel 
channels running northward to the Rio Talampaya. South of 
this plain is the Rio Chaiiares drainage, of which the lower part 
is notable for a considerable development of sand dunes which 



1966 RIO CHAN ARES-RIO GUALO GEOLOGY 7 

obscure the underlying formations. A major southern branch 
of the river rises well to the east toward the sierra ; in the region 
studied there are numerous small branches which gather into 
north and south forks, the two uniting a short distance east of 
the entrance of the river into the plain. The main branch of the 
Rio Gualo arises at the western borders of the sierra, runs south- 
westward across the alluvial plain and, entering the region 
studied, flows along the northern margins of a flat, sandcovered 
plain, which we shall term the Piano del Gualo. Here it receives 
several short tributaries. One from the northeast has along its 
course the onlj' perennial spring of any note in the region ; a 
neighboring hill of red sandstone which serves as a landmark for 
the spring is the Mogote del Gualo. At the west border of the 
Piano del Gualo the river breaks through the sandstone cliffs 
which nearly surround this plain to enter a narrow north-south 
valley, bounded on the west by a ridge of red sandstone. This 
ridge is broken half-way along its length, and maps show the 
Gualo as turning westward through this gap. It may have done 
so at one time ; currently, however, it follows the valley north- 
ward to the end of the sandstone ridge before debouching onto 
the plain only a short distance south of the Rio Chanares. 

Near the southwestern border of the Piano del Gualo is a 
prominent peak of volcanic rock, the Mogote de Agua Eseondida. 
Arising in this region is a stream whose channel may be termed 
the Arroyo de Agua Eseondida. This runs nearly directly west- 
ward out into the campo, and carves a deep channel along its 
course. It is of geological importance, since its walls show an 
almost complete section through all the formations present in 
the region studied. 

The whole area is considerably faulted; onh^ a few of the 
faults seen during our preliminary studies are shown on the 
map. In general, the beds in the north and east portions are 
relatively horizontal ; to the west and south, dips in that direc- 
tion become, on the average, increasingly steep until (in the 
region of the canyon of the Arroyo de Agua Eseondida, as seen 
in Figure 2) the highest beds of the series are vertically placed 
and even somewhat overturned. 

EL CHACHO FORMATION 

Of the three successive stages into which Bodenbender divided 
his "Paganzo" series, the second, "Paganzo II," was applied 
to strata of presumably Permian age and, more especially, to a 



8 BREVIORA No. 252 

series of red sandstones widespread in La Rioja Province — 
prominent, for example, from Patquia northward to Los Colo- 
rados station on the Chilecito railroad branch. Frenguelli ( 1946, 
p. 315, etc.) called these redbeds the Estratos de Pat^inia ; I 
understand that Dr. Fidalgo plans to term them the El Chacho 
Formation — appropriately so, since the well-known cave, north 
of Patquia, where this war chieftain of the last century made his 
headquarters, is formed in this red sandstone series. There is 
no adequate fossil evidence to date these El Chacho beds, but 
there is no reason to doubt their Permian age. El Chacho red- 
beds are widespread in the Sierra de Sanogasta ; some have been 
carried by diastrophism to high elevations in the range. West of 
the Sierra de Sanogasta, El Chacho redbeds are abundantl}- rep- 
resented along the western slopes in the region of the Cuesta de 
Miranda and Puerto Alegre (de Alba, 1954), and are also pres- 
ent, in the northwestern portions of the basin, in Cerro Villa 
Union and Cerro Guandacol. But on the east side of the basin, 
from the Rio Talampaya southward, major faulting, as Fidalgo 
(1963) notes, appears to have been responsible for a steep west- 
ern face of the Saiiogasta range and absence of foothills of El- 
Chacho type. In consequence, these redbeds are not exposed in 
the area here studied, although they would presumably be en- 
countered sub-surface. 

talajvipaya formation 

Lowest of beds present in the area studied are those which we 
here term the Talampaya Formation — the name being given 
because of their extensive exposure in the valley of the Rio 
Talampaya. The beds consist of rather uniform, pale reddish 
buff, sandy sediments, generally fine grained and finely bedded. 
Occasionally present, in seemingly random fashion, are cobbles, 
generally smoothly rounded, of crystalline material. In the Rio 
Talampaya region they have a maximum size of about 80 milli- 
meters; to the south, near the Agua Escondida, they are more 
numerous and larger, attaining a maximum of 30 centimeters in 
diameter. 

The lower portion of the formation as exposed includes a 
cyclic repetition of soft and more resistant layers ; in the upper 
175 to 200 meters there are no resistant elements. Where the 
formation is broadly exposed at the surface, it weathers to give 
gently rounded to subangular profiles. Where, however, the 
upper beds are covered by the resistant basal conglomerate of 



1966 BIO CHANARES-RIO GUALO GEOLOGY 9 

the overlying Tarjados Formation, the results are striking; the 
soft nature of the Talanipaya beds results in the formation of 
sheer, vertical, fluted cliffs, the outlines of which follow pre- 
cisely those of the overlying conglomerate. Such cliffs form 
much of the margin of the Sierra de los Tarjados north of the 
Rio Talampaya ; for several kilometers in the lower course of 
that river sheer cliff's of this sort, approximately 180-200 meters 
high, form the spectacular Puerta de Talampaya. 

Presumably the Talampaya Formation succeeds the typical 
redbeds of "Paganzo II" in the Permo-Triassic sequence of 
the basin, but nowhere in the area visited by us is a normal 
contact between the two visible. The only contact seen during 
our exploration was that of about 1 kilometer along a normal 
fault, in alignment with the general fault system of the mapped 
area, but about 10 kilometers north of the upper boundary of 
the mapped area. Because of lack of knowledge of the base of 
the formation, its total thickness is unknown, but may be esti- 
mated to be at least 400 meters. 

The Talampaya Formation is extensively exposed, as men- 
tioned, in the valley of the Rio Talampaj'a to the northeast of 
the mapped area and along the flanks of the Sierra de los Tar- 
jados. West of the Puerta de Talampaya, a strip of exposure 
of the formation extends southward toward the Rio Chaiiares 
for several kilometers along the base of cliffs formed by the Tar- 
jados Formation. Apart from this, the Talampaya Formation is 
covered by younger beds over most of our region. To the south, 
however, southwest of the Piano del Gualo, it is present at the 
surface over a considerable area. 

It is obvious that the Talampaya Formation is part of the 
complex of beds included by Bodenbender in his Paganzo system. 
It does not, however, correspond closely to any specific beds 
described by him in 1911. Possibly these strata may correspond 
to the fine, ligbt brown, cross-bedded sandstones which Furque 
(1963) describes, in the Guandacol region, as part of the Ojo de 
Agua Formation, above the red sandstones which he considers 
equivalent to Bodenbender 's "Paganzo II." Again, these beds 
may (cf. Turner, 1960) be equivalent to the lutites and marls 
which Bodenbender (1916) considers to be the basal portion of 
his "Estratos Famatinenses. " Stipanicic (in litteris) suggests 
that the Talampaya, as well as the overlying Tarjados Forma- 
tion, can be considered as part of "Paganzo III" or "Famati- 
nense." It is possible that this set of beds may be comparable 



10 BREVIORA No. 252 

to the "areniscas bandeadas" which de la Mota, in his unpub- 
lished thesis, includes as part of the third element (above a 
"Paganzo II" equivalent) in the series of beds described l)y him 
at Cerro Bola (the remainder of this set of beds include erup- 
tives and porphyritic conglomerates). 

TARJADOS FORMATION 

Above the Talampaya Formation lie sandstones, of a coarse 
and resistant nature, which at the Arroyo de Agua Escondida 
have a thickness of about 385 meters. The name here given them 
is due to the fact that the lower part of these beds forms a broad 
covering over the Sierra de los Tarjados to the north of the 
area studied. South of the Rio Talampaya, the Tarjados Forma- 
tion is broadly exposed at the surface, covering perhaps half of 
the mapped area. In the southern part of the area there is gen- 
erally a sharp division of the formation into lower and upper 
members, the lower mainh^ of red-colored sandstones, the upper 
predominantly white. However, the contrast betAveen the two 
members is not so evident to the north. The lower, red member 
has a thickness of 130 meters in the measured section along the 
Arroyo de Agua Escondida. Throughout the area there is a 
basal conglomerate of variable thickness, but with a maximum 
of 2 to 3 meters, which lies unconformably above the Talampaya 
Formation. In the measured section there follow 10 meters of 
bright red, cross-bedded sandstone, then 35 meters of dull, red- 
dish brown shale ; this in turn is followed by 85 meters of bright 
red cross-bedded sandstones very similar to the basal section. 
Transitional here to the upper division, there are 4 meters of 
variegated sandstone, a thin limey nodular zone, and a further 
thin layer of sand and clay pebbles ; above this are about 118 
meters of sandstones, which are, for the most part, white in color. 
Cliffs of this upper division form much of the boundary of the 
Piano del Gualo ; scraps of skull and postcranial materials, prob- 
ably of a dicynodont, were found here in the transitional beds at 
two localities. A kilometer or so north of the boundary of the 
Piano del Gualo, a west-flowing northern tributary of the Rio 
Gualo has excavated a deep canyon in the massive white sand- 
stones of the upper division. 

In the region of drainage into the Rio Talampaya, at the 
north of the mapped area, the Tarjados beds are predominantly 
red in color. Here the lower sediments, above the basal con- 
glomerate, are a dull red sandstone, 40 to 45 meters thick, with 



1966 RIO CHANARES-RIO GTTALO GEOLOGY 11 

alternate soft and resistant zones. They are overlain by 80 
meters or so of bright ochre-red sandstones with almost no re- 
sistant strata. These erode in spectacular fashion to form fluted 
cliff faces, similar to the folds in a heavy drape. The appearance 
of these cliffs, apart from the difference in color, is similar to 
that of the cliff's of the underlying Talampaya Formation. 

Whenever the entire thickness of the Tar j ados surface is pre- 
served, it is seen to terminate above in an uneven, undulating 
surface of hard, resistant materials containing a considerable 
quantity of chert, suggesting that this layer, typically about a 
half meter thick, results from some type of hydrothermal activ- 
ity. This surface is frequently seen to be penetrated from above 
by small silica-filled tubules, averaging 2.5 centimeters in diame- 
ter, with a maximum of 5 centimeters, and penetrating to a 
depth as great as 3 meters. Their appearance suggests that we 
are dealing with root S3^stems of plants which grew on the un- 
eroded surface of the Tarjados before the beginning of the 
deposition of the overlying Chaiiares beds. 

Boclenbender 's ' ' Paganzo III ' ' was none too clearly defined by 
him, and, further, he appears to have been uncertain whether or 
not his "Famatinense" (Bodenbender, 1916) is an equivalent of 
''Paganzo III" (cf. Bodenbender, 1924; Turner, 1960). At all 
events, it seems jn-obable that his intent was to apply the term 
"Paganzo III'" primarily to beds of red sandstones lying above 
"Paganzo II" and below his "Rhaetic." It thus seems certain 
that the sandstones here named the Talampaya Formation are 
in the category of formations which he would term "Paganzo 
III." Stipanicic {in litteris) confirms our belief in this regard. 

Below the "carboniferous" beds in the Ischigualasto region, 
which were termed Los Rastros and Ischichuca by Frenguelli 
and Heim, lie a series of red sandstones which are as yet im- 
perfectly studied. Frenguelli (1948) considered them to be 
"Paganzo II," his Patquia Formation; Heim (1949) believes 
them to include both "Paganzo II" and "Paganzo III." I un- 
derstand that Dr. Ortiz considers these beds to belong entirely to 
"Paganzo III," and that "Paganzo II" is lacking there. Quite 
surely, these beds are, in part at least, equivalent to our Tar- 
jados Formation. 

No comparable beds appear to have been present in Cerro 
Bola or the neighboring region of Cerro Guandacol, as studied 
by de la Mota (Frenguelli, 1946, 1948). 



12 BREVIOKA No. 252 

CHANARES FORMATION 

Laid down on the undulate upper surface of the Tarjados 
beds are some 75 meters of light-colored beds, mainly fine vol- 
canic tuffs, M'hich form a striking contrast to all others in the 
area. 

The basal beds of this formation, for a few meters overlying 
the Tarjados, are somewhat variable in nature, consisting of 
dull, pale green to dark buff or pinkish tuffs in w'liich are in- 
cluded occasional thin pebble conglomerates with fossil ver- 
tebrate scraps and silicified plant materials, random beds of 
abundant limestone nodules with major diameters of up to 30 
centimeters, and various compact nodular lenses composed of 
coarse sands, limestone, and chert, cemented into resistant masses 
of up to 8 meters in horizontal extent. 

A short, if variable, distance above the base, the sediments be- 
come light-colored fine grained tuft's, with a somewhat bluish 
tinge, particularly in the lower part of the formation. They are 
well stratified ; possibly aeolian, possibly, in part at least, lacus- 
trine. With one exception, all of the numerous vertebrate fossils 
found in the Chaiiares Avere from the low^est 10 meters or so of 
the formation. A fraction of them were found directly embedded 
in the stratified tuffs; a large proportion, however, were found 
in grey-brown concretions which in many areas are abundant 
at this level ; these consist of coarser tuff's, infiltrated by calcite 
and secondary- silica.^ These are flattened spherical structures, 
often with irregular outlines, and vary from 0.3 to 2.5 meters in 
diameter. In some areas they appear to be lacking in fossil 
content; in others (notably in the most northwesterly portion 
of the area of Chafiares exposure) nearly every concretion con- 
tains reptilian material, sometimes with all or parts of two or 
three individuals in a single concretion. The remains are not 
always restricted to the limits of the concretion, and parts of a 
skeleton may extend into the surrounding stratified tuffs. 

Above the fossil-bearing portion of the Chanares, the beds are 
nearly pure white in color, with a white concretionary zone well 
toward the top which tends to produce vertical cliff's. 

At their top, the Chanares tuffs are followed without any un- 
conformity by the l^eds here assigned to the Los Rastros Forma- 
tion. 



1 We wish to thank Dr. Raymond Siever for studying samples of Chanares For- 
mation sediments. 



1966 RIO CHANARES-RIO GUALO GEOLOGY 13 

A main series of outcrops of tlie Chaiiares beds, somewhat 
interrupted by faults, follows the base of the Los Rastros For- 
mation from the northwest part of the Rio Chaiiares drainage, 
eastward and southward to the Rio Gualo just beyond the point 
where it leaves the Piano del Gualo ; faulting has caused the 
series to appear again to the eastward of this line, well up the 
courses of l)oth the Rio Chaiiares and Rio Gualo. 

Because of lack of unconformity, it would be possible to con- 
sider the Chaiiares beds a basal member of the Los Rastros. We 
believe, however, that they merit distinction as a separate forma- 
tion, because of their distinctive lithology and because of their 
paleontological importance. I find in the available literature no 
description of Triassic beds which correspond at all closely to 
those of the Chaiiares. Presumably they may be equivalent to 
some portion of the sediments classed in other areas as part of 
the Ischichuca Formation (and which are presumably part of 
the Los Rastros Formation, as here defined). Our Chaiiares beds 
do not, however, compare with the basal part of the Ischichuca 
in the Rio de la Peiia region as described by Frenguelli (1948, 
p. 191 and lig. 29) on the basis of data furnished him by 
Ramaccioni, or as described by Heim (1949), nor do they com- 
pare closely with any part of the Ischichuca Formation in the 
type region of that formation, as described by Frenguelli (1948, 
pp. 200-208) mainly on the basis of de la ]\Iota's unpublished 
work. 

Future study of the fauna will presumably give evidence as to 
the age of the Chaiiares Formation. As seen in the field, the 
material suggested a somewhat later evolutionary stage than that 
seen in the Cynognaihus Zone of South Africa. This zone seems 
certainly to be of early Triassic, Scythian, date (Watson, 1942, 
Lehman, et al., 1959). Hence the Chaiiares may be considered 
as either late Scythian or more probably Anisian in age. 

LOS RASTROS FORMATION 

The beds which we here include in the Los Rastros Formation 
occupy a considerable area in the lower parts of the Rio 
Chaiiares and Rio Gualo valleys east of the point at which these 
rivers debouch into the Campo de Talampaya. Over a large part 
of this area, however, these beds are covered by sand dunes, and 
they are variably tilted due to faulting. A further exposure of 
some extent is present, due to faulting, north of the main chan- 
nel of the Rio Gualo near the east edge of the mapped area. 



14 BREVIOKA No. 252 

The conformable contact between the Los Rastros and the under- 
lying- Chaiiares Formation is exposed over very considerable 
areas ; on the other hand, a Los Rastros-Ischignalasto contact is 
visible only in the section seen in the Arroyo de Agua Escondida. 
Here the thickness of the formation is 216 meters (as contrasted 
with an estimated thickness of 600 meters for the combined Los 
Rastros-Ischiehnca in the western part of the basin). 

In general, the Los Rastros sediments consist of brown and 
tan sandstones interbedded with shales and clays, of various pale 
colors, and carbonaceous layers. Lack of continuovis exposures 
due to sand dune coverage, and difficulty of correlation between 
various areas due to faulting and variable tilting, make it diffi- 
cult to give a general description ; it seems, however, certain 
that there is considerable variation in the beds from one region 
to another. Even over a short distance, such variations can be 
seen in the thickness of sandstone members. In some regions the 
sandstones are very prominent, in others — notably in the 
drainage of the south fork of the Rio Chaiiares — shales and 
clays dominate; in this area, for example, shale and clay beds 
may reach a thickness of 30 meters or more without intercalation 
of any sandstone. There are a number of carbonaceous layers 
of variable thickness. Some of them apparently persist over a 
considerable portion of the mapped area; these contain abundant 
plant materials. In a zone in the upper portion of the formation 
there are several widespread beds of haematite which range in 
thickness from 2 to 10 centimeters. Occasional ironstone con- 
cretions in these layers contain fossil plant or fish remains of 
poor quality. In the upper Gualo area most of the beds in the 
formation appear to be thinner than they are farther to the west, 
suggesting (together with a relatively high ratio of sandstones to 
clays) a region of marginal deposition. On the south side of the 
Rio Chaiiares canyon, near its mouth, there is a notable display 
of a cyclical type of deposition which appears to be to some 
degree characteristic of the formation as a whole. Here four 
cycles can be seen, each consisting, in order from top to bottom, 
of sandstones, sandy shales, carbonaceous shales, and clays, fol- 
lowed below by the sandstone of the next cycle. These cycles 
have thicknesses of 10 to 25 meters. 

In the northwestern part of the Talampaya Basin, de la Mota 
has distinguished an Ischichuca Formation; Heim (1949), in his 
description of the Rio de la Pena region, somewhat arbitrarily 
divides the carbonaceous beds lying between the Ischigualasto 
Formation above and the redbeds below into the Ischichuca and 



I 



1966 RIO CHANARES-RIO GUALO GEOLOGY 15 

Los Rastros formations. This is also done by Frenguelli (1948) 
who, it seems, Avas personally familiar only with the upper por- 
tion of the Los Rastros Formation in the Cerro Bola region. I 
am informed that current belief is that the whole series of 
carbon-bearing beds in the La Pena region should be considered 
a single unit, the Los Rastros Formation. We likewise see no 
reason why the similar (and much thinner) beds in our area of 
interest should not be considered a unit, although, as mentioned 
earlier, the underlying Chanares Formation may be in some 
fashion equivalent to part of the Ischichuca. 

ISCHIGUALASTO F0R:\IATI0X 

In the western portion of the Talampaya basin, the Ischi- 
gualasto Formation is extensively exposed; it was described in 
some detail by Frenguelli in 1948, and has become well known 
through the discovery in it of an abundant vertebrate fauna. 
On the east side of the basin, however, it is seen only in the 
walls of the Arroyo de Agua Escondida ; here the Ischigualasto 
is present in proper sequence between the Los Rastros and Los 
Colorados Formations, the beds tilted at an angle close to the 
vertical. Presumably they continue to the north, in the valley in 
which the Rio Gualo flows northward for some distance east of 
the ridge formed by the Los Colorados. It can be assumed that 
this valley formed because the lack of resistant layers in the 
Ischigualasto beds resulted in greater erosion of them than of 
the sandstones of the Los Colorados; there is no Ischigualasto 
exposure here, due to a covering by Recent materials. In con- 
trast to the much greater thickness, of perhaps 400-500 meters, in 
the Ischigualasto Valley, the beds as measured in the Arroyo are 
only 175 meters thick. We found no fossils in the limited avail- 
able exposures. The sediments, however, compare closely with 
those in the type area, except that there is an almost complete 
absence here of the resistant sandstone layers encountered at 
intervals in the type area. As at Ischigualasto, the beds contain 
pale purple, grey, and tan joint-clays interbedded with pale ma- 
roon, green, and yellow siltstones. Two or three thin beds of 
nodular limestone are present, one of which includes some 
poorh^ preserved plant material. 

Below, there is a sharp contrast between the Ischigualasto 
sediments and the interbedded hard sandstones and shales of the 
Los Rastros Formation. Aljove (as in the Ischigualasto Valley) 



16 BREVIORA No. 252 

there is, over a short vertical distance, a transitional zone be- 
tween the Ischigualasto Formation and the overlying Los Colo- 
rados of interbedded sands and silts grading in color from greys 
and yellow-tans into dull reddish brown. It was apparently such 
transitional beds, between Cerro Morado and the southern end 
of the Los Colorados in the Ischigualasto Valley, which led 
Bodenbender (1911) to insert a "Jurassic" phase into his se- 
quence in that region ("perfiles III y IV" and p. 97). 

As noted in the previous paper in this series, the fauna of the 
Ischigualasto Formation is essentially Middle Triassic ; it is, 
however, difficult to determine whether it should be considered 
as parallel to the Ladinian or to the Carnian of the marine 
Triassic sequence. 

LOS COLORADOS FORMATION 

The massive red sandstones which bound the Ischigualasto 
Valley to the east and form a nearly continuous range of hills 
along the Avestern side of the Campo de Talampaya were termed 
the Estratos de Gualo by Frenguelli (1948, pp. 170-174), on 
the assumption that the red sandstones forming the Mogote del 
Gualo were of similar age. However, this is not the case, as was 
pointed out by de la Mota to Groeber and Stipanicic ; at his 
suggestion, they renamed these sandstones the Los Colorados 
Formation (1953, p. 88). In contrast to the prominence of the 
Los Colorados on the western side of the Campo de Talampaya, 
their exposure on the eastern side is limited to a low ridge ex- 
tending only from the Arroyo de Agua Escondida north to the 
point where the Rio Gualo debouches onto the plain. As seen at 
the Arroyo de Agua Escondida, the Los Colorados sandstones 
are almost vertical. Only 95 meters of sediments are present; 
they are innnediately followed by Tertiary deposits having an 
angular discordance of about 20°. Since the thickness here is 
much less than in the Ischigualasto region across the campo, it 
seems certain that much of the thickness of the formation was 
eroded at some time before the deposition of the Tertiary, and 
that it is only the basal portion of this sandstone series that is 
preserved. The beds consist of soft, dull red sandstones inter- 
bedded with silts and clays of the same color. One thin but re- 
sistant layer of lime nodules forms the highest portion of the 
ridge. 

Bodenbender (1911), for no strong reason, considered these 
redbeds to be Cretaceous in age — " Cretacea superior andina." 



1966 BIO CHANARES-RIO GUALO GEOLOGY 17 

Their earlier age was recognized by later authors ; Frenguelli 
assigned them, together with still lower formations, to the Rhae- 
tic (1948, p. 302), and Groeber and Stipanicic likewise con- 
sidered them as Rhaetic in 1953. No fossils have been described 
from these beds; however, Sr. Bonaparte of the Instituo Lillo 
of Tuciiman has recently discovered dinosaur remains of Upper 
Triassic type in tlie Los Colorados beds on the west side of 
the Campo de Talampaya. It thus seems reasonable to compare 
them in general with other redbeds, generally considered Xorian, 
which conclude the Triassic series in various other parts of the 
world, such as the redbeds of South Africa, the Lufent series 
of China, the Upper Keuper of Europe, and part, at least, of the 
North American Triassic redbeds. 

TERTIARY 

Bodenbender (1911, perfil III, etc.) assumed, apparently be- 
cause the red Los Colorados sediments appear on both margins 
of the Campo de Talampaya, that these rocks directly underlay 
the Campo. This is not the case. At the mouth of the canyon 
of the Arroyo de Agua Escondida there is seen a thick series 
of Tertiary sediments to the west of the Los Colorados which are 
not merely turned, like them, into a vertical position, but are 
even overturned beyond the vertical to about 20° as a result of 
major late Tertiary or Pleistocene diastrophism. If there is 
any continuity between the Los Colorados on the two sides of 
the campo, this must be at a considerable depth. We have not 
studied tliese Tertiary sediments, which, like those of the Loma 
de los Loros farther north, are presumably of the sort cus- 
tomarily termed "Estratos Calchaquenos." 

ERUPTIVE ROCKS 

In the northwestern part of tlie Permo-Triassic basin, volcanic 
rocks are prominent. De la ]\lota, for example, in his unpub- 
lished works on Cerro Bola describes thick intrusions of mela- 
phyrite; Frenguelli (1944a) likcAvise describes similar intrusions 
in the sediments of Cerro Guandacol; and farther to the south 
on the western basin margin, Cerro Rajado owes its name to 
its partial bisection by a conspicuous but eroded dyke ; still fur- 
ther eruptives are present in tlie Cerro ^lorado area. In the 
area here studied, in contrast, there are few evidences of such 
activity, but there are present three dykes : 



18 BREVIORA No. 252 

(1) In the southwestern portion of the mapped area there 
appears a dyke, running in an ENE direction, which forms a 
notable elevation, the Mog'ote de Agua Escondida ; a lesser ele- 
vation lies just to the east of this and, following a gap, two fur- 
ther exposed sections lie in the Piano del Gualo. The dyke is 
composed of green crystalline material Avhich is less resistant to 
erosion than the metamorphosed sediments immediately adjacent. 
The Agua Escondida is not a true spring, but a catch-basin for 
water in the hollow formed in the dyke by erosion in the low 
spot between the mogote and the lesser elevation to the east of it. 
Part way up the side of the mogote there adheres a mass of 
modified sedimentary rock including a portion of the contact 
between the Talampaya and Tarjados formations, thus giving 
a maximum date for dyke formation. However, the mogote is 
capped by a basalt-like rock, dense, brown in color, and giving a 
conchoidal fracture, indicative of its extrusive nature and sug- 
gesting the formation of the dyke posterior to erosion of 
higher, Triassic sediments in the basin, presumably in Tertiary 
times. 

(2) In the northeastern portion of the Gualo Basin shown in 
our map are modest exposures of a SW-NE trending dyke pene- 
trating Los Rastros sediments. 

(3) Along a high divide between the Gualo and Chaiiares 
drainages, a few kilometers east of the Campo de Talampaya, are 
two conspicuous peaks, close together, and rising abruptly about 
100 meters above the Los Rastros beds at their bases, which may 
be termed Los Mogotes Mellizos. These have superficially the 
appearance of volcanic plugs, but seem to be in reality portions 
of a dyke. The upper portion of the intrusive material, appar- 
ently basaltic in nature, contains considerable fragments of 
sedimentary rocks apparently originating in surrounding beds. 
The baking of the sediments about the base of the dyke is not 
extensive ; the situation as a whole suggests that little if any ex- 
trusive surface action occurred. 

LITERATUEE CITED 

Alba, E. de 

19.")4. Descripeion geologica de la Hoja 16c, Villa Union (Provincia de 
La Rioja). Bol. Dir. Nac. Min. (Argentina), 62: 1-81. 

BODENBENDER, G. 

18!)6. Beobachtnngen iilier Devon- niid Goiidwaii.'i-Seliieliten in der 
argentinisehen Republik. Zeitschr. Deutscli. Geol. Ges., 48: 743- 
807. 



1966 RIO CHANARES-RIO GUALO GEOLOGY 19 

1897. Devono y Gondwana en la Eepublica Argentina. Bol. Acad. 
Nac. Ciene. Cordoba, 15: 201-252. 

1902. Contribuciou al eonocimiento de la PrecordiUera de San Juan, 
de Mendoza, j de las Sierras centrales de la Eepublica Argen- 
tina. Bol. Acad. Nac. Cienc. Cordoba, 17: 203-261. 

1911. Constitucion geologica de la parte mei'idional de la Eioja y re- 
giones limitrofes (Eepublica Argentina). Bol. Acad. Nac, 
Cienc. Cordoba, 19: 1-220. 

191G. El Nevado de Famatina. Bol. Acad. Nac. Cienc. Cordoba, 
21: 100-182. 

192-i. El Calcliaqueiio y los cstratos de la Puna de Penck. Bol. Acad. 
Nac. Cienc. Cordoba, 27: 405-468. 

FiDALGO, F. 

1963. Algunos rasgos tectonicos y geoniorfologicos de la Sierra de 
Sanogasta-Vilgo (Prov. de la Eioja). Eev. Asoc. Geol. Argen- 
tina, 18: 139-153. 
Feenguelli, J. 

1944a. Apuntes acerca del Paleozoic© Superior del noroeste argentine. 
Eev. Mus. La Plata, (NS) Sec. Geol., 2: 213-265. 

1944b. La serie del llamado Eetico en el oeste argentine. Notas Mus. 
La Plata, Geol., 9: 261-269. 

1944c. Excursiones al territorio del Neuquen y a las provincias de 
Mendcza y La Eioja. Eev. Mus. La Plata, (NS) Sec. Oficial, 

1943 (1944): 87-102. 

1945. Yiaje a la PrecordiUera. Eev. Mus. La Plata, (NS) Sec. Oficial, 

1944 (1945) : 157-169. 

1946. Consideraciones acerca de la "serie de Paganzo" en las provin- 
cias de San Juan y La Eioja. Eev. Mus. La Plata, (NS) Sec. 
Geol., 2: 313-376. 

1948. Estratigrafia y edad del llamado Eetico en la Argentina. Gaea, 
An. Soc. Argentina Est. Geogr., 8: 159-309. 
FURQUE, G. 

1963. Descripcion geologica de la Hoja 17b-Guandacol. Bol. Dir. Nac. 
Geol. Min. (Argentina), 92: 1-104. 
Groeber, p. F. C. 

1940. Descripcion geologica de la Provincia de la Eioja. In : Aguas 
Minerales de la Eepublica Argentina, 6: 17-29. 
Groeber, P. F. C. and P. N. Stipanicic 

1953. Geografia de la Eepublica Argentina, II (Priniera Parte) : 
Triiisico. Bueros Aires. Pp. 13-141. 
Heim, a. 

1946. El carbon de la uiina "La Ncgra," Villa Union (La Eioja) y 
su posicion tectonica. Bol. Dir. Min. Geol. (Argentina), 60: 
1-19. 

1947. El carbon del Eio Huaco (Provineia de San Juan) y su posicion 
tectonica. Bol. Dir. Min. Geol. (Argentina), 62: 1-18. 



20 BREvroRA No. 252 

1949. Estudio geologico del carbon "Eetico" y del Valle de la Pefia 
(provincias de San Juan y La Rioja). Bol. Dir. Gen. Ind. Min., 
69: 1-31. 
Lehman, J. -P., C. Chateau, M. Laurain, M. Nauche 

1959. Paleontologie de Madagascar. XXVIII. Les poissons de la 
Sakamena moyenne. Ann. Paleont., 45: 177-219. 

Stelzner, a. W. 

1885. Beitrage zur Geologie und Palaeontologie der argentinischen 
Republik. I. Geologischer Theil. Cassel & Berlin, 329 pp. 
Turner, J. C. M. 

1960. Estratigrafia del tranio medio de la Sierra de Famatina y 
adyaciencias (La Rioja). Bol. Acad. Nac. Cienc. Argentina 
(Cordoba), 42: 77-126. 

Watson, D. M. S. 

1942. On Permian and Triassic tetrapods. Geol. Mag., 79: 81-116. 

(Received 6 May, 1966.) 



BREVIORA 

Mmseitaim of Comparative Zoology 

Cambridge, Mass. Xovkmbkk 4, 190(1 Number 253 

A NEW IIISPANIOLAX GECKO 

By Richard Thomas 

10,000 S.AA'. 84th St., Miami, Florida 33143 



Recent colleetino- in Haiti yielded a single specimen of an un- 
described species of S phacrodactijlus from the western part of 
the Massif de la Hotte on the Tiburon Peninsula. I wish to ex- 
press my appreciation to Dr. Albert Schwartz, Avho sponsored 
my collecting- in Haiti (February to April of 1966), which re- 
sulted in the discovery of this new species. The abbreviation 
MCZ denotes the Museum of Comparative Zoology, and ASFS 
denotes the Albert Schwartz Field Series. 

In reference to its most salient distinguishing character, this 
new gecko maj- be known as 

Sphaerodactylus elasmorhynchus ^ new species 

IIolotyp( : ]\ICZ 81119, a female, taken ca. 5 km (airline) SSE 
^larche Leon, Dept. du Sud, Haiti, at an elevation of 2600 feet 
(7f)0 meters), on 15 March 1!)()6 l)y Richard Thomas. Original 
number ASFS V9353. 

Diagnosis: A species of SpJiacruddcfj/lu.s distinguished by: 1) 
much enlarged sui)ranasals and postnasals- which abut against 
the rostral and labials to form a continuous, platelike covering 
over most of the snout; 2) a finely granular dorsal scalation; 3) 
large, smooth, imbricate ventral scales; and 4) a dorsal colora- 
tion of small orange ocelli on a dark brown ground color. The 
first character distinguished elasmorhyncluis from all other 
known species of Sphaerodactylus. 

Description of holotypc (Fig. 1) : Snout-vent length 17 mm; 
tail (unregenerated) 17 mm. Rostral with a median crease; post- 
nasals enlarged, roughly trapezoidal with bases towards anterior 



1 Fridii the (ireek. eluninos. plate, and rliynchos, snout. 

- .Mtlioufih I cniild not sec a definite nasal scale, even under high niagnitica- 
rion, I have called the larse scale behind the naris the post nasal because of its 
position (see Sinitli. 1'.14C., tig. '2 i } . 



BREVIORA 



No. 253 






Fig. 1. Holotype of Spliaerodactylus elasmorhyvchus: lateral and dorsal 
views of head and full dorsal view. 



edges of orbits and ventral long sides sutured with upper edges 
of first supralabials ; two large, platelike, roughly trapezoidal 
supranasals with their bases abutting against posterior edge of 
rostral and their lateral edges against dorsal edges of postnasals. 
Upper labials to mid-eye, two. Rostral, supranasals, postnasals 
and labials form platelike covering to snout. Scales of snout be- 
hind platelike covering swollen, subimbricate to imbricate, and 
keeled. Scales of top of head and on body granular, subimbri- 
cate, keeled, more swollen and more erect on flanks than mid- 
dorsally; dorsal scales, axilla to groin, 56 (an injury on the 
venter prevents the taking of midbody and longitudinal ventral 
counts) ; 1-2 large hair-bearing (3 hairs, at most) scale organs on 
the tip of each scale. Scales on anterodorsal parts of limbs flat- 
tened and imbricate. Gular scales granular; ventral scales large, 
rounded, flattened, smooth and imbricate ; transition between the 
two on posterior part of neck abrupt. Dorsal scales of tail flat- 
tened, flat-lying, acute, keeled and imbricate ; ventral scales of 



1966 NEW HISPANIOLAN GECKO 3 

tail flattened, flat-lying, rounded and smooth, niidventral row 
enlarged. Fourth toe lamellae 10. 

Coloration: In life the type was dark brown with dark-edged, 
orange-centered ocelli on body and tail. The ocelli are arranged 
in six irregular longitudinal rows on the body ; those of the 
dorsolateral rows are largest and most prominent ; those of the 
paramedian and ventrolateral rows are smaller and less distinct. 
The occiput bears a pair of transversely elongate light markings, 
and a faint pair of forked, dark, postocular stripes was noted in 
life. The sacrum has a pair of enlarged, diagonal light markings 
whose dark medial margins unite to form a U-shaped sacral 
figure. Two dorsal rows of ocelli occur on the tail. The throat is 
lightly pigmented and has faint lighter longitudinal streaks ; the 
venter is very heavily pigmented but has light central areas to 
many of the scales. 

Comparisons: The enlarged platelike scales of the snout dis- 
tinguish this species from all other known Sphaerodactyliis. Also 
distinctive are the size difference between the dorsal and ventral 
scales and the relatively aln-upt transition from gular granules 
to ventral scales. Head length (ear to tip of snout) has been 
plotted against snout-vent length for five species of SpJiaerodac- 
tylus in addition to the type of elasmorhynchus (Fig. 2). The 
value plotted for the type of elasmorhynchus falls with the 
values for adult specimens of small species (^S*. n. nicholsi Grant 
and 8. parthenopion Thomas), rather than with the values plot- 
ted for juvenile specimens of larger species {S. cinereus Wagler, 
S. roosevelti Grant, S. difficilis Barbour). Thus it appears that 
the type of dasmorhynchus is an adult or subadult, not a juve- 
nile, and represents a small-sized species. 

The distinctness of 8. elasmorhynchiis prevents an assessment 
of its relationships at this time ; it is not obviously a member of 
any of the recognized groups of the genus. iVlthough elas- 
morhyiicJiiis has granular dorsal scales, it dift'ers strongly from 
all members of the predominantly granular scaled dccoratus 
group in the characters mentioned above, and apparently in size 
also (for a review of the decoratus group, see Thomas and 
Schwartz, 1966). Of the known species of Hispaniolan sphaero- 
dactyls, *S'. cinereus is the only other one having granular dorsal 
scales, but it too has no evident affinities with elasmorhynchus. 
The large hair-bearing scale organs of elasmorhynchus are char- 
acteristic of other granular scaled Sphaerodactylus ; the small 
number per scale is probal)ly a reflection of the small size of the 
si)ecies. 



BREVIORA 



No. 253 





10 - 
9- 












*U 


s 


8 - 










♦ 




z 


7- 
6- 






♦♦ 

^ ,^o" 


♦ 
♦ 

X 


A A 




I 


5 






♦a^ 








H 
o 

z 

UJ 


4- 














_J 








o •> 








o 








0» 








< 






O 










Ul 






o 










X 


3- 
2- 
1- 


• 













10 



20 



30 



SNOUT-VENT MM 

Fig. 2. Scatter diagiain of head length (anterior border of ear to tip of 
snout) versus snout-vent length for six species of Sphaerodactylns: solid 
rhombs, S. ciiicn-Hs; hollow triangles, S. (lifficilis; crosses, <S'. rooscvelti ; 
hollow circles, >S'. n. iiidioisi ; solid circles, «S'. pariheno'pion ; solid hexagon, 
type of »S'. elasmorltynchu.s. For each species, aside from elasmorliynchtis, 
the range of values plotted includes near-hatchling juveniles through mature 
adults. 



Grant (1957) has shown that the enigmatic ^S*. nigropunctatus 
Gray, known from a singk^ type of unknown provenance, is a 
o'ranular scaled form. However, references to the snout scala- 
tion of nigropuiictafus in Grant (1957), and a drawing of its 
snout (Boulenger, 1885, pi. 18), indicate that it has the normal 
Sphacrodocfylufi snout scale configuration; the type is also large 
in size. 



1966 NEW HISPAXIOLAN GECKO 5 

Ronarks: The tyi)e of N. el(is))iorhyncJius was found among 
the roots of a standing,' but rotten tree on the side of a ravine 
filled with a jumble of limestone rocks and boulders. Although 
the surrounding countryside was much eut over, the ravine was 
overgrown with mesie vegetation. Also taken at this locality 
were specimens of the relatively little known Anolis mo)iticola 
Shreve. No other sphaerodactyls were found at this locality. Of 
the named Hispaniolan species, nS. copci Steindachner and N. 
cinereus are the only ones occurring in this general region of 
Haiti. 

LITERATURE CITED 

BOULENGER, GeORGE ALBERT 

ISS."). Catalogue of the lizards in the British Museum (Natural His- 
tory). Loiulou. 2d ed., vol. 1, pp. i-xii, 1-436. 
Grant, Chapman 

19.")7. The status of Spliacrodacti/Ius nigyopHUctatus, a neotropical 
gecko. Nat. Hist. Misc., Chicago Acad. Sci., No. 158, pp. 1-4. 
Smith, Hobart M. 

1946. Handbook of lizards. Comstock Pub. Co., Ithaca, New York, pp. 
i-xxi, l-.3.")7. 
Thomas, Richard, and Albert Schwartz 

1966. The Sphaerodactylus decora t us complex in the West Indies. 
Brigham Young Univ. Sci. Bull., Biol. Ser. (In press.) 



(Received 23 May, 1966.) 



E V I O R A 

Miaseiiioi of Comparative Zoology 

Cambridge, ^^Ias^. Xoven:ber 4, 19GG Number 254 

PRELIMINARY DESCRIPTIONS OF NEW 
HONEY-EATERS 

(AYES, MELIPHAGIDAE) 

By Finn Salomonsen ^ 



For many years I have been engaged in a study of the Meli- 
phagidae, mainly based on the outstanding collections in The 
American Museum of Natural History, New York (AMNH). 
During this study it appeared that a number of forms were un- 
described ; these forms are named and described in the present 
paper. The descriptions must be regarded as preliminary, and 
more detailed documentation will appear in a monograph of the 
Honey-Eaters, now in preparation. The descriptions had to be 
published now, however, in order to incorporate the new forms in 
my review of the Meliphagidae in Peters' Checklist of Birds of 
the World, Yolume 12, which is now in press. 

ilELILESTES MEGARHYNCIIUS BRUNNEUS UCW SubspecieS 

Type: AMNH No. 294369, adult male; Siwi, Arfak Moun- 
tains, New Guinea ; I\Iay 2, 1928 ; Ernst Mayr collector. 

Diagnosis: Darker and more lu'ownish than nominate mega- 
rhijuchns (type locality: Aru Islands). The upper parts are 
warm brown without the olive tinge of mcgarhynchus, the under 
parts are darker brownish grey than mcgarliyncJius, not so olive, 
and the abdomen and under tail-coverts are darker and warmer 
cinnamon brown. Also, the upper parts of the head are darker 
brown with broader blackish striation. 

Range: Misol and SalaAvati in the western Papuan Islands, 
Yogelkop and Onin Peninsula in northwestern New Guinea, in 
the Weyland ilountains tending towards nominate megarhyn- 
ch us. 



1 Zoolo,uical .^lu^^t■Ulll, Copwiliuguu. 



2 BREVIORA No. 254 

Oedistoma pygmaeum waigeuense new subspecies 

Txjpe: AMNH No. 301021, adult male; AVaigeu Island; June 
5, 1931 ; G. Stein collector. 

Diagnosis: Differs from nominate pygmaeum (type locality: 
Arfak Mountains, New Guinea) in having crown and nape dark 
grey, strongly contrasting with green upper parts (in nominate 
pygmaeum crown green, only slightly darker than upper parts), 
throat and breast paler grey than in nominate pygmaemn. Pro- 
portions smaller. Wing length of adult males 45-46 mm, in 
pygmaeum 47-51 mm; bill 11.8-12.8 mm, in pygmaeum 13-14 mm. 

Range: AVaigeu Island, in the western Papuan Islands. 

Oedistoma pygmaeum flavipectus new subspecies 

Type : AMNH No. 422241, adult male ; Wuroi, Oriomo River, 
British Papua (western division), southern New Guinea; Feb. 
10. 1934; R. Archbold and A. L. Rand collectors. 

Diagnosis: Upper parts distinctly paler green than in nomi- 
nate pygmaeum, crown and nape of the same green color as rest 
of upper parts, under parts strongly suffused with yellow, abdo- 
men and flanks with more yellow than in nominate pygmaeum 
and waigeuense. Proportions as in the nominate form. 

Range: Southern New Guinea, from Etna Bay to Milne Bay. 

Oedistoma pygmaeum olivascens new subspecies 

Type: Mus. Comp. Zool., Cambridge, Mass., No. 167910, adult 
male ; Upper AVatut River, southeastern New Guinea ; July 27, 
1932; H. Stevens collector. 

Diagnosis: Very similar to nominate pygmaeum, but under 
parts with a conspicuous greyish olivaceous tinge and upper 
parts slightly darker. 

Range: Northern coast of southeastern New Guinea, from 
Milne Bay to Iluon Peninsula. 

Lichmera incana mareensis new subspecies 

Type: AMNH No. 337641, adult male; Mare Island, Loyalty 
Islands; Nov. 8, 1937; L. Macmillan collector. 

Diagnosis: Yerj similar to L. i. poliotis (type locality: Loy- 
alty Islands, restricted here to Lifu Island), but under parts 
darker grey, and lower breast and abdomen with much more 
yellow. Not differing from poliotis in measurements. 

Range: Mare Island, Loyalty Islands. 



1966 NEW MELIPHAGIDAE 3 

LiCHMERA ixcAXA GRiSEOViRiDis iiew subspecies 

Type: AMNH No. 212924, adult male; Lopevi Island, New 
Hebrides; Au"'. 9, 1926; J. G. (\)rreia (Tbe Wbitney South Sea 
Expedition) collector. 

Diagnosis: Very similar to /.. i. flavotincfa (type locality: 
Erromanofa, New Hebrides), but paler throughout; crown paler 
srey, upper ]iarts paler and less brownish green, and, particu- 
larly, under parts paler with less yellowish olive on lower l)reast 
and abdomen. Differs from marecnsis in its much larger propor- 
tions, but plumage coloration very similar, although the under 
parts are slightly darker grey with the yellow tinge less pro- 
nounced. 

Range: Central New Hebrides (Efate, Makura, Mai, Tongoa, 
Epi, Lopevi, Pauuma, Ambrym, and Malekula). 

Myzomela nigbita ntgebrima new subspecies 

Type: AMNH No. 422677, adult male; Long Island, off the 
northeastern coast of New Guinea ; Dec. 2, 1933 ; W. F. Coultas 
(The Whitney South Sea Expedition) collector. 

Diagnosis: Similar to M. n. ernstmayri (type locality: ^lanu 
(= Allison) Island, west of Ninigo Islands), but darker through- 
out. Under wing-coverts black like the axillaries, inner linings 
of remiges darker greyish and almost indistinguishable from the 
greyish black ground color of the remiges, plumage of adult 
males more shining black and with darker greyish black feather 
bases. Proportions distinctly larger than in ernstmayri. Wing 
length of adult males 77-79 nun, in ernstmayri 15-11 ; adult fe- 
males 71 mm, in ernstmayri 66-69 mm. 

Range: Long Island, off the northeastern coast of New 
Guinea. 

Myzomela cardinalis asuncionis new subspecies 

Type: x\.MNH No. 692934, adult female; Asuncion Island, in 
the northern Marianas Islands; June 1904; Owston's Japanese 
collectors. 

Diagnosis: Adult males do not differ from M. c. saffordi (type 
locality: Guam Island, southern [Marianas Islands) except by 
larger proportions. Wing length of adult males 76-79 mm, in 
saffordi 70-75 mm. Adult females differ strikingly from saffordi 
by having the red color strongly reduced, and by having light, 
greyish yellow under parts. Upper jiarts dark olive brown, with 



4 BREVIORA No. 254 

the feathers of back and rump (but not mantle and neck) 
broadly edged with red, feathers of forehead and nape edged 
with dull red, chin dull red, under parts greyish yellow with a 
reddish wash on throat and chest. The only adult female ever 
collected (the type specimen) in its color pattern somewhat re- 
sembles immature birds of saffordi, but in all respects appears to 
be an adult bird. 

Range: Northern Marianas Islands (Asuncion, Agrihan, Pa- 
gan, Alamagan), on Saipan being intermediate between this 
form and saffordi. 

Meliphaga mimikae rara new subspecies 

Type: Ai\INH No. 342965, adult male; Bernhard Camp, Iden- 
burg River, 50 meters altitude, northwestern New Guinea ; April 
26, 1939; The Archbold Expedition collection. 

Diagnosis: Differs strikingly from the two other known forms 
of this species, granii (type locality: Mafulu, mountains of 
southeastern New Guinea) and nominate mimil-ae (type locality: 
Mimika River, southern New Guinea) in having uniform pale 
yellowish grey under parts (in mimikae and granti dark grey 
with longitudinal yellow streaks), abdomen and flanks much 
paler greyish yellow, and upper parts distinctly clearer and 
paler green, not so brownish olive-green. Proportions, particu- 
larly tarsus, much smaller. Wing length of rara (adult male) 84 
mm, of iniinilae 86-88 mm, of granti 89-95 mm; tarsus length in 
rara 19 mm, in mimikae 21-22 mm, in granti 22-23.5 mm (only 
adult males measured ) . 

Range: Only the type known. 

Remarks: This form belongs to the difficult analoga species 
group. It agrees with M. mimikae in its general diagnostic 
characters (dark coloration, especially on upper parts, ochra- 
ceous under wing-coverts and pale huffish inner linings on 
remiges), but in other respects differs so much that it may pos- 
sibly represent a separate species. 

Meliphaga analoga papuae new subspecies 

Tijpe: AMNH No. 422342, adult male; AVuroi, Oriomo River, 
British Papua (western division), southern New Guinea; Jan. 
28, 1934; The Archbold Expedition collection. 

Diagnosis: Yevy similar to nominate analoga (t^'pe locality: 
Triton Bay, western New Guinea), but under parts distinctly 
paler grey with a somewhat stronger yellowish tinge ; upper 



1966 NEW MELIPHAGIDAE 5 

parts paler, more clear greyish green, not so olive green ; ear- 
tufts paler, yellowish white ; under wing-eoverts paler whitish. 
Proportions slightly smaller. Wing length of adult males 78-83 
mm, in nominate analoga 81-87 (one 91) mm. 

Range: Southern New Guinea from the Fly River district 
eastward to Hall Sound. 

Meliphaga axaloga connectens new subspecies 

Type: AMNH No. 267966, adult male; Madang, Astrolabe 
Bay, northeastern New Guinea ; Aug. 30, 1928 ; R. H. Beck col- 
lector. 

Diagnosis: Differs from flavicla (type locality: Japen Island) 
in having under parts paler grey with much less yellow, upper 
parts paler and clearer green, not so bright citrine green, and 
ear-tufts slighth^ paler yellow. Also smaller proportions than 
topotypical flavida. Differs from stevensi (type locality: Upper 
Watut River, northeastern New Guinea) by greyish white, not 
yellowish inner linings on remiges, paler yellow under wing- 
coverts, and dusk}' olive green, not blackish, lores and postocular 
spot, while the general coloration of the plumage is very similar 
to that of stevensi. Proportions slightly larger than in stevensi, 
the new subspecies in this respect forming the connecting link 
between jiavida and stevensi. Wing length of adult males of 
topotypical flavicla, 82-85 mm, of connectens 79-83 mm, of 
stevensi 77-80. 

Bayige: Lowlands of northern New Guinea from Wewak east- 
ward to Huon Gulf. 

Remarks: The new form comes nearest in coloration to the 
remote nominate analoga from Vogelkop Peninsula, but has the 
upper parts slightly clearer and paler grey; it is also smaller, 
and has a smaller bill. 

Meliphaga versicolor vulgaris new subspecies 

Type: AMNH No. 268962, adult male; Finschhafen, Huon 
Peninsula, northeastern New Guinea ; Feb. 7, 1929 ; R. H. Beck 
collector. 

Diagnosis: Similar in coloration to sonoroides (type locality: 
Waigeu Island), but much smaller. Wing length of adult males 
100-108 mm, in sonoroides 110-114 mm. Differs from intermedia 
(type locality: Samarai Island) and nominate versicolor (type 
locality: Cape York, northern Queensland), which have similar 
small proportions, by the pale, whitish, not yellow under parts. 



6 BREVIORA No. 254 

Range: Japen Island in Geelvink Bay; coastal zone of north- 
ern New Guinea from Geelvink Bay to Huon Gulf; Fergusson 
Island in the D'Entreeasteaux Archipelago. 

Meliphaga fusca deserticola new subspecies 

Type: AMNH No. 695517, adult male; Margaret River, Kim- 
berley Division, northern Western Australia ; April 20, 1902 ; 
I. T. Tunney collector. 

Diagnosis: Palest of all forms within the flavescens group of 
M. fusca, differing from flavescens (type locality: Derby, north- 
ern AVestern Australia) by having paler grey upper parts with- 
out any brownish tinge, paler yellow under parts with very faint 
striation, paler yellow front and crown (and yellow color more 
restricted), and duller olive edges on outer web of remiges. 
Differs from zanda (type locality: Normanton, Gulf of Carpen- 
taria) especially by the much paler grey upper parts and much 
duller olive edges on remiges ; under parts only slightly paler. 

Range: Margaret River, in arid interior of northern Western 
Australia. 

Meliphaga penicillata interioris new subspecies 

Tijpe: AMNH No. 343828, adult male; Ward River, south of 
Charleville, southwestern Queensland. Australia; March 1, 1940; 
L. Macmillan collector. 

Diagnosis: Throughout paler than nominate penicillata (type 
locality: interior of New South Wales, restricted here to Wagga- 
Wagga) ; under parts with much paler greyish throat and chest, 
and with whitish, not greyish, abdomen and flanks; upper parts 
paler and colder greyish brown, forehead and crown only slightly 
suffused with pale yellow, cheeks and ear-coverts yellow, without 
the olive greenish tinge found in nominate penicillata; black ear- 
stripe reduced or absent. xVlso slightly smaller. Wing length of 
adult males of topotypical penicillata 85-92 mm, of interioris 
83-85, adult females of penicillata 79-83, of interioris 76 mm. 

Range: Northwestern New South Wales, northward to south- 
central Queensland. 

Meliphaga flaviventer tararae new subspecies 

Type: AAINH No. 428247, adult male; Tarara, Wassi Kussa 
River, 90 miles west of Daru, British Papua (western division), 
southern Ncav Guinea; Dec. 8, 1936; The Archbold Expedtition 
collection. 



1966 NEW MELIPHAGIDAE 7 

Diagnosis: Much paler throughout than saturatior (type lo- 
cality: Aru Islands) and (jiulianettvi (type locality: Aroa River, 
southeastern New Guinea), and, further, differing strikingly by 
possessing a broad and conspicuous streak of white feathers lead- 
ing from gape to the white supraauricular spot. This white 
streak is completely missing in giulianettii and in the mainland 
populations of saturatior, but occasionally is present in Aru 
Islands birds. In tararae the upper throat is pale grey, lower 
throat and chest pale greyish brown, breast and abdomen pale 
buff (not rich cinnamon), flanks cinnamon (not chestnut), 
mantle and back distinctly paler earth-brown than in saturatior 
and giulianettii, and upper parts of head not so dark brown. 

Range: Southern coast-land of New Guinea, between middle 
and lower Fly River, westward to about Digul River. 

Meliphaga obscura viridifroxs new subspecies 

Type: Acad. Nat. Sci. Philadelphia No. 132851, adult female 
(labelled male) ; Bamoskaljoe, Karoon, Tamrau Mountains, Vo- 
gelkop Peninsula, 2300 feet altitude, northwestern New Guinea; 
March 1, 1938; D. Ripley (Denison-Crockett South Pacific Ex- 
pedition) collector. 

Diagnosis: Dift'ers from nominate ohscura (type locality: Mt. 
Scratchley, southeastern New Guinea) in having front and crown 
green, like nape and upper parts, while in ohscura front and 
crown are grey, contrasting Avith nape. 

Range: Mountains of Vogelkop Peninsula, northwestern New 
Guinea. Specimens from the Weyland Mountains are interme- 
diate between viridifrons and nominate ohscura. 

Melithreptus laetior normantoniensis new subspecies 

Type: AMNH No. 691532, adult male; Normanton, northwest- 
(M-n Queensland, Australia; Jan. 31, 1914; R. Kemp collector. 

Diagnosis: Dift'ering from nominate laetior (type locality: 
Lake Eyre district, central Australia) only in its smaller propor- 
tions. Wing length of adult males 81-83 mm, in nominate laetior 
84-90 mm ; adult females 77-79 mm, in nominate laetior 82-84 
mm. 

Range: Northwestern Queensland, south of Gulf of Carpen- 
taria. Specimens from Cooktown and Cairns area, northeastern 
Queensland, are probably referable to this form. 



8 BREVIORA No. 254 

Melithreptus laetior parus new subspecies 

Type: AMNH No. 691546, adult male; Exmouth Gulf, mid- 
Western Australia; June 21, 1902; T. Carter collector. 

Diagnosis: Differing from nominate laetior (type locality: 
Lake Eyre district, central Australia) in its larger and more 
robust bill and, on the average, slightly longer wings. Bill 
length (measured from skull) in adult males 18.0, 18.5 mm, in 
nominate laetior 16-17 mm. 

Range: Only known from the type locality. 

Pycnopygius ixoides cinereifrons new subspecies 

Type: AMNH No. 428278, adult male; two miles below junc- 
tion of Black and Palmer rivers, 100 meters altitude, British 
Papua (western division), southern New Guinea; July 13, 1936; 
The Archbold Expedition collection. 

Diagnosis: Differs strikingly from nominate ixoides (type 
locality: Sorong, Vogelkop Peninsula) in having grey feather 
edges on front, crown, and occiput, contrasting with the color of 
the upper parts (in ixoides olive green feather edges like those 
on upper parts) ; further, in having lighter brown ground color 
and paler olive-green feather edges on upper parts, light cinna- 
mon throat contrasting with greyish brown breast and abdomen 
(in ixoides throat greyish brown like breast), and deeper and 
brighter cinnamon axillaries, under wing-coverts and inner lin- 
ings on remiges, the axillaries and smaller under wang-coverts 
being almost bright rusty, almost as colorful as in the south- 
eastern form finschi. Proportions as in nominate ixoides. 

Range: Southern New Guinea from Mimika River eastward 
to upper Fly River. The population of the Weyland Mountains 
belongs to nominate ixoides, although slightly tending towards 
cinereifrons. 

Philemon citreogularis carpentariae new subspecies 

Type: AMNH No. 696823, adult male; Normanton, north- 
western Queensland, Australia ; May 1914 ; R. Kemp collector. 

Diagnosis: In coloration, hardly differing from nominate 
citreogularis (type locality: interior of New South Wales) and 
johnstoni (type locality: Johnston River, northern Queensland), 
only very slightly paler generally, but differing from nominate 
citreogularis in having a much larger bill, and from johnstoni in 
having much greater wing length. Wing length of adult males: 



196G NEW MELIPHAGIDAE 9 

citreogidaris 136-140 mm, carpentariae 135-141 mm, johnstoni 
127-131 mm : bill length : cifreogularis 30.5-31.5 mm, carpen- 
tariae 33-34 mm, johnstoni 33-34 mm. 

Range: Southern coast of Gulf of Carpentaria, northwe.stern 
Queensland. 

Remarks: It is possible that the name pseudonymus Mathews 
and Neumann, 1939 (tj^pe locality: Cloncurry, northern Queens- 
land) can be used for this form. The subspecies pseudonymus 
was based on irrelevant color characters and no measurements 
were given. I have seen only one specimen (adult male) from 
Cloncurrj". It had a wing length of 130 mm, but was, unfortu- 
nately, worn, and may have measured 132 mm in fresh plumage. 
The bill was broken, but appeared to have been large. 

Philemon novaeguineae fretensis new subspecies 

Type: AMNH No. 330277, adult male; Delena, Hall Sound, 
British Papua (central division). New Guinea; May 28, 1929; H. 
Hamlin collector. 

Diagnosis: Differs from nominate novaeguineae (type lo- 
cality: Triton Bay, southwestern New Guinea), 'brevipennis 
(type locality: Utakwa River, southern New Guinea) and aruen- 
sis (type locality: Aru Islands) in being distinctly paler slate 
grey on upper parts and slightly paler grey on under parts, and, 
in addition, differs from the neighboring hrevipennis in being 
much larger. Wing length of adult males: topotypical hrevi- 
pennis 142-144 mm, fretensis 158 mm (same length in three 
specimens). 

Range: Southern New Guinea from middle Fly River east- 
ward along south coast of southeastern New Guinea to Milne 
Bay. 

Philemon novaeguineae trivialis new subspecies 

Type: AMNH No. 697135, adult male; Collingwood Bay, 
north coast of southeastern New Guinea ; June 28, 1894 ; A. S. 
^leek collector. 

Diagnosis: Same general coloration as suhtuberosus (type lo- 
cality: Fergusson Island, D'Entrecasteaux Archipelago), but 
slightly lighter (the suhtubcrosus from the Trobriand Islands 
are the darkest ) , with distinctlj^ longer bill and, especially, with 
larger casque on bill. Length of bill in adult males of trivialis 
47.5, 50.2 mm, in siihtnherosus from the D'Entrecasteaux Archi- 
pelago 43-44 mm, and from the Trobriand Islands 44-47.5 mm. 



10 BREVIORA No. 254 

Range: Northern coast of southeastern New Guinea, known 
from Collingwood Bay and Knmnsi River. 

Philemon corniculatus clamans new subspecies 

Type: Los Angeles County Museum No. 15117, adult male; 
Wallaroo Gorge, Carnarvon Range, southeastern Queensland, 
Australia; July 2, 1954; K. E. Stager and J. B. Davidson col- 
lectors. 

Diagnosis: Intermediate in size between the large nominate 
corniculatus (type locality: New South Wales) and the small 
ellioti (type locality: Mt. Elliot, northern Queensland). Wing 
length of adult males: corniculatus 159-166 mm, clamans 154- 
156 mm, ellioti 137-147 mm. 

Range: Southeastern Queensland. 

Melidectes puscus gilliardi new subspecies 

Type: AMNH No. 705793, adult male; ^It. Wilhelm, Bismarck 
Range, 11,000 feet altitude, east-central New Guinea; June 7, 
1950; E. T. Gilliard collector. 

Diagnosis: Same coloration as occidentalis (type locality: 
Wichmann Mountains, central New Guinea) and nominate fus- 
cus (type locality: Wharton Range, southeastern New Guinea), 
differing from occidentalis in longer wing and bill, and from 
nominate fuscus in shorter bill and tarsus. The bill/tarsus ratio 
in the three forms is noteworthy; in occidentalis the bill is 
much shorter than the tarsus, in gilliardi it is equal in length 
with the tarsus, and in nominate fuscns it is longer than the 
tarsus. Measurements of adult males (in mm) : 





WING 


BILL 


tarsus 


occidentalis 


108-113 


29.0-30.0 


31.0-33.0 


gilliardi 


114-116 


31.5-33.0 


32.5-33.5 


fuscus 


113-117 


34.0-37.0 


33.0-35.0 



Range: Bismarck ^Mountains, eastern New Guinea. 
Remarks: Named after the collector, the late Dr. E. Thomas 
Gilliard. 

Myza celebensis parvirostris new subspecies 

Type: AMNH No. 300232, adult male; Mt. Tanke Salokko, 
2000 meters altitude, ]\lengkoka ]\Its., southeastern Celebes ; July 
22, 1931; G. Heinrich collector. 



I 



1966 NEW MELIPHAGIDAE 11 

Diagnosis: Differs from nominate celehensis (type locality: 
Bone Mts., Gorontalo, northern Celebes) in its much smaller 
proportions, especially its smaller bill. Wing length of adult 
males : nominate celehensis (from Latimodjong Mts., central 
Celebes) 85-87 mm, parvirostris 81-83 mm; bill length: nominate 
celehensis 32-34 mm, parvii'ostris 28-30 mm. No material from 
the type locality of celehensis has been examined, but a series 
from Ile-Ile, northern Celebes, is similar to the Latimodjong 
birds in measurements, although not quite so big. 

Bange: Mengkoka ]\Iountains, southeastern Celebes. 

PtIYLIDONYRIS NOVAEHOLLANDIAE CAUDATA neW SubspecieS 

Type: AMNH No. 692106, adult male; King Island, Bass 
Strait, Australia; April 25, 1914; T. Tregellas collector. 

Diagnosis: Differs from all other forms of this species by its 
distinct reduction of the yellow color in wings and tail, and by 
its larger proportions, particularly the longer tail. Length of 
tail in adult males of caudata 83-91 mm, compared A^dth 75-83 
mm in all other subspecies. 

Range: King Island and Flinders Island in Bass Strait, 
Australia. 

ACANTKCRHYNCHUS TENUIROSTRIS TROCHILOIDES UCW SUbspCcicS 

Type: AMNH No. 703283, adult male; Bunya Mountains, 
2000 feet altitude, southeastern Queensland, Australia ; Oct. 3, 
1940; L. Macmillan collector. 

Diagnosis: Differs from nominate tenuirostris (type locality: 
New South Wales) by having under parts (below the black pec- 
toral band) much paler cinnamon, the band across nape dull 
rusty, not deep chestnut, and the mantle olive greyish green 
with a rusty tinge, not dark warm rusty brown; differs from 
cairnsensis (type locality: Cairns, northern Queensland) by 
having throat chestnut, not pale vinous pink. 

Range: Kestricted to the isolated pockets of rainforest in 
southeastern Queensland. 

ACANTPIOEIIYNCHUS TENUIROSTRIS REGIUS IICW SUbspCcicS 

Type: AMNH No. 693497, immature male; King Island, Bass 
Strait, Australia; April 22, 1914; T. Tregellas collector. 

Diagnosis : Only the type specimen known, an immature male, 
differing from all other subspecies by the very long bill, measur- 
ing 32 mm, and the extremely long tarsus, measuring 21.5 mm. 



12 BREVIORA No. 254 

with a wing length of 71 mm. Only one other immature male 
has been examined, belonging to nominate tenuirostris, with bill 
24.5 mm, tarsus 18.2 mm, wnng 68 mm. The largest bill of adult 
birds of any subspecies is 31 mm, reached in nominate tenuiros- 
tris; the liill of the Tasmanian form measures in adult males 
25-26.5 mm. The tarsus of adult males of the nominate form, 
which is the largest of all, measures 18-19 mm. 

IlaiKje: King Island in Bass Strait; probably also on Flinders 
Island. 

(Kec-eivcd 'IZ May liHRi.) 



BREVIORA 

Miiseem of Comparsitive Zoology 

Cambridge, Mass. December 6, 1966 Number 255 

A REVISION OF THE FOSSIL SELENODONT 

ARTIODACTYLS FROM THE MIDDLE MIOCENE 

THOMAS FARM, GILCHRIST COUNTY, FLORIDA 

By 
Vincent Joseph Maglio 

INTRODUCTION 

Since its discovery in 1931 the fossil locality at Thomas Farm 
in Gilchrist County, Florida, has proven to be the most prolific 
Miocene deposit east of the Mississippi River. The vertebrate 
fauna is impressive, both in numbers of individuals and in variety 
of forms present, and has greatly increased our knowledge of the 
Florida Miocene and its correlation with western faunas of com- 
parable age. By far the most numerous animals are the horses, 
which may be used in such correlations more successfully than any 
other group. However, it is in the Artiodactyla that the most 
highly specialized forms are to be found, some of which show no 
close relationship to known genera of the western plains. 

The strange combination of characters found in Floridatragulus 
dolichanthereus, for example, has led authors in the past to place 
this animal in three distinct families, the Hypertragulidae, the Pro- 
toceratidae, and the Camelidae. White (1940) described the first 
jaw of this species as a new genus of Hypertragulidae without dis- 
cussing its relationship or similarities with other known hyper- 
tragulids. Additional specimens (referred to this species in the 
present paper) were subsequently named as a second species of 
Floridatragulus (White, 1947), a second genus of Hypertragulidae, 
Hypermekops olseni (White, 1942), and, in the case of a partial 
palate, as a species of Syndyoceras, a protoceratid (White, 1941). 
Following White's determinations, Romer (1947) retained these 
generic names in his discussion of the mammalian fauna of Thomas 
Farm. It was not until 1957, when Professor Bryan Patterson pre- 
pared a manuscript catalogue of the Thomas Farm vertebrates in 



2 BREVIORA No. 255 

the collections of the Museum of Comparative Zoology, that a 
major reorganization of the taxa became available. In this refer- 
ence, Patterson considered all of the above specimens as a single 
camelid species. Although never published, this list has become 
incorporated, partially or completely, into almost every major 
taxonomic reference concerning late Tertiary Floridian mammals, 
most notably Ray, 1957, and Puri and Vernon, 1964. 

A similarly confused story is associated with the history of 
Nothokemas ftoridanus, which was originally made the nominate 
genus of a new hypertraguloid family, the Nothokemadidae 
(White, 1947). This, also, was referred to the Camelidae by 
Patterson. 

No new material has been added to the MCZ collection since 
1957, but it became clear that a restudy of these artiodactyls in 
the light of Patterson's list was necessary. This is the purpose of 
the present work, which, in addition, presents a more detailed de- 
scription of the species involved than has hitherto been available. 
A concurrent study of the postcranial remains has contributed to 
our knowledge of these forms. 

The present study confirms Patterson's findings except for the 
change in rank of Synthetoceras australis and the interesting addi- 
tion of an oreodont to the fauna. 

A discussion of the perplexing geological problems involving 
the bone-bearing sediments of the Thomas Farm is beyond the 
purpose and scope of this report, but a detailed review is available 
in Puri and Vernon (1964). Older accounts of the quarry itself 
may be found in Simpson (1932), and White (1942). 

I am indebted to Professor Bryan Patterson for suggesting the 
problem and for his helpful discussions and suggestions throughout 
the course of the research. I also wish to thank Dr. Malcolm Mc- 
Kenna for permitting the examination and loan of specimens in 
the American Museum and Frick collections, Mr. Morris F. Skin- 
ner for his help in the identification of the oreodont remains, and 
Mr. Beryl E. Taylor for his examination of some indeterminate 
camel molars. 

Abbreviations used in this paper are as follows: American Mu- 
seum of Natural History (AMNH), Carnegie Museum (CM), 
Florida State Geological Survey (FSGS), Museum of Comparative 
Zoology (MCZ), and Yale Peabody Museum (YPM). 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 3 

DESCRIPTION OF SPECIES 

CAMELIDAE 

FLORIDATRAGULINAE new subfamily i 

Floridatragulus dolichanthereus White 

Floridatragiihis dolichanthereus White. 1940, p. 35, pi. VII. 
Syndyoceras austral is White, 1941 (in part), p. 97, pi. XV, figs, la, lb. 
Hypermekops olseni White, 1942. pp. 11-12, pi. 8. 
Floridatragulus barbouri White, 1947, pp. 505-507, fig. 4. 

Hypodigm: MCZ 3635, type, incomplete mandible with M-^ to 
Mg and alveoli for P^ to P-, anterior crescent of right Mi and 
hypoconulid of right Mo missing, both ascending rami and the 
symphysis anterior to the C-Pj diastema missing; MCZ 4086 
(type of F. barbouri), right mandible with partial symphysis and 
complete ascending ramus, left canine, right P2 to M3 and the 
alveoli for Pi; MCZ 3711 (type of Hypermekops olseni), incom- 
plete skull with right Ii and I-, PMo M^, left P, P-i to M^, and 
the alveoli for the remainder of the dentition; MCZ 3642 (para- 
type of Syndyoceras australis), left palate with P^ to M^; MCZ 
4291, right mandible with Po to Mo and alveoH for Pi and P2; 
MCZ 7780, 30 astragah; MCZ 7781, 11 calcanea; MCZ 7782, fs 
cuboids; MCZ 7783, 11 naviculars; MCZ 7784, 41 proximal 
phalanges; MCZ 7785, 10 second phalanges; MCZ 7786, 6 ungual 
phalanges; MCZ 7787, 4 metapodials. 

Horizon and locality: Middle Miocene, Hawthorne Formation, 
Thomas Farm, 8 miles north of the town of Bell, Gilchrist 
County, Florida. 

Redescription: The referred skull (MCZ 3711) is a reconstruc- 
tion of a badly broken and incomplete specimen designated as the 
type of the supposed hypertragulid, Hypermekops olseni, by White 
(1942, pi. 8). Its most striking character is the narrow and 
very long snout, the region from P- to the anterior tip of the pre- 
maxillary being more than twice the length of the continuous pre- 
molar-molar series. The anterior teeth from P nearly to P- form 
straight rows on each side of the snout, parallel to each other. 
Just anterior to P- the palate begins to widen, becoming, at the 
level of M'^ more than twice the snout width. The skull is very 
low throughout, with the nasal bones nearly parallel to the palate 
below. Above, the frontals are expanded and send down broad, 
fiat, postorbital processes which do not reach the zygomatic arch, 
the orbit being open behind. The cranium is largely missing, but 

^ See page 9 for diagnosis. 



4 BREVIORA No. 255 

parts of the occipital and temporal regions are preserved. The 
supraoccipital forms a very high lambdoidal crest, considerably 
higher than in Paratylopus primaevus and more vertical in orien- 
tation. Although the parietals are missing, there are indications 
that the sagittal crest was also quite prominent. The occipital 
condyles are relatively narrower than in Paratylopus primaevus; 
these are confluent in MCZ 7764 but separated by a median 
groove in MCZ 3711. The auditory region is poorly known, only 
the posterolateral portion of the right bulla being present. It is 
clear that this fragment was erroneously oriented on the recon- 
structed skull and was almost certainly closer to the condyles, with 
the tympanohyal recess and groove directed posterolaterally rather 
than laterally, as restored. The bullae were also probably some- 
what larger than figured and filled with spongy bone. The molar 
series is situated well forward, with the posterior edge of M^ 
some distance in front of the orbit; the internal nares open behind 
this tooth. 

The lower jaw of F. dolichanthereus shows many specializations 
similar to those of the skull, especially in the great elongation and 
in the reduction of the premolars. The mandible is narrow an- 
terior to Pi and expands to approximately five and a half times 
that width at the angle of the jaw. The symphysis is long, ending 
some distance behind Pj, with the jaws solidly fused along its 
length. The angle of the jaw is strongly hooked in typical camelid 
fashion, and the coronoid process is long and narrow. The jaw is 
very deep posterior to Po, as in Myotylopus bathygnathus. 

The dental formula is' If, C}, P|, Mf. In MCZ 3711 U is 
a large, caniniform tooth worn posteriorly along its entire height 
and along a broader area lingually. It is situated on the side of 
the snout near the posterior margin of the elongated incisive fora- 
men. 

A long diastema separates P and P; this is the longest spacing 
within the dental row. P is also caniniform, smaller than I^ and 
somewhat compressed laterally. Its anterior and posterior mar- 
gins are raised to form a blade-like ridge. It is strongly recurved, 
and except for a small area at the very tip, is unworn. 

P is separated from P by a diastema equal to somewhat less 
than half that separating the first two incisors. This tooth is miss- 
ing on both sides but the alveoli indicate a tooth similar to, but 
smaller than P. The alveolus for the canine follows after a very 
short diastema equal to little more than its own diameter, and sug- 
gests a tooth no larger (and probably smaller) than I". After 
another equally short diastema comes the alveolus for Pi. This 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 5 

was a single-rooted tooth and could not have been more than a 
mere vestige judging from the size of the socket. 

A very long diastema nearly equal to that separating P and 1 2 
separates Pi and P-. The second premolar (Fig. 1 ) is a long, 
narrow, two-rooted tooth similar to that in Poebrotherium wilsoni 
and in its relation to the molars it is essentially unreduced from 
the condition in that species. Its three cusps are in a straight line 
and are connected to each other by a sharp, high ridge. A weak 
basal cingulum is present along the anterior and posterior of the 
lingual surface and a well developed parastyle runs the height of 
the tooth. 

PS is preserved on the referred palate, MCZ 3642, and is only 
slightly reduced, three cusped, and three rooted. It is shorter and 
wider than P^ with the internal basal cingulum similarly developed, 
but less so than in Poebrotherium. Externally, the tooth is simple 
with the parastyle nearly indistinguishable and the mesostyle rep- 
resented only by a low convex ridge. 

The fourth premolar is a small triangular tooth with a shorter 
inner crescent that remains separate from the outer crescent until 
the tooth is considerably worn. The difference in length between 
the inner and outer crescents is about as in Paratylopus primaevus 
and, as in that species, the inner crescent is skewed anteriorly. 
The outer face is relatively simple with low antero- and postero- 
external styles and a low median convex ridge. Short weak cingula 
are present on the anterior and posterior sides of the inner crescent. 
This tooth is somewhat more reduced than in Oxydactylus longipes 
or Paratylopus primaevus, which are more nearly like Poebrothe- 
rium in this respect. 

The molars increase in both length and width from Mi to M^, 
as in Paratylopus primaevus, and are as brachyodont as in that 
species. M^ is wider than it is long resembling the nearly square 
first molar of P. primaevus and M. bathygnathus, more so than the 
elongated Mi of O. longipes. This tooth is only moderately worn 
in MCZ 3642 and has a strong parastyle, as in "Paratylopus" 
cameloides." The mesostyle is directed perpendicular to the outer 
face of the tooth and is as strong as in O. longipes. The metastyle 
is absent. The median ridges on the outer face of the tooth are 
very prominent, especially on the anterior crescent, and widen 
somewhat near the base of the crown. This last feature is similar 
to the condition found in P. primaevus and quite unlike that in 



- I am not convinced that cameloides belongs in this genus. 



BREVIORA 



No. 255 



Poebrotherium or Oxydactylus. A prominent internal pillar rises 
from the bases of the protocone and hypocone, particularly the 
latter. A similar pillar is found in M. bathygnathus and "P." 
cameloides and incipiently on M-' of P. primaevus. The protocone 
remains separated from the mesostylar region until worn nearly to 
the base of the crown, whereas the hypocone fuses to this area 
somewhat earlier. A very weak basal cingulum marks the pos- 
terior edge of the hypocone. 



I' 



O 






Figure 1. Floridotraguhis dolicliantliereiis. a. Diagram of upper denti- 
tion, X 4/5. b, c, Composite crown view of left cheek teeth; b, upper P- 
to M3, c, lower Po to M.;, XI. 



M- is as long as it is wide in MCZ 3642, but slightly wider than 
long in MCZ 3711. It is similar to M^ with the para- and meso- 
style more prominent. The median ridge of the anterior crescent 
is considerably stronger than that of the posterior one. M?' is 
similar to M^, but with a somewhat weaker and less compressed 
mesostyle. 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 7 

The lower incisors are unknown. A large, strongly recurved 
canine is present in MCZ 4086 and is heavily worn posteriorly to 
a point below the crown. The tooth is slightly compressed laterally 
and emerges from its socket at a lateral inclination of about 20 
degrees from the vertical. 

Pi is separated from the canine by a long diastema somewhat 
exceeding the combined length of Mi and Mo in the type, but 
variable in referred specimens. Pj is not preserved, but the alve- 
oli indicate a small, one-rooted tooth probably similar to Pi and 
roughly equivalent to it in size. 

After another long diastema the second premolar follows (Fig. 
Ic) and is a simple, two-rooted, three-cusped, trenchant, and later- 
ally compressed tooth. The cusps are essentially in a straight line, 
the anterior cusp being very slightly inflected medially as in Poe- 
brotherium eximium, but the tooth is generally much more robust 
than in that species. This tooth did not occlude with the upper 
dentition, as shown by its completely unworn condition. 

The diastema between Po and Po is quite variable; that of MCZ 
4086 is 52 per cent smaller than that of the type. P3 is similar to 
P2 but more robust. The anterior cusp is more inflected produc- 
ing a well defined parastylid which does not continue to the base 
of the crown. Only a suggestion of a posterior intermediate crest 
is present internal to the median cusp. The outer surface is simple 
and gently convex. 

P4 foflows immediately behind P3 and is shorter and broader 
than the latter. This tooth strongly contrasts with its homologue 
in O. longipes, being somewhat more reduced and morphologically 
simplified. The anterior cusp is more inflected than in P3, but 
does not form the prominent right angle of Poebrotherium wilsoni, 
nor the sharply bent and somewhat expanded anterior column of 
Oxydactylus. The lake enclosed between the posterior and poste- 
rior intermediate crests is relatively small and shallow. Laterally 
the surface is simple, lacking prominent ridges and troughs. 

The molars become larger from front to back, are simple and 
generally similar to each other. The lingual faces of the inner 
crescents are gently convex and for the most part lack distinct 
stylids and median ridges. A parastylid is only suggested on the 
type, but is absent on MCZ 4291, and present, although weak, on 
MCZ 4086. A distinct median pillar rises from the bases of the 
external crescents of each molar. 

The metaconid remains isolated from the other crescents until 
worn to the base, whereas the anterior crest of the hypoconid fuses 



8 BREVIORA No. 255 

early to the entoconid and protoconid. The hypoconuUd is di- 
vided, forming two grinding surfaces; the space between these is 
continuous with the space between the posterior pair of crescents 
in young individuals. The two grinding surfaces quickly fuse at 
their anterior extremity and the fusion proceeds posteriorly with 
wear. 

Discussion: From the material available, it appears that only 
one species of Floridatragulus is represented in the Thomas Farm 
fauna. As mentioned above, the P1-P2 and Po-P.s diastemata are 
quite variable, but this may be due to the functionless condition 
of P2. Similar variation is observed in Lama with respect to the 
non-occluding canine. The referred jaw, MCZ 4291, is not as 
deep as in the type or in MCZ 4086 by about 30 per cent. This 
kind of variation is also seen in Lama, although not to the same 
extent. MCZ 4291 also differs in the position of the posterior 
extent of the symphysis which is almost directly under Pi rather 
than about 14 mm behind this tooth as in the type. McKenna 
(1966) notes two species of Floridatragulus, but does not discuss 
the reasons for this decision. The differences noted above pos- 
sibly indicate a second species, but until additional material is 
available for study it is preferable to refer this specimen (MCZ 
4291), at least tentatively, to F. dolichanthereus. 

The camelid affinities of this species are inferred from such typi- 
cal characters as the hooked angle of the jaw, auditory bullae filled 
with spongy bone and divided by a tympanohyal pit and groove, 
relatively simple premolars, and the similarity of the dentition as 
a whole to those of such early camels as Poebrotherium and 
Paratylopus. ^ 

The upper dentition of F. dolichanthereus is more nearly like 
that of Paratylopus primaevus than like the Miocene Oxydactylus- 
like camels. The open orbit, squared molars, retention of promi- 
nent mesostyles, presence of median internal pillars, and graded 
increase in size from M^ to M^ indicate an origin close to P. 
primaevus. In the elongated snout and deep jaws this form re- 
sembles Miotylopus bathygnathus, but this similarity is probably 
no more than a parallel development of habitus characters. Other 
characters which Miotylopus shares with Floridatragulus, such as 
median pillars, reduced upper canine, and (probably) an open 



3 G. M. Allen, in a personal communication to T. Barbour, expressed 
this vievv' more than 20 years ago (correspondence in the Harvard Archives). 



J 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 9 

orbit, more likely indicate retention and development of the prim- 
itive characters of a Paratylopus-like ancestor than any close phy- 
logenetic relationship. 

The many distinctive characters which separate this genus from 
other late Tertiary Camelidae have justifiably confused past work- 
ers. The greatly elongated snout is a peculiar habitus character, 
which is only distantly approached in such specialized forms as 
O. longirostris and "P." cameloides, and the large caniniform in- 
cisors decreasing in size from I^ to 1^ are peculiar to this genus. 
In the mandible the long symphysis and exceptionally narrow jaw 
strongly divergent in the region of the cheek teeth are not seen in 
other camels. Also, the very simplified premolars and divided 
hypoconulid of M3 tend to set this form off from other Miocene 
camels. 

A number of astragali, calcanea, and proximal phalanges are 
referred to this species. The tarsal bones are similar to those of 
other Miocene camels and show no distinctive characters. As is 
common in other camels with an elongated snout, the phalanges 
are long and slender. 

Two distinct morphological groups of phalanges are present; 
these are differentiated by the symmetry and dorsal extent of the 
distal articular facet. Differences of this type are present between 
the manus and pes of the Recent species of Dorcatheriiim, Mos- 
chus, and Lama and probably represent manus-pes differences in 
the present species also. 

The referred metapodial fragments show that the metatarsals 
are closely appressed, with some fusion near the proximal end; 
the plantar processes are completely coossified. Metatarsal II is 
fused with metatarsal III, whereas metatarsal V is entirely separate 
from metatarsal IV and articulates with it by a small facet. Meta- 
carpals III and IV are separate. 

Because of the very peculiar characters of this genus and be- 
cause of its distant relationship to contemporary camelids, it is pro- 
posed that a new subfamily, the Floridatragulinae, be established 
to receive it, with the subfamilial diagnostic characters being those 
of the genus until such time as additional members of this group 
are found. 

Diagnosis of the Subfamily Floridatragulinae 

Differing from Miocene Alticamelinae in retention of many 
primitive characters such as complete brachyodont dentition, open 
orbits, and nearly square upper molars with median pillars; upper 



10 



BREVIORA 



No. 255 



incisors caniniform, lateral in position, separated from each other 
by diastemata, unlike those of Camelinae; premolars tending toward 
reduction and simplification, but less so than in Stenomylinae; 
lower molars simple with pillars, hypoconulid of Mg divided. 



TABLE 1 

Comparative measurements of Floridatragulus 
dolichanthereus in mm 



Skull length to condyles 
Width of palate at P' 
Width of palate at M2 
Ratio: snout (Pmx to orbit) 

to skull length 
Tip of Pmx to V- 
II length 
II width 
P length 
P width 

II to \- diastema 
P to canine 
Canine to Pi diastema 
Pi to P2 diastema 
Pi length 
Pi width 
P3 length 
P3 width 
P4 length 
P-i width 
Ml length 
Ml width 
M2 length 
M2 width 
M3 length 
M3 width 



Floridatragulus 
dolichanthereus 



MCZ 3711 
"385 
45 
•76 

"0.70 
23 

9 

7 

7 

4.5 
37.5 

9 

9 
28 
11.5 

4 



9 

8.5 

ni 

16 

17 

18.5 

18.5 



MCZ 3642 



10 

5 

9 

8.5 
13.5 
15 
17 
17 



Paratylopus 
primaevus 

AMNH 9806 
=200 
21 

52 

0.51 



3 
4 
7.5 



'10 
'12 
'13 
'13 

16 

14.5 



I 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 11 

Floridatraguliis dolichdnthereus 
MCZ 3635 MCZ 4086 MCZ 4291 



Jaw length 


— 


^■300 


— 


Length of symphysis behind Pi 


13 


14 


3 


Width of jaw at Pi 


18.7 


19.5 


— 


Width at M:; 


— 


■■■70 


— 


Depth of jaw under Pi 


21.5 


21 


14 


Depth under Pi 


•32 


32 


23 


Depth under M;; 


"38 


37 


»32? 


Canine to Pi diastema 


— 


28.5 


— 


Pi to Po diastema 


33 


37 


24 


P- length 


— 


10.5 


— 


Po width 


— 


3.7 


— 


P;.. to P:; diastema 


21 


10.5 


19.5 


P:; length 


— 


10.5 


=•10.5 


P;j width 


— 


4.5 


M.5 


P4 length 


— 


10 


no 


P4 width 


— 


6 


6 


Ml length 


•13 


10 


10.5 


Ml width 


"9 


9 


8.5 


Mo length 


15.5 


15 


15.5 


Mo width 


11 


11 


11 


M?, length 


25.5 


13 


13 


M:i width 


12.5 


13 


13 



" estimate based on incomplete specimen. 
"" worn. 



ALTICAMELINAE 

NoTHOKEMAS FLORIDANUS (Simpson) 1932 

Oxydoctylus floridaniis Simpson, 1932, pp. 35-37, figs. 20, 21. 
Paratylopus grandis White, 1940, p. 33, pi. V. 
Nothokemas grandis White, 1947, pp. 508-515, figs. 5, 6. 

Hypodigm: FSGS V-5247, type, incomplete right maxilla with 
P- to M3; FSGS V-5238, paratype, right lower jaw with P3 to 
M3; MCZ 3636 (type of N. grandis), incomplete right mandible 
with P3 to M3 and alveolus for Po ; MCZ 4322, right maxilla with 
Po to M3; MCZ 4323, left mandible with P4 to M2 and anterior 
crescent pair of Mg ; MCZ 4324, partial left mandible with Po to 



12 BREVIORA No. 255 

P4; MCZ 4325, left mandible with Mj to M. and alveoli for P2 
to P4; MCZ 4326, left mandible with dP.. to Mo; MCZ 4329, 
crushed skull with poorly preserved P^ to M^, rostrum anterior 
to Pi-P2 diastema missing; MCZ 4541, crushed skull with dP- to 
M^, anterior snout missing; MCZ 7297, left mandible with dP4 
to M3; MCZ 7592, incomplete left mandible with Mo to M3; 
AMNH 22672, left mandible with Po to Mo; MCZ 7767, 22 
astragah; MCZ 7768, 10 calcanea; MCZ 7769, 17 cuboids; MCZ 
7770, 3 naviculars; MCZ 7771, 15 proximal phalanges; MCZ 
7772, 23 second phalanges; MCZ 7773, 8 ungual phalanges. 

Horizon and locality: Middle Miocene, Hawthorne Formation, 
Thomas Farm, Gilchrist County, Florida. 

Redescription: The present description is based on Simpson's 
account (1932, pp. 35-37) and on the referred material in the 
Museum of Comparative Zoology and American Museum of Nat- 
ural History collections. 

This species is about the size of Oxydactylus brachyodontus 
Peterson 1904 in measurements of the dentition, but the maximum 
skull length was doubtless considerably greater due to the elonga- 
tion of certain of the diastemata. The rostrum is deep and very 
narrow. A prominent, quadrangular lacrimal vacuity lies in the 
maxillary bone and is as large as the orbit. The latter is com- 
pletely closed behind by a wide process from the frontal bone. 
The last molar is somewhat anterior to the orbit, but less so than 
in Floridatragulus. 

The incisors and canine are unknown; the anterior portion of 
the snout is missing on both referred skulls. A small depression 
is present in some broken bone on the anterior edge of MCZ 4329 
which may be the socket for Pi. If this is correct this tooth was 
single rooted as in O. brachyodontus, but was very much smaller, 
being only a tiny and almost certainly functionless vestige. Be- 
hind Pi is a long diastema approximately equal to the combined 
lengths of P^, M^ and M-. 

P2 is a long, narrow, trenchant tooth essentially unreduced. 
Proportionately, it is narrower than in O. longipes, being more 
similar in this respect to O. longirostris. A complete internal 
basal cingulum originates anteriorly at the base of a style. A 
similar anterior style is present on the external face. The external 
and internal surfaces are somewhat folded and expanded into a 
convex longitudinal ridge in the region of the median cusp. The 
posterior cusp is strongly skewed externally in MCZ 4329, but 
less so in 4322. 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 13 

P3 is larger than P^, considerably wider, and has a much 
stronger internal cingulum. A strong parastyle forms a wide con- 
vex fold and is continuous around the front of the tooth with a 
very narrow anterointernal style. 

P^ has the internal crescent fully expanded and somewhat 
skewed anteriorly, as in "Paratylopus" cameloides. It is nearly as 
wide as it is long, resembling in this respect "P." cameloides and 
O. longirostris rather than O. longipes. A well developed cingulum 
extends along the posterior base of the internal crescent and a 
weaker one is present on the anterior side. The external surface 
is ribbed by a prominent anterior style, which widens toward the 
base, and a variable posterior style, which is moderately developed 
on MCZ 4329, but nearly absent on MCZ 4322. 

The upper molars are brachyodont, with M^ the widest of the 
series, as in O. longipes. They are proportionately longer and 
wider than in "P." cameloides and resemble O. longipes more 
than any other species in this respect. A prominent internal pil- 
lar, V-shaped in cross-section, is present on all the molars, but 
decreases in size from M^ to M^. In this character Nothokemas 
resembles "P." cameloides and contrasts with the pillarless molars 
of Oxydactylus. 

Ml has a moderate parastyle and strong mesostyle, the latter 
directed somewhat anteriorly. The metastyle is very weak to ab- 
sent and the median ridges of the external faces are prominent, 
especially on the metacone. A strong basal cingulum extends 
around the base of the protocone becoming weak to nearly absent 
on the most lingual part of the crescent. A similar crescent is 
present along the posterior edge of the hypocone. The internal 
crescents remain separated from the outer ones until an advanced 
stage of wear is reached. 

M.- is similar to M^, but is longer and has more prominent 
styles and ridges. The basal cingula are generally weaker and less 
extensive around the lingual curves. M-^ is proportionately more 
elongated than M-, the styles more prominent, and the cingula 
nearly absent. In contrast to O. longipes the parastyle is weak as 
in "P." cameloides. Also as in cameloides the protocone is some- 
what narrower anteroposteriorly than the hypocone. 

The mandible is long and relatively shallow as compared with 
that of O. longipes. The anterior portion of the symphysis, from 
Ij to Pi, is unknown. MCZ 4325 has a broken symphyseal area 
which contains what may be the socket for Pi. If it is, this tooth 
was only slightly larger than P^ and situated well in front of the 
posterior border of the symphysis. The long diastema between Pi 



14 BREVIORA No. 255 

and P2 is comparable to that in the upper dentition, being at least 
43 mm in length in MCZ 4325. 

The premolars are large and unreduced, and increase in size 
from front to back. P2 is expanded medially with its internal face 
strongly convex. A small anterior cusp is connected with a high 
and pointed median cusp by a blade-like anterior crest. The pos- 
terior crest is strongly deflected laterally before turning posterad 
so that the posterior cusp lies external to the straight line through 
the anterior and median cusps. A strong posterior intermediate 
crest extends back from the central cusp for about one-half the 
distance to the posterior margin of the tooth, forming a shallow 
open valley between itself and the posterior crest. 

P3 has the anterior cusp very strongly inflected internally, more 
so than P4. The posterior intermediate crest is somewhat more 
developed than in Po, enclosing with the posterior crest a deeper 
but still medially open valley. P4 is similar to P-j, but with the 
posterior intermediate crest long and completely enclosing a deep 
vaUey, which is confined to the posterior one-third of the tooth. 
A well defined cusp is present on the crest near its posterior mar- 
gin, and a slightly oblique longitudinal ridge extends along the 
length of the crown on the lingual surface. 

The lower molars are long and narrow, as in O. brachyodontus 
and less so than in O. longipes. A strong median pillar, variable in 
size, rises primarily from the protoconid, at least on Mo. The 
metaconid remains isolated to the base and somewhat overlaps 
the entoconid, whereas the latter fuses with the protoconid early 
in wear. The entoconid overlaps the hypoconulid for about one- 
fourth of the latter's length and remains separated from it nearly 
to the base of the crown. A very weak parastylid is present on M2. 

The tarsal bones and phalanges that may be referred to Notho- 
kemas on the basis of size and relative abundance are very Oxy- 
dactylus-VikQ, and the cuboid and navicular are separated, unlike 
those of hypertragulids. It is surprising that the limb material does 
not show the stilt-like character that might be expected to be asso- 
ciated with the elongated snout. These elements are somewhat 
more slender than those of O. longipes, but not nearly as stilt-like 
as in O. longirostris. 

Discussion: Nothokemas floridanus differs from the hypertrag- 
ulids, to which it was first referred (White, 1947), principally in 
the structure of the premolars, but also in that of the molars. In 
Hypertragulus, P3 is a simple, single-rooted tooth with a very 
weak posterointernal fold, while the comparable tooth in Notho- 
kemas is three cusped and much more complex in its folds and 



1966 



SELENODONT ARTIODACTYLS FROM THOMAS FARM 



15 



TABLE 2 



Comparative measurements of Nothokemas floridanus in mm 

Oxydactyliis "Paratylopus" 
Nothokemas floridanus longipes caineloides 

MCZ 4329 MCZ 4322 MCZ 4541 CM 918 



Skull length 


^'340 


Pi-P2 diastema 


^53 


P2 length 


13 


P2 width 


6 


P3 length 


"13.3 


P3 width 


8.5 


P^ length 


■M3 


P4 width 


11 


Ml length 


— 


Ml width 


— 


M- length 


— 


M- width 


— 


M^^ length 


23 


M-^ width 


19 



12.5 — 
5.2 — 

13.4 — 
7.5 — 

13 — 

12 — 

18.6 19.7 

18.5 18.5 
"22 22 

21.5 20.5 

23 23 

19.5 18.8 



I 918 


YPM 10917 


340 


"278 


18 


— 


11 


— 


6 


— 


14 


— 


8 


— 


14 


11 


10 


10 


19 


13 


16 


— 


23 


17.3 


20 


16.5 


25 


17.4 


20 


17.3 



Oxydactyliis "Paratylopus" 
Nothokemas floridanus longipes cameloides 

MCZ 3636 MCZ 4325 MCZ 4324 CM 918 AMNH 8197 



P1-P2 diastema 
P2 length 
P2 width 
Pa length 
P:i width 
P4 length 
P4 width 
Ml length 
Ml width 
Mi> length 
M2 width 
Ma length 
Ma width 



13.8 
6.4 

15 
7.5 
M8 

10.7 

21.5 

14 

35 

16.5 



45.5 



19.2 
10.7 
23.6 
13.2 

33 
14.1 



10.8 

4.5 

12.8 

5.5 

14.5 

7.3 



17 
10 

5 
12 

5 
12 

7 
18 
10 
23 
14 
31 
13 



21 
10 

12.5 
4.3 

13 
5.9 

14 

14 

17.5 

11 

25 

12 



" estimated from incomplete specimen, 
''estimated from figure, Dougherty, 1940. 



16 BREVIORA No. 255 

crests. P4 is marked by a very strongly inflected parastylid in 
Hypisodus and Hypertragidus, more prominent than in Notho- 
kemas and extending farther towards the base of the crown. A 
strong stylid extends along the internal surface of the median 
cusp in the above mentioned hypertragulids so that the depression 
formed in front and behind this is very much deeper in the Florida 
species. 

The lower molars of Hypertragulus are complicated by very 
strong cingula and strong pillars rising primarily from the proto- 
conid. In hypertragulids generally, the hypoconulid is strongly 
divided, a condition not seen in Nothokemas. The upper denti- 
tion is also quite different, and generally more complex in the 
Hypertragulidae. 

There can be no question that the dentition as described above 
is indicative of camelid affinities, a conclusion supported by the 
referred postcranial elements. Although highly specialized, Noth- 
okemas floridanus presents no characters which could not easily 
be derived from an early Oxydactylus-Wkt form, from which "P." 
cameloides may also have arisen. In many of its characters Noth- 
okemas stands somewhere intermediate between typical Oxydacty- 
lus (as represented by longipes and brachyodontus) , on the one 
hand, and "P." cameloides, on the other, and it is likely that both 
A^. floridanus and "P." cameloides represent early specialized off- 
shoots from the basal Oxydactylus line that possibly arose in earli- 
est Arikareean time. 

Camelidae gen. et sp. index. 

Several isolated teeth and bones suggest the presence of a third 
camelid in the Thomas Farm fauna. Three upper molars, MCZ 
7811, are generally comparable to those of Oxydactylus from the 
Marsland deposits of western Nebraska and eastern Wyoming, but 
are higher crowned (Beryl Taylor, personal communication), 
ps is represented by two specimens, MCZ 7812, which are very 
similar to P"^ of Alticamelus in structure, but are considerably 
narrower. Four proximal phalanges, MCZ 7813, are long and 
slender with ungrooved distal articular facets and a very shallow 
proximal surface. 

All of these specimens suggest the presence of a camel of a 
somewhat more advanced type than Floridatragulus or Nothoke- 
mas, but only additional material will reveal its relationships. 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 17 

PROTOCERATIDAE 

Synthetoceras (Prosynthetoceras) rileyi australis 

(White), 1941 new rank as subspecies 

Syndyoceras australis White, 1941 (in part), p. 97, pi. XV, figs. 2, 2a. 
Synthetoceras {Prosynthetoceras) douglasi White, 1947, pp. 504-505, fig. 3a. 

Hypodigm: MCZ 3654, type, right mandible with P^, P4 to M3, 
and alveoli for Po and P.;, coronoid process and symphysis an- 
terior to Pi missing; MCZ 3708, right mandible with P.., to Mo 
and alveoli for I^ to Po; MCZ 3655, right mandible with Pj to M^ 
and partial alveoli for lo to the canine; MCZ 3656, right mandible 
with Pi to Mg and partial alveoH for lo to the canine; MCZ 3666, 
partial right mandible with Po to Mo; MCZ 4065 (type of Synthe- 
toceras douglasi), palate with right and left P^ to M-'-; MCZ 7635, 
right mandible with P4 to Mo and alveoli for I2 to P;.; MCZ 7636, 
incomplete left mandible with P4 to M^; MCZ 7637, left mandible 
with P3 to Mo and alveoh for Pi and P2; MCZ 7774, 13 astragali; 
MCZ 7775, 3 cuboids; MCZ 7776, 4 naviculars; MCZ 7777^ 22 
proximal phalanges; MCZ 7778, 19 second phalanges; MCZ 
7779, 5 ungual phalanges. 

Horizon and locality: Middle Miocene, Hawthorne Formation, 
Thomas Farm, 8 miles north of the town of Bell, Gilchrist 
County, Florida. 

Diagnosis: Equal in size to Synthetoceras rileyi rileyi"^ Frick, 
but P3 is 13 per cent smaller, and median cusp of P4 somewhat 
narrower, so that internal depression is flatter and deeper. 

Description: The mandible is long and slender, remaining rather 
shallow beneath the molars. The angle of the jaw is a smooth, 
continuous curve, as in Protoceras. Anterior to Po^ the jaw begins 
to flare outward so that Pi is inclined at about 35 degrees from the 
vertical. 

The alveoli for the incisors indicate that they were small, 
closely spaced, and quite procumbent. That for the canine is 
larger and also procumbent and close behind L., suggesting that 
it was a functional member of the incisor series, as in Protoceras. 
Pi follows after a short diastema equal to Mi in length. In the 



4 Prick's collection data is not precise, but the type specimen appears to 
have been collected in the Catahoula sandstone in Walker County, Texas. 
Although the age of this formation is still not settled, Fisk (1940) considers 
it of Miocene age, and Gidley (see Renick, 1936) of Middle Miocene age. 



18 BREVIORA No. 255 

type and MCZ 3656 and 7637, Pj is a large, single-rooted, re- 
curved, caniniform tooth laterally compressed and with blade- 
like anterior and posterior edges. In MCZ 3708, 7635, and 3655 
the alveolus for the first premolar indicates a very tiny vestigial 
tooth which could not have been functional to any great extent in 
the living animal. In size and dental morphology these two groups 
are identical. Since dimorphs of this kind are known among artio- 
dactyls, and with no evidence to the contrary, it is preferable to 
consider these as sexual morphs rather than two distinct taxa, the 
latter group being the females. 

A long diastema separates Pi and Po. The latter is a two-rooted, 
simple, robust tooth equal to P3 in length (Fig. 2). P3 is propor- 
tionately smaller than the corresponding tooth in S. rileyi rileyi, 
but similar to it in structure. The anterior edge is somewhat re- 
curved onto the internal face forming a moderate stylid. The high 
median cusp is connected with the anterior and posterior cusps by 
narrow crests. A posterior intermediate crest extends backward 
nearly to the edge of the tooth, but fails to fuse with the posterior 
crest, resulting in a partially closed lake which is open postero- 
medially. 

P4 is similar to Po with a stronger parastylid and a lake which 
is completely closed near the base of the crown. It differs from 
S. rileyi rileyi in having a narrower median cusp. 

The molars are indistinguishable from those of S. r. rileyi. M^ 
and Mo have narrow crescents, strong parastylids, and weak 
metastylids. The median ridges are well developed, especially on 
M2. M3 is similar but with the posterior edge of the metaconid 
slightly overlapping the entoconid. The hypoconulid is C-shaped 
forming two functional grinding surfaces, and is quite variable in 
size. 




MCZ 3660 MCZ 3656 



Figure 2. Synthetoceras {Prosy nthetoceras) rilexi anstralis, composite 
crown view of right lower cheek teeth, XI. 



In the upper dentition, the referred palate most closely resem- 
bles that of Synthetoceras jrancisi Frick. M-. is longer than broad 
with the posterior pair of crescents narrower than the anterior pair. 
The metastyle is weak and the mesostyle prominent, but less so 



I 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 19 

than in S. francisi. The parastyle is as strong as the mesostyle. 
The internal crescents fuse early and extend as a narrow process 
between the external crescents, which they do not fuse with until 
late in wear. A median pillar, check-shaped in cross section, is 
present between the internal crescents, with the longer arm along 
the protocone. A prominent elongate pillar rises from the cingu- 
lum along the anterior side of the protocone. 

M- is smaller than M^ and shorter anteroposteriorly than in 
Synthetoceras tricornatus; the styles are as prominent as in that 
species and the median ridges are low and indistinct. M^ is the 
smallest of the molar series and, although worn, appears to be 
similar to M- in character. 

?•* is triangular, as in S. francisi, with a very strong anteriorly 
curved metastyle. Unlike conditions in the corresponding tooth 
of that species, however, the parastyle is nearly absent. P^ re- 
sembles that of S. francisi, but has a stronger metastyle and a 
strong and complete internal cingulum. A small wear facet on 
the anterior edge of P^ indicates that P- was present, as in S. 
francisi. 

On the basis of relative size, a number of foot bones are re- 
ferred to this species. The phalanges are short and heavy as in 
Protoceras. 

Discussion: As can be seen from Table 3 the range of variability 
of S. australis is sufficient to include the type of S. rileyi. In the 
character of the molars there are no differences which could not 
be attributed to intraspecific variation. The premolars, except for 
the differences already noted, are nearly identical in the two forms. 

The subspecific rank of S. rileyi australis is proposed in view of 
the premolar differences, and the distance between Walker County, 
Texas, and Gilchrist County, Florida (nearly 800 miles), and be- 
cause of the possibility of a slight difference in horizon between 
these localities. Since both forms are so poorly known at present 
it is advisable to retain a subspecific distinction, at least until the 
extent of variation in each is better known. 

Frick (1937) was apparently uncertain as to the subgeneric 
reference of 5. rileyi, questionably placing it in the subgenus Syn- 
thetoceras. The subgeneric distinction between S. {Synthetoceras) 
tricornatus and S. (Prosynthetoceras) francisi is based on the 
smaller size of the latter, its shorter crowned teeth, the retention 
of P-, and the prominence of the upper canine alveolus. 

S. rileyi is referred to the subgenus Prosynthetoceras because of 
its small size and lower crowned teeth as compared with S. tri- 
cornatus, and because of the retention of P- in the referred palate, 
MCZ 4065. 



20 



BREVIORA 



No. 255 



TABLE 3 



Comparative measurements of Synthetoceras 
(Prosy nthetoceras) rileyi australis in mm 







Synthetoceras (P.) 


rileyi 


australis 








MCZ 4065 




Width of palate between Mi 




64.5 






P'^ to M'^ length 






64.8 






P3 length 






9.0 






P- width 






6.1 






P^ length 






9.0 






Pi width 






9.0 






M' length 






13.6 






Ml width 






14.8 






M- length 






17.2 






M- width 






15.9 






M-' length 






17.0 






M'^ width 






16.8 








'^> 


nthetoceras (P.) 


Syn 


hetoceras (P.) 






rileyi australis 




rileyi 


rileyi 










AMNH 34181, 




MCZ 3654 


MCZ 3708 MCZ 7635 




type 


Depth of jaw 












under Pj 


21.2 


22.7 


22.5 




19.3 


Depth under P4 


23.6 


— 


23.0 




20.5 


Depth under M^ 


29.1 


27.5 


28.4 




— 


P2 to M3 length 


85.4 


76.6 


76.7 




77.4 


Pi to P2 diastema 


42.0 


46.0 


46.1 




— 


Po length" 


— 


— 


— 




8.2 


P2 width" 


— 


— . 


— 




3.1 


P:! length 


— 


8.5 


— 




9.7 


Ps width 


— 


4.0 


— 




4.3 


P4 length 


10.5 


10.5 


10.1 




10.4 


P4 width 


6.5 


6.3 


5.7 




6.4 


Ml length 


13.0 


12.2 


14.4 




12.4 


Ml width 


10.0 


9.2 


9.4 




9.4 


M2 length 


18.0 


16.1 


15.4 




16.7 


M2 width 


11.0 


10.5 


21.6 




22.4 


M3 length 


25.5 


25.0 


21.6 




22.4 


M;; width 


11.4 


11.1 


10.5 




10.1 



" Known only from MCZ 3656; length, 8.2 mm, width, 4.0 mm. 



1966 



SELENODONT ARTIODACTYLS FROM THOMAS FARM 



21 



MERYCOIDODONTIDAE 
MERYCHYINAE 
cf. Merychyus sp. 

An oreodont is represented in the Thomas Farm fauna by a 
single left upper molar, MCZ 7765, and an incomplete tarsus, 
MCZ 7766. In structure and size the molar (Fig. 3) most closely 
resembles Meiychyus minimus, but is also quite similar to M. 
elegans of a later age. The external faces of the outer crescents 
are transversely concave and longitudinally convex, so that their 
apices overhang the internal crescents in typical oreodont fashion. 
The posterior crescents are narrower than the anterior pair and are 
relatively longer anteroposterioriy, as in M^ of M. elegans. A 
strong cingulum is present on the anterior side of the protocone 
and a weaker one on the posterior side of the hypocone. 





Figure 3. Oreodont upper left molar; cf. Merychyus. External view 
(left), and crown view (right), XlVi. 



There can be little doubt of the oreodont affinities of the tarsal 
elements, which probably beiong to a single individual. The cal- 
caneum is relatively short, strong, and laterally compressed with 
a strongly truncated proximal end which bears a deep groove for 
the tendon of the gastrocnemius. The sustentaculum is very small 
and only slightly projecting beyond the medial surface of the tuber. 
The astragalar facet on the sustentaculum is relatively long and 
narrow. 

The astragalus is wide, low, and quite asymmetrical. The proxi- 
mal trochlear is inclined at an angle to the distal one, and is placed 
more lateral in position as is typical of oreodonts. The sustentacu- 
lar facet is narrow, occupying little more than half of the plantar 
surface. 



22 BREVIORA No. 255 

These fragments are tentatively referred to the genus Merychyus 
because of the similarities to M. minimus in both size and morphol- 
ogy. However, a more certain reference must await more com- 
plete material. 

The discovery of oreodont remains in Florida is a significant, 
although not a completely unexpected find. The apparent rarity 
of these animals in this region may not reflect their actual abun- 
dance. Schlaikjer (1935, p. 173) suggested that a gregarious habit 
combined with localized distribution and/or species habitat re- 
quirements of oreodonts could account for their great abundance 
in some areas, and relative rarity in others of the same age. As 
suggested by Thorpe (1937, p. 9) and Patterson (1949, p. 270), 
the absence of sediments deposited under appropriate conditions 
or their subsequent removal by erosion could be responsible for a 
meager oreodont fauna in certain areas. The idea, as proposed 
by Romer (1947, p. 10), that Florida was ecologically unfavor- 
able for these animals is less attractive since the environment of 
the Florida Miocene was inferred partially by the absence of oreo- 
donts. The rarity of this group from Florida beds is probably due 
to a combination of the first two factors: localized distribution and 
scarcity of sediments. 

CERVIDAE 

Elastomer Y\ (Parablastomeryx) floridanus White, 1940 

Hypodigf.:: MCZ 3626, type, partial left mandible with P4, 
Ml, the ante: or external crescent of Mo, and the alevoli for Po 
and P3; MCZ 3627, paratype, upper left molar; MCZ 3706, right 
mandible with dP.> to M;.; MCZ 3707, left mandible with dPo to 
M3; MCZ 3912, left mandible with dP4 to M.; MCZ 4220, left 
mandible with Pj, dP. to M-.; MCZ 4221, right mandible with 
P3 to M3; MCZ 4222, right mandible with Po, Mi to M3; MCZ 
7795, 23 astragali; MCZ 7796, 24 calcanea; MCZ 7797, 17 cubo- 
naviculars; MCZ 7798, 103 proximal phalanges; MCZ 7799, 53 
second phalanges; MCZ 7800, 24 ungual phalanges; MCZ 7801, 
6 metapodiais; MCZ 7802, left tibia; MCZ 7803, right radius. 

Horizon and locality: Middle Miocene, Hawthorne Formation, 
Thomas Farm, Gilchrist County, Florida. 

Diagnosis: White (1940, p. 34) diagnosed this species as fol- 
lows: "A large Parablastomeryx, paraconid of P4 with very small 
stylid, metaconid large and placed about the mid-length of the 
tooth, entoconid projecting at nearly right angles to the axis of 
the tooth, crest of hypoconulid extending along posterior border 
of the tooth nearly to the inner border." 



1966 



SELENODONT ARTIODACTYLS FROM THOMAS FARM 



23 



Discussion: Since White's diagnosis, many new specimens have 
been collected. These demonstrate a remarkable degree of vari- 
ation in the premolars and reveal the character of the lower mo- 
lars and milk dentition, which were not available to him. The 
molars (Fig. 4) tend to confirm White's observation that B. flori- 
danus is similar to B. primus. 





Figure 4. Blastomeryx floricianiis. a, Composite crown view of left lower 
cheek teeth, XI 1/2. h, MCZ 3706, right dPi, X3. 



The hypoconulid of M-. has a prominent anterointernal cusp, 
and a strong palaeomeryx fold is present on the protoconid. The 
internal stylid of P4 which White described as being "very small" 
on the type varies from strong on MCZ 7763 to absent on MCZ 
4221. In P;; of most of the specimens the entoconid is joined to 
the hypoconid by a semicircular crest around the posterior margin 
of the tooth, forming an enclosed lake. However, in MCZ 4349 
the lake remains open posteriorly and is divided from the ento- 
conid by a strong median ridge, which extends posteriorly to the 
end of the tooth and, after turning medially, ends in a minute cap- 
sule. These and other variants must be considered to be within 
the range of variation of this species since intermediate types link 
every extreme. 

The lower milk teeth, dp2 to dP4 are known from two speci- 
mens, MCZ 3706 and 3707? DP4 (Fig. 4b) is five cusped with 
an elongated pillar between the anterior inner cusps and a conical 
pillar between the posterior inner cusps. DPo is very similar to 
P3 in structure and length, but is much narrower. There is little 
difference between dP-. and P.. 



24 



BREVIORA 



No. 255 



Because of the palaeomeryx folds, hypoconulid cusp, and rela- 
tively large premolars, this species is referred to the subgenus 
Parablastomeryx. 

Referred postcranial material includes numerous tarsal bones, 
phalanges, a tibia, a radius, and several cannon bones. The tarsus 
is typically cervid. The tuber of the calcaneum is light and later- 
ally compressed; the cuboid and navicular are solidly fused. The 
proximal phalanx is lightly built and considerably more slender 
dorsoventrally than in Recent cervids. 

The metatarsals are fused along their entire length and there is 
a deep longitudinal groove on the dorsal side between metatarsal 
III and metatarsal IV. The distal facets are completely keeled, 
and together are considerably wider than the shaft. The metacar- 
pals are fused and somewhat shorter than the metatarsals. The dor- 
sal groove is shallow and indistinct. The tibia is long and slender, 
but less curved laterally than in Recent forms. 

TABLE 4 
Comparative measurements of Blastomeryx (P.) fioridanus in mm 





MCZ 


MCZ 


MCZ 


MCZ 




7763 


4221 


4349 


3626 


Jaw and lower dentition: 








Jaw length 


— 


"132 


— 


— 


Po to Mo 


— 


60.8 


66.9 


— 


Po length 


5.6 


— 


— 


— 


?2 width 


2.6 


— 


— 


— 


P3 length 


7.3 


8.0 


8.2 


— 


Ph width 


4.3 


4.3 


4.6 


— 


P4 length 


8.9 


8.4 


8.7 


8.2 


P4 width 


5.4 


5.0 


5.0 


4.9 


Ml length 


— 


lO.l 


10.3 


9.0 


Ml width 


— 


7.3 


7.0 


6.3 


M2 length 


— 


11.8 


11.7 


— 


M2 width 


— 


8.2 


7.9 


— 


M3 length 


— 


16.2 


17.3 


— 


M3 width 


— 


8.2 


7.7 


— 



' estimated from incomplete specimen. 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 25 

Machaeromeryx gilchristensis White, 1941 

Hypodigm: MCZ 3651, type, partial right mandible with P4 to 
Mo and alveoli for P2, P;^ and M^; MCZ 3652, right mandibular 
fragment with Mj, and a left upper molar; MCZ 3709, partial 
right mandible with P3 to M^ and part of Mo,; MCZ 3710, partial 
right palate with Mi and M^; MCZ 7804.' 37 astragali; MCZ 
7805, 1 calcaneum; MCZ 7806, 3 cubo-naviculars; MCZ 7807, 
24 proximal phalanges; MCZ 7808, 5 second phalanges; MCZ 
7809, 2 ungual phalanges; MCZ 7810, 1 metacarpal. 

Horizon and locality: Middle Miocene, Hawthorne Formation, 
Thomas Farm, Gilchrist County, Florida. 

Diagnosis: Premolar series relatively longer than in M. tragulus, 
Mo shorter, upper molars less square with outer crescents consid- 
erably wider than inner ones, median pillars of upper molars only 
vaguely suggested in contrast to M. tragulus. 

Discussion: No new information can be added to the diagnosis 
, . . . ^ 

smce this species remains so poorly represented. Aside from the 

differences in proportion, the teeth are very similar to M. tragulus. 

The few tarsal remains and a referred metcarpal are essentially 

like those of Blastomeryx, but much smaller. 

CONCLUSION 

The fossil selenodont artiodactyls from Thomas Farm, Gilchrist 
County, Florida, indicate faunal communication with other parts 
of the North American continent during the Lower Miocene. 
Synthetoceras r. australis is at most subspecifically distinct from 
the western S. r. rileyi, and Blastomeryx fioridanus and Machaero- 
meryx gilchristensis are close to western species. The specialized 
camelids, Floridatragulus and Nothokemas, are so far unknown 
in the West, but they or closely related forms have been recorded 
from the Oakville of the Texas coastal plain (Wilson, 1962). 
This contradicts White's contention that north-central Florida was 
an island until the end of Oak Grove time. Continuity, even if 
intermittent, between Florida and the mainland certainly seems 
to have occurred during the earlier Miocene. On geological 
grounds, Cook (1945) suggests that a narrow peninsula persisted 
through much of this period, and Bader (1956), on the basis of 
his comparisons of the Thomas Farm equids with those of Texas, 
states that isolation was, at best, very ephemeral. The opportunity 
for faunal movement between these areas, then, must have been 
considerably greater than suggested by White (1942). 



26 BREVIORA No. 255 

REFERENCES 

Bader, R. S. 

1956. A quantitative study of the Equidae of the Thomas Farm 
Miocene. Bull. Mus. Comp. Zool., 115: 49-78. 

Cook, C. W. 

1945. Geology of Florida. Bull. Florida Geol. Surv., No. 29: 1-339. 
Dougherty, J. F. 

1940. Skull and skeletal remains of the camel Paratxiopiis cameloides 
from the John Day, Oregon. Carnegie Inst. Washington Publ. 
No. 514: 49-58. 
FiSK. H. N. 

1940. Geology of Avoyelles and Rapides parishes. Bull. Louisiana 
Geol. Surv., 18: 1-240. 
Frick, C. 

1937. Horned ruminants of North America. Bull. Amer. Mus. Nat. 
Hist., 69: 1-669. 
McKenna, M. C. 

1966. Synopsis of Whitneyan and Arikareean camelid phylogeny. 
Amer. Mus. Novit. No. 2253: Ml. 
Patterson, B. 

1949. Rates of evolution in taeniodonts. Pp. 243-278. //; G. L. 
Jepsen, G. G. Simpson, and E. Mayr eds.. Genetics. Paleon- 
tology, and Evolution, Princeton Univ. Press. 
PuRi, H. S. and R. O. Vernon 

1964. Summary of the geology of Florida and a guidebook to the 
classic exposures. Florida Geol. Surv., Special Publ. No. 5: 
1-312. 
Ray. C. E. 

1957. A list, bibliography, and index of the fossil vertebrates of 
Florida. Florida Geol. Surv., Special Publ. No. 3: 1-175. 

Renick, B. C. 

1936. The Jackson group and the Catahoula and Oakville formations 
in a part of the Texas Gulf coastal plain. Univ. Texas Bull. 
No. 3619: 1-104. 
Romer, a. S. 

1947. The fossil mammals of the Thomas Farm. Gilchrist County, 
Florida. Quart. Jour. Florida Acad. Sci.. 10: 1-10. 
Schlaikjer, E. M. 

1935. Contributions to the stratigraphy and paleontology of the 
Goshen Hole area, Wyoming, Part 4. Bull. Mus. Comp. Zool., 
74: 97-189. 
Simpson, G. G. 

1932. Miocene land mammals from Florida. Bull. Florida Geol. 
Surv., No. 10: 11-14. 



1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 27 

Thorpe, M. R. 

1937. The Merycoidodontidae, an extinct group of ruminant mam- 
mals. Mem. Peabody Mus. Nat. Hist., 3: 1-428. 
White, T. E. 

1940. New Miocene vertebrates from Florida. Proc. New England 
Zool. Club, 18: 31-38. 

1941. Additions to the Miocene fauna of Florida. Proc. New Eng- 
land Zool. Club. 18: 91-98. 

1942. The Lower Miocene mammal fauna of Florida. Bull. Mus. 
Comp. Zool., 92: 1-49. 

1947. Additions to the Miocene fauna of northern Florida. Bull. 
Mus. Comp. Zool., 99: 497-515. 
Wilson, J. A. 

1962. Field trip no. 10, Tertiary formations between Austin and 
Houston, with special emphasis on the Miocene and Pliocene. 
In Geology of the Gulf Coast and Central Texas and a guide- 
book of excursions. Houston Geol. Soc, 1962: 342-354. 

(Received 1 November, 1966.) 



Errata 

Page 6, Fig. \a, for X4/5 read X2/5. 

Page 10, Table 1. for P^ length and width read P-. 

Page 11. Table 1. M, length. MCZ 4086, for 13 read 25; MCZ 

4291, for 13 read 24. " 
Page 11, Nothokemas, line 8 (MCZ 4322) for Po to M,.; read P- 

to M3. 
Page 15, Table 2. M^ width, AMNH 8197, for 14 read 9.5. 
Page 17, lines 9, 10. MCZ 3655, for Pi to M. and partial alveoh 

for lo to canine, read M^-Mo and alveoli for P1-P4. 
Page 17, line 1 1. for MCZ 3666 read 3660. 
Page 20, Table 3. M^ width, MCZ 7635, for 21.6 read 10.3; 

AMNH 34181, for 22.4 read 10.4. 
Page 22, Blastomcryx, Hypodigm. Add MCZ 7763, partial left 

jaw with P:.-P4, and MCZ 4349, left jaw with P^-M.. 



BREVIORA 

MiLiseiLijrii of Compsirsitive Zoology 

Cambridge, Mass. 3 February, 1967 Number 256 



ANOUS CHOCORUM, A NEW PUNCTATUS-UKE ANGLE 
FROM DARIEN, PANAMA (SAURIA, IGUANIDAE) 

By Ernest E. Williams 
and William E. Duellman' 



INTRODUCTION 

In the course of a survey of the herpetofauna of Panama, the 
junior author, accompanied by Charles W. Myers, Tomas Quin- 
tero, and Linda Trueb, travelled in a dugout canoe from Santa 
Maria de El Real, Darien, to the upper part of the Rio Tuira Basin 
in July, 1965. Most of the month of July was spent working out 
of a base camp at the confluence of the Rio Tuira and the smaller 
tributary, the Rio Mono. Collections also were made at the 
mouth of Quebrada La Plata lower on the Rio Tuira, and on 
Cerro Quia on the Panamanian-Colombian border. Six specimens 
of a previously undescribed species of anole are included in the 
collection; these plus one specimen obtained by G. B. Fairchild 
at Tacarcuna, Darien, and three specimens obtained more recently 
from the Serrania de Pirre, form the basis for the present descrip- 
tion. A long previously collected specimen from Colombia in 
the American Museum of Natural History (AMNH) proves to be- 
long to the same species. 

The junior author is grateful to his field companions, whose 
combined efforts made the trip a great success. The survey of the 
herpetofauna of Panama is being conducted in cooperation with 
the Gorgas Memorial Laboratory in Panama and is supported by 
a grant (NIH GM- 12020) from the National Institutes of Health. 
This paper is one of a series of studies of the genus Anolis pre- 
pared by the senior author with the support of National Science 
Foundation Grant GB-2444. 

The material of the new species has been divided between 
the Museum of Natural History, Kansas University (KU) and 

1 Museum of Natural History, University of Kansas, Lawrence, Kansas. 



2 BREVIORA No. 256 

the Museum of Comparative Zoology (MCZ). We name the new 
anole for the Choco Indians of the region: 

Anolis chocorum new species 

Type. KU 96934, adult $ , Rio Tuira at Quebrada La Plata, 
100 m, Darien, Panama, collected by a native for Charles W. 
Myers, 26 July 1965. 

Paratypes. Panama: Darien. KU 76027, young 6 , Tacarcuna 
Village on Rio Tacarcuna, 550 m, G. B. Fairchild coll., 9 July 
1963; KU 96931, Rio Tuira at Rio Mono, 130 m, William E. 
Duellman coll., 12 July 1965; MCZ 82546, same locality, William 
E. Duellman coll., 13 July 1965; MCZ 82547, same locality, 
Charles W. Myers coll., 13 July 1965; KU 96932, Cerro Quia, 
740 m, Charles W. Myers coll., 9 July 1965; KU 96933, same 
locality, Charles W. Myers coll., 26 July 1965; KU 98520, north 
end, Serrania de Pirre, 320 m, C. W. Myers coll., 15 January 
1966; KU 98521, south ridge, Cerro Cituro, Serrania de Pirre, 
1100 m, C. W. Myers coll., 23 January 1966; KU 98522, same 
locality, C. W. Myers coll., 25 January 1966. Colombia: Choco. 
AMNH 18235, Atrato Drainage, Quesada River, R. D. O. John- 
son coll., 6 November 1920. 

Diagnosis. An Anolis related to A. punctatus Daudin and A. 
transversalis A. Dumeril but differing from the first in coloration, 
and in having a large dewlap in the female, and the snout not 
swollen in the male; it differs from the second in lacking sexual 
dichromatism in dorsal pattern, and in a higher number of loreal 
rows, and in coloration. It differs from both species in a lower 
number of lamellae under phalanges 2 and 3 of fourth toe. See 
also Tables 1 and 2. 

Description. (Paratype variation in parentheses.) Head. Head 
scales moderate to small, smooth. Fourteen (11-14) scales across 
snout between second canthals. Weak frontal ridges outlining a 
shallow frontal depression. Scales in frontal depression equal to 
or distinctly smaller than surrounding scales. Seven (7-9) scales 
border rostral posteriorly. Nasal scale separated from rostral by 
one to two scales. Six (5-7) scales between supianasals. 

Supraorbital semicircles separated by three (1-3) scales, sepa- 
rated partly or wholly from the weakly differentiated supraocular 
disks by one (1-2) row of scales. One (1-3) elongate supraciliary 
continued posteriorly by granules. Canthus distinct, nine (8-10) 
canthal scales, the anteriormost very small. Loreal rows eight 
(7-9), the uppermost abruptly larger. 



1967 



NEW ANOLIS FROM PANAMA 




Fig. 1. Anolis chocorwn, new species. Head scales of type, KU 96934. 
Actual snout-vent length 79 mm. 



Temporal and supratemporal scales finely granular, several lines 
of weakly enlarged granules at the angle between temporal and 
supratemporal areas. Interparietal larger (or equal to, or smaller) 
than ear, separated from the supraorbital semicircles by four (2-4) 
somewhat enlarged scales. Laterally, slightly smaller scales grade 
into the supratemporal granules. Behind the interparietal the still 
smaller scales grade very quickly into granules as minute as those 
of the supratemporal area. 

Three to four (2-4) suboculars in contact with the supralabials 
anteriorly, in contact with (or separated by 2-3 scales from) the 
canthus, posteriorly decreasing abruptly in size. Nine (8-9) supra- 
labials to the center of the eye. 

Limbs and digits: Hind foot scales multicarinate. Scales on 
anterior face of upper and lower arm, thigh and lower leg uni- 
carinate, those at knee multicarinate. The larger scales of the 
hind limb as large or larger than the ventrais. About 20 (17-20) 
lamellae under phalanges 2 and 3 of fourth toe. 

Tail: Tail slightly compressed. No dorsal crest, lateral and 
dorsal scales subequal. Two ventral rows larger, keeled. Verticils 
not evident. Postanal scales enlarged in male. Scales immediately 
behind vent smooth. 



BREVIORA 



No. 256 




Fig. 2. Anolis chocoriim, new species. Side view of head of type, 
KU 96934. Actual snout-vent length 79 mm. 



Size (snout/ vent length): c^ (Type) 79 mm; longest 9 73 



mm. 



Color. Color in life is summarized in Table 3. The blue gray 
belly spotted with white is quite distinctive and differentiates this 
species immediately from any previously known. 

As indicated in Table 3, the sexes differ most prominently in the 
color of the dewlap; it is in fact uncertain whether any other 
sex-correlated color differences exist. Both sexes vary somewhat 
in color; the occurrence in both of uniform and patterned dorsa 
is especially conspicuous. It is possible, however, that the pat- 
tern when present is feebler in females than in males; it is true 
in the specimens as preserved. Whether it was true in life is not 
recorded. Whether also some peculiar color variants (the light 
middle of the belly in some males and the gray tongue of one 
female) imply a range of variation in one sex not found in the 
other is unknown. 

Ecology. The Darien localities from which A. cliocorum is 
known are in a broad expanse of relatively undisturbed tropical 



1967 NEW ANOLIS FROM PANAMA 5 

evergreen forest that is moderately well stratified and characterized 
by a lower story of slender palms in many^ places. The area has a 
definite, but not severe, dry season. 

Anolis cJiocorum is arboreal. All individuals observed by day 
were in trees. One was obtained from the branches of a large 
tree immediately after it was felled. Another individual dropped 
from a tree during a storm at night. Two individuals were sleep- 
ing on bushes at night, one in a large fern on the steep bank of a 
mountain stream. Several species of Anolis are associated with 
A. chocorum in eastern Darien; these include two large arboreal 
species {A. biporcatus and A. frenatus), the moderate-sized arbo- 
real A. capito, the smaller bush-dwelling A. chloris, limijrons, 
tropidogaster, and vittigerus, and the semi-aquatic A. poecilopus. 

On July 10, 1965, Charles W. Myers found a striated egg partly 
concealed under loose bark on top of a rotting log on Cerro Quia. 
The egg was 16 mm x 12 mm; it hatched on July 26. The hatch- 
ling (KU 96933) had a snout-vent length of 30 mm and a total 
length of 92 mm. In coloration the hatchling resembled adults of 
A. chocorum by being green above with lateral diagonal rows of 
blackish brown spots. The tail was green with indistinct grayish 
brown crossbands. The venter was white with a faint bluish cast 
to the belly and a pale dull yellow cast to the median part of the 
dewlap. 

Relationships. The series to which A. chocorum belongs is 
clearly an endemic South American one (see discussion of the 
alpha and beta series of Etheridge, 1960, in Williams, 1965). Its 
discovery in series in Darien is probably only an accident of 
activity and attention: similarly thorough and careful collecting 
has not been done in the adjacent areas of Colombia. However, 
a single specimen (AMNH 18235) records its presence there. 
This is a distribution resembling that of a number of other anoles. 
A. chloris, for example, associated in Darien with A. chocorum, is 
likewise a South American autochton, well known in Chocoan, 
Colombia (there known under the names A. palmer i and A. gor- 
gonae), and in trans-Andean Ecuador. The distribution of A. 
chocorum may well be more like that of A. chloris than we now 
know and extend into western Ecuador. However, the range of 
A . chocorum, as at present known, is only eastern Darien Province 
in Panama, where it occurs at elevations of from 100-1100 m, in 
the upper Rio Tuira Basin and on the Pacific slopes of Cerro Quia 
and the Serrania del Darien, and in the Colombian Choco. 

Of the endemic South American group, Anolis chocorum is very 
cieariy most similar to Anolis punctatus Daudin of Brazil, the 



BREVIORA 



No. 256 



Guianas, Peru and Ecuador. The most immediately striking dif- 
ferences are in the coloration of the belly (blue gray with white 
spots) and in the lack of any snout swelling in the male. The 
presence of a much lower lamellar count and a large dewlap in 
the female are soon discovered as further differences. 

A. punctatus is a species of the Amazonian-Guianan forests and 
the closest approach to the Darien-Choco form thus far known is 




Fig. 3. Map of the distribution of Anolis choconim, new species. 



1967 NEW ANOLIS FROM PANAMA 7 

represented by a specimen from the Rio Apaporis in eastern 
Colombia. Over most of its enormous range A. pimctatus has 
smooth ventrals like A. chocorum. Some of the Peruvian, all the 
Ecuadorian, and the Colombian (Apaporis) populations have 
keeled ventrals. For these the subspecific name boulengeri is 
available. 

Overlapping a substantial part of the western range of A. punc- 
tatus, all of boulengeri, and part of Peruvian typical punctatus is 
A. transversalis. Structurally, this differs from A. chocorum in 
lower loreal and toe lamellar counts and in the peculiar mental- 
postmental scale pattern. The very striking throat pattern of the 
female is very different from the condition of either sex in A. cho- 
corum and the bold crossbanding of the dorsal surface of the 
female transversalis is also very different. The dorsal pattern of the 
male transversalis (originally described as a distinct species, buck- 
leyi), however, does somewhat resemble the irregular spotted 
condition mentioned above as seen in some A. chocorum. 

A. transversalis, so far as known, gets geographically no closer 
to A. chocorum than does A. punctatus. There is the very sub- 
stantial gap in Andean Colombia. 

The similarity in many scale characters of the three species 
is seen in Table 1 and is reinforced by the general similarity in 
habitus and size. A. chocorum, as shown in Tables 1 and 2. in many 
ways connects A. punctatus with A. transversalis. 

The oblique rows of dark blotches on the dorsum of some 
specimens (well shown in the type. Fig. 1) resemble the dorsal 
pattern of the giant anoles, A. frenatus, A. princeps, A. squamu- 
latus, A. latifrons. All of these have considerably higher scale 
counts in all respects than A. chocorum, and differ also in their 
much larger size (over 100 mm snout- vent length rather than 
70-80 mm). They are, however, quite clearly closely related, and 
in mental shape, for example, are more like A. chocorum than are 
either A. punctatus or A. transversalis. 

These apparent affinities of A. chocorum in several directions 
seem to effectively subvert the recent attempt of the senior author 
to define a punctatus group from which A. transversalis was 
expressly excluded as a "fringe species" and from which the giant 
anoles were considered still more remote. This newly discovered 
anole clearly and firmly connects the so-called "fringe species" 
(and the giant anoles as well) with the punctatus group. Thus 
we seem to be faced with a single series of species which, how- 
ever, exhibit striking and peculiar trends in several adaptive lines. 
The presence of still surviving intermediates helps to connect up 
and to indicate the affinities of the more peculiar species. 



BREVIORA 



No. 256 



TABLE 1 

Scale Counts 
Anolis chocorum and Relatives 





piinctalus 








(including 








boulengeri) 


chocorum 


iransversolis 


Scales across snout 








between second canthals 


8-14 


10-14 


4-8 


Scales between semicircles 


0-2 


1-3 


0-1 


Scales between interparietal 








and semicircles 


0-4 


2-4 


0-3 


Loreals 


4-7 


6-8 


3-6 


Labials to center of eye 


6-10 


8-10 


6-9 


Lamellae, 4th toe 


23-30 


17-20 


22-27 



TABLE 2 

Qualitative Characters 
Anolis chocorum and Relatives 





piinctatus 








(including 








boulengeri) 


cliocoritni 


transversalis 


Snout 


Swollen in $ 


Not swollen in 


Not swollen in 






either sex 


either sex 


Ventrals 


Smooth (piinc- 
tatus) or keeled 
(boulengeri) 


Smooth 


Smooth 


Mental 


At least as deep 
as wide 


Wider than deep 


Deeper than wide 


Sublabials 


+ 


— 


+ (large) 


Dewlap ( 6 ) 


Moderate (lateral 


Large (lateral 


Large (lateral 




scales large, elon- 


scales small. 


scales small, 




gate, in close- 


weak, in multiple 


weak, in multiple 




packed rows) 


rows widely sep- 


rows widely sep- 






arated by naked 


arated by naked 






skin) 


skin) 


Dewlap ( ? ) 


Absent 


Large as in male 


Large as in male 






but difTerently 


but differently 






colored 


colored 


Dorsal pattern 


Not sexually 


Not sexually 


Strong sexual 




dichromatic 


dichromatic 


dichromatism 



1967 



NEW ANOLIS FROM PANAMA 



TABLE 3 
Color in life of A . chocorum 



DEWLAP 

Edge anteriorly white, 

posteriorly orange 

Sides pale orange with several 

faint bluish white or green 

lines 

Base pale green 

DORSUM 

Green, uniform or with 
oblique rows of dark green 
blotches on flanks 

VENTER 

Green of dorsum continued 
on edge of belly. Center of 
belly light or blue gray. Both 
blue and green areas heavily 
spotted with white 

EYELIDS 

Edges yellow 

IRIS 

Pale brown (coppery) with- 
out dark markings 

TONGUE 

Yellow orange, sometimes 
with tip dark gray 

LINING OF THROAT 
Black 



Edge white 

Sides green with white lines or 
pale green 

Base pale yellow or gray 



As in S 



As in $ , but center of belly not 
light in any specimen 



As in 3 



As in S , or reddish brown with 
gray periphery 



As in $ , or pale flesh gray or 
pale greenish yellow or pale 
yellow 



As in $ 



REFERENCE CITED 

Williams, Ernest E. 

1965. South American Anolis (Sauria, Iguanidae): Two new species 
of the piinctatiis group. Breviora, No. 233:1-15. 



(Received 5 April, 1966.) 



10 



BREVIORA 



No. 256 




1967 



NEW ANOLIS FROM PANAMA 



11 




12 



BREVIORA 



No. 256 




lu 



BREVIORA 

Museitaiin of Coampsirsitive Zoology 

Cambridge, Mass. 3 February, 1967 Number 257 

A REVIEW OF THE CLARK FORK VERTEBRATE FAUNA 
By Roger C. Wood 

INTRODUCTION 

The Clark Fork fauna, hitherto regarded as the standard for 
one of the provincial ages of the North American Continental Ter- 
tiary (Wood et al., 1941, p. 9 and plate I), is known from only 
one area in the Bighorn Basin of northwestern Wyoming. In con- 
trast to other Paleocene faunas representing provincial ages, no 
Clarkforkian quarry or pocket has yet been discovered. Con- 
sequently, this fauna has never been as well characterized as others 
in the standard sequence, and the validity of this provincial age 
has not been very well substantiated. This paper reviews previ- 
ous work on the fauna and discusses some associated problems. 
Conclusions are based on field observations, a re-evaluation of 
the described collections, inspection of the original field notes, 
and unpublished annual reports of the Department of Vertebrate 
Paleontology of the American Museum of Natural History. 

Professor Glenn L. Jepsen suggested this problem as a senior 
thesis topic while I was an undergraduate at Princeton. His 
generosity in permitting me access to Princeton's valuable collec- 
tion of Paleocene fossils is here gratefully recognized. Dr. Mal- 
colm C. McKenna of the American Museum of Natural History 
most kindly made available for study the specimens upon which 
the original Clarkforkian faunal description was based; I have 
also profited from discussions with him. I am much obliged to 
Professors Bryan Patterson, George Gaylord Simpson, Albert Wood, 
and Dr. Leigh Van Valen for criticisms and suggestions concerning 
this manuscript. In addition I am grateful to Dr. Giles Maclntyre, 
who found the map used by Walter Granger in the Bighorn Basin 
during the field seasons of 1910 through 1912. A portion of this map 
is reproduced in Figure 2. The hospitality of the Churchill family 
of Powell, Wyoming, added considerably to the enjoyment of 



2 BREVIORA No. 257 

doing field work during the summers of 1961 and 1962. I would 
also like to thank Mrs. Frances Wood and Miss Margo Hayes 
for typing the numerous revisions of this paper. 

Financial assistance provided from the John Boyd Fund of 
Princeton University enabled me to spend the latter part of the 
summer of 1961 and the entire field season of 1962 in the Big- 
horn Basin, working in the type areas of the Clark Fork fauna. 
Support from the National Science Foundation training grant for 
evolutionary biology at Harvard University was helpful during 
the final stages of my research. 

Abbreviations used in this paper are as follows: AMNH, 
American Museum of Natural History; PU, Princeton Univer- 
sity; FMNH, Field Museum of Natural History; USNM, United 
States National Museum. 

HISTORY OF STUDY 

During the summers of 1910 through 1913, and again in 1916, 
field parties sponsored by the American Museum of Natural His- 
tory conducted extensive investigations of the Tertiary sediments 
of the Bighorn Basin. Some of the results were published by 
Sinclair and Granger in 1912; of interest to the present study 
was the recognition of a new vertebrate fauna occurring near the 
top of the Fort Union.' Specimens comprising the fauna were 
obtained from three different localities whose approximate posi- 
tions, determined from the following descriptions, are labelled 
A, B, and C on the accompanying map (Fig. 1). These areas 
were described (Sinclair and Granger, 1912, pp. 59-60) as being 
". . . on the southwest slopes of McCulloch Peak . . . about a 
mile due east of the point where the Fort Union-Wasatch contact 
line crosses the Shoshone River, 245 feet stratigraphically below 
the contact with the red-banded beds [A] ; on the north side of 
the Shoshone River in the bluffs opposite Ralston station [B]; 
to the northwest of Ralston on Big and Little Sand Coulee [C]." 
Only the McCulloch Peak locality (A) was regarded as being 
unquestionably Paleocene; some uncertainty was expressed con- 
cernmg the exact stratigraphic position of the other two areas. 
This doubt led the authors to conclude ". . . we feel that further 
examination of the stratigraphy is desirable. Should the beds in 
question prove to be older than the Knight [early Eocene], and 
it be deemed advisable to give them a formation name, they 
may be referred to as the Ralston beds or Ralston formation." 

1 The Paleocene rocks in this area have been referred to as the Polecat 
Bench Formation; see Jepsen, 1940. 



1967 



CLARK FORK VERTEBRATE FAUNA 




4 BREVIORA No. 257 

A more precise definition of this fauna was subsequently pub- 
lished by Granger in 1914. Further collecting had increased the 
number of specimens to fragments "representing nearly 200 in- 
dividuals" (Granger, 1914, p. 204). Characterized by the absence 
of perissodactyls, artiodactyls, rodents and primates (the last two 
not then known from Paleocene deposits), the fauna was com- 
posed predominantly of representatives of two genera, Phena- 
codus and Ectocion, which constituted over three-fourths of the 
collection. The rest of the fauna included some Paleocene genera, 
whose range was known to extend into the overlying Eocene Gray 
Bull beds, and genera of Gray Bull age. Assignment of the 
fauna to an appropriate epoch was supposedly aided by the abun- 
dant presence of the reptilian genus Champsosaurus, which was at 
the time considered to be a distinctly Cretaceous and Paleocene 
genus. This evidence, coupled with the absence of the four orders 
of mammals previously mentioned, inclined Granger to regard 
these beds as being of late Paleocene rather than early Eocene age. 

Because the name "Ralston" was preoccupied by a group of 
Pennsylvanian rocks. Granger substituted in its place the term 
"Clark Fork" beds. These sparsely fossiliferous strata were esti- 
mated to attain a maximum thickness of 500 feet. Besides the 
McCulloch Peak locality (A), exposures were prominent "In the 
bluff [Polecat Bench] in the northern part of the Bighorn Basin" 
where "characteristic fossils were found as far east as a point 
north of Powell [i.e. in beds extending northeast from B]." In 
addition, two new localities were described: "In the Clark Fork 
basin the fossils were obtained from both sides of the wagon road 
where it drops down to the Big Sand Coulee from the Bighorn 
Basin divide, also from . . . the opposite side of Clark Fork River 
between the mouths of Line and Little Rocky Creeks" {Ibid., 
p. 204). These last areas, not mentioned in Sinclair and Granger's 
paper, are marked on the map as localities D and E. 

Descriptions of the various new types ascribed to the Clark 
Fork fauna appeared in a series of papers by Matthew ( 1915a, b, c; 
1918) and Granger (1915), and in one by Simpson (1929). It is 
interesting to note Simpson's remark (p. 2) that some of the 
specimens being described might be from the lowest Eocene Sand 
Coulee horizon rather than from the Clark Fork. However, no 
reason was given for this statement. 

Not until 1930 was a complete faunal list published (Jepsen, 
1930b, pp. 492-493), which included several species collected by 
Princeton University field parties subsequent to the American 
Museum's initial discoveries. Three new forms were described, 



1967 CLARK FORK VERTEBRATE FAUNA 5 

the most significant for this study being Dipsalodon matthewi, the 
only genus restricted to the Clark Fork. In addition, note was 
made of the fact that Champsosaiirus could no longer be used to 
demarcate the upper limit of Paleocene beds, thus invalidating 
Granger's premise that this genus could be used for distinguishing 
between Paleocene and Eocene sediments. 

Revision of the Clark Fork fauna was undertaken by Simpson 
(1937). Several forms from the original collection were for the 
first time described. In addition, one species, Ambloctonus pris- 
cus Matthew (= Palaeonictis occidentalis Osborn; Denison, 1938, 
p. 175), was removed from the fauna on the grounds that field rec- 
ords associated with the specimen clearly indicated that it belonged 
with the Sand Coulee fauna. A further deletion from the fauna was 
made by Jepsen, who removed Parectypodus, the only multi- 
tuberculate that had been regarded as part of the fauna, because 
its original inclusion had resulted from "an error in locality identi- 
fication" (Jepsen, 1940, p. 324). No additions or deletions to the 
fauna have subsequently been published. 

INTERPRETATION OF MAPS 

Since Granger used only verbal descriptions, rather than sec- 
tion, township, and range data for indicating the locations of 
specimens and type areas, great value must be attached to the 
recent finding of his original field map, part of which is reproduced 
as Figure 2. Its discovery permits much greater confidence in the 
determination of some of the areas where the original Clark Fork 
collections were made. No date appears on the map, which was 
taken from Fisher's 1906 report. Perusal of the field notes, how- 
ever, indicates that Granger was in this part of the Bighorn 
Basin only in 1911 and 1912. Therefore, it seems reasonable to 
infer that the hachures sketched on his map, undoubtedly intended 
to indicate badlands, correspond to areas prospected in those 
years. These badlands, consequently, must represent the type 
localities for the Ciark Fork and Sand Coulee beds described by Sin- 
clair and Granger (1912) and Granger (1914). Some of these 
hachures coincide with the placement of letters B, C, D, and E on 
Figure 1, thus confirming the positions of most of Granger's type lo- 
calities. Unfortunately, uncertainty must still remain as to the exact 
location of the area on the southwest slopes of McCulloch Peak (in- 
dicated approximately in Figure 1 by the letter A) because no 
hachures were drawn on this part of Granger's map. 

Confusion may be averted by noting the following problems 
concerning the same name for different places on various maps. 



BREVIORA 



No. 257 




N 9S I 



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1967 



CLARK FORK VERTEBRATE FAUNA 







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8 BREVIORA No. 257 

On Fisher's map (1906; see Fig. 2), Bennett Creek is shown as a 
tributary of Little Rocky Creek. All subsequent maps, however, 
depict Little Rocky Creek as being a tributary of Bennett Creek 
(see Fig. 1). Furthermore, Fisher's map indicates Little Sand 
Coulee as being a branch of Big Sand Coulee, but in the Annual 
Report of the Department of Vertebrate Paleontology for 1912, 
Granger remarks that ". . . What is known locally as Little Sand 
Coulee [is] not the Little Sand Coulee of Fisher's map." The 
actual location of the coulee is written on Granger's field map (see 
Fig. 2), and later maps all follow Granger's positioning of the 
coulee. 

A final problem is the question of where the Paleocene-Eocene 
contact in the Clark Fork Basin is situated. The first geological 
map of the area was published by Fisher in 1906. Presumably the 
contact there drawn (refer to Figure 3 for position of this and 
subsequent contact lines) represented only a rough estimate, as 
the line across the basin is unnaturally straight, and no discussion 
of the means by which the contact was detected are included in 
Fisher's report. Sinclair and Granger (1912) apparently did not 
take Fisher's contact very seriously, since, with the exception of 
the McCuUoch Peak locality, the beds they describe as having 
yielded their presumably Paleocene Clarkforkian material are 
located several miles to the southwest of Fisher's line, well into his 
Eocene, as can be seen by referring to Granger's field map (Fig. 
2). Their 1912 paper, however, includes a sketch map which 
shows a contact coinciding with that of Fisher's map. In 1930 Jep- 
sen published a map whose Paleocene-Eocene contact lay several 
miles southwest of and parallel to that recognized by Fisher. The 
line was drawn more precisely on the basis of paieontological evi- 
dence, using the conventional criterion that the lowest Eocene 
beds are marked by the first appearance of Hyracotherium. Agree- 
ing relatively closely with Jepsen's contact is one drawn by Stow 
(1938) on the basis of the presence or absence of certain types 
of heavy minerals that he thought permitted differentiation be- 
tween Paleocene and Eocene sediments. Although apparently un- 
aware of Jepsen's and Stow's works (as their papers are not indi- 
cated as being sources of data), Andrews, Pierce, and Eargle 
(1947) drew a contact closely paralleling these previous ones, but 
generally slightly to the northeast. For that part of the basin 
shown on the map. Pierce (1965) indicates minor deviations from 
his previously drawn contact except in the area directly east of 
the mouth of Little Sand Coulee, where his boundary diverges 
appreciably from the earlier one. Thus, the boundary between 



1967 CLARK FORK VERTEBRATE FAUNA 9 

Paleocene and Eocene beds has been reasonably clearly estab- 
lished within rather narrow limits. The area where the greatest 
divergence exists between the various lines, however, is to the 
east of the mouth of Little Sand Coulee, a fact which is of some 
importance concerning the provenance of about half the specimens 
in the Princeton collection (see p. 21 ) . 

THE AMERICAN MUSEUM COLLECTION 

Data concerning the American Museum's Clark Fork collection 
are summarized in Table I. One fact readily apparent from this 
table is that there are only 89 specimens in the original collection, 
not the "nearly 200" mentioned by Granger. This observation may 
indicate that many specimens, in addition to the one discussed by 
Simpson (1937; see above, p. 5), have been at one time or 
another removed from the fauna without this action having been 
noted in any publication. On the other hand, it may merely indi- 
cate that Granger had overestimated the size of the collection. At 
any rate, the basis available for defining and characterizing the 
type collection is less than half as large as published data indicate. 

An additional feature worthy of note is the nature of the speci- 
mens from the McCuUoch Peak area (A, Fig. 1); these are 
described in Table I, footnote 10. The identifiable fragments in- 
clude no elements restricted to the Paleocene. This fact lends 
itself to some interesting speculation, as this locality is the only 
one of the three originally described by Sinclair and Granger as 
representing unquestionable Paleocene sediments. In this connec- 
tion some of Granger's comments in the 1911 Annual Report of 
the Department of Vertebrate Paleontology are most informative: 
"These beds [at Ralston] proved to be of the same age as those 
[from the base of McCuUoch Peak] described above and although 
fossils were scarce some 50 individuals were obtained. The small 
phenacodonts constituted about Vi the entire number and there 
still was no trace of Perissodactyla, of Artiodactyla, nor Rodentia. 
The presence of Bathyopsis, Limnocyon, and a Metacheiromid were 
surprising, considering that the beds lie below the Wasatch. . . . 
The dip of the [Ralston] beds where exposed would carry them 
below the Wasatch at the base of McCuUoch Peak, allowing no 
fault or change in dip." In other words. Granger apparently con- 
sidered that the fauna contained in the beds north of Ralston 
included various forms that he would have expected to find in 
typical early Eocene sediments. Thus the Clark Fork beds were 
defined on the basis of negative evidence, i.e., what had not been 



10 



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12 BREVIORA No. 257 

Footnotes for Table I (continued) 

SDenison (1938, p. 175) states: "Amer. Mus. No. 16068 from the Clark 
Fork horizon, described by Matthew ... as 'lOxyaena sp. innom.', is 
referred here doubtfully to Dipsalodon. It includes much worn upper teeth 
of approximately the same size as D. matthewi." Van Valen (1966, p. 82) 
has referred this specimen to D. matthewi. 

^AMNH No. 18668 has been tentatively referred to this genus, but specific 
identification is difficult because of the scrappy nature of the specimen. 
4 Simpson (1937) recognizes three, or possibly four, species of Esthonyx 
from the Clark Fork beds: E. bisulccitiis, E. grangeri, E. latidens, and per- 
haps E. spatulariiis. AMNH No. 16065 is labelled as "E. cf. spatulariiis 
(type)," but is figured in Simpson (1937, p. 6, fig. 1) as £. ?bisidcatns. How- 
ever, this specimen could not be the type for either species since it was 
found in 1912, while the two species in question were described by Cope in 
1874 and 1880, the type for E. spatulariiis being AMNH No. 4809 and that 
for E. hisiilcatus being USNM No. 1103 (Gazin, 1953, pp. 17, 21). None 
of the specimens in the collection are presently labelled as representing E. 
bisulcatiis. 

•"•Jepsen (1930b, p. 493) listed this species as being included in the Ameri- 
can Museum's collection although Simpson (1937) does not include it in 
his Clark Fork faunal list. However no mention or reason for its deletion 
is given. Gazin (1962, p. 67) notes that "the type of M. prisciim from the 
Clark Fork beds could not be located in the American Museum collections." 
Personal investigation has confirmed this report. 

"Denison (1938, p. 167) referred Matthew's type of Dipsaladictis platypus 
(AMNH No. 15857) to this species, stating that "it is definitely an 
oxyaenine, which differs from other primitive species of Oxyaena only in 
its smaller size." One additional specimen (AMNH No. 18667) is labelled 
as belonging to this species. 

^This specimen (AMNH No. 15850) was listed as "Nyctitheriidae genus 
and sp. indet." by Simpson (1937, p. 4), but has been described as Palac- 
oryctes punctatus by Van Valen (1966, p. 56). 

'^Many Phenacodus specimens are represented as disarticulated bones or 
miscellaneous tooth and jaw fragments rather than as associated sets of 
teeth or bones. Individual fossils were labelled variously as Phenacodus, 
P. primaevus, P. intermedins, and P. hemiconns, and thus do not concur 
with the two subspecies included in Simpson's 1937 faunal list. 
^Only two specimens in the collection are labelled as Probathyopsis prae- 
cursor. Four other specimens in the collection are labelled Bathyopsis or 
Bathyopsidae. The latter fossils were clearly intended by Simpson to be 
included in the new genus since one of them comes from opposite the mouth 
of Little Rocky Creek, one of the localities mentioned as an area where 
specimens of this species had been found, and it is in fact the only speci- 
men of any kind from this area. For another of the four localities listed 



1967 CLARK FORK VERTEBRATE FAUNA 13 

as having yielded P. praecursor remains, three miles east of the mouth of 
Pat O'Haia Creek (locality 2, Fig. 3), no Clark Fork specimens could be 
located. Simpson suggested (1929, p. 2) that "Some of this material may 
be from the Sand Coulee horizon." Four specimens, including the type, are 
from locality 1 (Fig. 3), confirming the idea that at least some of 
the specimens are part of an early Eocene fauna rather than a late Paleo- 
cene Clark Fork one. 

^" In addition to the specimens listed in the chart, the following material is 
included in the collection: three boxes of "miscellaneous bones and teeth" 
from the west slope of McCulloch Peak, containing fragments identifiable 
only as Champsosaurus, Phenacodus, Ectocion, and a pantodont; four boxes 
of miscellaneous fragments labelled only as being from the "Clark Fork 
Basin," and another from "north of Ralston." None of this fragmentary 
material would appear to affect the conclusions based on analysis of the 
better specimens in the collection, and hence has not been discussed. 

found in them, rather than on fossils actually contained in them 
which could be characterized as having a distinctive Paleocene 
aspect. Since the type fauna was principally found in strata whose 
position in the section was by no means certain, and since the 
fauna from the only site regarded by Granger as being definitely of 
Paleocene age had no obvious Paleocene affinities, it is not at 
all certain whether or not a Clark Fork fauna can in fact be 
distinguished. 

Examination of the labels accompanying the specimens in the 
American Museum's Clark Fork collection reveals that over 85 
per cent of the fossils, acquired in the years 1910-1916, were 
found at three sites. These localities were: 

(1) the head of Big Sand Coulee (Area D, Fig. 1); 

(2) east, or three miles east, of the mouth of Pat O'Hara 
Creek (Area C, Fig. 1 ) ; 

(3) north, or three miles north of Ralston; or bluff, or base 
of bluff, northeast of Ralston (Area B, Fig. 1). 

For convenience of reference, American Museum specimens from 
these areas will henceforth be designated merely as being from 
locality number 1, 2, or 3 (see Fig. 1). 

Locality number 1 is in the region characterized by Granger as 
the type area of the lowest Eocene Sand Coulee beds (1914, 
p. 205). "Near the head of Big Sand Coulee in the Clark Fork 
basin there is a series of about 200 feet of red-banded shales lying 



14 BREVIORA No. 257 

between the gray shales^ of the Clark Fork and the Systemodon- 
bearing gray shales above which are referred to the Gray Bull 
beds. The outcrop of these red-banded beds extended, from the 
high bluff on the south side of the Coulee, in a northwesterly direc- 
tion across the basin for several miles." Furthermore, in this same 
paper, Granger describes (p. 203) the Gray Bull beds and men- 
tions various areas where they outcrop. "In the Clark Fork basin 
they are exposed in the southwestern part along the heads of Big 
and Little Sand Coulees, where they are of a uniform gray color 
instead of the usual gray, red, and yellowish banding." Reference 
to all the geologic maps of this part of the basin shows that the 
Paleocene-Eocene contact line crosses the southern branch of Big 
Sand Coulee approximately two miles below its head, and the 
western branch nearly five miles from its head. The eastern 
prong of the coulee, of which the one on the map is only one of 
many, is probably not that referred to on the labels, since its head 
originates at the base of Polecat Bench and such a fact would 
have made a convenient reference for the labels. As Granger 
noted the misidentification of Little Sand Coulee on his field 
map (refer to p. 8 and Fig. 2), he may have been talking about 
either the southern or western branches of Big Sand Coulee. 
However, whichever of these two is chosen, the locality would 
be well into Eocene Gray Bull or Sand Coulee strata. In view 
of the fact that Granger's field map indicates an extensive area 
of prospected exposures at the head of the southern branch of 
Big Sand Coulee, and none are marked near the head of its 
western fork, it is most probable that this area is the one in 
question. There is no doubt at present that these are Gray 
Bull beds. 

In addition, many specimens in the American Museum collec- 
tions labelled as "Wasatch," "Sand Coulee," or "Gray Bull" are 
noted also as having come from "the head of Big Sand Coulee." 
For example, the label accompanying the type of Apheliscus nitidus 
(AMNH No. 15849, listed in Table I) reads "Clark Fork forma- 
tion, Clark Fork, Wyoming, 1911, head of Big Sand Coulee." 
That accompanying another specimen of the species, AMNH No. 
16925, reads "Wasatch formation. Sand Coulee Beds, Clark Fork, 



' Jepsen has pointed out that it is impossible to use color banding for 
distinguishing Clark Fork from overlying beds, since ". . . red strata 
occur well down in the Clark Fork beds" (1940, p. 237; see also 1930b, 
p. 493). 



1967 CLARK FORK VERTEBRATE FAUNA 15 

Wyoming, 1913, head of Big Sand Coulee." All other specimens 
of A. nitidus are similarly labelled as being from Gray Bull beds. 
It would therefore appear that the two cited specimens, although 
labelled as being from two different horizons, are most probably 
from the same one, the Gray Bull. Consequently, based on the 
evidence revealed by the specimen labels, it is probable that all 
supposed Clark Fork fossils labelled as having been found at the 
"head of Big Sand Coulee" should more properly be referred to 
the lower Gray Bull or Sand Coulee beds. Since half of Sinclair 
and Granger's original Clark Fork fauna (see Table I) was 
obtained from the vicinity of the head of the Coulee, it is likely 
that these specimens are part of one or more early Eocene faunas 
rather than of a late Paleocene Clark Fork fauna. 

"East [or three miles east] of the mouth of Pat O'Hara Creek" 
(locality 2) is an area lying in an extensive series of Willwood 
exposures. This locality, as is true for locality 1 also, was never 
mentioned by Granger as one of the five type areas (A through E, 
Fig. 1) for the Clark Fork beds. Granger's field map (Fig. 2) 
shows two sets of badlands sketched along a north-south axis in 
this region, the southern one being labelled "red-banded beds" 
and the northern one "below red-banded beds." These may be 
interpreted in the light of Granger's remarks in the 1913 Annual 
Report of the Department of Vertebrate Paleontology: "Section 
across Clark Fork Basin from head of Little Sand Coulee northeast 
to mouth of Big Sand Coulee: 

Wasatch — gray shales — 500+ feet 
[Sand Coulee] — red-banded shales — 200 feet 
Ralston — gray shales — 500+ feet." 
Clearly, the red-banded beds must represent some of Granger's 
Sand Coulee exposures. Consequently, the beds farther north can 
only be regarded as Clark Fork beds, and it seems reasonable to 
assume that locality 2 corresponds to this area, which has always 
been mapped as being covered by Eocene beds (Fig. 3). It seems 
curious that part of a fauna purported to be of latest Paleocene 
age should be derived from sediments whose presumed early 
Eocene age has never been questioned. With respect to this para- 
dox, an examination of the specimens from locality 2 is most 
instructive. Reference to Table II reveals that all of the Clark 
Fork fossils from locality 2 are also known from the Sand Coulee 
or Gray Bull, and often from both of these horizons. In fact, the 
one specimen of Phenacodus hemiconus from this locality was 
apparently regarded by Granger as being of Gray Bull provenance, 



16 BREVIORA No. 257 

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1967 CLARK FORK VERTEBRATE FAUNA 17 

as he states (1915, p. 339): "Eleven specimens from the Gray 
Bull beds and one from the Lost Cabin are referable to this small 
variety." Thus there is nothing distinctive about the specimens 
from this locality to suggest their association with the Clark Fork 
fauna; on the contrary, based on the presence of Phenacodm 
hemiconus, there is a slight suggestion that these specimens may 
actually be of Gray Bull derivation. At any rate, since this area 
was not described as one of Granger's type localities, and since 
its faunule does not differ significantly from those of other Sand 
Coulee or lower Gray Bull sites, it seems highly probable that 
"east" or "three miles east of Pat O'Hara Creek" is in Sand Coulee 
or lower Gray Bull, and not Clark Fork, strata. 

Specimens from locality 3 are too vaguely labelled to be of any 
value in this study. Faunas from the southeastern side of Polecat 
Bench have been found ranging from the Silver Coulee through 
the lower Gray Bull. Without knowledge of section, township, and 
range, it is virtually impossible to infer to what fauna these Ameri- 
can Museum fossils belong. It is very likely that elements from 
several faunas are actually represented by these fossils. There- 
fore, on the basis of the foregoing inferences, none of the Ameri- 
can Museum materials from localities 1, 2, or 3 can certainly be 
ascribed to what have been regarded as Clarkforkian strata. 
Rather, those from localities 1 and 2 can be considered as per- 
taining in all probability to one or more early Eocene faunas, 
while specimens from locality 3 cannot be definitely assigned 
either to a late Paleocene or to an early Eocene fauna. 

No specimens in the collection are labelled as having come 
from either side of the wagon road dropping into Sand Coulee 
Basin (area D on map). However, any such specimens would 
definitely be of early Eocene age. 

Listed in Table I is another group of specimens that may be 
considered as a unit for the purposes of this review. In contrast 
to the previous three groups discussed, which were character- 
ized by similarities in locality data, this fourth group is distin- 
guished by a total lack of useful locality information. Labelled 
variously as coming from "Clark Fork beds" or "formation," or 
even "Clark Fork Period," these fossils clearly can be of no use 
in defining the composition of the fauna for the following reasons. 
First, as has already been shown, if these fossils came from locali- 
ties 1, 2, or 3, they probably should not be included in the Clark 
Fork fauna. Second, if these specimens were not found at locali- 
ties 1, 2, or 3, then they could have been picked up anywhere by 



18 BREViORA No. 257 

chance in a basin where four faunal horizons (Silver Coulee, Clark 
Fork, Sand Coulee, and Gray Bull) have been described, as the 
labels associated with these specimens give no guidance as to the 
specific area in which they were collected. 

The five remaining specimens from the type collection, not 
excluded from being Clarkforkian by virtue of the considerations 
already discussed, are listed in Table I under the column headed 
"other." Of the fossils in this category, only two specimens, one 
of Coryphodon sp., and one of Ectocion osbornianus ralstonensis, 
were found at one of Granger's type areas, "between Little Rocky 
and Line Creeks," marked as locality E on the map. The hachures 
in section 3, T 57 N, R 102 W of Granger's field map probably 
represent the exposures in which these specimens were collected. 
Andrews, Pierce, and Eargle (1947) showed both Paleocene and 
Eocene sediments in this region. However, Pierce (1965) indi- 
cates that only Eocene and Quaternary deposits are found within 
this section. Very probably, therefore, the two specimens under 
consideration should be associated with the Sand Coulee or Gray 
Bull faunas. 

Also in this last category is one specimen of Phenacodus pri- 
niaevus that was found "6 miles north and one mile east of Powell, 
Wyoming, at base of bluff," unquestionably an area of Paleocene 
deposits. This specimen may be regarded as coming from the 
Poiecat Bench Formation. 

Accompanying a single specimen of ProbatJiyopsis praecursor 
(AMNH 16063) are the following locality data: "southeast side 
of Clark's Fork, opposite mouth of Little Rocky Creek, Ralston 
[Clark Fork] formation." The most recent maps show Little 
Rocky Creek as being a tributary of Bennett Creek, and the sedi- 
ments opposite its mouth, on the west side of the Clarks (or 
Clark) Fork River are mapped as Quaternary in age. However, 
Fisher's map shows Bennett Creek as being a tributary of Little 
Rocky Creek (see p. 8), and the rocks opposite the latter creek's 
mouth on the east bank of the Clarks Fork River were mapped as 
being of Eocene age. Therefore, it would seem that the locality 
designated by the label refers to the area opposite the mouth of 
Bennett Creek as its course is presently mapped. Hachures in this 
vicinity on Granger's field map (Fig. 2) would appear to confirm 
this deduction. Excluding the Quaternary deposits immediately 
to the east of the Clarks Fork channel, Jepsen (1930b), Stow 
(1938), Andrews, Pierce, and Eargle (1947), and Pierce (1965) 
have all regarded the sediments of this area as being of Paleocene 
age. In this review, therefore, this one specimen of ProbatJiyopsis 
will be regarded as being from upper Paleocene beds. 



1967 CLARK FORK VERTEBRATE FAUNA 19 

In the case of the only fossil not yet discussed, the locality data 
are relatively specific. The type of Oxyaena aequidens is from 
"Clark Fork beds, Clark Fork Basin, 1912, about seven miles east 
of Pat O'Hara Creek." Rocks in this part of the Clark Fork Basin 
are extremely flat-lying, and it is rather difficult to establish the 
contact between the conformable Paleocene and Eocene strata, as 
can be seen from the fact that the contact lines drawn by different 
mappers differ markedly one from another (see Fig. 3). The 
Oxyaena specimen was probably found somewhere in this contact 
region. However, considering both the fact that the distance esti- 
mate made in 1912 could have been at best only an approxima- 
tion, and that there is no agreement as to exactly where the contact 
lies in this region anyway, it can only be a matter of speculation as 
to whether this specimen belongs to a late Paleocene or an early 
Eocene fauna. Thus, the specimen is of no help in defining the 
Clark Fork fauna. 

Of the original Clark Fork collection of 89 specimens, therefore, 
only two, one of Phenacodus primaevus and one of Probathyopsis 
praecursor, may be certainly regarded as being from the Polecat 
Bench Formation. The remainder of the specimens must be ex- 
cluded from this category for one of two reasons. Either the fos- 
sils are of a difi'erent age from what they were originally con- 
sidered to be, or the locality data associated with the specimens 
are not specific enough to be of any stratigraphic utility. As it 
would be unwarranted to try to define a fauna on the basis of 
only two species, each represented by only one specimen, it is 
necessary to analyze other evidence pertaining to this problem. 

THE PRINCETON UNIVERSITY COLLECTION 

Comparison of Tables I and III reveals several marked dif- 
ferences between the Princeton and the American Museum Clark 
Fork collections. First, there are only about one-fifth as many 
specimens in the Princeton collection as in that of the American 
Museum. Second, there are somewhat better documented locality 
data for the Princeton material. Still another contrasting factor 
is that the preponderance of the American Museum specimens was 
collected during the two field seasons of 1910 and 1911, while 
Princeton parties have been adding to the collection continuously 
for over 30 years. Only genera added to Clark Fork fauna by 
Jepsen in 1930 (with the exception of Parectypodus, see above, 
p. 5) will be considered in the present review, as these elements 
are the ones that have formed part of the basis on which the 
fauna has usually been defined. 



20 BREVIORA No. 257 



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1967 CLARK FORK VERTEBRATE FAUNA 21 

Over one-half of the specimens in the Princeton collection were 
found in an area designated on the labels as "east of [or near] the 
mouth of Little Sand Coulee" (see Table III). How restricted an 
area this description represents is a moot point since the topog- 
raphy of this region offers few outstanding features to serve as 
convenient reference points and, as noted in the previous section, 
various mappers have differed in their opinions of exactly where 
the contact between the Paleocene and Eocene should be drawn 
in this region. Consequently, these fossils may represent strictly 
Paleocene forms, strictly Eocene forms, or some of both ages. In 
this case, not only a lack of accurate locality data, but also a lack 
of agreement as to where the contact is, hinder any useful evalua- 
tion of this part of the collection with respect to defining the Clark 
Fork fauna. The remainder of the specimens in the Princeton 
collection represent isolated finds scattered across a broad region. 
Nearly fifty miles separate the farthest spaced of the specimens 
attributed to the Clark Fork. 

Representatives of two species, Ectocion osbornianus ralsto- 
iiensis and Phenacodus primaevus, comprise roughly one-third of 
the Princeton Clark Fork collection. Both of these species are 
found in overlying faunal horizons. In addition, the presence of 
Phenacodus sp. and Ectocion sp. has been reported from the Silver 
Coulee faunal horizon immediately below the Clark Fork beds 
(Jepsen, 1930b, p. 491). Reference of most of the American 
Museum specimens, previously regarded as being of Clarkforkian 
age, to one or more of the early Eocene faunas was shown in the 
previous section to be virtually certain. Some aspects of the 
phylogeny of these species, originally reconstructed from studies 
of American Museum materials, have thus been deduced on the 
basis of erroneous stratigraphic assumptions. Restudy is required 
before they can be used in the definition of any fauna. Conse- 
quently, specimens of Phenacodus and Ectocion are at present of 
no value in helping to define a characteristic Clark Fork fauna. 

The specimen originally designated as Coryphodon (species 
undetermined) by Jepsen (1930b, p. 493) has been described by 
Simons (1960, p. 13) as Coryphodon proterus. Only the type 
can be assuredly ascribed to this species. Definite inclusion of 
this specimen in the Clark Fork fauna is not possible. It was 
found at the foot of the Beartooth Mountains where unconformities 
between Tertiary and Paleozoic rocks prevent accurate determi- 
nation of stratigraphic position. No other fossils were found asso- 
ciated with Coryphodon proterus, so that no faunal clues are 
available to help make stratigraphic inferences. In fact, the only 



22 BREVIORA No. 257 

reason for referring this specimen to a Paieocene horizon is that it 
was found in rocks mapped, presumably on non-paleontological 
grounds, as being of Paieocene age. Whether C. proterus is of 
Paieocene or Eocene age cannot be determined on the basis of 
available evidence. 

Simnarly, the one specimen of Esthonyx sp. known from 
the Princeton collection must be excluded from further con- 
sideration as an element of the Clark Fork fauna, since it is im- 
possible to determine whether the specimen came from Paieocene 
or Eocene beds (see Table III, text p. 21). However, since all 
the American Museum Clarkforkian representatives of this genus 
have been shown to be most probably of Lower Eocene derivation, 
it seems reasonable to assume, contrary evidence being lacking, 
that the Princeton Esthonyx is likely to be of the same age. 

A single specimen recovered from the sediments "east of the 
mouth of Litde Sand Coulee" was identified by Jepsen (1930b, 
p. 493) as Probathyopsis sp. Since its stratigraphic position re- 
mains in question (see above p. 21), it is unsuitable in helping to 
define any fauna. For the same reason, the two specimens of 
Dissacus praenuntius listed in Table III can not be used to dis- 
tinguish between different faunal horizons. 

The specimens of Carpolestes dubius attributed to the Clark 
Fork horizon have been recovered from two widely spaced locali- 
ties. Although the age of the rocks at one of these localities is 
problematical, the other site is undoubtedly of Paieocene age. 
However, Carpolestes dubius is known from horizons both above 
and below the Clark Fork level (Jepsen, 1930b, p. 491; Van 
Houten, 1945, p. 450), the type being from the Silver Coulee 
faunal horizon. Thus this species is so wide-ranging through time 
that its presence is of no use as an indication of Clarkforkian age. 

There is no reason to question that the type specimen of Dip- 
salodon matthewi was found in late Paieocene sediments. Unfor- 
tunately, the only other specimen was collected in the problematical 
region "east of the mouth of Little Sand Coulee." However, Pat- 
terson (personal communication) informs me that a specimen 
of ?Dipsalodon sp. is present in the Plateau Valley fauna of Tif- 
fanian age. Patterson's specimen (FMNH No. P26095) is a frag- 
ment of the right maxilla with P^, the broken stump of M\ and 
the root of M-. It can be compared with AMNH No. 16068, con- 
sisting of P^ and M\ which Denison (1938, p. 175) has doubt- 
fully referred to D. matthewi. (This specimen is listed as an 
oxyaenid in Table I.) But since Jepsen's two specimens of D. 



1967 CLARK FORK VERTEBRATE FAUNA 23 

matthewi are both represented by lower teeth only, no direct com- 
parison can be made between them and Patterson's specimen. 
(PU No. 13152 (type) has Pi 4, Mi 2; PU No. 13311 is an Mi.) 
Patterson's manuscript notes state that his "specimen is smaller 
than the type of D. matthewi and AMNH No. 16068. It probably 
represents a new species." In spite of not being able to make 
direct comparisons between Patterson's and Jepsen's specimens, 
however, it is possible to say that they are roughly of the same 
size and morphology, as might be expected in forms related at 
the generic level. Consequently, Dipsalodon can no longer be 
regarded as the only genus restricted to the Clark Fork fauna. 

Precise locality data are associated with only one of the sup- 
posedly Clarkforkian representatives of Plesiadapis cookei (Table 
III). But this specimen, the type, was found in the region east of 
the mouth of Little Sand Coulee where disagreements exist con- 
cerning the location of the contact between Paleocene and Eocene 
sediments. The specimen's uncertain stratigraphic derivation pre- 
cludes its being clearly assigned either a Paleocene or an Eocene 
age. The two other specimens of P. cookei were also found in 
the same general area, and the same uncertainty applies to them. 
Thus, on the basis of published evidence it is not possible to 
include P. cookei definitely in the Clark Fork fauna. 

As already stated, over half of the Princeton collection must be 
eliminated from consideration as elements of the Clark Fork fauna 
solely on the basis of insufficiently precise locality data. Because 
of wide-ranging stratigraphic distribution and a lack of reliable 
knowledge regarding phyletic relationships, other specimens are 
similarly not suitable for defining the fauna. Only two described 
specimens in the Princeton collection, the type of Dipsalodon 
matthewi and one of Carpolestes dubius, may still be considered 
as having certainly come from the latest Paleocene strata in this 
area. Together with one specimen of Phenacodus primaevus and 
one of Probathyopsis praecursor in the American Museum collec- 
tion, these fossils represent the only certain components of what 
has been described as the Clark Fork fauna. As previously men- 
tioned, however, Phenacodus is at present of no use for fine age 
discriminations between rocks in the mid-Paleocene to early 
Eocene time span. The single specimen of Probathyopsis prae- 
cursor can hardly be diagnostic of a latest Paleocene faunal assem- 
blage when all the other described specimens of this species are 
probably of early Eocene age. Carpolestes dubius is already 
known to range from Silver Coulee to Gray Bull. Since nothing 



24 BREVIORA No. 257 

is known about the rates of deposition of the Polecat Bench sedi- 
ments, it is possible that the "Clark Fork" specimens of Phena- 
codus primaevus, Dipsalodon matthewi, Carpolestes dubius and 
Probathyopsis praecursor may be either essentially contemporane- 
ous with or of a younger age than the fossils in the Silver Coulee 
fauna. But until a larger number of fossils from the latest Paleo- 
cene sediments are found, it is not possible in practice to differen- 
tiate a Clark Fork fauna from those of under- and overlying strata. 
At any rate, to base one of the provincial ages in the standard 
section of the North American Continental Tertiary on a "fauna" 
consisting of only four specimens representing four species seems 
unwarranted. As a direct consequence there also do not seem to 
be sufficient grounds for continuing to regard the Clark Fork as a 
discrete uppermost Paleocene faunal zone or as a member of the 
Polecat Bench Formation. ^ 

SOME RELATED PROBLEMS 

If it is accepted that a definitive Clark Fork fauna is not a 
reality, then a number of questions arise. For example, most of 
the American Museum material of Probathyopsis praecursor, in- 
cluding the type, clearly is from lower Gray Bull beds. In addi- 
tion, there is no certainty as to whether the one specimen of 
Probathyopsis sp. was found in Paleocene or Eocene deposits. 
These facts complicate the evolutionary sequence proposed by 
Jepsen (1930a, p. 129) of (1) P. sp., (2) P. praecursor, and (3) 
P. successor. The evidence strongly suggests that P. praecursor 
and P. successor actually existed contemporaneously rather than 
that the former gave rise to the latter. Furthermore, it is entirely 
possible that P. sp. might also have coexisted with the other two 
forms. Wheeler (1961, p. 21) states that P. newbilli from the 
Plateau Valley beds "is quite distinct from both P. praecursor and 
P. successor and is not closer to one than to the other." Such a 
statement suggests that P. newbilli could be ancestral to the two 
later, divergent forms. However, this scheme would not clarify 
the relationship of P. sp. to the other forms with which it may 
have been contemporaneous. Jepsen (1930a, p. 129) felt that P. 



^ The possibility of course exists that a post-Tiffanian Paleocene provin- 
cial age may be recognized in the future. In that event the age would 
require a new name. "Clarkforkian" could be resurrected only if a real 
fauna of such age were to be found in the Clark Fork area. In view of the 
intensive search for one that has been made there over the past thirty-five 
years I regard this possibility as most unlikely. 



1967 CLARK FORK VERTEBRATE FAUNA 25 

sp. is more similar to P. praecwsor than to P. successor, but until 
there is better stratigraphic control it will be difficult to resolve the 
problem of Probathyopsis taxonomy. 

The classification of Ectocion has received attention from 
Granger, Simpson, and McKenna. From the material available to 
him. Granger (1915, pp. 348-354) described four species; E. 
parvus, E. ralstonensis, E. osbornianus, and E. superstes. Simpson 
(1937, pp. 19-22; 1943, pp. 174-176), however, felt that Granger's 
four species should be regarded as successive subspecies grouped 
under the specific name E. osbornianus. More recently, McKenna 
(1960, pp. 102-103) discussed Ectocion classification in his paper 
on the Four Mile fauna. His remark (p. 103) that "the locality 
data of practically all existing collections of Willwood mammals 
are inadequate for detailed stratigraphic (and hence evolutionary) 
analysis" is equally applicable to specimens attributed to the Clark 
Fork fauna. But neither Simpson's nor McKenna's proposed clas- 
sification is clearly acceptable because of the probability that most 
of the American Museum's Clarkforkian specimens are of Eocene 
rather than Paleocene age. E. osbornianus ralstonensis, for exam- 
ple, presumably existed contemporaneously with E. o. complens, 
formerly thought to be the evolutionary successor of this species. 

Restudy of early representatives of the genus Phenacodus will 
also be required. Two species, P. primaevus and P. intermedius, 
whose ranges were listed as extending down into Clark Fork beds, 
were described by Granger (1915, pp. 337, 342). Subsequently 
Jepsen (1930b, p. 491) reported the presence of P. sp. in his 
Silver Coulee fauna. Phenacodus primaevus grangeri is also 
known from the Plateau Valley fauna (Patterson, personal com- 
munication). Simpson (1937, pp. 17-19) published a statistical 
analysis of the lower Gray Bull and Clark Fork phenacodonts in 
the American Museum collection. He thought that possibly the 
Clark Fork and some of the Gray Bull specimens represented a 
single species. Further, he felt that perhaps the two groups might 
be differentiated on a subspecific level, concluding (p. 19) "that 
the Clark Fork specimens are a nearly or quite homogeneous 
sample of one subspecies and that this subspecies may also occur 
in the Gray Bull ... or may be distinct. These alternatives cannot 
at present be adequately checked." In view of the probability that 
most of the supposed Clarkforkian phenacodonts were actually 
found in Gray Bull beds, it would not be surprising if the two 
samples represented only one species. 

Conceivably the type and only known specimen of Plesiadapis 
dubius (AMNH No. 16073) could have come from either late 
Paleocene or early Eocene strata, having been found at locality 



26 BREVIORA No. 257 

number 3. In appearance it most closely resembles P. fodinatus, 
originally described (Jepsen, 1930b, p. 515) from the late Paleo- 
cene Silver Coulee horizon. The character separating these two 
species was noted by Jepsen as follows: "The anterior crest of P4 
is developed into a minute cuspule, and a small ridge appears on 
the anterolingual part of the protoconid, quite in contrast to the 
distinct metaconid on P4 of P. diibius.'" Unfortunately, P4 of the 
type of P. fodinatus (PU No. 13278) has been broken off and is 
now missing, so that it is no longer possible to make the compari- 
son except from Jepsen's illustrations. The abundant quarry ma- 
terial of P. fodinatus, most of which has been obtained since 1930, 
reveals, however, that the presence or absence of a distinct meta- 
conid is a variable feature and not a specific character. That only 
one specimen of P. dubius has ever been found seems curious, in 
view of the rather extensive amount of field work in the upper 
Paleocene and lower Eocene beds of this region. Since the range 
of P. fodinatus has been, subsequent to 1930, extended up into 
the lower Gray Bull by continued collecting, it appears entirely 
possible that P. fodinatus is a synonym of P. dubius. 

The Sand Coulee beds were defined by Granger (1914, p. 205) 
as follows: "This horizon does not contain Systemodon but it does 
contain the Perissodactyl genus Eohippus in abundance as well 
as Artiodactyls, Rodents, and Primates, and marks the first ap- 
pearance of these four orders. It also marks the last appearance 
of the primitive order Multituberculata which is represented in 
this horizon by a genus of Plagiaulacids." Many of the criteria 
used to characterize this faunal zone have in the past fifty years 
been shown not to be definitive. Rodents and primates were later 
found in the Paleocene. Granger's "plagiaulacids," now classified 
as ptilodonts, are known from levels higher than the Sand Coulee. ^ 
When Homogalax (Systemodon) was found in the Sand Coulee 
beds subsequent to Granger's definition of them, Jepsen (1930a, 
p. 119) suggested suppression of this term and included the beds 



^ However, multituberculates are very rare in Gray Bull and later deposits. 
Van Houten (1945, p. 448) reports one species, Ectypodus simpsoni, extend- 
ing into middle Gray Bull beds. Robinson, Black, and Dawson (1964, 
p. 810) have described a multituberculate from upper Eocene deposits 
near Badwater, Wyoming. However, their specimens, representing "an 
undescribed species related to Ectypodus hazeni, . . . comprise a small per- 
centage of the fauna, certainly less than 1 percent . . ." These two cases 
are the only known occurrences of multituberculates above the Sand Coulee 
strata. 



1967 CLARK FORK VERTEBRATE FAUNA 27 

in the Gray Bull. Simpson (1937, p. 1), however, noted that "the 
Sand Coulee fauna was not wholly defined ... on the absence 
of Homogalax but also by the generally slightly less advanced 
character of its mammals." Now that most of the American Muse- 
um's Clarkforkian material can be probably relegated to lower Eo- 
cene beds, it may be possible to redefine a Sand Coulee fauna. For 
example, this might be done on the basis of the presence of 
Plesiadapis cookei and a species of Probathyopsis, the last abun- 
dant appearance of the multituberculates, and a more primitive 
aspect in those forms common to both the Sand Coulee and Gray 
Bull. Reappraisal of the Sand Coulee fauna is desirable in order 
to determine conclusively whether or not such a horizon can be 
distinguished. 

SUMMARY 

Some of the published data concerning the American Museum's 
Clark Fork collection do not agree with the locality information 
on specimen labels. For example, there are no fossils from either 
side of the road descending into the Sand Coulee Basin from the 
Bighorn Basin divide (area D), although it was described as one 
of the Clarkforkian type localities. The majority of the American 
Museum collection comes from three localities (1, 2, and 3, Fig. 
3 ) . Review shows that it is doubtful that most of these specimens 
from localities 1 and 2 should be referred to a late Paleocene Clark 
Fork horizon, but that they are rather of early Eocene age. Other 
specimens (those from locality 3 or labelled as being from Clark 
Fork "beds," "period," or "formation") are associated with such 
vague stratigraphic information that it is not possible to determine 
whether they are from upper Paleocene or lower Eocene beds. 
Because of insufficient locality data it is also impossible to say 
whether or not approximately one-half of the described Princeton 
cohection belongs to a late Paleocene or an early Eocene fauna. 
A wide ranging stratigraphic distribution or a lack of adequate 
knowledge regarding phyiogenetic relationships precludes using 
most of the remaining fossils to characterize the fauna. 

Thus, of 107 specimens (representing at least 23 and perhaps 
26 species) in both the American Museum and Princeton collec- 
tions that have been described as elements of the Clark Fork 
fauna, a total of only four specimens representing four species 
{Carpolestes dubius, Phenacodus primaevus, Probathyopsis prae- 
cursor, and Dipsalodon matthewi) can be surely ascribed to a 
possible latest Paleocene faunal horizon in the Polecat Bench 
Formation. However, only one of these species {Dipsalodon 



28 BREViORA No. 257 

matt hew i) may be unique to what has been described as the Clark 
Fork fauna. Carpolestes dubhis is known from both the Silver 
Coulee and the Gray Bull. Specimens of Phenacodus primaevus 
are known from the Gray Buil, and Phenacodus sp. is present in 
the Silver Coulee. Until the phylogeny of this genus is reviewed, 
it is not possible to use species of Phenacodus for horizon markers. 
Probathyopsis praeciirsor is much more commonly found in early 
Eocene than late Paleocene beds. Thus this species can hardly 
be considered as diagnostic of the Clark Fork fauna. If ?Dipsal- 
odon sp. from the Tiffanian Plateau Valley deposits of Colorado 
is correctly referable to this genus, then it invalidates the supposed 
restriction of this genus to the Clark Fork. 

Therefore, the Clark Fork "fauna" consists of four specimens 
referable to four mammalian genera, each from a different locality. 
None of the genera (and not more than one of the species) are 
restricted to a lithically distinguishable Clark Fork horizon. Such 
evidence scarcely warrants recognition of the Clark Fork as a pro- 
vincial age, faunal zone, or member of the Polecat Bench Formation. 

LITERATURE CITED 

Andrews, D. A., W. G. Pierce, and D. H. Eargle. 

1947. Geologic map of the Bighorn Basin, Wyoming and Montana, 
showing terrace deposits and physiographic features. Oil and 
Gas Investigations Preliminary Map 71. 
Cope, E. D. 

1874. Report on the vertebrate fossils discovered in New Mexico, 
with descriptions of new species. Extract from appendix FF 
of the Annual Report of the Chief of Engineers, 1874, Wash- 
ington, Gov. Print. Office, pp. 4-18. 
1880. The northern Wasatch fauna. Amer. Natur., vol. 14, pp. 
908-909. 
Denison, R. H. 

1938. The broad-skulled Pseudocreodi. Ann. N. Y. Acad. Sci., 
vol. 37, pp. 163-256. 
Fisher, C. A. 

1906. Geology and water resources of the Bighorn Basin, Wyoming. 
U. S. Geol. Surv. Prof. Pap. No. 53, III-VI. pp. 1-72, 1 map. 
Gazin, C. L. 

1953. The Tillodontia: an early Tertiary order of mammals. Smith- 
son. Misc. Coll., vol. 121, no. 10, pp. I-llO. 
1962. A further study of the lower Eocene mammalian faunas of 
southwestern Wyoming. Smithson. Misc. Coll., vol. 144, no. 1, 
pp. 1-98. 



1967 CLARK FORK VERTEBRATE FAUNA 29 

Granger. W. 

1914. On the names of lower Eocene faunal horizons of Wyoming 
and New Mexico. Bull. Amer. Mus. Nat. Hist., vol. 33, 
pp. 201-207. 

71914. Annual Report of the Department of Vertebrate Paleontology, 
American Museum of Natural History, for the years 1911. 
1912, and 1913. MS on file in the department. 

1915. A revision of the lower Eocene Wasatch and Wind River 
faunas. Part III. Order Condylarthra, families Phenacodonti- 
dae and Meniscotheriidae. Bull. Amer. Mus. Nat. Hist., vol. 
34, pp. 329-361. 

Jepsen, G. L. 

1930a. New vertebrate fossils from the lower Eocene of the Big- 
horn Basin, Wyoming. Proc. Amer. Phil. Soc. vol. 69, pp. 
117-131. 

1930b. Stratigraphy and paleontology of the Paleocene of northeastern 
Park County, Wyoming. Proc. Amer. Phil. Soc. vol. 69, pp. 
463-528. 

1 940. Paleocene faunas of the Polecat Bench formation. Park 
County, Wyoming, Part I. Proc. Amer. Phil. Soc, vol. 83, 
pp. 217-341'. 
Matthew. W. D. 

1915a. A revision of the lower Eocene Wasatch and Wind River 
faunas. Part I. Order Ferae (Carnivora), suborder Creo- 
donta. Bull. Amer. Mus. Nat. Hist., vol. 34, pp. 4-103. 

1915b. A revision of the lower Eocene Wasatch and Wind River 
faunas. Part II. Order Condylarthra, family Hyopsodontidae. 
Bull. Amer. Mus. Nat. Hist., vol. 34, pp. 311-328. 

1915c. A revision of the lower Eocene Wasatch and Wind River 
faunas. Part IV. Entelonychia, Primates, Insectivora (part). 
Bull. Amer. Mus. Nat. Hist., vol. 34, pp. 429-483. 

1918. A revision of the lower Eocene Wasatch and Wind River 
faunas. Part V. Insectivora (continued), Glires, Edentata. 
Bull. Amer. Mus. Nat. Hist., vol. 38, pp. 565-657. 
Matthew, W. D. and W. Granger. 

1915. A revision of the lower Eocene Wasatch and Wind River 
faunas. Introduction. Bull. Amer. Mus. Nat. Hist., vol. 34, 
pp. 1-4. 
McKenna, M. C. 

1960. Fossil Mammalia from the early Wasatchian Four Mile 
fauna, Eocene of northwest Colorado. Univ. Calif. Publ. 
Geol. Sci., vol. 37, pp. 1-130. 
Pierce, W. G. 

1965. Geologic Map of the Clark Quadrangle, Park County, Wy- 
oming. U. S. Geol. Surv. Quadrangle Map GQ-477. 
Robinson, P., C. C. Black, and M. R. Dawson. 

1964. Late Eocene multituberculates and other mammals from Wy- 
oming. Science, vol. 145, pp. 809-811. 



30 BREVIORA No. 257 

Simons, E. L. 

1960. The Paleocene Pantodonta. Trans. Amer. Phil. Soc, vol. 50, 
pp. 1-98. 

Sfmpson, G. G. 

1929. A new Paleocene uintathere and molar evolution in the 
Amblypoda. Amer. Mus. Novit., No. 387, pp. 1-9. 

1937. Notes on the Clark Fork, Upper Paleocene fauna. Amer. 
Mus. Novit., No. 954, pp. 1-24. 

1943. In Bogert, et al.. Criteria for vertebrate subspecies, species, 
and genera. Criteria for genera, species, and subspecies in 
zoology and paleontology. New York Acad. Sci., vol. 44, 
pp. 145-178. 
1945. The principles of classification and a classification of mammals. 
Bull. Amer. Mus. Nat. Hist., vol. 85, pp. 1-350. 
Sinclair, W. J., and W. Granger. 

1912. Notes on the Tertiary deposits of the Bighorn Basin. Bull. 
Amer. Mus. Nat. Hist., vol. 31, pp. 57-67. 
Stow, M. H. 

1938. Dating Cretaceous-Eocene tectonic movements in Bighorn 
Basin by heavy minerals. Bull. Geol. Soc. Amer.. vol. 49, 
pp. 731-761. 

Van Houten, F. B. 

1945. Review of latest Paleocene and early Eocene mammaUan 
faunas. Jour. Paleont., vol. 19, pp. 421-461. 
Van Valen, L. 

1966. Deltatheridia, a new order of mammals. Bull. Amer. Mus. 
Nat. Hist., vol. 132, pp. 1-126, 8 plates. 
Wheeler, W. H. 

1961. Revision of the uintatheres. Bull. Peabody Mus. Nat. Hist. 
Yale Univ., No. 14, iii-vi. pp. 1-93, 14 plates. 

Wood, H. E., 2nd et al. 

1941. Nomenclature and correlation of the North American con- 
tinental Tertiary. Bull. Geol. Soc. Amer., vol. 52, pp. 1-48, 
1 plate. 



BREVIORA 

Mmsenamii of Coioipsirsitive Zoology 

Cambridge, Mass. 3 February, 1967 Number 258 



BIOLOGY OF THE PARTHENOGENETiC FUNGUS BEETLE 
CIS FUSaPES MELLIE (COLEOPTERA: CIIDAE) 

By Jolin F. Lawrence 

Thelytoky, or female-producing parthenogenesis, has been re- 
ported for a large number of animal groups, and a considerable 
amount has been written on its occurrence in various groups of 
insects, such as the Psychidae. Aphididae, Tettigoniidae, Phasmi- 
dae, and Curculionidae (White. 1954; Suomalainen, 1962). Aside 
from the well known cases in Ptinus clavipes and several European 
weevils, there have been few references to thelytoky in the Coleop- 
tera; yet this type of reproduction is probably fairly common 
throughout the order, judging from the number of groups in 
which unusual sex ratios have been noted. The step from the pre- 
liminary observation to experimental evidence is often a big one. 
however, because of the difficulties encountered in rearing many 
insects. While endeavoring to straighten out a taxonomic problem 
concerning some small fungus-feeding beetles in the family Ciidae, 
I discovered an apparent case of thelytoky in the species Cis 
juscipes Mellie, and was able to confirm this by a series of rear- 
ing experiments discussed below. To my knowledge, this is the 
first record of its occurrence within the large superfamily Cucu- 
joidea. 

In connection with a study on the systematics and biology of 
western North American Ciidae. an attempt was made to distin- 
guish between two apparent sibling species, Cis juscipes Mellie 
and Cis impressus Casey. The two species are widespread in North 
America, have broadly overlapping distributions, and feed on the 
same species of fungi. The only characters given in the literature 
to separate the two are the form of the male pronotum and ab- 
domen; in C. impressus the pronotum of the male is impressed 
anteriorly and the abdomen bears a setigerous pore on the first 
visible sternite, while both characters are apparently absent in the 
"male" of C. juscipes. Females of the two species are apparently 
indistinguishable (Casey, 1898; Dury, 1917). 



2 BREVIORA No. 258 

In an attempt to solve this problem, I examined about 1500 
museum specimens from various parts of North America, collected 
large series of adults and larvae from both dimorphic and mono- 
morphic populations in California, and dissected series from Cali- 
fornia and Minnesota, in order to examine genital characters. No 
characters could be found to separate females or larvae of the two 
species, nor could any external sexual characters be found in C. 
fuscipes. When dissections were made of relatively large samples, 
it was found that the monomorphic series consisted entirely of 
females. This led to the hypothesis that thelytoky was occurring in 
C. fuscipes and that the name impressus had been applied to male 
specimens of the same species. The following rearing experiments 
were then conducted. 

REARING EXPERIMENTS 

Larvae of Cis fuscipes and fresh pieces of Polyporus versicolor 
were collected in Marin Co. and Alameda Co., California. Each 
larva was isolated in a petri dish with a piece of fungus, and water 
was added from time to time. Each resulting female was kept 
in isolation and observed periodically for the presence of eggs and 
larvae. Several rearings were attempted, but only one will be dis- 
cussed here. One larva was isolated on December 9, 1962, and by 
January 26, 1963, it had pupated and eclosed, producing an adult 
female. By February 7, eggs had been laid and early instar larvae 
were seen boring in the fungus. By April 1,15 females were seen 
in the medium; 7 of these were preserved, while the other 8 were 
isolated in separate dishes. Eggs were seen in most of these dishes 
by May 4, and on May 31, all the adult females were removed and 
preserved, leaving only the ¥2 larvae in the dishes. By July 19, 
adult Fo females were present in all 8 dishes. Five of the clones 
were preserved, and the numbers of individuals contained in each 
were as follows: 51, 50, 43, 42, and 47. The remaining clones 
produced an F3 generation by October 24, and one of these was 
retained to produce F4's by the end of the year. 

Because of intermittent field work, the exact generation time 
could not be recorded, but the Fi generation was produced in 
about 60 days, which appears to be a reasonable figure when 
compared with observations made on several other ciid species. 

Two attempts were made to cross females from these partheno- 
genetic clones with males collected in the same area; all of these 
were unsuccessful in that only female progeny resulted. Mating 
probably did not take place, but many more trials will be necessary 



1967 PARTHENOGENETIC FUNGUS BEETLE 3 

before one can conclude that parthenogenetically-produced females 
will not cross with males from bisexual populations. 

It is obvious that some form of thelytoky is occurring in this 
species, but the actual cytological mechanisms involved have not 
been studied. Chromosome counts made by Mr. Norihiro Ueshima 
showed that the diploid number is 14 both in the male and in the 
parthenogenetic female. Thus polyploidy does not occur in the 
California populations examined. 

GEOGRAPHICAL AND ECOLOGICAL RANGE 

Cis fuscipes is the most widespread and common species of 
Ciidae in North America, ranging from northern British Columbia 
to southern California on the Pacific Coast, east across southern 
Canada to Nova Scotia, and south throughout the eastern and mid- 
western United States (east of the lOOth meridian) to Browns- 
ville, Texas, and Dunedin. Florida (Fig. 1). It is apparently 
absent from the more arid parts of the continent, such as the Great 
Plains, the Great Basin, and the southwestern deserts. A single 
record from Provo, Utah, indicates its possible occurrence in the 
poorly collected Rocky Mountain region. Specimens have also 
been seen from Mexico (no specific locality), Cuba, Hawaii, and 
from the island of Madeira in the eastern Atlantic. These isolated 
records will be discussed below. 

Throughout its range, the species occurs in association with sev- 
eral species of bracket fungi (Basidiomycetes: Polyporaceae), 
where both adults and larvae feed within the fruiting bodies. It 
appears to be restricted to members of the Polyporus versicolor 
group (Paviour-Smith, 1960), which have small, relatively thin 
sporophores with whitish, punky context and a trimitic hyphal sys- 
tem and which usually cause white rot in dead or dying angio- 
sperms. It is especially common in Polyporus versicolor, which 
maybe considered its "headquarters" (Elton, 1949; Paviour-Smith, 
1960), but also feeds on related fungi. Out of the 92 host records 
which I have accumulated for this species, 72 are from P. versi- 
color, 1 from P. hirsLitus, 6 from Lenzites betulina, 2 from P. con- 
chifer, and a single record each from P. squamosus, Trametes 
suaveolens, Fomes jraxineus, Ganoderma brownii, and P. gilvus. 

In the western part of its range, Cis fuscipes is usually found 
associated with Cis versicolor Casey, Cis vitulus Mannerheim. 
Sulcacis curtulus (Casey), Ennearthron calijornicum Casey, and 
Octotemnus laevis Casey. In California, extensive collecting has 
revealed that C. fuscipes (as well as C. vitulus and O. laevis) 



4 BREVIORA No. 258 

requires a somewhat more humid environment than C. versicolor, 
S. curtulus, or E. californicum, and it may be replaced by these 
species in drier environments. In the eastern part of its range, 
the species occurs with a number of other ciids, including Cis pis- 
toria Casey, Cis falli Blatchley, Sulcacis lengi Dury, Strigocis opaci- 
collis Dury, and Octotemnus laevis Casey, and other fungivorous 
beetles, such as Neomida bicornis (Fabricius) and Euparius mar- 
moreus (Olivier). It is relatively common in most areas and 
appears to be successfully competing with other species inhabiting 
the same fungi. 

Like many species of polypores, P. versicolor and its relatives 
are widely distributed and are found on many different tree species, 
although they are normally restricted to angiosperms. One might 
expect a fairly continuous distribution of these fungus species 
within any deciduous forest, but the actual occurrence of fruiting 
bodies will depend upon the forest composition, number of dead 
trees, and physical factors affecting the establishment of the fungus 
and the production of sporophores. Even in a continuous forest, 
one may find P. versicolor replaced in certain areas by P. gilvus 
or P. porgamenus, each of which support different ciid species. 
This breaking up of the habitat will be more evident, of course, 
where the hardwoods themselves are scattered or rare. The 
absence of Cis juscipes from the more arid part of western North 
America may be due to the inability of the species to tolerate drier 
environments, as was suggested by observations on California pop- 
ulations, rather than to the lack of hardwood forests, or to physical 
factors affecting the host fungi. The relative rarity of the beetle in 
the more humid parts of the Sierran and Cordilleran ranges, how- 
ever, may be related to the scattering of the habitat, since large 
stands of conifers separate riparion situations in which alders and 
other angiosperms provide a suitable medium for the host fungi. 

DISTRIBUTION OF BISEXUAL POPULATIONS 

The abundance of male specimens varies considerably through- 
out the range of this species. Since intensive population sampling 
was not attempted, it is not possible to present a detailed analysis 
of the sex ratios, but a general idea of the distribution of bisexual 
populations may be derived from the data recorded for over 2500 
specimens (from 405 localities) collected by myself or bor- 
rowed from various museums. For the entire range, 12 per cent 
of the specimens examined were males, and these represented 21.7 
per cent of the localities. In Figure 1, the localities from which 



1967 



PARTHENOGENETIC FUNGUS BEETLE 




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6 BREVIORA No. 258 

males were taken are represented by stars, while those from which 
only females were recorded are represented by black dots. Two 
things that are immediately apparent are: 1) the preponderance 
of male records in the northwestern part of the continent, as com- 
pared to their rarity on the east coast and complete absence in 
the Midwest, and 2) the scattered occurrence of males through- 
out the range as far south as Florida and Texas. From California, 
Oregon, Washington, British Columbia, and Alberta, 918 speci- 
mens (36.2%) from 77 localities (19.0%) were examined, and 
of these 221 specimens (24.1%) from 51 localities (66.2% ) were 
males. From the remainder of Canada and the United States, 1618 
specimens (63.8%) from 328 localities (81.07c) were exam- 
ined, and of these 84 specimens (5.2%) from 37 localities 
(12.7%) were males. Of the total male specimens examined, 
72.4 per cent were collected in the Pacific states and western 
provinces. 

Although the sample sizes vary greatly and were often small, 
(decreasing the reliability of negative evidence), I think there is 
little doubt that bisexual populations are more abundant in the 
Northwest and become increasingly rare to the east and south. 
Parthenogenetic populations occur throughout the range, and in 
northern coastal California both types appear to be common in 
any one area. Males have been taken at several localities along 
the east coast and in the Gulf Coast states, but I have seen no 
records of their occurrence in the Midwest. 

SYSTEMATIC RELATIONSHIPS AND PROBABLE ORIGIN 

Cis fuscipes is closely related to a group of Old World species 
which were placed in the subgenus Hadraule Thomson by Reitter 
(1902), Chujo (1939), Miyatake (1954), and others. The name 
Hadraule, however, does not apply to the group, since its type 
species, Cis elongatulus Gyllenhal, has been excluded (Lohse, 
1964; Lawrence, 1965). The species seem to form a natural unit 
and are characterized as follows : stout, oblong to subparallel body 
form; pronotum with fairly broad lateral margins and produced 
and rounded apical angles, which are preceded by a lateral in- 
flexion of the disc; dual and subseriate eiytral punctation; vesti- 
ture of stout, erect bristles; prosternum slightly tumid but not cari- 
nate; apex of the protibia strongly dentate; and the male with 
two weakly developed clypeal piates, anteriorly impressed pro- 
notum, setigerous pore on the third abdominal sternite, and rela- 
tively simple aedeagus. The species group extends from eastern 



1967 PARTHENOGENETIC FUNGUS BEETLE 7 

Siberia through southeastern Asia and India to Africa and Mada- 
gascar, being absent in northern Africa and Europe, but the species 
which most closely resemble C. jiiscipes occur mainly in Siberia 
and Japan. These include Cis seriatopilosus Motschulsky (Siberia: 
Amur, Sakhalin; Japan); Cis seriatiilus Kiesenwetter (Japan); Cis 
heiroglyphicus Reitter (Siberia: Khabarovsk; Japan); Cis tai- 
wanus Chujo (Taiwan, Loo Choo Islands, Japan?); Cis subrobus- 
tus Miyatake (Japan); and Cis japonicus Nobuchi (Japan). 

Those species which have been studied biologically occur on 
the same fungi as C. fuscipes. Cis seriatopilosus and C. taiwanus 
have both been recorded from Polyporus versicolor, while C. sub- 
robustus has been collected on P. hirsutus (Chujo, 1939; Fukuda, 
1940; Miyatake, 1954). Although most of these beetles appear to 
be specifically distinct from the North American species, a single 
Siberian specimen identified by Reitter as C. seriatopilosus is very 
similar to individuals of C fuscipes. Perhaps further collecting in 
eastern Siberia will reveal the presence of fuscipes there. 

The presence of several close relatives in Asia and the abun- 
dance of males in the northwestern part of the range suggest that 
Cis fuscipes or its ancestor dispersed into North America across 
the Bering Strait from a point of origin in the eastern Palaearctic. 
Since there is no evidence of unusual sex ratios among the Japanese 
and Siberian species, it is probable that the parthenogenetic mode 
of reproduction originated in northwestern North America and 
the species subsequently spread east and south across most of the 
continent. 

In the absence of a fossil record, there is no direct evidence 
relating to the time of dispersal of C. fuscipes into North America. 
Some indirect support may be derived from Linsley's analysis of 
the cerambycid beetle fauna (1958), in which he considers all 
of the northern or Holarctic elements of the fauna to be post- 
Pleistocene in origin. Several of the examples of theiytoky in 
animals have been demonstrated to be glacial phenomena; this is 
discussed by Suomaiainen (1962) for insects, and by Darewski 
and Kulikova (1961) for lacertid lizards. The recent origin of 
parthenogenesis has been argued by some authors on theoretical 
grounds, because of the inherent long-term disadvantages of this 
type of genetic system. 

SPREAD AND OVERSEAS DISPERSAL 

The ability of a parthenogenetic species to colonize and spread 
rapidly throughout a new area is well known and has been dis- 
cussed in various texts, such as White (1954), and Mayr (1963). 



8 BREVIORA No. 258 

A system which suspends sexual reproduction not only makes pos- 
sible the perpetuation of a successful genotype as soon as it is 
formed, but doubles the fecundity by eliminating the "reproductive 
wastage" characteristic of a population in which roughly half of 
the individuals are males. Where the suitable habitat for a species 
is broken up, a parthenogenetic form will be able to spread more 
rapidly, since only a single female need reach a favorable micro- 
environment. In an area which is initially unsuitable for a bisexual 
species, the development of a parthenogenetic system may speed 
up the process of adaptation to the new environment by imme- 
diately fixing and reproducing a favorable genotype. Stebbins 
(1950) and others have stressed the importance of apomixis in 
the rapid colonization of new habitats by plant species. 

A comparison of the ranges of North American Ciidae shows 
that only Cis juscipes occurs throughout the northern part of the 
continent and yet is also an important element in the fauna of the 
southeastern and midwestern United States. Several other species 
have northern distributions and similar affinities to Palaearctic 
species. Some of these, such as Cis horridulus Casey and DoU- 
chocis indistinctus Hatch extend southward only in montane 
regions and may occur in the southern Appalachians or in the 
mountains of Arizona. Sulcacis ciirtulus (Casey) extends into 
southern California but is rare in the northeastern and northern 
midwestern states. Eridaulus levettei (Casey) is fairly wide- 
spread in the eastern part of the continent, but does not occur 
in the West, while Octotemnus laevis Casey is common along both 
coasts and in parts of the Midwest. Neither of these species is as 
common or widespread as C. juscipes, and both are absent from 
the Southern Coastal Plain and Gulf Coast. Two other wide- 
ranging species, Ennearthron calijornicum Casey and Enneorthron 
thoracicome (Ziegler), occur in the western and eastern states 
respectively. Both of these, however, are of southern origin, hav- 
ing their closest relatives in the Neotropical Region, and neither 
extends far into Canada. The comparison of C. juscipes with 
other northern or Holarctic members of the North American fauna 
may seem to imply that all of these have similar relationships to 
Palaearctic species and thus represent Old World invasions of 
roughly the same age. This does not appear to be the case. All 
of the other northern species have Old World counterparts which 
extend throughout the Palaearctic from Europe to Siberia and 
Japan, whereas only C. juscipes belongs to a group restricted to 
the eastern Palaearctic and Oriental regions and having some 
relationships with palaeotropical forms. It may be that C. juscipes 



1967 PARTHENOGENETIC FUNGUS BEETLE 9 

represents the most recent element in our fauna, but there is no 
direct evidence for this. 

I think there is little doubt that the evolution of thelytoky in 
C. jiiscipes is responsible for the apparently rapid spread of the 
species and for its present wide distribution in North America. 
In addition to the obvious advantages of the parthenogenetic sys- 
tem to dispersal and colonization, the increased fecundity has 
probably contributed to its success in competing with the large 
and diverse fauna of southern origin occupying the southern peri- 
phery of the range. The only large area in North America which 
is suitable for C. fuscipes, but in which the forest cover is con- 
siderably broken up, due partly to agriculture, is the Midwest. It 
is interesting that this is the only region in which males are 
totally absent. 

The presence of males in scattered localities along the east 
coast and at the southern extremities of the range raises several 
questions which can be answered only by more detailed popula- 
tion studies. Is the parthenogenetic form the result of a single 
evolutionary event, an obligate thelytokous form being completely 
independent of the bisexual form? Is a residual bisexual popula- 
tion continually giving rise to parthenogenetic clones? Is partheno- 
genesis facultative in this species, so that occasional females can 
give rise to normal males? It is hoped that future studies will 
provide the answers. 

Mellie's original series of Cis fuscipes included several speci- 
mens from the island of Madeira. It would seem improbable that 
the Madeiran series represents the same species, but an examina- 
tion of Mellie's and Wollaston's specimens revealed that they are 
conspecific with the North American fuscipes, and that they are 
all females. According to Wollaston (1854, 1865), the species 
has become well established in cultivated areas at low elevations 
around Funchal and is likely to be an accidental introduction. 
Although it is possible that a specimen of fuscipes could have 
rafted from the Caribbean to Madeira, it is more reasonable to 
assume that the species was introduced by man. 

Recently, a series of female specimens of fuscipes were collected 
at Olinda on the island of Maui, Hawaii, and presented to the 
Commonwealth Institute of Entomology. Although Perkins, 
Swezey, and others have collected extensively in Hawaii, this is 
the first record of this species there, and probably represents a 
recent human introduction. In addition to this, two female speci- 
mens in the Reitter collection were apparently collected in Cuba. 
Whether by natural means or by human transport, Cis fuscipes 



10 BREVIORA No. 258 

has dispersed from North America to these various islands, and 
has become estabhshed on at least one of them. The advantages 
of parthenogenesis in the establishment of a species having been 
introduced into a new area by long-distance dispersal have been 
discussed by several authors, including Longhurst (1955) for 
Crustacea, and Baker (1955) for plants. 

TAXONOMIC STATUS 

Because of its variability and wide distribution, Cis fuscipes 
has been given a number of names by different authors. Mellie 
(1848) described the species from a series of 5 specimens from 
Boston in the Chevrolat collection and 4 more collected by Wollas- 
ton on Madeira. In the same paper, three other names were 
applied to the species: Cis atripennis, also from Boston, Cis 
chevrolatii from "Nouvelle-Orleans," and Cis dubius from the 
latter locality. Leconte, Horn, and others applied the name 
fuscipes to the common North American species, and it has been 
used consistently by later authors. Casey (1898) described two 
more species, Cis carolinae (North Carolina) and Cis pallens 
(Montana), which were distinguished from fuscipes on the basis 
of color and relative lengths of antennal segments. He also 
described a third species, Cis impressus (Pacific Coast), differing 
from fuscipes only in the form of the male pronotum, which was 
said to be "broadly impressed at apex." Dury (1917) noted that 
Cis fuscipes occurred throughout North America, that C. impres- 
sus occurred on both coasts, and that the females of the two 
species could not be distinguished from one another. He also 
considered chevrolatii and carolinae to be synonyms of fuscipes. 

During the summer of 1963, I examined the Casey types at 
the United States National Museum, Washington, D. C, and in 
the spring of 1966 I had the opportunity to study the types of 
Mellie in the Salle and Pic collections at the Museum National 
d'Histoire Naturelle, Paris, and in the Wollaston collection at the 
British Museum (Natural History), London. All of the above 
names, with the exception of Cis impressus Casey, are based on 
female specimens of the variable Cis fuscipes. The name fuscipes 
is here selected as the senior synonym because of its continual 
usage in the North American literature. Being based on a bisexual 
population, Casey's name impressus presents certain difficulties. 
Since it has not yet been established whether this species is faculta- 
tively parthenogenetic or rather composed of a bisexual species 
and one or several obligate parthenogenetic clones, it could be 



1967 PARTHENOGENETIC FUNGUS BEETLE 11 

argued that the name impressus should be applied to populations 
in which males occur. The true nature of the biological situation 
will be made clear only after intensive population analyses, rear- 
ing experiments, and cytological studies. In any case, I prefer 
to consider impressus a synonym of fuscipes on purely practical 
grounds, since no characters have been found to distinguish the 
larvae or females of the two forms. I therefore propose the fol- 
lowing synonymy: 

Cis FUSCIPES Mellie 

C is fuscipes Mellie, 1848: 271. pi. 2, fig. 23; Wollaston, 1854: 281; Woliastor, 
1865: 234;Cas2y, 1898: 78; Blatchley, 1910: 898; Dalla Torre, 1911: 11: 
Dury, 1917: ll;'Leng. 1920: 246; Weiss, 1920: 138 (host); Weiss and 
West, 1920: 8 (host); Weiss and West, 1921b: 169 (host); Boving and 
Craighead. 1931: 270-71, pi. 92. figs. K-R (larva); ArneU, 1962: 827, 
fig. r.98; Hatch, 1962: 231; Lawrence, 1965: 279. Type locality: "Bos- 
ton." Lectotype, female, Pic collection, Mus. Natl. Hist. Nat., Paris. 

Cis atripennis Mellie. 1848: 258. pi. 2, fig. 15; Casey. 1898: 77: Dalla Torre, 
1911: 6; Leng, 1920: 246. Type locality: "Boston." Holotype. female. 
Pic collection, Mus. Natl. Hist. Nat., Paris. NEW SYNONYMY. 

Cis chcvrohitii Mellie. 1848: 249; Blatchley, 1910: 898; Dalla Torre, 1911: 
8; Dury, 1917: 11; Leng, 1920: 246. Type locality: "Nouvelle-Orleans." 
Lectotype, female, Pic collection, Mus. Natl. Hist. Nat., Paris. 

Cis diibiiis MtWiQ. 1848: 273; Dalla Torre, 1911: 10; Leng, 1920: 247. Type 
locality: "Nouvelle-Orleans." Lectotype, female, Salle collection, Mus. 
Natl. Hist. Nat., Paris. NEW SYNONYMY. 

Cis carolinae Casey, 1898: 78; Dalla Torre, 1911: 8; Dury, 1917: 11; Leng, 
1920: 246. Type locality: "North Carolina (Asheville)." Holotype, 
female, Casey collection, U.S. Nat. Mus., Washington, D.C. 

Cis impressa Casey, 1898: 79; Dalla Torre, 1911: 12; Dury, 1917: 11; Blatch- 
ley, 1918: 54; Leng, 1920: 246; Weiss and West, 1921a: 61 (host); 
Blatchley, 1923: 19; Hatch, 1962: 231. Type locality: "Vancouver 
Island." Holotype, male. Casey collection, U. S. Nat. Mus., Washington, 
D. C. NEW SYNONYMY. 

Cis pollens Casey, 1898: 78; Dalla Tone. 1911: 15: Leng, 1920: 246. Type 
locality: "Montana (Missoula)." Holotype, female, Casey collection, 
U. S. Nat. Mus.. Washington. D. C. NEW SYNONYMY. 

ACKNOWLEDGMENTS 

This investigation was supported in part by a grant from the 
National Science Foundation (GB-4743), grants-in-aid from the 
Woodrow Wilson Foundation, Society of the Sigma Xi, and Uni- 
versity of California Committee on Research, and fellowships 
from the National Science Foundation and Woodrow Wilson 



12 BREVIORA No. 258 

Foundation. Sincere thanks are extended to E. G. Linsley, R. 
L. Usinger, C. D. MacNeill, and H. V. Daly for their advice dur- 
ing the early part of the study, to I. Taveres, R. L. Gilbertson. 
and J. A. Stevenson for the identification of host fungi, to N. 
Ueshima for the preparation and interpretation of cytological 
material, and to E. Mayr and P. J. Darlington, Jr. for their criti- 
cal review of the manuscript. I am indebted to the curators and 
staff of the following institutions for making their collections 
available to me: Academy of Natural Sciences, Philadelphia, Pa.; 
American Museum of Natural History, New York, N. Y.; Brigham 
Young University, Provo, Utah; British Museum (Natural His- 
tory), London, England; California Academy of Sciences, San 
Francisco, Calif.; California Insect Survey, University of Cali- 
fornia, Berkeley, Calif.; Canadian National Collection, Entomology 
Research Institute, Ottawa, Canada; Carnegie Museum, Pitts- 
burgh, Pa.; Chicago Natural History Museum, Chicago, 111.; Cin- 
cinnati Museum of Natural History, Cincinnati, Ohio; Cornell 
University, Ithaca, N. Y.; Illinois Natural History Survey, Urbana, 
111.; Museum of Comparative Zoology, Cambridge, Mass.; Museum 
National d'Histoire Naturelle, Paris, France; Oregon State Uni- 
versity, Corvallis, Oregon; United States National Museum, Wash- 
ington, D. C; University of Alberta, Edmonton, Alta.; University 
of California, Davis, Calif.; University of Washington, Seattle, 
Wash.; University of Kansas, Lawrence, Kansas; Utah State 
University, Logan, Utah. Thanks are also extended to the num- 
erous individuals who have collected material for me or made 
their own collections available for study. 

LITERATURE CITED 

Arnett, R. H. 

1962. The beetles of the United States (A manual for identification). 

Part V. Fascicle 98. Cisidae. Catholic Univ. Amer. Press, 

Washington, D. C. Pp. 827-830, pi. 1.98. 
Baker, H. G. 

1955. Self-compatibility and establishment after "long-distance" dis- 
persal. Evolution, 9(3): 347-348. 
Blatchley, W. S. 

1910. An illustrated descriptive catalogue of the Coleoptera or beetles 

(exclusive of the Rhynchophora) known to occur in Indiana. 

Bull. Indiana Dept. Geol. Nat. Res., 1, 1385 pp., 590 figs. 
1918. On some new or noteworthy Coleoptera from the west coast of 

Florida. — IV. Canad. Ent., 50: 52-59. 
1923. Notes on the Coleoptera of southern Florida with descriptions 

of new species. Canad. Ent., 55: 13-20, 30-36. 



1967 PARTHENOGENETIC FUNGUS BEETLE 13 

BoviNG, A. G. and F. C. Craighead 

1931. An illustrated synopsis of the principal larval forms of the order 
Coleoptera. [part] Entom. Amer.. N.S.. 11(4): 258-351. pis. 
86-125. 
Casey, T. L. 

1898. Studies in the Ptinidae, Cioidae, and Sphindidae of America. 
New York Ent. Soc. 6(2): 61-93. 
Chujo, M. 

1939. On the Japanese Ciidae (Coleoptera). Mushi. 12( 1 ) : 1-10. 
Dalla Torre, K. W. von 

1911. Cioidae. Pars 30. //; W. Junk and S. Schenkling, edit.. Coleop- 
terorum Catalogus. Junk, Berlin. 32 pp. 
Darewski. I. S. and W. N. Kulikowa 

1961. Natiirliche Parthenogenese in der polymorphen Gruppe der 
kaukasischen Felseidechse {Lacerta saxicola Eversmann). Zool. 
Jahrbuch., 89(1 ): 1 19-176. 35 figs. 

DURY, C. 

1917. Synopsis of the coleopterous family Cisidae (Cioidae) of 
America north of Mexico. J. Cincinnati Soc. Nat. Hist.. 22(2): 
1-27. 
Elton, C. 

1949. Population interspersion: an essay on animal community pat- 
terns. J. Ecol.. 37: 1-23. 

FUKUDA, A. 

1940. .A study of the Japanese Ciidae, Coleoptera (II). [In Japanese] 
Trans. Nat. Hist. Soc. Formosa, 30: 48-53. 5 figs. 

Hatch, M. H. 

1962. The beetles of the Pacific Northwest. Part III: Pselaphidae and 
Diversicornia I. Univ. Washington Press. Seattle, ix + 503 pp.. 
66 pis. 

Lawrence, J. F. 

1965. Comments on some recent changes in the classification of the 
Ciidae (Coleoptera). Bull. Mus. Comp. Zool., 133(5): 273- 
293. 
Leng. C. W. 

1920. Catalogue of the Coleoptera of America north of Mexico. Sher- 
man, Mount Vernon, N. Y. x -\- 470 pp. 
Linsley, E. G. 

1958. Geographical origins and phylogenetic affinities of the ceram- 
bycid beetle fauna of western North America. In C. L. Hubbs, 
edit., Zoogeography. Amer. Assoc. Adv. Sci., Publ. 51. Pp. 
299-320. 
LoHSE, G. A. 

1964. Die in Mitteleuropa vertretenen Gattungen der Cisidae. (1. 
Beitrag zur Kenntniss der mitteleuropaischen Cisidae.) Ent. 
Blatter, 60(2): 116-122. 



14 BREVIORA No. 258 

LONGHURST, A. R. 

1955. Evolution in the Notostraca. Evolution, 9: 84-86. 
Mayr, E. 

1963. Animal species and evolution. Harvard Univ. Press, Cambridge, 
Mass. xiv + 797 pp. 
Melli^, J. 

1848. Monographie de I'ancien genre Cis des auteurs. Ann. Soc. Ent. 
France, Scr. 2, 6: 205-274,313-396, pis. 9-12. 

MiYATAKE, M. 

1954. Studies on the Japanese Ciidae, I (Coleoptera). Sci. Rep. 
Matsuyama Agr. Coll.. 14: 40-67, 11 pis. 
Paviour-Smith, Kitty 

1960. The fruiting-bodies of macrofungi as habitats for beetles of the 
family Ciidae (Coleoptera). Oikos, lid): 43-71, 1 pi. 
Reitter, E. 

1902. Analytische Uebersicht der palaearctischen Gattungen und 
Arten der Coleopteren-Familien: Byrrhidae (Anobiidae) und 
Cioidae. Verb. Naturforsch. Ver. Briinn, 40: 1-61. 
Stebbins, G. L. 

1950. Variation and evolution in plants. Columbia Univ. Press. New 
York, xix -|- 643 pp. 

SUOMALAINEN, E. 

1962. Significance of parthenogenesis in the evolution of insects. Ann. 
Rev. Ent., 7: 349-366. 
Weiss, H. B 

1920. Coleoptera associated with Poly poms versicolor L. in New 
Jersey. Psyche, 27: 137-139. 
Weiss, H. B. and E. West 

1920. Fungous insects and their hosts. Proc. Biol. Soc. Washington. 

33:1-20, 1 pi. 
1921a. Additional fungous insects and their hosts. Proc. Biol. Soc. 

Washington, 34: 59-62. 
1921b. Additional notes on fungous insects. Proc. Biol. Soc. Washing- 
ton, 34: 167-172, 1 pi. 
White, M. J. D. 

1954. Animal cytology and evolution. 2d ed. Cambridge Univ. Press. 
454 pp. 
Wollaston, T. V. 

1854. Insecta Maderensia; being an account of the insects of the 
islands of the Madeiran group. Van Voorst, London, xliii -f 
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(Received 5 July, 1966). 



BREVIORA 

Mmseiiwii of Comiipsirsitive Zoology 

Cambridge, Mass. 3 February, 1967 Number 259 

EXPANDING THE PALPI OF MALE SPIDERS' 

By William A. Shear^ 

INTRODUCTION 

Genitalia and secondary sexual structures are important in the 
determination of spiders to the species level. This is especially 
true in the male, where the palpus has become modified as an 
intromittent organ. In many genera, the females are virtually in- 
separable into species, and small differences in shape, size, rela- 
tive position, and complexity of the parts of the highly modified 
and specialized male palpus are used to distinguish between 
closely related species of a genus. 

In families where the male palpus is highly specialized, the 
palpal organ has become so folded and convoluted that some 
parts are hidden in the normal retracted palpus. While these parts 
are not always of taxonomic importance, they nevertheless need 
to be examined in detail, especially in characterizing new species. 

Many publications concerned with spider taxonomy include 
figures of male palpi that have been treated in order to unfold or 
to expand the organ. In other publications, figures are of un- 
treated and unexpanded palpi and, as a result, confusion and un- 
certainty arise in comparing species and in making identifications, 
because frequently there are striking differences between ex- 
panded and unexpanded palpi of the same species. In an ex- 
panded palpus, sclerites, which are hidden in an unexpanded pal- 
pus, are revealed and the serial relationship of the sclerites be- 
comes more apparent. 

My purpose is not to decide what parts of the palpus are of 
value in taxonomy, but merely to determine what additional parts 
can be seen when the palpi of representative species are expanded. 

' Part of a thesis submitted to the faculty of the University of New Mexico 

for the degree of Master of Science 

' Department of Biology, Concord College, Athens, West Virginia 



2 BREVIORA No. 259 

I wish to express my indebtedness to Dr. C. Clayton Hofif for 
his guidance throughout the period of this study, and especially to 
Dr. Herbert W. Levi of the Museum of Comparative Zoology at 
Harvard University, Cambridge, Massachusetts, for many helpful 
suggestions. I appreciate the use of space and facilities made 
available by Dr. James S. Findley, Dr. William C. Martin, and 
Dr. William G. Degenhardt, curators of the Museum of South- 
western Biology, Albuquerque, New Mexico. Some of the speci- 
mens used for this study were donated by Dr. Andrew A. Weaver 
of the College of Wooster, Wooster, Ohio, and by Mr. Joseph A. 
Beatty of Southern Illinois University, Carbondale, Illinois. 

METHODS 

In The Spider Book, Comstock (1940, p. Ill) described a 
process for expanding the complex palpal organs of male spiders. 
This method was probably the one used by Comstock in prepar- 
ing specimens for a 1910 paper on spider palpi, because the text 
of the 1910 publication is the same as the section on palpi in 
The Spider Book. The method he described involves immersing 
the palpus in boiling 10 per cent "caustic" for 10-15 minutes. 
After treatment, the palpus is rinsed in water and preserved in 
glycerine. 

Engelhardt (1910) challenged this method on the grounds that 
the boiling solution was somewhat destructive. He was working 
on the more delicate female genitalia at that time. In 1925. 
Petrunkevitch, in a similar study, concluded that Engelhardt's ob- 
jections were not substantiated by results, and further stated 
that palpi prepared by boiling corresponded closely to those he 
observed in actual use by the male spider. 

Gering (1953) prepared a large series of palpi using boiling 
potassium hydroxide solution, but for studies of the natural mode 
of expansion placed preserved palpi in concentrated potassium 
hydroxide solution for a few minutes before a rapid transfer to 
distilled water, where the palpi were inflated by osmotic pressure. 

Levi (1961) used hot sodium hydroxide solution, followed by 
distilled water. Osmotic pressure expanded the organ. In 1965, 
he suggested the use of boiling 85 per cent lactic acid solution, on 
the grounds that it is far less destructive than sodium hydroxide 
solution. He also mentioned that satisfactory results can be ob- 
tained by using fine needles and forceps to expand the parts of the 
palpal tarsus. 



1967 PALPI OF MALE SPIDERS 3 

The method used in this study was similar to that used by 
Gering (1953) and Levi (1961). Palpi removed from specimens 
preserved in 70 per cent ethanol solution were placed in 10 per 
cent KOH solution for 6-8 hours at room temperature and were 
completely expanded in distilled water. The expanded palpi were 
then returned to alcohol. 

Twenty-nine species, distributed among 22 genera and 10 fam- 
ilies of higher spiders, were selected as test samples on the basis 
of the number of specimens available, completeness of taxonomic 
definition, and number of available references. Also, each of 
these species has palpi similar in degree of complexity to the palpi 
of a number of other species and genera in their respective families. 

Before expanding the palpus of one of the species, a number 
was assigned and placed in the vial containing the specimens. In 
most cases, the right palpus of an individual was removed and 
treated, leaving the left as a control, but in some small species, 
the whole male was treated to avoid damage to the palpus in re- 
moval. After expansion, the palpus (or the whole specimen) was 
placed in alcohol in a separate vial and labelled with the same 
number as was previously assigned. The vial containing the pal- 
pus was then bound to the original vial of specimens with a rub- 
ber band. These specimens in alcohol were used in making the 
freehand drawings and in writing the descriptions. 

TERMINOLOGY 

The course taken by the evolution of secondary sexual struc- 
tures in spiders is open to speculation. Comstock (1940), Bar- 
rows (1925), Savory (1928), Gertsch (1949), Alexander and 
Ewer (1957), and Levi (1961) have expressed ideas on the sub- 
ject. Until we can be reasonably certain of the phylogeny of the 
sclerites of the male palpus, names applied to these scierites do 
not necessarily indicate homologies. 

Comstock (1940) effectively reviewed previous research on 
spider palpi and devised a system of terminology widely accepted 
by most authors. Comstock's terminology will be followed in this 
study. 

The following description refers to the expanded palpus of 
Araneus sericatus (Clerck) (Figs. 20, 21). 

The segments of the palpus proximal to the tarsus are only 
rarely modified. In most spiders, the coxa of the palpus bears an 
endite that functions as a mouth part, crushing and straining food 
material. The trochanter can bear a small projection of unknown 



4 BREVIORA No. 259 

function. The femur of the palpus of a male spider is sometimes 
furnished with sclerotic picks, which rub against stridulating areas 
on the sides of the chelicerae, producing squeaking or barking 
noises. Gering (1953) demonstrated in the genus Agelenopsis 
that apophyses on the tibia and patella can serve as locking de- 
vices in copulation. 

It is the palpal tarsus, especially the tarsal claw, that has been 
strongly modified (Barrows, 1925). The body of the tarsus has 
become modified in males of the more specialized families to form 
a cup-shaped cymbium (cy), the cavity of which is the alveolus. 
The cymbium is basal in position and the palpal organ itself folds 
into the alveolus when at rest (Figs. 18, 19). The most proximal 
portion of the organ is called the basal division. Actually attached 
to the wall of the alveolus is a hollow, bladder-like structure 
termed the basal hematodocha (bh). The cavity of the basal 
hematodocha is continuous with that of the cymbium, allowing 
the hematodocha to be inflated by increasing blood pressure. At 
the distal end of the basal hematodocha is a smooth or annulated 
ring of sclerotic material, the subtegulum (st). 

The middle division consists of the tegulum (/), an inappropri- 
ately named ring of heavily sclerotized tissue connected to the 
subtegulum by the middle hematodocha (nih). Articulated with 
the tegulum is a usually small sclerite, the median apophysis (ma). 
In some male spiders, this appears as a knob or process on the 
tegulum. 

The apical division is the most complex of the three divisions. 
The embolus (e) is the functional part of the organ, bearing at its 
tip the opening of the ejaculatory duct. The conductor (c) is a 
usually membranous outgrowth from the base of the embolus, 
serving to protect the embolus. The radix (r) and stipes (s) are 
small, and are often absent in less complex palpi. Both radix and 
stipes arise near the base of the embolus, although they may ap- 
pear more distal. The palpal organs in some families do have 
parts of the organ actually distal to the embolus. These parts in- 
clude a distal hematodocha (dh) and the lateral subterminal 
apophysis (Isa). The terminal apophysis (to) is located at the 
very tip of the organ. 

OBSERVATIONS 

Order ARANEAE 
Family THERIDIIDAE 

The theridiids, or comb-footed spiders, have recently been the 
subject of an extensive revision by Levi from 1952 to 1964. In 



1967 PALPI OF MALE SPIDERS 5 

1961, Levi published his ideas on the evolution of the palpal organ 
in this large and complex family, and concluded that the original 
type of palpus was probably simple, that most members of the 
Theridiidae have a complex palpal organ, and that secondary 
simplification can be seen in some otherwise highly specialized 
theridiids. 

For my purposes, I selected four species that represent various 
degrees of complexity within the family. 

Steatoda borealis (Hentz) 

Figs. 1, 2, 3 

Levi (1957a) gives excellent figures of the palpus of Steatoda 
borealis in his revision of the genus Steatoda. Both expanded and 
unexpanded palpi are shown, and some individual sclerites are 
figured. 

In a mesal view of the unexpanded right palpus (Fig. 1), the 
embolus {e) can be seen, but the basal portion is hidden. The 
conductor (c), and the radix (r), both of which support the em- 
bolus in this species, are fairly obvious, but the shape of the radix 
is obscured by other sclerites lying over its ectal part. In a mesal 
view of an expanded right palpus (Fig. 2), the rotate base of the 
embolus {e) is exposed, and its true relationship to the conductor 
and radix (r) is revealed. In an ectal view of the same palpus 
(Fig. 3), expansion has caused a general ectad rotation of all the 
sclerites, bringing the median apophysis {ma) from behind over- 
lying sclerites. 

Enoplognatha tecta (Keyserling) 
Figs. 4, 5 

When Levi (1957b) revised the genus Enoplognatha, he gave 
two figures of an unexpanded palpus and one of an expanded 
palpus of this species. 

In an ectal view of an unexpanded left palpus (Fig. 4), the 
median apophysis and subtegulum are completely covered by 
overlying sclerites, although the shape of the embolus (e) and its 
relationship to the conductor (c) and radix (r) are obvious. In 
an ectal view of an expanded palpus (Fig. 5), the median apophy- 
sis {ma) has been rotated ectad and ventrad and the subtegulum 
{st) is visible. In both Figures 4 and 5, the hooklike projection 
of the anterior portion of the cymbium {cy) is seen. This is the 
paracymbial hook, which in some families is developed as a sep- 
arate sclerite. 



6 BREVIORA No. 259 

Theridion differens Emerton 
Figs. 6, 7 

Levi (1957b) reviewed the genus Theridion, a very large and 
geographically widespread assemblage of, in many cases, remark- 
ably similar species. Levi did not figure an expanded palpus of 
Theridion differens, although he figured expanded palpi of sev- 
eral other species of the genus. 

When an unexpanded right palpus in ventral view (Fig. 6) is 
compared with an expanded right palpus (Fig. 7) seen from the 
same aspect, it is clear that little more is visible after expansion. 

Theridion frondeum Hentz 
Figs. 8, 9 

This species is figured by Levi in his revision of Theridion 
(1957b). As in Theridion differens, an expanded palpus (Fig. 9) 
shows very little that is not readily visible in the unexpanded pal- 
pus (Fig. 8). 

Family LINYPHIIDAE 

Recent literature on this family has been restricted to a few 
occasional papers describing new species and giving new distribu- 
tional records (Chamberlin and Ivie, 1943; Gertsch and Davis, 
1946; Gertsch, 1951; and others). Blauvelt (1936) studied the 
palpus of Linyphia. 

Because the palpi of members of this family are somewhat uni- 
formly complex, two of the more common species were selected 
for this study. 

Drapetisca aheranda Chamberlin 
Figs. 10, 11 

In a mesal view of an unexpanded right palpus (Fig. 10) of 
Drapetisca alteranda, the paracymbium {pc) is seen as a free 
sclerite, loosely articulated with the cymbium {cy). The lateral 
subterminal apophysis {Isa) is highly developed and forms a 
shield covering the entire palpal organ. In a mesal view of an 
expanded right palpus (Fig. 11), the general ectad rotation 
caused by expansion is readily noticed. Parts, including the sub- 
tegulum {st) and tegulum (0, which were completely hidden 
before treatment, are now easily seen. The bulky base of the 
embolus (e) is now visible, and the shape of the lateral sub- 
terminal apophysis {Isa) is more satisfactorily observed. 



1967 PALPI OF MALE SPIDERS 7 

Linyphia margimita C. L. Koch 
Figs. 12, 13 

In this species, studied by Blauvelt (1936), the lateral sub- 
terminal apophysis {Isa), as seen in an ectal view of the unex- 
panded right palpus (Fig. 12), is not so highly developed as it is 
in the complex shield of Drapetisca. The tegulum (/) and sub- 
tegulum {st) are at least partially visible, but the conductor (c) 
and embolus (e) are obscured. In an ectal view of an expanded 
right palpus (Fig. 13), an ectad rotation brings into clearer view 
the position of the median apophysis {ma), which seems much 
more distal in the unexpanded palpus, and the exact shape and 
size of the unusually heavy embolus (e). The conductor (c) is 
readily observed, with its supporting sclerotic bar along one side. 
The lateral subterminai apophysis {Isa) is reflexed, appearing as 
if it were proximal, rather than distal, to the embolus. 

Family MICRYPHANTIDAE 

This family has not been studied as a unit since the publication 
of a long series of papers by Crosby, Crosby and Bishop, and 
Bishop and Crosby from 1905 to 1938 (listed by Kaston, 1948). 
Because of the seemingly simplified structure of the male palpus, 
there is some controversy about the proper taxonomic position of 
the Micryphantidae. The family is such a large one and shows 
so many degrees of palpal complexity that it would be impossible 
to treat the Micryphantidae satisfactorily in a paper of this nature. 
A recent study by Merrett (1963) of the palpal anatomy of this 
family treats the Micryphantidae as the subfamily Erigoninae of 
the Linyphiidae, but Merrett's work was confined to British species. 

The species I studied were selected on the basis of their inter- 
mediate type of palpal organ. 

Soulgas corticarius (Emerton) 
Figs. 14, 15 

This species was treated by Crosby and Bishop (1936), who 
gave a figure of an unexpanded palp, along with the characteristic 
cephalic modification found in this group. 

An ectal view of an unexpanded right palpus (Fig. 14) shows 
the obviously simple structure, including a free paracymbium 
(pc), subtegulum {st), tegulum {t), and embolus {e). The shape 
of the tibia {ti) is unusual, having a modification that in at least 
some degree occurs in all male micryphantids. In mesal view, an 



8 BREVIORA No. 259 

expanded right palpus (Fig. 15 shows two features that cannot 
be seen in an unexpanded palpus. The great development of the 
basal hematodocha (bh) is obvious. A small sclerite (ma) pos- 
sibly represents a rudimentary median apophysis. 

Ceratinopsidis jormosa (Banks) 
Figs. 16, 17 

Bishop and Crosby (1930) published a figure of the unex- 
panded palpus of Ceratinopsidis formosa. The unexpanded pal- 
pus (Fig. 16) has features similar to those of Soulgas corticarius. 
In an expanded palpus (Fig. 17) observed in dorsoectal view, 
the greatly developed basal hematodocha (bh) is seen, but no 
sclerite corresponding to a median apophysis is apparent. 

FamUy ARANEIDAE 

Archer (1951, p. 3) defined this family as follows: 
"Paracymbium vertical or divergent from the axis of the cym- 
bium, but its basal face not resting on the apical face of the tibia, 
instead usually being separated from it by a distinct gap. Attach- 
ment of the genital bulb to the cymbium ranging from universal- 
median and then by all degrees of migration to frankly basal (as 
in the Theridiidae). Tegulum greatly overbalancing the sub- 
tegulum, the latter ranging from ring-like to a vestigial knob. 
Position of the cymbium and genital bulb normal." 

If the Araneidae are considered in the strict sense, excluding 
the Tetragnathidae and the Theridiosomatidae, then all members 
of the family have complex palpi, but some do show various de- 
grees of reduction in certain sclerites. I selected eight fairly rep- 
resentative species. 

Araneus sericatus Clerck 
Figs. 18, 19, 20, 21 

The complicated palpus of this species was described in the 
section on TERMINOLOGY. In a ventral view of the unex- 
panded right palpus (Fig. 18), little can be seen except the sub- 
tegulum {st), tegulum {t), terminal apophysis {ta), and median 
apophysis {ma). In mesal view (Fig. 19), some additional struc- 
tures are visible, but their relationships to definitely identifiable 
structures are obscure. A number of interesting structures are 
revealed in an ectal view of an expanded right palpus (Fig. 20). 
The anterior portion of the subtegulum is completely unsclero- 
tized, so that the basal and middle hematodochae {bh, mh) are 



1967 PALPI OF MALE SPIDERS 9 

fused anteriorly. The tegulum (/) is a heavy rounded structure 
articulating closely with the large median apophysis (ma). The 
parts of the embolic subdivision are visible distal to the tegulum. 
The spatulate stipes (s) and small conductor (c) articulate with 
the base of the embolus (e), but the radix (r) has been dis- 
placed proximad. In a mesal view of the same palpus (Fig. 21), 
the hook-like lateral subterminal apophysis (Isa) is fused with 
the large plate of the terminal apophysis (ta), and both are sep- 
arated from the embolic subdivision by the distal hematodocha 
(dh). 

Araneus nordmanni (Thorell) 

The palpus of Araneus nordmanni is very similar in general 
plan to that of Araneus sericatus, differing only in the shape and 
position of some of the sclerites. Expansion in this species has 
the same effect as expansion has on the palpus of Araneus seri- 
catus. 

o 

Argiope trifasciata (Forskal) 
Figs. 22, 23 

The main difference between the palpus of this species and 
those of species of the genus Araneus lies in the reduction of some 
of the sclerites of the embolic subdivision. 

In a mesal view of an unexpanded right palpus (Fig. 22), the 
subtegulum is not seen, but in its place a portion of the basal 
hematodocha (b/i) appears as the most basal part. The tegulum 
(t), embolus (e), and well-developed conductor (c) are all read- 
ily visible. The median apophysis (tna) is large and has a ser- 
rated ventral edge. In a dorsomesal view of an expanded right 
palpus (Fig. 23), the subtegulum (st) has been moved into view. 
The ectad rotation of the median apophysis (ma) has revealed a 
small hook on the dorsal surface. The embolus (e) and con- 
ductor (c) are fused for about two-thirds of their length. Other 
sclerites of the embolic subdivision seem to be absent, although 
the swollen base of the embolus possibly represents a reduced 
radix. 

Araniella displicata (Hentz) 
Figs. 24, 25 

Chamberlin and I vie (1942) removed this species from Ara- 
neus and made it the type species of their new genus Araniella. 
Unfortunately, they did not present figures of the type species. 



10 BREVIORA No. 259 

I studied the palpus in detail. In an ectal view of an unexpanded 
right palpus (Fig. 24), the subtegulum (st) appears cupped, as 
if it has taken on the function of the conductor in this species. 
The very large tegulum (0 is easily visible. The basal part of 
the embolus (e) seems to be fused to the tegulum. The basal part 
of the median apophysis (ma) is hidden by the cymbium (cy). 
In an ectal view of an expanded right palpus (Fig. 25), the re- 
duction of the subtegulum (st) characteristic of this family, as 
described by Archer (1951), is clearly seen. The basal part of 
the median apophysis (ma) is visible and there is a small "hema- 
todocha" around the base of the embolus (e) and median apophy- 
sis. This area of lightly sclerotized tissue may represent a vesti- 
gial distal hematodocha, but due to the absence of any sclerites 
morphologically distal to the embolic subdivision, this "hema- 
todocha" is best considered an articulating membrane between 
the tegulum (t) and the median apophysis. 

Singa pratensis Emerton 
Figs. 26, 27 

The members of the genus Shiga are structurally close to the 
theridiids (Kaston, 1948), but the structure of the palpus is 
closer to Araneiis than are some species formerly considered to 
be in Araneus. According to Kaston (1948), who figures the pal- 
pus and epigynum, the species of Singa resemble one another so 
closely in form and appearance that they can be separated only 
by examining the external genitalia. 

By comparing a mesal view of an unexpanded right palpus 
(Fig. 26) with the palp of Araneus sericatus (Fig. 19), one sees 
that the relative sizes of the structures correspond closely. The 
subtegulum (st) is reduced. The tegulum (0. median apophysis 
(ma), and terminal apophysis (ta) are all visible. The expanded 
right palpus in ectal view (Fig. 27) bears close resemblance to the 
expanded palpus of Araneus sericatus (Fig. 20). The sclerites of 
the embolic subdivision are all visible in essentially the same posi- 
tions as in Araneus sericatus, but the radix of Araneus sericatus 
is in a slightly different position (Fig. 21). 

Neoscona arabesca (Walckenaer) 

The genus Neoscona is closely allied to Araneus. The males 
bear complex palpal organs built around the general Araneus 
plan. Expansion delineates the relationships of the various scler- 
ites in much the same way as in Araneus sericatus, Araneus nord- 
manni, and Singa pratensis. 



1967 PALPI OF MALE SPIDERS 11 

Mangora gibberosa (Hentz) 
Figs. 28, 29 

Mangora gibberosa has a complex palpus that, due to the dis- 
placement of the sclerites, seems even more complex than it really 
is. In an ectal view of an unexpanded right palpus (Fig. 28), 
the subtegulum {st) is visible and is large for a member of the 
family Araneidae. The tegulum (0 and terminal apophysis {ta) 
are separated by a large radix (r), the sheet-like, membranous 
conductor (c), and the enlarged base of the embolus (e). In a 
ventroectal view of an expanded right palpus (Fig. 29), the hema- 
todochae seem to appear very much reduced, and most of the 
sclerites occupy positions different from those of the previously 
described araneids. In addition, the conductor is much enlarged. 

Cyclosa conica (Pallas) 

Kaston (1948) gave a figure of the palpus of the male of 
Cyclosa conica. In the degree of palpal complexity and reduction 
of hematodochae, this species lies somewhere between Argiope 
trifasciata and Mangora gibberosa. As in Mangora, the sclerites 
are much displaced, making them difficult to identify. Expansion 
causes an ectad rotation and further displacement, but, after 
treatment, individual sclerites can be seen as entities and at least 
tentatively identified. The median apophysis is bifurcate, as in 
many araneids, and the distal lobe is sigmoid. As in Mangora, the 
conductor is large and membranous and interferes with identifica- 
tion of the sclerites in the unexpanded palpus. 

Family AGELENIDAE 

The genitalia of some agelenids have been studied intensively 
with regard to their form and function (Gering, 1953). The 
palpus is moderately complicated, with a reduced subtegulum and 
a well-developed conductor that serves, at least in Agelenopsis, 
as a locking device during copulation (Gering, 1953). 

Wadotes calcaratus (Keyserling) 
Figs. 30, 31, 32 

Muma (1947) revised the genus Wadotes and figured only 
individual sclerites rather than complete palpal organs. He con- 
sidered the terminal apophysis to be of unusual value in taxo- 
nomic decisions. 



12 BREVIORA No. 259 

In a ventral view of an unexpanded right palpus (Fig. 30), the 
subtegulum (st), tegulum (/), and embolus (e) are easily seen, 
as is the terminal apophysis (ta). In an ectal view of an ex- 
panded right palpus (Fig. 31), the distal portion of the embolus 
(e) is extensive and threadlike. Muma (1947) refers to the em- 
bolus as lying along a distinct conductor, but it seems to me that 
the only structure one can suggest as a conductor is solidly fused 
to the terminal apophysis. In a mesal view of an expanded right 
palpus (Fig. 32), the feature most noticeable is the distinctly 
annulated subtegulum (st). 

Coras lamellosus (Keyserling) 

Muma (1946) revised the North American members of the 
genus Coras and figured the palpi of a number of species, includ- 
ing Coras lamellosus. The palpus is very much like that of 
Wadotes calcaratus in the general arrangement of sclerites. After 
expansion, a general ectad rotation brings the long, thin embolus 
into full view. The conductor can be seen as separate from the 
terminal apophysis, and there is a twisted median apophysis that 
before expansion was concealed by the conductor and terminal 
apophysis. 

Agelenopsis utahana (Chamberlin and Ivie) 
Figs. 33, 34 

This is one of the species used by Gering (1953) in a monu- 
mental study of the structure and function of the external geni- 
talia of the genus Agelenopsis, and he gives several very useful 
figures of the expanded palpus. 

In an ectal view of the unexpanded right palpus (Fig. 33), the 
embolus {e) is seen to be long, heavy, and spirally coiled. The 
tegulum (0 is somewhat indented to accommodate the embolus. 
In a mesal view of an expanded right palpus (Fig. 34), the fused 
basal and middle hematodochae {bh-mh) are visible. The sub- 
tegulum {st) has been pushed into view, and the conductor (c) 
has rotated mesad. Gering (1953) refers to the basal part of the 
embolus as the radix, and this may be morphologically correct. 

Tegenaria domestica (Clerck) 
Figs. 35, 36 

Tegenaria domestica is a common house spider often referred 
to as Tegenaria derhami (Scopoli), and its habits have been 
studied extensively. The legs and palpal segments of this species 
are elongate. 



1967 PALPI OF MALE SPIDERS 13 

In a ventral view of the unexpanded right palpus (Fig. 35), the 
cymbium (cy) bears a long distal finger. The tegulum (0, median 
apophysis (ma), and embolus (e) are fused into one sclerite 
capping the hematodochae. In an ectoventral view of an ex- 
panded right palpus (Fig. 36). the fused basal (bh) and middle 
hematodochae (mh) are inflated. The "cap" has been rotated 
ectad, and the hook-like conductor (c) has been extended on a 
short articulating membrane. No subtegulum was evident in this 
palpus, but an area on the basal hematodocha that is more heavily 
sclerotized than the rest of the hematodocha may correspond to 
the subtegulum. 

Cicurina robusta Simon 

Figs. 37, 38 

Exline (1936) and Chamberlin and Ivie (1940) examined rep- 
resentatives of the genus Cicurina, and both gave excellent figures, 
concentrating on the tibia, which in males of this genus is uniquely 
developed. Frequently they neglected to provide figures of the 
palpal organ itself. 

In an ectal view of an unexpanded left palpus (Fig. 37), the 
hooked conductor (c), the radix (r), and the embolus {e) are 
all visible, at least in part. The unusual tibial apophysis (//) ex- 
tends up the ectal side of the cymbium (cy). In an ectal view of 
an expanded right palpus (Fig. 38), the flattened subtegulum {st) 
has become visible, and the cupped tegulum (0 accommodates 
the radix (r) and the base of the embolus {e). The long and 
spirally wound embolus tapers to a hairlike tip. The hooked con- 
ductor (c) is extended. 

Family LYCOSIDAE 

This is a large and homogeneous family of which Lycosa is the 
principal genus. The palpi of most lycosids are similar to one 
another in general plan. 

Lycosa ammophila Wallace 

Wallace (1942) studied the lenta group of the genus Lycosa 
and described a number of new species, including Lycosa ammo- 
phila, from southeastern United States. Wallace gave figures of 
the palpus in ventral view, but emphasized the median apophysis, 
which he also figured separately in outline. In the unexpanded 
palpus, most of the features are readily visible. Expansion does, 
however, clarify relationship among the sclerites. 



14 BREVIORA No. 259 

Family CLUBIONIDAE 

The palpi of this family are simple, except for those of certain 
genera belonging to the subfamily Liocraninae and none of these 
are considered here. The palpal organs of most liocranines are 
similar to those of the Agelenidae. 

Clubiona bryantae Gertsch 
Figs. 39, 40 

Gertsch (1941) proposed Clubiona bryantae as a new name 
for Clubiona agreslis Emerton, but he gave no figures. The new 
name was accepted by Edwards (1958), who gave several figures 
of the male palpus. 

In an ectal view of an unexpanded right palpus (Fig. 39). the 
subtegulum (st), tegulum (r). and embolus {e) are subequal. 
The expanded right palpus (Fig. 40) shows the ectad rotation of 
the embolus (e), exposing its fingerlike tip. 

Ays ha gracilis (Hentz) 
Figs. 41, 42 

Aysha and a number of related genera (Anyphaena, Anyphae- 
nella, Gayenna, and others) are often placed in a separate family, 
the Anyphaenidae, on which no recent work has been done. 

In a ventral view of an unexpanded right palpus (Fig. 41), the 
tegulum (0 is large and extends below the alveolus or cavity of 
the cymbium (cy). The embolus (e) has a large, rotate, flattened 
base, and then tapers to a fine point, cupped by a projection of 
the cymbium. In a mesal view of an expanded right palpus 
(Fig. 42), all the parts appear to have been pushed ectad, and the 
subtegulum (st) is now visible. The track of the internal sperm 
duct through the tegulum is prominent. 

Family SALTICIDAE 

The Salticidae is considered by some authorities to contain the 
most advanced of spiders (Kaston, 1948), but the palpi are 
highly simplified. 

Metacyrba undata (De Geer) 
Figs. 43, 44 

In a ventral view of an unexpanded right palpus of Metacyrba 
undata (Fig. 43), little is visible except the tegulum {t), which 
is enlarged and greatly extended proximally into a hollow on the 



1967 PALPI OF MALE SPIDERS 15 

ventral surface of the palpal tibia (//), and the fused embolus- 
conductor (e, c). In the ectodorsal view of an expanded palp 
(Fig. 44), the basal hematodocha (b/i) is large and displaces all 
the sclerites ectad. The subtegulum (st) is visible as a small, 
sclerotic ring. 

Tutelina elegans (Hentz) 
Figs. 45, 46 

Kaston (1952) gave figures showing the general appearance 
of this species, but he did not illustrate the genitalia. In a ventral 
view of an unexpanded right palpus (Fig. 45), the general ap- 
pearance is essentially the same as in Metacyrba undata, but no 
recognizable conductor is present on or near the embolus (e). 
Expansion of the palpus (Fig. 46) brings about an ectad rotation 
and. in an ectodorsal view, the subtegulum ist) is exposed. 

Salticus scenicus (Clerck) 

This species is distributed throughout the United States and 
Europe and has been widely studied. The palpus is very simple 
and follows the general plan of the two previously described spe- 
cies of the family Salticidae. In the expanded palpus, the basal 
hematodocha is greatly expanded, and the subtegulum is brought 
into view. 

Family ULOBORIDAE 

This family was recently revised by Muma and Gertsch (1964), 
and a number of new species were described. Figures were given 
of unexpanded palpi and individual segments, but an error was 
made in the figure titles. 

The palpi of these spiders are remarkable for the extreme 
development of certain sclerites. The habits of the Uloboridae 
are unique, but among some species the modes of web-building 
converge with those of the Araneidae. This behavioral converg- 
ence is not reflected in the morphology of the palpal organs of the 
males. 

Hyptiotes cavatus Hentz 
Figs. 47, 48, 49 

Muma and Gertsch (1964) illustrate the entire palpus of Hyp- 
tiotes cavatus and also give a figure of what they call the '"tip of 
the embolus." Close examination of actual specimens and of the 
textual description given by Muma and Gertsch leaves little doubt 



16 BREVIORA No. 259 

that this figure actually represents the tip of the median apophysis. 
In an ectal view of an unexpanded right palpus (Fig. 47), the 
most striking feature is the large, elongated median apophysis 
(ma) with its membranous tip. The tegulum (0 and subtegulum 
(st) are almost completely covered by the enlarged membranous 
conductor (c). The embolus (e) originates mesally and coils 
around the other sclerites of the palpal organ. In a mesal view of 
the same palpus (Fig. 48), the origin of the embolus (e), as well 
as its attenuated distal part entering the conductor (c), is easily 
seen. The conductor is accompanied by the long, hooked radix 
(r). An ectoventral view of an expanded right palpus (Fig. 49) 
shows that expansion has resulted in little change, but the whole 
of the distal part of the embolus (e) can now be seen and its 
relationship to the median apophysis (ma) is clear. 

Family DICTYNIDAE 

Chamberlin and Gertsch (1958) revised this family, of which 
Dictyna is the largest genus. While the palpal organs of all North 
American species of Dictyna are similar, Chamberlin and Gertsch 
separated species on the basis of the structure of the tip of the 
embolus. 

Dictyna sublata (Hentz) 
Figs. 50, 5 1 

In a mesal view of an unexpanded right palpus (Fig. 50) of 
Dictyna sublata, the embolus (e) appears heavy and coiled and 
distally enters the large shieldlike conductor (c). The path of the 
sperm tube can be traced through the tegulum (0 and the base of 
the embolus. A ventral view of an expanded right palpus (Fig. 
51) shows that an almost complete ectad rotation of the base of 
the embolus has occurred. The subtegulum (st), as well as the 
basal and middle hematodochae (bh, mh), is visible. 

DISCUSSION 

For purposes of discussion, the males of species used in this 
study can be separated into three groups: those with very complex 
palpi, those with moderately complex palpi, and those with simple 
palpi. 

Drape tisca alteranda, Linyphia marginata, Araneus sericatus, 
Araneus nordmanni, Neoscona arabesca, Cyclosa conica, Singa 
pratensis, Mangora gibberosa, and Hyptiotes cavatus fall in the 



1967 PALPI OF MALE SPIDERS 17 

group having very complex palpi. In this group almost all pos- 
sible sclerites are present in a more or less highly developed form. 
These species are especially characterized by the number of scle- 
rites distal to the embolus. Males of species of Drapetisca and 
Linyphia have palpi in which the lateral subterminal apophysis is 
developed into a large shield covering the whole organ. In species 
of Araneus and Shiga, the male palpus has many structures con- 
cealed by other sclerites and by the manner in which the palpus 
is folded. In males of Mangora, Cyclosa, and Hyptiotes displace- 
ment of the sclerites has caused confusion when unexpanded palpi 
are used in the description of species. Expanding the palpus can 
be of great value in this group. 

In the group with moderately complex male palpi are: Enoplo- 
gnatha tecta, Theridion differens, Theridion jrondeum, Steatoda 
borealis, Argiope trijasciata, Araniella dispUcata, Wadotes calcara- 
tiis, Agelenopsis utahana, Coras lamellosus, Cicwina robusta, 
Lycosa ammophila, and Dictyna sublata. This group is character- 
ized by the approximately equal development of most sclerites. 
The apical division is poorly represented, and some of the sclerites 
of the embolic subdivision are reduced. Usually the basal hema- 
todocha is well developed. When the palpus is moderately com- 
plex, one has to use his own judgment whether or not expansion 
will be worthwhile. The process of expansion has the value of 
making relationships clear and exposing portions of sclerites that 
may be hidden, because in most species in this group the full com- 
plement of sclerites (except the subteguium) is usually visible be- 
fore expansion. 

The third group, species with simple male palpi, includes: 
Soulgas corticarius, Ceratinopsidis jormosa, Tegenaria domestica, 
Clubiona bryantae, Aysha gracilis, Metacyrba undata, Salticus 
scenicus, and Tutelina elegans. These spiders have palpi charac- 
terized by the loss, fusion, or reduction of sclerites. Expansion 
seems to be of little utility in this group. 

The process of expansion seems to have its greatest value in 
studying species with complex palpi. The complex palpus may 
indicate specialization and occurs in three very highly specialized 
families (Theridiidae, Linyphiidae, Araneidae). However, spiders 
of a fourth very specialized family, the Salticidae, have a simple 
type of palpus. 

If the hematodochae of a complex palpus are reduced, expan- 
sion does not show structures that were not visible in the unex- 
panded palpus. In such cases, a delicate dissection of the palpus 



18 BREVIORA No. 259 

may be indicated. The need for expansion may also be partly ob- 
viated by the presence of other taxonomically useful sexual modi- 
fications, such as the ornate third leg of some salticid males or the 
tibial apophyses and cephalic modifications of the micryphantids. 

SUMMARY 

1. Several methods have been suggested for expanding the palpi 
of male spiders for taxonomic purposes. The method used in 
this study involved immersing the palpus for 6-8 hours in a 
10 per cent KOH solution, followed by a quick transfer to 
distilled water. Expanded palpi were stored in 70 per cent 
ethanol solution. 

2. The expanded palpus of each of 29 species of spiders was 
compared with the unexpanded palpus of the same species to 
determine if any parts not observable in the unexpanded pal- 
pus could be seen in the expanded palpus. 

3. The process of expansion was found to be most useful in 
species with complex palpi and least useful in species with 
simple palpi. 

4. Under certain conditions, such as the reduction of hemato- 
dochae, many features of complex palpi were still obscure. 

5. The experimenter must use his own judgment in deciding when 
the process of expansion is worthwhile, but certainly more 
detailed examination of palpi after expansion can be of benefit 
in the present confused state of spider taxonomy. 

LITERATURE CITED 

Alexander, A. J., and D. W. Ewer. 

1957. On the origin of mating behavior in spiders. Amer. Natur. 
91: 311-317. 
Archer, A. F. 

1951. Studies in the orbweaving spiders (Argiopidae). 1. Amer. 
Mus. Novit. No. 1487: 1-52. 
Barrows, W. M. 

1925. Modification and development of the arachnid palpal claw, 
with especial reference to spiders. Ann. Entomol. Soc. Amer. 
18: 483-516. 
Bishop, S. C, and C. R. Crosby. 

1930. Studies in American spiders: genera Cemtinopsls, Ceratinop- 
sidis, and Tiitaiho. J. New York Entomol. Soc. 38: 15-34. 
Blauvelt, H. H. 

1936. The comparative morphology of the secondary sexual organs 
of Linypliia. Festschr. Embrik Strand 2: 81-171. 



1967 PALPI OF MALE SPIDERS 19 

Chamberlin, R. v., and W. J. Gertsch 

1958. The spider family Dictynidae in America north of Mexico. 
Bull. Amer. Mus. Natur. Hist. 116: 1-152. 
Chamberlin. R. V., and W. Ivie 

1940. Agelenid spiders of the genus Ciciiriiui. Bull. Univ. Utah, 
Biol. Ser. 5(9): 1-108. 

1942. A hundred new species of American spiders. Bull. Univ. Utah, 
Biol. Ser. 7(1): 1-117. 

1943. New genera and species of North American linyphiid spiders. 
Bull. Univ. Utah, Biol. Ser. 7(6): 1-39. 

COMSTOCK, J. H. 

1940. The spider book, revised and edited by W. J. Gertsch. Com- 
stock Publishing Company, New York. 721 pp. 

Crosby. C. R., and S. C. Bishop 

1936. Studies in American spiders: miscellaneous genera of Erigo- 
neae. Festschr. Embrik Strand 2: 52-64. 
Edwards, R. J. 

1958. The spider subfamily Clubioninae of the United States, Can- 
ada and Alaska (Araneae: Clubionidae). Bull. Mus. Comp. 
Zool. 118: 365-436. 
Engelhardt, v. 

1910. Beitrage zur Kenntnis der weiblichen Copulationsorgane eini- 
ger Spinnen. Z. Wiss. Zool. 96: 32-117. 

EXLINE, H. 

1936. Nearctic spiders of the genus Ciciirina Menge. Amer. Mus. 
Novit. No. 8S0: 1-26. 
Gering, R. L. 

1953. Structure and function of the genitalia in some American 
agelenid spiders. Smithson. Misc. Collec. 121(4): 1-84. 
Gertsch, W. J. 

1941. New American spiders of the family Clubionidae. 11. Amer. 
Mus. Novit. No. 1148: 1-18. 

1949. American spiders. D. Van Nostrand Publishing Company, Inc., 

New York. 285 pp. 
1951. New American linyphiid spiders. Amer. Mus. Novit. No. 

1514: 1-11. 
Gertsch, W. J., and I. Davis 

1946. Report on a collection of spiders from Mexico. V. Amer. Mus. 

Novit. No. 1313: 1-11. 
Kaston, B. J. 

1948. Spiders of Connecticut. Connecticut State Geol. Natur. Hist. 

Surv. Bull. 70: 1-874. 



20 BREVIORA No. 259 

Kaston, B. J. 

1952. How to know the spiders. Wm. C. Brown Company, Dubuque, 

Iowa. 220 pp. 
Levi, H. W. 

1957a. The spider genera Crust iilinu and Steatoda in North America. 

Central America, and the West Indies (Araneae, Theridiidae). 

Bull. Mus. Comp. Zool. 117: 367-424. 
1957b. The spider genera Enoplognatha, Theridion, and Pcndisca in 

America north of Mexico (Araneae, Theridiidae). Bull. Amer. 

Mus. Natur. Hist. 112: 1-123. 
1961. Evolutionary trends in the development of palpal sclerites in 

the spider family Theridiidae. J. Morphol. 108: 1-9. 
1965. Techniques for the study of spider genitalia. Psyche 72(2): 

152-158. 
Merrett, p. 

1963. The palpus of male spiders of the family Linyphiidae. Proc. 
Zool. Soc. London 140(3): 347-467. 

MUMA, M. 

1946. North American Agelenidae of the genus Coras Simon. Amer. 
Mus. Novit. No. 1329: 1-20. 

1947. North American Agelenidae of the genus Wadotes Chamberlin. 
Amer. Mus. Novit. No. 1334: 1-12. 

MuMA, M., and W. J. Gertsch. 

1964. The spider family Uloboridae in North America north of 
Mexico. Amer. Mus. Novit. No. 2196: 1-43. 

Petrunkevitch, a. 

1925. External reproductive organs of the common grass spider, 
Agelena uaevici Walckenaer. J. Morphol. 40: 559-573. 
Savory, T. H. 

1928. The biology of spiders. Sidgwick and Jackson, London. 376 pp. 
Wallace, H. K. 

1942. A study of the lenta group of the genus Lycosa with descrip- 
tions of new species (Araneae, Lycosidae). Amer. Mus. Novit. 
No. 1185: 1-21. 



1967 PALPI OF MALE SPIDERS 21 



ABBREVIATIONS USED IN FIGURES 



bh 


basal hematodocha 


c 


conductor 


cy 


cymbium 


dh 


distal hematodocha 


e 


embolus 


Isa 


lateral subterminal apophysis 


ma 


median apophysis 


ink 


middle hematodocha 


pc 


paracymbium 


r 


radix 


s 


stipes 


St 


subtegulum 


t 


tegulum 


ta 


terminal apophysis 


ti 


tibia 



22 



BREVIORA 



No. 259 




Figs. 1-3. Steatoda boiealis. 1, Mesal view of an unexpanded right pal- 
pus. 2, Mesal view of an expanded right palpus. 3, Ectal view of an 
expanded right palpus. Figs. 4, 5. Enoplogiwtha tecta. 4, Ectal view of an 
unexpanded left palpus. 5, Ectal view of an expanded left palpus. Figs. 

6, 7. Theridion difjerens. 6, Ventral view of an unexpanded right palpus. 

7, Ventral view of an expanded right palpus. Figs. 8, 9. Theridion 
frondeum. 8, Ventral view of an unexpanded right palpus. 9, Ventral 
view of an expanded right palpus. 



1967 



PALPI OF MALE SPIDERS 
bh 



23 




Figs. 10, 11. Drape tisca alteranda. 10, Mesal view of an unexpanded 
right palpus. 11, Mesal view of an expanded right palpus. Figs. 12, 13. 
Linyphia marginata. 12, Fetal view of an unexpanded right palpus. 13, 
Ectal view of an expanded right palpus. Figs. 14, 15. Soiilgas corticarius. 
14, Ectal view of an unexpanded right palpus. 15, Mesal view of an ex- 
panded right palpus. Figs. 16, 17. Ceratinopsidis formosa. 16, Ectal view 
of an unexpanded right palpus. 17, Dorsoectal view of an expanded right 
palpus. 



24 



BREVIORA 



No. 259 





Figs. 18-21. Araneus sericatiis. 18, Ventral view of an unexpanded right 
palpus. 19, Mesal view of an unexpanded right palpus. 20, Ectal view of 
an expanded right palpus. 21, Mesal view of an expanded right palpus. 
Figs. 22, 23. Argiope trifasciato. 22, Mesal view of an unexpanded right 
palpus. 23, Dorsomesal view of an expanded right palpus. Figs. 24, 25. 
Araniella displicata. 24, Ectal view of an unexpanded right palpus. 25, 
Ectal view of an expanded right palpus. 



1967 



PALPI OF MALE SPIDERS 



25 






bh-mh 




34 

Figs. 26, 27. Singa prateiisis. 26, Mesal view of an unexpanded right 
palpus. 27, Ectal view of an expanded right palpus. Figs. 28, 29. Mangom 
gibberosa. 28, Ectal view of an unexpanded right palpus. 29, Ventroectal 
view of an expanded right palpus. Figs. 30-32. Wadotes calcaratus. 30, 
Ventral view of an unexpanded right palpus. 31, Ectal view of an expanded 
right palpus. 32, Mesal view of an expanded right palpus. Figs. 33, 34. 
Agelenopsis iitahana. 33. Ectal view of an unexpanded right palpus. 34, 
Mesal view of an expanded right palpus. 



26 



BREVIORA 



No. 259 





37 




40 






43 



Figs. 35, 36. Tegenuria domestica. 35, Ventral view of an unexpanded 
right palpus. 36, Ectoventral view of an expanded right palpus. Figs. 37. 

38. Cicwina robiista. 37, Ectal view of an unexpanded right palpus. 38, 
Ectal view of an expanded right palpus. Figs. 39, 40. Cliibiona bryantae. 

39, Ectal view of an unexpanded right palpus. 40, Ectal view of an ex- 
panded right palpus. Figs. 41, 42. Aysha gracilis. 41, Ventral view of an 
expanded right palpus. 42, Mesal view of an unexpanded right palpus. Fig. 
43. Metacyrba undata, ventral view of an unexpanded right palpus. 



1967 



PALPI OF MALE SPIDERS 



27 




Fig. 44. Metacyrbu iindata, ectodorsal view of an expanded right palpus. 
Figs. 45, 46. Tutelina elegans. 45, Ventral view of an unexpanded right 
palpus. 46, Ectodorsal view of an expanded right palpus. Figs. 47-49. 
Hyptiotes cavatiis. 47, Ectal view of an unexpanded right palpus. 48, Mesal 
view of an unexpanded right palpus. 49, Ectoventral view of an expanded 
right palpus. Figs. 50, 51. Dictyna siiblata. 50, Mesal view of an unex- 
panded right palpus. 51, Ventral view of an expanded right palpus. 



BREVIORA 

Meseimi of Coimparative Zoology 

Cambridge, Mass. 3 February, 1967 Number 260 



MONOGRAPH OF THE GENUS SPIROCERAMUS 
(MOLLUSCA: PULMONATA: UROCOPTIDAE) 

By William J. Clench 

The last complete monograph of the family Urocoptidae was 
that of H. A. Pilsbry, 1902-1904, in the second series of the Man- 
ual of Conchology. At that time only a single species was cred- 
ited to the genus Spiroceramiis. This species was S. ampins 
(Pfeiffer) from the region of Bayamo in the south of Oriente 
Province in eastern Cuba. Since then, C. G. Aguayo has de- 
scribed a species from the northern portion of Oriente Province, 
and I have described two species from the Bahamas, one from 
Little Abaco in the northern part, and one from Acklin's Island 
in the southern part of the Bahamian Archipelago. Two new 
species are described in this report from the Sierra de Cubitas, an 
isolated mountain system in north-central Camaguey Province in 
central Cuba. 

With comparatively few exceptions the West Indian Urocop- 
tidae are strictly calciphiles, living on limestone rocks where they 
feed on the encrusting lichens. A very few are arboreal. The 
Bahamian species were living on or under stones; the Cuban spe- 
cies on the limestone ledges. Under optimum conditions they can 
exist in incredible numbers, usually preferring shade, but they also 
exist on the limestone with no shade at all. They are active at 
night and during periods of rain. 

UROCOPTIDAE 

Genus Spiroceramus Pilsbry and Vanatta 

Spiroceramiis Pilsbry and Vanatta. 1898, Proc. Acad. Nat. Sci. Philadel- 
phia, 50: 281 (type species, Macroceramiis ampins Pfeiffer, original desig- 
nation). 

Shells thin, cylindrical, and composed of many narrow whorls. 
Spire acute, the apex entire, not deciduous. Colored a pale brown- 
ish with patches of white. Sculptured with fine oblique striae. 



2 BREVIORA No. 260 

Whorls 9 to 15, convex, early whorls straight or slightly angular. 
Aperture somewhat oblique, subcircular, and the lip narrowly re- 
flected. Suture indented, smooth or crenulate. Axis twisted or 
encircled with a thin spiral lamella. 

Pilsbry (1904, p. 172), on the basis of the then known single 
species S. ampins, stated that it had the shape of Holospira, the 
axis of Arangia, and the aperture and protoconch of Micro- 
ceramus. This statement can still stand with only the slight modi- 
fication that all species do not have a spiral lamella on the axis 
and certain species possess a crenulated suture. 

Subgenus Spiroceramus Pilsbry and Vanatta 

Species with or without the spiral lamella on the axis and hav- 
ing the oblique sculpture of fine striae similar on all whorls. The 
suture with or without crenulations. 

Spiroceramus (Spiroceramus) amplus (Pfeifi"er) 
Plate 1, figures 6, 7 

Macroceramiis amplus Gundlach, 1856, [//;] F. Poey, Memorias Sobre 
la Historia Natural de la Isla de Cuba, Habana 2: 8, no. 378 (Cuba). 
[Noinen niidiun.] 

Macroceramiis amplus 'Gundlach' Pfeiffer, 1858, Malakozoologische 
Blatter 5: 44 (Guisa [Bayamo, Oriente], Cuba); Pfeiffer, 1859, Mono- 
graphia Heliceorum Viventium 4: 689; Pfeiffer, 1868, Novitates Con- 
chologicae 3: 383, pi. 89, figs. 12-14; Arango, 1878, Contribucion a la 
Fauna Malacologica Cubana, Habana, p. 84 (Guisa and San Andres, 
Bayamo, Cuba). [Syntypes, MCZ 39318.] 

Microceramiis (Spiroceramus) amplus Pfeiffer. Pilsbry, 1904, Man. of 
Conch. (2) 16: 172, pi. 26, figs. 24, 29. 



Measurements 



Height Width 
mm mm 



10.5 3.9 Syntype. Guisa, Bayamo, Cuba 

11.1 4. Syntype. Guisa, Bayamo, Cuba 

11.1 3.3 Syntype. Sao Arriba, Holguin, Cuba 

Description: Shell reaching 11.1 mm in height, minutely um- 
bilicate, cylindrical, attenuate, the upper third rapidly tapering 
to the early whorls and sculptured. Whorls 14 to 15, convex, 
those on the upper one-third slightly angled. Color a light grayish 
brown and mottled with a dull white. Spire extended, the lower 
two-thirds nearly parallel sided, the conic one-third forming an 



1967 THE GENUS SPIROCERAMUS 3 

angle of about 35°. Aperture subcircular, with the outer lip 
slightly reflected. Inner or parietal lip reflexed over the minute 
umbilicus. Axis twisted and forming a broad lamella. Columella 
short and slightly angled. Suture deeply impressed. Sculpture con- 
sisting of numerous and fine, oblique striae. Protoconch with 1 V^ 
whorls and sculptured with fine, oblique striae. 

Remarks: This is rather a remarkable species for any member 
of the Urocoptidae as it occurs in two distinct and well separated 
areas. Holguin is about 70 kilometers NE of Bayamo with much 
relatively flat country between them. 

Specimens examined: Cuba. Oriente Prov.: Guisa, Bayamo; 
Cerro Moncada and Cerro San Juan. Sao Arriba, both Holguin. 

SPIROCERAMUS (Spiroceramus) barbouri Aguayo 
Plate 1, figure 5 

Spiroceramus ampins barbouri Aguayo, 1935, Mem. Soc. Cubana Hist. 
Nat., 9: 126, pi. 9, figs. 3-4 (Paradones junto a la Curva de la Campana, 
Gibara [Oriente], Cuba). [Holotype, MCZ 237877. Additional paratypes 
from the same locality, MCZ 110570.] 





Measurements 


Height 


Width 




mm 


mm 




5.5 
4.5 
4.75 
5.8 


2 
1.8 

2 
2 


Holotype 
Paratype 
Paratype 
Paratype 



Description: Shell reaching 5.8 mm in height, imperforate, 
cylindrical, attenuate, the upper third rapidly tapering to the early 
whorls and sculptured. Colored a light brown with most of the 
oblique costae and crenulations white. Whorls 9 to 11 and con- 
vex, the early 5 whorls straight-sided and vertical. Spire extended, 
the lower two-thirds nearly parallel sided, the conic one-third 
forming an angle of about 40°. Aperture subcircular, the outer 
lip simple and but slightly reflected. Axis twisted and forming 
a narrow lamella. Columella short and slightly angled. Suture 
impressed. Sculpture consisting of numerous and fine, oblique 
striae, many of which terminate above with crenulations at the 
suture. Protoconch with IVz whorls and sculptured with fine 
and straight striae. 

Remarks: Originally described as a subspecies of S. amplus, 
this present entity is a distinct species. S. barbouri is only one-half 



4 BREVIORA No. 260 

the height of amplus and has a crenulated suture which ampins 
does not have, and the axial lamella is much narrower. So far as 
now known, this species is limited to the type locality. 
Specimens examined: Holotype and paratypes. 

Spiroceramus (Spiroceramus)pilsbryi new species 
Plate 1, figure 3 

Holotype: MCZ 188845, from Cerro de Tuabaguey, Sierra de 
Cubitas, Camagiiey Province, Cuba. 

Paratypes: MCZ 256082, from the type locality, and MCZ 
188847 from Cueva del Circulo, Sierra de Cubitas, Camagiiey 
Province, Cuba. 

Measurements 

Height Width 
mm mm 

15.1 4.8 Holotype 

13 4.1 Paratype MCZ 188847 

Description: Shell reaching 15 mm in height, minutely umbili- 
cate, tapering from the ninth whorl to the protoconch and sculp- 
tured. Whorls 13 to 14 and slightly convex. Color a light horn 
with numerous, irregular patches of white. Spire extended and 
forming an angle of about 36°. Aperture subcircular, the outer 
lip slightly reflexed and the inner or parietal lip reflected over the 
minute umbilicus. Axis twisted and forming a broad lamella. 
Columella angled and somewhat broadened. Suture indented and 
coarsely crenulated. Sculpture consisting of exceedingly fine, ob- 
lique striae. Protoconch with two very finely lirate whorls. 

Remarks: S. pilsbryi and S. vanattai, both from the Sierra de 
Cubitas are very different from one another. S. pilsbryi differs 
from vanattai by being much larger, having the broad axial lamella, 
coarse sutural crenuiations and very much finer sculpture. The 
conic taper toward the protoconch starts from near the center 
(ninth whorl) in pilsbryi, while in vanattai the entire shell from 
the body whorl tapers toward the protoconch. 

This must represent quite a rare species, as only a very few 
specimens are known. 

Specimens examined: Holotype and paratype. 



1967 THE GENUS SPIROCERAMUS 5 

Spiroceramus (Spiroceramus) vanattai new species 
Plate 1 , figure 4 

Holotype: MCZ 256083, from Los Cangilones, Sierra de Cubitas, 
Camaguey Province, Cuba. 

Paratypes: From the same locality (MCZ 131387; MCZ 
80765), and from El Cercado de Cubitas, near Cueva del Circuto 
(MCZ 131388; MCZ 188846); Finca Santa Gertrudis, Minas 
(MCZ 80766); Corrales de Cangilones (MCZ 131390); Paso de 
los Burros (MCZ 131386); Paso Paredones (MCZ 131391); El 
Tuabagiiey, near the Cueva del Indio (MCZ 188844), all Sierra 
de Cubitas. Camaguey Province, Cuba. 

Measurements ^ 



-leight 


Width 




mm 


mm 




9.4 
10 
8.6 
9 


4.6 
4.5 
4.5 

4.2 


Holotype 
Paratype 
Paratype 
Paratype 



Description: Shell reaching 10 mm in height, minutely umbili- 
cate, conic, tapering from the body whorl to the protoconch and 
sculptured. Whorls 10 to 11 and moderately convex. Color 
light horn with irregular patches of white which cover mainly the 
oblique striae. Spire extended and forming an angle of 30°. 
Aperture subcircular, the outer lip slightly reflexed. Inner or 
parietal lip reflected over the minute umbilicus. Axis twisted but 
lacking a lamella. Columella short and slightly angled. Suture 
moderately indented and finely crenulate. Sculpture consisting 
of numerous and prominent oblique striae. Protoconch with 2 
whorls and smooth. 

Remarks: See Remarks under S. pilsbryi. Named for E. G. 
Vanatta who had been associated with H. A. Pilsbry at the Acad- 
emy of Natural Sciences, Philadelphia. 

Specimens examined: Holotype and paratypes. 

Subgenus Insulaceramus new subgenus 

Species with the axis twisted but not encircled with a spiral 
lamella. Early non-protoconch whorls rather coarsely lirate, later 

^ All specimens measured are from the type locality. 



6 BREVIORA No. 260 

whorls finely Urate. Suture noncrenulate. 

So far this subgenus is limited to the Bahama Islands. Speci- 
mens are exceedingly rare, as only three are known to represent 
the two species. 

Type species: Microceramus {Spiroceramus) greenwayi Clench. 

Spiroceramus (Insulaceramus) robertsoni (Clench) 
Plate 1, figure 2 

Microceramus (Spiroceramus) roliertsoni Clench, 1963, Bull. Mus. 
Comp. Zool. 128: 406. pi. 3, fig. 1 (Delectable Bay Settlement. Acklins 
Island, Bahama Islands). [Holotype, MCZ 225313.] 





Measurements 


Height 


Width 


mm 


mm 



6.7 1.0 Holotype 

Description: Shell reaching 6.7 mm in height, thin, imperforate, 
dull and sculptured. Color probably gray, whorls 1 1 and convex. 
First 5 to 6 whorls forming the conic portion of the spire. Aper- 
ture subcircular. Lip simple. Columella nearly straight. Axis 
simple and not twisted. Suture indented. Sculpture consisting of 
numerous, fine, oblique axial riblets. These riblets are somewhat 
stronger on the first 5 to 6 whorls. 

Remarks: This species is closely related to S. greenwayi Clench 
from Fox Town, Little Abaco Island, Bahamas. It differs by being 
a little more coarsely sculptured and having the protoconch a 
litde smaller. 

Spiroceramus (Insulaceramus) greenwayi (Clench) 
Plate 1, figure 1 

Microceramus (Spiroceramus) greenwayi Clench, 1938, Mem. Soc. Cu- 
bana Hist. Nat., 12: 329, pi. 25, fig. 1 (Fox Town, Little Abaco Island, 
Bahama Islands). [Holotype, MCZ 116688.] 

Measurements 

Height Width 
mm mm 

6.5 1.9 Holotype 

Description: Shell thin, small and imperforate. Color a dull 
isabelline (true color, however, cannot be given as the single 



1967 THE GENUS SPIROCERAMUS 7 

Specimen was found dead). Whorls 11, early 6 whorls which 
form the cone strongly convex; later whorls much less so. Aper- 
ture subcircular, lip simple and slightly flaring. Columella con- 
cavely arched, not distinct but forming the inner margin of the 
aperture. Axis simple with only a very slight twist. Sculpture: 
nuclear whorl faintly costate; next five strongly costate with 
oblique somewhat sinuous riblets, generally whitish; remaining 
whorls finely costate. Suture deeply indented but not crenulate. 
Aperture cast at an angle of 40° from a base line. 

Remarks: See under S. robertsoni (Clench). 

Specimens examined: Holotype. 

REFERENCES 

Arango, Rafael 

1878. Contribucion a la Fauna Malacologica Cubana, Habana, pp. 

1-315. 

PiLSBRY, H. A. 

1904. Manual of Conchology. (2) 16: 1-329. 
(Received 2 August, 1966.) 



BREVIORA No. 260 



Plate 1 

Fig. 1. Spiroceramus (Insulaceramus) greenwayi (Clench), Fox Town, 
Little Abaco Island, Bahama Islands (18.4 X). Holotype, MCZ 116688. 

Fig. 2. Spiroceramus (Insulaceramus) robertsoni (Clench), Delectable Bay, 
Acklins Island, Bahama Islands (10 X). Holotype, MCZ 225313. 

Fig. 3. Spiroceramus {Spiroceramus) pilsbryi n. sp., Cerro de Tuabagiiey, 
Sierra de Cubitas, Camaguey, Cuba (5.3 X). Holotype, MCZ 188845. 

Fig. 4. Spiroceramus (Spiroceramus) vanattai n. sp., Los Cangilones, Sierra 
de Cubitas, Camaguey, Cuba (5.3 X). Holotype, MCZ 256083. 

Fig. 5. Spiroceramus (Spiroceramus) barbouri Aguayo, Curva de la Com- 
pana, Gibara, Oriente, Cuba (5.3 X). Holotype, MCZ Til%ll . 

Figs. 6-7. Spiroceramus (Spiroceramus) amplus (Pfeiffer), Guisa, Bayamo, 
Oriente, Cuba (5.3 X). Syntypes, MCZ 39318. Figure 7 shows the 
axis and the spiral lamella. 



1967 



THE GENUS SPIROCERAMUS 




10 



BREVIORA 



No. 260 




'=^Z> 



Plate 2 

Map showing distribution of the species. 1. Spiroceramus (Spiroceramiis) 
ampins (Pfeiffer), Guisa, Bayamo, and Sao Arriba, Holguin, both Oriente, 
Cuba. 2. Spiroceramus (Spiroceramus) barbouri Aguayo, near Curva de 
la Campana, Gibara, Oriente, Cuba. 3. Spiroceramus {Spiroceramus) 
pilsbryi n. sp., Cerro de Tuabagiiey, Sierra de Cubitas, Camagliey, Cuba. 
4. Spiroceramus (Spiroceramus) vanattai n. sp., Los Cangilones, Sierra de 
Cubitas, Camaguey, Cuba. 5. Spiroceramus (Insulaceromus) robertsoiii 
(Clench), Delectable Bay, Acklins Island, Bahama Islands. 6. Spiroceramus 
(Iiisulaceramus) greenwayi (Clench), Fox Town, Little Abaco Island, 
Bahama Islands. 



BREVIORA 

MeseiuiitM of Compsirative Zoology 

Cambridge, Mass. 31 March, 1967 Number 261 

THE MONTICOLA GROUP OF THE LIZARD GENUS 
ANOUS IN HISPANIOLA 

By 

Richard Thomas and Albert Schwartz^ 



In 1962, Williams summarized the then available data on three 
allied species of Hispaniolan anoles: A. monticola Shreve, A. chris- 
tophei Williams and A . etheridgei Williams { = A. darlingtoni Coch- 
ran). Of the three species, monticola was named on the basis of 
a single male from "the northern and eastern foothills, Massif de 
la Hotte, 1000-4000 feet, Haiti." Additional specimens of this 
species were reported by Williams from the general region about 
and to the north of Camp Perrin and from He Grande Cayemite 
off the north coast of the Tiburon Peninsula of southwestern Haiti. 
The third species (described as A. darlingtoni by Cochran, but 
later changed to A. etheridgei by Williams [1962], for nomencla- 
tural reasons) was named from seven specimens collected by P. 
J. Darlington at four localities in the Cordillera Central of the 
Republica Dominicana. Finally, A. christophei from the Citadelle 
Laferriere in northern Haiti was based upon two females, both of 
which had been rather long in preservative. Williams (1962) 
reported on the coloration and pattern in life of A. monticola, and 
described the males of A. christophei. The latter species was still 
known only from the vicinity of the type locality. As for A. ether- 
idgei, the coloration and pattern of this species were still unknown, 
and its range remained as delimited by the original holotype and 
paratypes. 

The three species in the monticola group share a community 
of scale characteristics; those that distinguish them from other 
Hispaniolan anoles are the ventrals in transverse rows plus the 



1 10,000 SW 84th St., Miami, Florida 33143. 



2 BREVIORA No. 261 

subocular scales separated from the supralabials by a row of inter- 
vening scales (Williams, 1962:6). There is also agreement in 
other and more minor details of scalation, but the limits given by 
Williams (1962) for these data have been somewhat expanded 
by our examination of far more material than was available to him 
at the time of that paper. 

Perhaps the most significant observation on the monticola 
group is that of Williams (1962:7), who stated that, "Whether 
there are any more members of this small sub-group of Hispanio- 
lan anoles will have to be determined by more thorough search 
of the island ... If related species are equally local in distribu- 
tion, they may well have been missed." In the spring of 1966, 
Thomas secured a series of still another member of the monticola 
group in north-central Haiti, rather close to the type locality of 
A. christopliei. Since we now have large series of the members of 
this group and much additional information on variation, distribu- 
tion, and natural history, it is appropriate to summarize all of our 
data in the present paper and to describe the new northern Haitian 
species. 

We are very grateful to Miss Patricia A. Heinlein. Donald W. 
Buden, Ronald F. Klinikowski and David C. Leber for assistance 
in the field in the Republica Dominicana. It also gives us very 
great pleasure to acknowledge Mr. Leber's work on our behalf in 
the careful execution of the color portraits reproduced herein; 
with three exceptions (the races of A. monticola and the new 
species) these were done in the field from living lizards. We most 
readily acknowledge the assistance of Ernest E. Williams in the 
present endeavor, not only for suggesting that we name the new 
species and for the loan of recently accumulated material in the 
Museum of Comparative Zoology of a group in which he is per- 
sonally interested but also for his guiding our way through the 
at times impenetrable mass of Hispaniolan anoles. We have fol- 
lowed his techniques in taking of data and in description, so that 
the present information may be more readily compared with his 
own descriptions of several Hispaniolan anoles. Finally, Dr. 
Williams has graciously allowed that portions of a projected paper 
by himself and Schwartz be withdrawn and utilized herein, so 
that information on all the members of the monticola group might 
be more compactly available. 

Our data are based primarily on specimens in the Albert 
Schwartz Field Series (ASFS), but we have also borrowed ma- 
terial from the Museum of Comparative Zoology (MCZ), the Field 
Museum of Natural History (FMNH), and the American Museum 



1967 



ANOLIS MONTICOLA GROUP 



of Natural History (AMNH); for the courtesy extended us in 
these matters, we wish to thank Ernest E. Williams, Robert W. 
Inger, Hymen Marx, Charles M. Bogert, and Richard G. Zweifel. 
Dr. Zweifel's assistance in supplying copies of pertinent field 
notes is also much appreciated. William G. Hassler kindly allowed 
us to examine his photographs and field sketches of some of the 
anoles concerned. 

This study has been supported in part by National Science 
Foundation grant GB 2444 to Dr. Ernest Williams. 



Anolis christophei Williams 

The apparently most primitive member of the christophei- 
etheridgei-monticola trio of Hispaniolan anoles, A. christophei, was 
described from two female specimens from the Citadelle Laferriere, 
Dept. du Nord, Haiti (Williams, 1960). Later, this description 
was amplified by a series of 20 additional specimens from the 
type locality (Williams, 1962). The species has thus been known 
only from topotypical specimens and its distribution has otherwise 
been unknown. In March 1963, four of these lizards were taken 
at Paraje La Palma, La Vega Province, Republica Dominicana, 
by C. E. Ray and R. Allen; the species was thus known to inhabit 
the eastern Cordillera Central as well as the Bonnet-a-rEveque in 




Figure 1. Dorsal view of head of Anolis christophei, ASFS X9193. 
Snout-vent length 45 mm. 



4 BREVIORA No. 261 

the Chaine de Marmelade in the north. The elevation of the type 
locality is 2840 feet (865 meters); the elevation of the La Palma 
specimens is unknown. The coloration and pattern of christophei 
have not been described from living specimens; additional material 
collected by ourselves and associates in 1963, 1964 and 1966, and 
in 1964 by J. D. Lazell, Jr. has contributed information not only 
on these characters but also on the ecology and habits of the 
species. 

A. christophei is now known to occur in four regions, and 
doubtless is distributed in the intervening areas, with one prob- 
able exception. It is a fairly common lizard in the Cordillera 
Central in the Republica Dominicana. Specimens have been col- 
lected from the northeast of Jarabacoa, south to west of Con- 
stanza, but not in the Valle de Constanza itself. Not only does the 
species occur in the uplands, but it is also to be found on the 
steep eastern escarpment of the Cordillera between El Rio and 
Jayaco. Elevations of occurrence in this general area vary be- 
tween about 2000 feet (610 meters) and 4250 feet (1296 meters) 
(Fig. 14). A second major area of occurrence is in San Cristobal 
Province, near El Cacao. This locality lies on the southern slopes 
of the Cordillera Central; the lizards were taken at elevations of 
1200 to 1400 feet (366 to 427 meters). The third area of occur- 
rence is in the Cordillera Septentrional; at only one locality in 
this range, A. christophei was collected at an elevation of 2200 
feet (671 meters). The final region whence A. christophei has been 
taken is the Chaine de Marmelade in Haiti, at elevations of 2840 
feet (865 meters) to 3500 feet (1068 meters). The altitudinal 
distribution of the species is known to encompass elevations of 
from 1200 to 4250 feet (366 to 1296 meters) and its geographi- 
cal distribution includes the Massif du Nord (presumably) in 
northern Haiti, the Cordillera Central in central and southern 
Republica Dominicana, and the Cordillera Septentrional in the 
northern portion of the latter country. The first three areas are 
presumably confluent, whereas the lizards in the Septentrional are 
apparently completely cut off from their southern and western rel- 
atives by the arid portion of the Valle de Cibao. It is barely pos- 
sible that A. christophei occurs to the east in the Republica Dom- 
inicana and thus circumvents the inhospitable part of the Valle de 
Cibao, thereby bridging the apparent gap between the Cordillera 
and Septentrional populations. Certainly the valley of the Rio 
Yuna is sufficiently mesic, and even now reasonably well forested, 
to support A. christophei. However, we consider it unlikely that 
these wet lowlands are presently inhabited by such a confirmedly 
montane lizard. 



1967 



ANOLIS MONTICOLA GROUP 



A. christophei, although observed with some frequency during 
the day, is much more readily collected at night. Most specimens 
were taken sleeping on ferns (especially tree-ferns), herbs, and 
shrubs along the margins of streams in gallery forest in the pine- 
clad uplands of the Cordillera Central. The greenish brown dorsal 
coloration renders them relatively inconspicuous under such cir- 
cumstances, but the long dark almost filamentous tail often re- 
veals their presence. The hindlimbs are flexed in the sleeping 
posture, and the lizards sleep more often across the leaflets and/or 
leaves rather than aligning themselves along the stems of branch- 
lets; often they give the appearance of having been suddenly 
overtaken by the lowering temperatures of nightfall and of having 
stopped abruptly in their tracks. They give the impression of 
being draped across the greenery rather than having deliberately 
chosen a resting place for the night. This posture is in direct 




Figure 2. Lateral view of head of Anolis christophei, ASFS X9193. 
Snout-vent lencth 45 mm. 



contrast to that of such larger anoles as A. cy botes, A. distichus, 
and A . ricordi which are encountered with them, but is quite simi- 
lar to that of A. etheridgei which occupies similar situations at 
night. 

The habitat of this anole is moist, shady, montane woods, par- 
ticularly stream-associated forests and mesic ravines. During the 
day A. christophei was observed on tree trunks (four to six feet 
above the ground) along wooded edges of the Rio Jimenoa below 
Paso Bajito, and on tree trunks and fallen but sloping logs near 
Paso Bajito itself. One was found on a fence post in riparian 



6 BREVIORA No. 261 

woods near El Rio. The Haitian specimens were collected in 
dense second-growth woods, where they were found on small 
trees, two to five feet above the ground. At night along a very 
narrow rivulet adjacent to a platanal east of El Rio, A. christophei 
occupied the usual stream-side plants, and was not encountered 
on larger shrubs and trees nor on the banana plants themselves. 
The specimens from near El Cacao in the southern Cordillera 
Central were collected at night as they slept on bushes and ferns 
along the roadside in a region of coffee and cacao plantations. In 
the Cordillera Septentrional, specimens were also taken at night 
from stream-side shrubs and vines (the only place where they 
were encountered on the latter type of plants); on the vines, the 
lizards slept on the leaves and not on the woody stems. In no 
instance was there any attempt at concealment. 

Perhaps the most noteworthy characteristic of A. christophei 
is the extremely large dewlap for so small and relatively slender 
a lizard (PI. 1. upper left). The dewlap is pale, grayish purple (PI. 
47E2, Maerz and Paul, 1950, for Haitian specimens) or violet 
in life, with widely separated rows of pale yellow to whitish, or 
grayish, or bluish scales. The dewlap skin usually has a strangely 
metallic lustre which is distinctive. The dorsum is greenish brown 
to yellow-brown, with the heads distinctly greener than the back. 
There are four pairs of darker brown, bronzy or distinctly green- 
ish brown paramedian blotches which often are fused across the 
midline of the back to give four butterfly-shaped figures between 
the shoulders and the sacrum. The spaces between blotches are 
not clear but are variously marbled or marked with darker green 
or brown, so that the whole elTect is extremely cryptic. There is 
a yellow-green to yellow-buff shoulder stripe which continues 
down the sides as a vague, paler, lateral stripe. There is a yellow 
to greenish yellow subocular semicircle and a black to very dark 
brown postocular smudge. The chin and throat are dark gray; 
the ventral color is yellow to greenish yellow, and the tail is ter- 
minally black. In the dark phase, the dorsal ground color is dark 
gray with a purplish tinge, and the markings are very dark gray 
(nearly black) with a reddish or bronzy tinge. There is neither 
sexual dichromatism nor ontogenetic change in coloration or pat- 
tern; a single tiny juvenile (snout-vent length 21 mm) is colored 
and patterned like adults. The iris is blue. Although A. christophei 
is a small anole, it is not obviously slim and attenuate; the long 
tail adds to the impression of attenuation. 

A. christophei is not strongly sexually dimorphic in size, al- 
though males reach a slightly larger size (49 mm snout-vent 



1967 ANOLIS MONTICOLA GROUP 7 

length; ASFS X8835) than females (45 mm snout-vent length; 
ASFS VI 957). There are 6 to 10 (mode 8) scales across the 
snout at the level of the second canthal, and 4 to 8 (mode 6) rows 
of loreals. The supraorbital semicircles are separated by one or 
two scales (mode 1), and there are 3 to 7 (mode 4) scales be- 
tween the interparietal and the semicircles. Fourth toe lamellae on 
phalanges II and III vary from 18 to 25 (mode 22, with 16 speci- 
mens, but 15 specimens have 23 lamellar scales). 




Figure 3. Mental region of head of AnoUs christophei, ASFS X9193. 
Snout-vent lenizth 45 mm. 



Specimens examined: HAITI, Dept. du Nord, Citadelle Lafer- 
riere, 20 (MCZ 66900-19); Dept. de I'Artibonite, 8 to 9 km W 
Marmelade, 3500 feet (1068 m), 13 (ASFS V9900-12); REPUB- 
LICA DOMINICANA, La Vega Province, Municipio Constanza, 
Paraje La Palma (not mapped), 15 (MCZ 75142-43, 79349-51, 
79353-62); 6 km W Constanza, 4250 feet (1296 m), 2 (ASFS 
X8834-35); 11 km NE Jarabacoa, ca. 2000 feet (610 m), 6 
(ASFS V4198-201, V4326-27); 10 km NE Jarabacoa. 1 (ASFS 
V4216); 4 km E Paso Bajito, Rio Jimenoa, 2700 feet (724 m), 
10 (ASFS X8850-57, VI 866-67); 4 km SW El Rio, 4000 feet 
(1220 m), 5 (ASFS X8552-55, X8737); 4 km SW El Rio. 3500 
feet (1068 m), 3 (ASFS X8886-88); 6 km E El Rio, 3600 feet 
(1098 m), 7 (ASFS X9205-11); 7.1 mi. (11.4 km) E El Rio, 
3500 feet (1068 m), 1 (ASFS X81 13); 23 km E El Rio, 3050 
feet (930 m), 3 (ASFS X9 193-95); Espaillat Province, 2 km N 
Puesto Grande. 2200 feet (671 m), 6 (ASFS V1956-61); San 
Cristobal Province, 15.5 km SE El Cacao, 1400 feet (427 m), 
2 (ASFS V2498-99); 2.1 km SE El Cacao, 1200 feet (366 m), 
6 (ASFS V2492-96, V2502). 



BREVIORA 



No. 261 



Anolis etheridgei Williams 

Anolis etheridgei Williams (= Anolis darlingtoni Cochran, 
1939) was described from four localities in the Cordillera Central 
in the RepubUca Dominicana: Loma Vieja, south of Constanza; 
Loma Rucilla; Valle Nuevo; and Constanza. The included eleva- 
tions for these four localities are from 3000 feet (915 meters) to 
8000 feet (2440 meters). The type series comprised a total of 
seven specimens. From this it might be assumed that A. etheridgei 
is uncommon; this is far from the case since it is one of the more 
abundant anoles of these interior uplands, although its total dis- 
tribution is still somewhat circumscribed and surely incompletely 
known. No previous information on coloration and pattern in life 
has been reported. 

New specimens (three of which were taken by J. D. Lazell, Jr., 
in 1964, and the balance by ourselves and associates in 1963) 
indicate that the known distribution of A. etheridgei is confined 
to the interior highlands of the Cordillera Central in the Republica 
Dominicana, from Loma Rucilla on the west to east of El Rio on 
the east, from Paso Bajito on the north to southeast of Constanza 
on the south (Fig. 14). Although one of the paratypes is re- 
corded from Constanza (and this town is within the limits of the 
range of etheridgei as noted above), no additional specimens have 




Figure 4. Dorsal view of head of Anolis etheridgei, ASFS X9146. Snout- 
vent length 42 mm. 



1967 



ANOLIS MONTICOLA GROUP 



been taken from the Valle de Constanza, despite careful search 
for it there. Recently collected specimens indicate an altitudinal 
range of 3050 feet (930 m) to 6100 feet (1890 m); these 
elevations are embraced by the data for the type series, although 
in almost every case of the latter, the elevation for each locality 
is an inclusive range. 

A . etheridgei is similar to A . christophei in habitat preferences, 
and although it was encountered frequently sleeping on tree ferns, 
shrubs and herbs in gallery forest at night, it is not confirmedly re- 
stricted to this sort of situation. Specimens were collected in a pro- 
tected and secluded depression (about 7 meters in diameter) in 
rainforest north of Constanza; the depression was protected by a 
dense thicket along its margins, and the lizards were sleeping on 
bushes in the center of the depression. During the day occasional 
specimens were encountered in forest on bushes or small trees ad- 
jacent to paths, but many more were observed and collected with 
facihty while sleeping at night. The discussion of the sleeping 
habits of A. christophei applies equally well to A. etheridgei. 
Sleeping lizards are found draped over the leaflets of ferns and 
tree-ferns, the leaves of herbs and shrubs, and are not aligned 
along the branches or twigs. Their long tails reveal their pres- 
ence with ease. None was encountered sleeping on vines, and 
usually the lizards were seen within two feet of the ground sur- 
face. The legs are flexed in the sleeping posture. 




Figure 5. Lateral view of head of Anolis etheridgei, ASFS X9146. 
Snout-vent length 42 mm. 



10 BREVIORA No. 261 

A. etheridgei, in contrast to A. christophei, is sexually dichro- 
matic. Males are transversely crossbanded with four darker cross- 
bands between the neck and sacrum. These bands may be fairly 
prominent or much obscured due to interband pigmentation. The 
neck is brown to bronzy above, with bright pea-green on the sides 
of the head and neck, blending into a duller green on the sides. 
The back is some shade of tan to brown, always dull in hue. The 
sides of the head and neck may be pale powder-blue rather than 
green. The ventral color is tannish opalescent to opalescent. The 
general coloration of males is a combination of tans, browns, and 
greens of varying hues. The dewlap is small, all white or white 
with a grayish basal area (Pi. 1, upper right). Females have a 
dorsal longitudinal bronzy zone, bordered dorsolaterally by rich 
dark brown longitudinal bands which extend onto the postorbital 
area. The dorsal bronzy band expands on the head and forms a 
U-shaped bronzy nuchal figure which abuts against the upper eye- 
lids. Females at times have a series of four or five middorsal 
diamonds superimposed on the dorsal band; the halves of each 
diamond may be staggered to give a more complex dorsal pat- 
tern. The sides of the neck are white, and often females lack green 
or any greenish tints at all. The venters are like those of males 
except that occasional specimens have the venter light brown with 
a pinkish tinge. Both sexes have a cream subocular spot (which 
is more prominent in males) and the iris is blue. The chin and 
throat are very pale green in males and cream with brown striae 
in females. 




Figure 6. Mental region of head of Anolis etheridgei. ASFS X9146. 
Snout-vent length 42 mm. 



1967 ANOLIS MONTICOLA GROUP 11 

Of the seventy-eight lizards available to us, the two largest 
males and the largest female (all with snout-vent lengths of 43 
mm) come from a series of four lizards from 9.1 mi. (14.6 km) 
N of Constanza. Despite the long series from elsewhere in the 
Cordillera, strangely the three largest lizards are from the same 
small series; the difference in bulk of animals in this small lot of 
lizards, compared to those from elsewhere, was rather striking in 
life as well. 

There are from 8 to 15 (mode 10) scales across the snout, and 
there are 5 to 10 (mode 6) rows of loreals. The scales between 
the supraorbital semicircles vary from (semicircles in contact) 
to 4 (mode 2), and there are from 3 to 6 (mode 4) scales be- 
tween the interparietal scale and the semicircles. Fourth toe lamel- 
lae on phalanges II and III vary from 15 to 21 (mode 18). 

Specimens examined: REPUBLICA DOMINICANA. La Vega 
Province, Municipio Constanza, Paraje La Palma (not mapped), 
3 (MCZ 79345-47); 7.2 mi. (11.4 km) S Constanza, 5000 feet 
(1525 m), 1 (ASFS X8241);Loma Vieja, 6000 feet (1830 m), 
south of Constanza, 1 (FMNH 73378); 12.6 mi. (20.2 km) SE 
Constanza, 6100 feet (1891 m), 1 (ASFS X9146); 16 km N 
Constanza, 6000 feet (1830 m), 1 (ASFS X8950); 9.1 mi. 
(14.6 km) N Constanza, 3500 feet (1068 m), 4 (ASFS X8791- 
94); 4 km SW El Rio, 4000 feet (1220 m), 30 (ASFS X8522- 
51); 4 kmSWEl Rio, 3500 feet (1068 m), 3 (ASFS X8889-91 ); 
6 km E El Rio, 3600 feet (1098 m), 1 (ASFS X9212); 23 km 
E E! Rio, 3050 feet (930 m), 14 (ASFS X9179-92); 22 km 
NW Bonao. 3900 feet (1189 m), 11 (ASFS V4282-92); 23 
km NW Bonao, 4100 feet (1251 m), 1 (ASFS V4272); 19.6 km 
NE Bonao, 3300 feet (1007 m), 6 (ASFS V4296-301); 11 km 
E Paso Bajito, 4500 feet (1372 m), 1 (ASFS X8849). 

Anolis MONTICOLA Shreve 

In 1962 Williams reviewed the then available material of A. 
monticola and recorded its colors in life, based on notes taken by 
W. G. Hassler in 1935, and by A. S. Rand and J. D. Lazell in 
1960. He then indicated that, on the basis of very recent material 
not included in that study, there appeared to be differences be- 
tween populations of the northern and southern slopes of the 
Massif de la Hotte. Our own material, plus additional speci- 
mens (MCZ 74866-70), shows that there are indeed two forms 
of monticola: one possessing the four prominent dark ocellar 
patches as shown by Williams (1962: fig. 2), and one lacking 
the pair of nape patches. 



12 BREVIORA No. 261 

Williams (1962) cited five specimens collected by W. G. Hass- 
ler (AMNH 49818, 49845, 50108-09, MCZ 65139, formerly 
AMNH 50110) with the locality given as "25 miles north of 
Aux Cayes on Jeremie Road" and MCZ 56140 from "mountains 
on Jeremie road about 8 miles from Camp Perrin." Reference to 
the map of Haiti published by the Service de Geodesic et Cartog- 
raphic 1:100,000 shows that 25 miles north of Les Cayes lies 
well onto the northern slope, as does the locality 8 miles from 
Camp Perrin. We have consulted Hassler's original field notes 
and find that the true situation is otherwise: AMNH 49845 is in 
actuality listed as being from "high Mts. on Jeremie road 32 
miles from Aux Cayes 2000-3000' approx."; AMNH 49818, 
50108-10 are said to be from "about 4 miles from Camp Perrin." 
It is thus evident that these localities have suffered in the transcrip- 
tion to the catalogue. 

The specimens from 4 miles north of Camp Perrin are from 
the south slope, if the distance is accurate; the Tombeau Cheval 
specimens cited by Williams, 1962, are apparently from south of 
the high point on the Jeremie road but near it. The specimens of 
these two series agree in having the four ocellar patches. 

Recent material from farther west, obtained by Thomas, lacks 
the nape ocellar patches and agrees in this respect with MCZ 
74866-70 from Trou Bois, ly^ miles south of Beaumont on the 
Les Cayes-Jeremie road, and with AMNH 49845 from 32 miles 
from Les Cayes on the Jeremie Road (which is probably in the 
vicinity of the locality for the Trou Bois series). The type speci- 
men of monticola, although with the color pattern obscured by 
preservation and age, lacks the pair of nape patches (the neck 
patches and dorsal dark bands can still be seen), and thereby 
agrees with the material just mentioned. The locality for this 
specimen is not precisely known, but "northern and eastern foot- 
hills of the Massif de la Hotte" is in reality the northern and 
eastern foothills of Pic Macaya (P. J. Darlington, pers. comm. to 
E. E. Williams), the highest peak of the range, which lies to the 
west of the Tombeau Cheval region (Fig. 10). We are convinced 
that two distinct populations of Anolis monticola have been 
sampled: one, of the northwestern and western extreme of the 
Massif de la Hotte, and another farther to the east (and possibly 
to the south ) . 

We regard the Grande Cayemite record for monticola (MCZ 
58026, a female, collected by W. J. Eyerdam) as dubious. Thomas 
has visited this island and the habitat is for the most part very 
arid, not at all similar to habitats occupied elsewhere by monticola. 



1967 



ANOLIS MONTICOLA GROUP 



13 



This is not in itself incontrovertible evidence against the occurrence 
of the species on that island, and it may be noted that Diploglossus 
sepsoides Gray, normally an inhabitant of distinctly moist situa- 
tions, does occur there. However, the single Eyerdam specimen 
of Spluierodactylus copei Steindachner was noted as being indis- 
tinguishable from S. copei picturatus Carman from the nearby 
mainland (Schwartz and Thomas, 1964:326), yet the recently 
obtained series of S. copei from Grande Cayemite is undeniably 
a distinctive new subspecies. The Eyerdam collection is also re- 
sponsible for the aberrant locality of the Citadelle Laferriere for 
specimens of Anolis hendersoni Cochran (Williams, 1963), a 
trenchantly south island species. We will not be apodictic but 
regard the Grande Cayemite specimen with suspicion. 




Figure 7. Dorsal view of head of Anolis ?nonticola qiiadrisartiis, type, 
MCZ 62998. Snout-vent length 43 mm. 



Anolis monticola shows strong sexual dimorphism in size; males 
reach a maximum of 55 mm and females 39 mm. There are 7-11 
(mode 9) scales across the snout at the level of the second canthal 
and 6-9 (mode 7) loreal rows. The supraorbital semicircles are 
separated by 2-4 (mode 3) scales, and there are 3-6 (mode 4) 
scales between the interparietal and the semicircles. Fourth toe 
lamellae in phalanges II and III vary from 17-21 (mode 18 or 
19). 



14 BREVIORA No. 261 



Plate 1. Lateral views of heads of males of four species of Hispaniolan 
A no' is, as follows: tipper left, Anolis christophei, ASFS X8113, 7.1 mi. 
E El Rio, La Vega Prov., Republica Dominicana, painted from living 
specimen; upper right, Anolis etheridgei, ASFS X8522, 4 km SW El Rio, 
4000 feet. La Vega Prov., Republica Dominicana, painted from living 
spec'msn; center left, Anolis monticola monticola, ASFS V9624, ca. 5 km 
SSE Marche Leon, Dipt, du Sud, Haiti, painted from Thomas" field notes 
and color transparencies of living specimens; center right, Anolis monticola 
qiiadrisartiis, MCZ 63004, paratype, Tombeau Cheval, Dept. du Sud, Haiti, 
painted from color noted by collectors, as quoted by Williams (1962:3-4); 
lower, Anolis rimartim, AMNH 96469, paratype. 8 to 9 km W Marmelade, 
Dept. de TArtibonite, Haiti, 3500 feet, painted from Thomas' field notes 
on living specimens. 



1967 



ANOLIS MONTICOLA GROUP 



15 



Anolis MONTICOLA MONTICOLA Shrcve 

Diagnosis: A subspecies of Anolis monticola characterized by 
the absence of paired, large, black, light-centered ocelli on the 
nape, and by a yellow to reddish orange dewlap; females have a 
relatively straight-sided (in contrast to wavy or scalloped) mid- 
dorsal zone. 




Figure 8. Lateral view of head of Anolis monticola qiiadrisartiis, type, 
MCZ 62998. Snout-vent length 43 mm. 



Color in life: The dorsal ground color of males is dull greenish 
yellow middorsally; the sides are brighter yellow-green to yellow- 
brown. The labials and the sides of the neck have an indistinct 
pastel blue to blue-green stripe. The saddles and the nuchal spots 
are black, and the ocelli in the nuchal spots are bright blue. The 
venter is faintly greenish yellow. The throat is blue to blue-green, 
becoming yellowish centrally. The dewlap is yellow to reddish 
orange (Pi. 1, center left). The limbs are greenish yellow with 
some brown suffusion and banding, and the tips of the digits, and 
palmar and plantar surfaces, are black. The nominate form is also 
known to assume a gray to brown ground color phase dorsally, 
with black markings and little or no bright dorsal color. The 
ground color of the sides below the middorsal zone may also be 
pale translucent green. The dark dorsal saddles continue diag- 
onally and posteriorly below the middorsal zone and are much 



16 BREVIORA No. 261 

invaded by light color, to the extent of appearing split longitu- 
dinally and giving the effect of trailing off. The ocelli in the black 
patches may be single and relatively large or multiple and 
smaller. The color portrait was executed from this information. 

Females are duller than males and have a simpler zonate pattern 
of a light (brown), relatively straight-edged middorsal zone (usu- 
ally with only one or two small undulations), and darker sides. 

Habitat: 1) ca. 7 km (airline) WSW Moron. This locality is 
in the foothills of the limestone massif (Monts Cartaches) of 
the northwestern Tiburon Peninsula. Specimens were collected on 
a hillside and ravine side of limestone rocks and boulders with 
mesic vegetation of coffee, some bananas, and a high shade cover 
of breadfruit and other trees. Specimens were seen only in shady 
areas, mostly in coffee, where they were found on rocks and 
ground and sometimes on low (usually dead and fallen) branches 
and twigs. They were most abundantly found in an area (a small 
ravine) which had low herbaceous plants covering the ground, 
where they appeared to be foraging. (The time of the visit to this 
locality was between 0900 and 1000.) These anoles were very 
agile and retreated, almost invariably, into rock crevices when pur- 
sued. Many more were seen than were collected. 2) ca. 10 km 
WSW Moron, 1500 feet (456 meters). This was a steep hill slope 
to the west of the road, the opposite side of which was covered 
with a dense almost rain forest-like vegetation; specimens of 
monticola were seen around some rock outcroppings in the 
wooded area. Five were collected. 3) ca. 5 km (airline) SSE 
Marche Leon, 2600 feet (793 meters). Specimens were collected 
in a ravine filled with a jumble of limestone boulders and over- 
grown with mesic vegetation; the surrounding area was much cut 
over. 4) ca. 8 km (airline) S Marche Leon, 3000 feet (915 
meters) (Castillon). No specimens of monticola were collected 
but many individuals were seen along the base of a hillside covered 
with slabs and fragments of limestone; the area was relatively open 
and without heavy shade, although with some ground vegetation, 
in contrast to other localities where this species was seen. An ad- 
jacent hillside on the opposite side of a ravine and stream had 
no extensive outcroppings of rock, and nowhere on this hillside, 
even in a few places where vegetation was moderately thick, were 
examples of A. monticola seen. 

Specimens examined: HAITI, Dept. du Sud : ASFS V91 96-211, 
ca. 7.5 km (airline) WSW Moron; ASFS V9270-74, ca. 10 km 
(airline) WSW Moron, 1500 feet (456 m); ASFS V9624-26, ca. 5 



1967 ANOLIS MONTICOLA GROUP 17 

km (airline) SSE Marche Leon, 2600 feet (793 m); MCZ 74866- 
70, Trou Bois, about 1 V2 km S of Beaumont, on Jeremie road; 
MCZ 38296 (type), northern and eastern foothills. Massif de la 
Hotte (= Pic Macaya), 1000-4000 feet (305-1220 m); AMNH 
49845, high in mountains on Jeremie road 32 miles from Les 
Cayes, 2000-3000 feet (610-915 m) approx.; MCZ 65026, "Grande 
Cayemite." 




Figure 9. Mental region of head of Anolis monticola qiiadrisartiis, type, 
MCZ 62998. Snout-vent length 43 mm. 

Anolis monticola quadrisartus^ new subspecies 

Holotype: MCZ 62998, collected at Tombeau Cheval between 
Camp Perrin and Beaumont, Dept. du Sud, Haiti, by A. S. Rand 
and James D. Lazell, Jr., 7 August 1960. 

Paratypes: MCZ 62999, 63001-04, same data as type; AMNH 
50108-09, AMNH 49818, MCZ 65139, about 4 miles from Camp 
Perrin, Dept. du Sud, Haiti. 

Diagnosis: A subspecies of A. monticola characterized by a pair 
of black, light-centered ocelli on the nape in addition to the pair 
on the neck, and a blue to bright yellowish green dewlap (PI. 1, 
center right). Females have edges of middorsal zone undulating 
and scalloped. 

Coloration: Like that of the nominate race except for the diag- 
nostic characters noted above. See Williams (1962) for color 
notes from specimens in life on the type series of quadrisartus; 
these notes also served as the source for the color portrait. 

1 From the Latin, meaning "patched four times." 



BREVIORA 



No. 261 



Range: Known definitely only from the type locality (see dis- 
cussion under the species above); possibly it is a race of the east- 
ern La Hotte, or possibly of the southern slopes, although the 
present data do not indicate the latter (Fig. 10). 

Habitat: Apparently identical to that of the nominate race; see 
Williams (1962) for habitat notes on the type series. 




LES CAYES 



Figure 10. Tip of the Tihuron Peninsula, Haiti, showing distribution of 
the subspecies of A. monticola, as follows: circles indicate locality records 
for A. m. monticola; triangles, locality records for A. m. qiiadrisartits. Line 
connecting Les Cayes and Jeremie indicates road. Arrow indicates highest 
point (840 meters) along road. Hatched areas approximate regions above 
1000 meter line. Large rectangle indicates northern and eastern foothills, 
Pic Macaya. 



1967 



ANOLIS MONTICOLA GROUP 



19 



Anolis rimarum' new species 

Holotype: MCZ 81128, an adult male, one of a series collected 
8 to 9 km (airline) W of Marmelade, Dept. de TArtibonite, Haiti, 
at an elevation of 3500 ft. (1068 m), 2 April 1966, by Elie Cy- 
phale and Richard Thomas. Original number ASFS V9896. 

Paratypes: AMNH 96469-70, ASFS V9886-91, ASFS V9898, 
MCZ 8 11 29, USNM 157914-16, same data as type. 

Diagnosis: An anole of the Hispaniolan christophei-etheridgei- 
monticola assemblage distinguished by: smooth dorsal head scales; 
smooth, squarish supraoculars; a "window" of enlarged squarish 
palpebral scales; a nearly vestigial dewlap; smooth, juxtaposed 
ventrals arranged in transverse rows; transversely enlarged an- 
terior femoral scales; somber coloration (browns, yellow-browns, 
dull greens); and black digits. 




Figure 11. Dorsal view of head of Anolis rimarum, type, MCZ 81128. 
Snout-vent length 43 mm. 



Description of type (variations of paratypes in parentheses): 
Head moderate, not especially elongate, snout pointed, slightly 
concave. Head scales smooth, at most with feeble keels; 7 (7-10) 
scales across snout at level of second canthal. Frontal depression 



1 From the Latin, rima, a crevice. 



20 BREVIORA No. 261 

shallow. Anterior nasal scale in contact with rostral. Supraorbital 
semicircles narrowly in contact (also in six paratypes; separated in 
seven), separated from supraocular disks by one row of granules 
(rarely in narrow contact). Supraocular disks composed of about 
7 enlarged, smooth, or only very faintly keeled scales and sep- 
arated from supraciliary scales by 3 (3-4) rows of granules. Supra- 
ciliaries end at level slightly posterior to mid-eye, continued pos- 
teriorly by double row of slightly enlarged granules. Canthus 
rostralis distinct, sharp-edged, canthal scales 5-7. Loreal rows 5 
(4-6), lower row largest. Temporals granular, supratemporals only 
slightly enlarged and with small granules between them and en- 
larged scales around interparietal. Interparietal large, about size 
of ear opening, separated from supraorbital semicircles by 2 (2-3) 
scales. Suboculars in contact with supralabials. Five supralabials 
to center of eye. Lower eyelids with window of enlarged squarish 
scales. Mental broader than long, in contact posteriorly with 3 
(2-4) gular scales. Infralabials narrow, in contact with first en- 
larged sublabial; sublabials continue posteriorly as moderately 
enlarged row (or may blend completely with other scales of lateral 
gular region). Throat granules small, swollen, weakly keeled, 
slightly elongate anteriorly. 

Trunk: Dorsal scales very small, granular, two middorsal rows 
enlarged; adjacent rows reduce gradually to normal flank scale 
size. Ventrals smooth, enlarged, squarish, juxtaposed and ar- 
ranged in transverse rows. 

Dewlap: Very small, principally confined to area between fore- 
arm insertion and angle of jaw, scales smooth, swollen, rounded, 
about size of ventrals or slightly larger (PI. 1, lower). 

Limbs and digits: Scales of upper surface of limbs imbricate, 
keeled; those of hands and feet multicarinate; one scale row of 
prefemoral surfaces, especially distally and onto knee, much en- 
larged transversely and multicarinate. Subdigital lamellae 18 
(16-23). 

Tail: Compressed, 4 middorsal scales per verticil. One pair of 
postanal scales well developed in males. 

Size: Holotype, a male, 43 mm snout-vent (largest male 45 
mm; largest female 40 mm). 

Color in life: The dorsal ground color is gray-brown with a 
pattern of four gray to black middorsal butterfly markings or wide 
transverse bands that fade out on the lower sides; a coppery 
tinge is present on the middorsal line. The head is gray-brown dor- 
sally with olive-green temporal stripes that meet on the occiput; 
another more ventral olive-green postocular stripe proceeds onto 



1967 



ANOLIS MONTICOLA GROUP 



21 



the neck and fades out. The ground color of the lower sides of 
the head, neck and flanks is pale yellow-green with olive-green 
markings, including dark edges to the longitudinal flank stripes 
and other small dashes and vermiculations which may be present. 
The chin is whitish, as is the retracted dewlap (due to crowding 
of scales); the venter is pale metallic yellow-green. The dewlap 
skin is dull greenish orange (about pi. 13 L7, Maerz and Paul, 
1950). The tail is dull yellow to orange, sometimes greenish on 
its basal half to two-thirds, and black distally. The digits of both 
hands and feet are black. The iris is pale blue and the lower eyelid 
is blue. Sexual dichromatism is not pronounced; the middorsal 
light stripe of females is broader and more conspicuous than that 
of males, and the lateral stripe is also more prominent. These color 
data were employed in making the portrait of A. rimarum. 




Figure 12. Lateral view of head of Anolis rimarum, type, MCZ 81128. 
Snout-vent length 43 mm. 



Habits and habitat: The type series of A. rimarum was col- 
lected in a steep, limestone boulder jumble, a talus formation near 
the crest of the Chaine de Marmelade. The area (only a few acres 
in extent) was covered with a dense, natural, second-growth 
woods, mostly of small trees, brush and viny tangles but also with 
some moderately large trees. The specimens were found on rocks, 
low twigs and branches, and occasionally near the ground on 
the trunks or exposed roots of trees. They were seen most abun- 
dantly in areas where there was little thick undergrowth but 



22 BREVIORA No. 261 

where there was considerable shade from taller plants. When ap- 
proached, the lizards retreated with agility into rock crevices; many 
more were seen than were collected. The situation and habits 
of these anoles were virtually identical to those of A. monticola. 
At precisely the same locality, A. christophei was also taken; 
specimens were found on the trunks of small trees about 2 to 5 feet 
above the ground. Less than half a kilometer to the west, A. chris- 
tophei was taken from trees along the banks of a small ravine; the 
area was not rocky, and no examples of A. rimarum were seen. 




Figure 13. Mental region of head of Anolis rimariiin. type, MCZ 81128. 
Snout-vent length 43 mm. 



Comparisons: See Table 1 for a comparison of the diagnostic 
morphological characters of rimarum, etheridgei, monticola and 
christophei. Of these four anoles, christophei is the most diver- 
gent and, if truly a member of this group, is possibly the most 
primitive (Williams, 1962:7). Among the other three (rimarum, 
etheridgei, and monticola), there is nothing in the way of clear- 
cut alliances; any two share certain characters, and each species 
has certain peculiarities of its own. Rimaru/n seems to have more 
scale characters special to itself than etheridgei or monticola (Table 
1:6, 8, 11, 12, 13, 14). All four species show certain similarities 
in body coloration, the basic pattern being that of dark crossbands 
or butterfly markings on a brown or gray ground color. A . rimarum 
may have either solid crossbars or butterfly markings; christophei 
has butterfly markings only. Christophei and rimarum are also 
similar in lacking strong sexual dichromatism (rimarum is slightly 
dichromatic; christophei is not); monticola and etheridgei are both 



1967 ANOLIS MONTICOLA GROUP 23 

Strongly dichromatic and dimorphic in size. In dewlap color, al- 
though not in dewlap size, rimarum and monticola are similar, both 
having some shade of orange as part of the variation; etheridgei has 
a whitish dewlap. The dewlap of christophei, which is altogether 
different from those of the others in size and scalation, has a purplish 
color. Rimarum and monticola are additionally similar in coloration 
in having black fingers and toes. Monticola is the most distinctive 
chromatically of the lot in having brighter colors and large, bold 
ocellar patches on the neck; etheridgei is next to monticola in 
brightness of coloration. The two agree further in the type of 
body banding: solid middorsally but trailing off into hollowed and 
punctulate bands on the sides. 

In habits, however, monticola and rimarum appear identical in 
being inhabitants of boulder jumbles overgrown with low vegeta- 
tion, where the hzards seek refuge in crevices. In contrast, the 
other two are tree (christophei) or tree and bush (etheridgei) an- 
oles, also of forested areas. 

Geographically, monticola is quite isolated from the other mem- 
bers of this group. The remaining three species are all inhabitants 
of the north island, etheridgei and rimarum being allopatric as 
far as is known, whereas the distribution of christophei encom- 
passes the ranges of both. 

In conclusion, we cannot offer a reasonable certain evolutionary 
analysis of this radiation of small Hispaniolan anoles. We follow 
Williams (1962) in regarding christophei, etheridgei, and monti- 
cola as members of a single assemblage; christophei is the most 
aberrant but agrees in general with the other three in habitus, 
dorsal coloration, blue iris color, and ventral squamation. We 
add rimarum to the group and note that it does little to bridge 
the gap between christophei and the others, although in some 
characters christophei and rimarum are slightly more similar to 
one another than to etheridgei and monticola (lack of sexual 
dichromatism, enlarged second canthal scale, generally smooth 
head scales.) 

Williams (1962:7) pointed out the inverse relationship between 
dewlap prominence and boldness of dorsal pattern in male 
monticola, and remarked on the possible significance of this 
in maintaining species recognition. We do not deny that there 
may be such causal connection between boldness of dorsal pat- 
tern and prominence of dewlap in some cases, but we note that 
rimarum has the smallest dewlap (albeit brightly colored) of 
this group of anoles and also a relatively dull pattern. A. eth- 
eridgei has a somewhat brighter coloration than rimarum and 



24 BREVIORA No. 261 

a larger but much less brightly colored dewlap. A. christophei 
has a dull dorsal coloration and a relatively dull dewlap (pur- 
plish) which is nonetheless distinctive because of its isolated rows 
of scales and its metallic hue. The presence of an incipient "win- 
dow" of flattened translucent scales in the lower eyelid of rimarum 
and, to a lesser extent, in monticola supports the "sunglasses" 
theory proposed by Williams and Hecht (1955) to explain simi- 
lar but more highly developed structures in two Cuban anoles {A. 
argenteolus and A. lucius). Both rimarum and monticola are in- 
habitants of shaded areas and, additionally, are associated with 
rocky crevices — conditions that may well be favorable to the 
development of a protective filter for the eyes. 

LITERATURE CITED 

Cochran, Doris M. 

1939. Diagnoses of three new lizards and a frog from the Dominican 
Republic. Proc. New England Zool. Club, 18: 1-8. 

Maerz, a., and M. Rhea Paul 

1950. A dictionary of color. New York, McGraw-Hill Book Co., 
pp. i-vii, 1-23, 137-208. 56 pis. 

Schwartz, Albert, and Richard Thomas 

1964. Subspeciation in Sphaerodactyhis copei. Quart. Jour. Florida 
Acad. Sci., 27(4): 316-332. 

Williams, Ernest E. 

1960. Notes on Hispaniolan herpetology. 1. Anolis christophei, new 
species, from the Citadel of King Christophe, Haiti. Breviora, 
Mus. Comp. Zool., No. 117: 1-7. 

1962. Notes on the herpetology of Hispaniola. 7. New material of 
two poorly known anoles: Anolis monticola Shreve and Anolis 
christophei V^illiams. Breviora, Mus. Comp. Zool., No. 164: 
1-11. 

1963. Notes on Hispaniolan herpetology. 8. The forms related to 
Anolis hendersoni Cochran. Breviora, Mus. Comp. Zool., No. 
186: 1-13. 

Williams, Ernest E., and Max K. Hecht 

1955. "Sunglasses" in two anoline lizards from Cuba. Science, 122 
(3172): 691-692. 

(Received 2 August, 1966.) 



1967 



ANOLIS MONTICOLA GROUP 



25 



Aiiolis cliristophei 

1. Snout most elongate, most con- 
cave. 

2. Canthus pronouncedly sharp- 
edged. 

3. Scales 6-10 (mode 8) across 
snout at level of second canthal. 



4. Loreal rows 4-8 (mode 6). 

5. Scales in frontal depression nu- 
merous, polygonal to rounded, 
smooth or slightly rugose. 

6. Supraoculars moderate to small 
in size, somewhat elongate, 
weakly keeled, ca. 9-11 in num- 
ber. 

7. Supraorbital semicircles keeled, 
separated by 1-2 scales (Fig. 
1). 

8. Lower eyelid covered with gran- 
ules, without "window" of 
squarish scales. 

9. Anterior canthal scales abruptly 
smaller. 

10. Ventrals rounded to squarish, 
smooth, slightly imbricate and 
arranged in transverse rows. 

11. Dewlap very large, extending 
from mid-gular region to mid- 
venter; scales at edge of dewlap 
smooth, imbricate, larger than 
ventrals. 

12. Anterior femoral scales keeled, 
imbricate, not tranversely en- 
larged. 

13. Interparietal small, ca. Vi or 
less size of ear opening. 

14. Scales between interparietal and 
supraorbital semicircles 3-7. 



AnoUs etheridgei 

1. Snout bluntest, not pronounc- 
edly concave. 

2. Canthus distinct but not prom- 
inently sharp-edged. 

3. Scales 8-15 (mode 10) across 
snout at level of second can- 
thai. 

4. Loreal rows 5-10 (mode 6). 

5. Scales in frontal depression nu- 
merous, polygonal, keeled. 



6. Supraoculars small to moder- 
ate, slightly elongate, keeled, ca. 
1 1 in number. 



7. Supraorbital semicircles keeled, 
separated by 0-4 scales (Fig. 
4). 

8. Lower eyelid covered with gran- 
ules, without "window" of 
squarish scales. 

9. Anterior canthal scales not ab- 
ruptly smaller., 

10. Ventrals rounded to squarish, 
subimbricate, smooth and in 
transverse rows. 

1 1 . Dewlap, small, extending from 
mid-gular region onto chest, 
scales keeled. 



12. Anterior femoral scales keeled, 
imbricate, only slightly enlarged 
distally. 

13. Interparietal small, less than Vj 
size of ear opening. 

14. Scales between interparietal and 
supraorbital semicircles 3-6. 



26 



BREVIORA 



No. 261 



Anolis monticola 

1. Snout elongate, somewhat con- 
cave. 

2. Canthus sharp-edged. 

3. Scales 7-11 (mode 9) across 
snout at level of second can- 
thai. 

4. Loreal rows 6-9 (mode 7). 

5. Scales in frontal depression nu- 
merous, polygonal, keeled. 

6. Supraoculars moderate in size, 
elongate, prominentaly keeled, 
ca. 9 in number. 

7. Supraorbital semicircles prom- 
inently keeled, separated by 2-4 
scales (Fig. 7). 

8. Lower eyelid with "window" of 
small flat granules. 

9. Anterior canthal scales not 
abruptly smaller. 

10. Ventrals acute to rounded, im- 
bricate, keeled, and in trans- 
verse rows but not so diagram- 
matically as in others. 

11. Dewlap small, extending from 
mid-gular region onto chest, 
scales keeled. 

12. Anterior femoral scales imbri- 
cate, multicarinate but not trans- 
versely enlarged. 

13. Interparietal usually small, ca. 
V2 size of ear opening but may 
be equal to ear opening in size. 

14. Scales between interparietal and 
supraorbital semicircles 3-6. 



Anolis rimariim 

1. Snout elongate, somewhat con- 
cave. 

2. Canthus sharp-edged. 

3. Scales 7-10 (mode 8) across 
snout at level of second can- 
thai. 

4. Loreal rows 4-6 (mode 5). 

5. Scales in frontal depression rel- 
tively few, polygonal, pavimen- 
tous. 

6. Supraoculars large, not elon- 
gate, smooth or only faintly 
keeled, ca. 5-7 in number. 

7. Supraorbital semicircles smooth 
or feebly keeled, in contact or 
separated by one scale only 
(Fig. 11). 

8. Lower eyelid with "window" of 
enlarged squarish scales. 

9. Anterior canthal scales abruptly 
smaller. 

10. Ventrals squarish, juxtaposed to 
subimbricate, smooth and in 
transverse rows. 

11. Dewlap very small, occupying 
only the region between the 
angle of the jaw and the chest, 
scales smooth, imbricate. 

12. Anterior femoral scales trans- 
versely enlarged, feebly multi- 
carinate. 

13. Interparietal large, about size 
of ear opening. 

14. Scales between interparietal and 
supraorbital semicircles 2-3. 



Table 1 . Comparison of the four species of the monticola group of 
Hispaniolan anoles. 



1967 



ANOLIS MONTICOLA GROUP 



27 




BREVIORA 

MmseiMii of Comparsitive Zoology 

Cambridge, Mass. 31 March, 1967 Number 262 

A PHYLOGENETIC SURVEY OF MOLLUSCAN SHELL 
MATRIX PROTEINS 

By 
Michael T. Ghiselin 

Systemafics-Ecology Program, Marine Biological Laboratory 
Woods Hole, Massachusetts 

Egon T. Degens and Derek W. Spencer 

The Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 

and 
Robert H. Parker 

Systematics-Ecology Program, Marine Biological Laboratory 
Woods Hole, Massachusetts 

INTRODUCTION 

Recent advances in biochemistry and cytology have vastly in- 
creased the availability of structures suited for phylogenetic re- 
search. Diverse and complicated chemical structures, such as cyto- 
chrome c (Margoliash and Smith, 1964) and chromosomes (Spen- 
cer, 1949), are amenable to a type of analysis based on formal 
properties analogous to that of traditional comparative anatomy. 
The growth of evolutionary biology has recently brought about 
a subtle, yet pervasive and fundamental, revolution in the aims 
and methods of systematics. The development of new techniques 
based on an understanding of the causes of evolution has so far 
been most pronounced in the study of species and speciation, but 
is being extended to higher levels (Cain, ed., 1959; Bock, 1959, 
1960). The present work is an attempt to continue these trends, em- 
ploying a new type of evidence made available by improved methods 

1 Contribution No. 91 from the Systematics-Ecology Program, Marine 
Biological Laboratory, Woods Hole, and Contribution No. 1834 from the 
Woods Hole Oceanographic Institution. 



2 BREVIORA No. 262 

of chemical analysis: the proteinaceous matrix which serves as a 
framework for the calcareous portion of the molluscan shell. The 
preliminary survey discussed here is a methodological experiment, 
evaluating the feasibility of new techniques (logical, chemical 
and statistical). It appears, from both the consistency of the 
results and reference to traditional comparative anatomy, that the 
approach used here is useful for classification. Until more data 
are collected, however, the detailed conclusions should be consid- 
ered tentative. 

The development of these techniques has depended upon an 
interdisciplinary approach which has allowed efficient use of the 
data that otherwise would have been impossible. Systematic meth- 
ods and results are stressed in this account. A fuller discussion of 
the chemical aspects of the study is being published elsewhere 
(Degens, et ai, 1966). A publication (Degens and Spencer, 
1966) giving raw data and details of analytical techniques and 
computer methods is available on request. 

ACKNOWLEDGMENTS 

Support for this research has been provided by a Ford Founda- 
tion grant, and grant GB 4509 from the National Science Foun- 
dation to the Systematics-Ecology Program, Marine Biological 
Laboratory, and grants from the National Aeronautics and Space 
Administration (NSR-22-014-001 ) and American Chemical So- 
ciety (PRF" 1943-A2) to the Woods Hole Oceanographic Insti- 
tution. Systematic methods were largely developed by the senior 
author during the tenure of a postdoctoral fellowship at the Mu- 
seum of Comparative Zoology, Harvard University, supported by 
National Science Foundation Grant NSF-G 18727. 

For providing specimens, we thank William Clench, Ruth D. 
Turner, A. Lee McAlester and Henning Lemche. 

Responsibility for content rests with the authors, although 
valuable suggestions were provided by John Allen, Melbourne R. 
Carriker, Vida C. Kenk, Howard Sanders and Ruth D. Turner. 

Drawings are by Ruth von Arx. 

ANATOMICAL AND CHEMICAL BACKGROUND 

The molluscan shell (cf. Wilbur, 1964; Glimcher, 1960) is 
produced by secretion of precursors from the epithelial tissue in 
specialized areas of the mantle, and may consist of several layers. 
The outer layer, or periostracum, is not calcified; it is thought to 
consist of quinone-tanned protein (Brown, 1952). The inner 



1967 MOLLUSCAN SHELL PROTEINS 3 

layers are calcareous, and include a proteinaceous matrix which 
serves as a nucleation site for specific cations and anions, and in 
which the mineral crystals are deposited. The crystalline structure 
(Bpggild, 1930) and protein configurations (Gregoire, 1961) are 
quite intricate, suggesting a highly organized system. In moUusks, 
the uncalcified matrix is laid down extracellularly; only after the 
matrix has been formed is inorganic material from the underlying 
tissue incorporated into the shell. 

We have been guided by the hypothesis that the shell matrix is 
derived from at least one structural protein which originally had 
some function other than that which it now serves. It seems rea- 
sonable that the evolution of a new function has occurred gradu- 
ally. With the change from a non-calcified to a calcified state, the 
protein should become increasingly reorganized, so as to facilitate 
the efi'ective binding and interconnection of the parts. In general, 
this should involve a proportionate increase in the amount of those 
organic components which facilitate the binding of important ions 
and a decrease in the quantity of superfluous organic materials. 
There should also be changes in the type of cross-linkages that 
interconnect the individual parts of the protein. These hypotheses 
may readily be verified by their implications for systematics; they 
predict that certain types of correlations will occur between taxo- 
nomic groupings, environmental and physiological relationships, 
and chemical composition. 

CHEMICAL METHODS 

The shell material was separated mechanically from all extrane- 
ous proteins and after being ground to a powder, decalcified, in 
the presence of 10';^ trichloracetic acid solution, by hydrochloric 
acid, added in quantities stoichiometrically necessary to dissolve the 
calcium carbonate. The organic remains were centrifuged, washed, 
and hydrolyzed with 6N hydrochloric acid for 22 hours in vacuo. 
The amino acids were analyzed by automatic ion-exchange chro- 
matography. The method gives reproducible results of better than 
l^c at the 10 '' molar level for identical runs. The overall varia- 
tion of two identical samples from the same collection site is less 
than 37c; this figure includes the variation between specimens, and 
the errors introduced by decalcification, chromatography and data- 
handling. (For details, see Degens and Spencer, 1966.) 

STATISTICAL METHODS 

For the comparison of a few forms, visual inspection of the 
measurements for each amino acid is adequate to place the various 



4 BREVIORA No. 262 

forms in sequences corresponding to hypothesized evolutionary 
lineages. With larger samples, however, the task of comparison is 
facilitated by statistical techniques using a computer. As certain 
amino acids tend to show a constant ratio relative to each other, 
they may be combined for the purpose of comparison. By means 
of a factor analysis, it was possible to reduce the data to a set of 
factor scores (8 in gastropods, 9 in bivalves), which explain 90^r 
of the variance in the data. The factor scores may then be com- 
pared independently, or in an 8 or 9-dimensional hyperspace. We 
found that separate factor analyses for gastropods and bivalves 
were desirable, as with increasing phylogenetic diversity the vari- 
ous amino acids show slightly different patterns in their tendencies 
to vary in correlation with others. Our attempt to combine all the 
data for the phylum did little more than increase the number of 
extraneous relationships. 

The statistical methods used here should not be confused with 
what has been called "numerical phenetics." Factor analysis, 
which has become the method of choice in psychology, but which 
has found little use in systematics, has several advantages over the 
clustering techniques employed, for example, by Sokal and Sneath 
(1963). A clustering technique tells nothing more than how "sim- 
ilar" things are, in terms of an arbitrary standard; what is called 
"overall similarity" is merely the summation of diverse properties 
which may be no more comparable than the shape and hardness 
of lightbulbs and pears. It is advantageous, when using any type of 
statistical technique, to work with a clearly formulated model. 
A factor analysis generates such a model, allowing the treatment of 
each individual factor as independent evidence, and its separate 
evaluation. It also enables one to formulate hypotheses and test 
them by reference to particular aspects of the data. 

At first we tried to relate the species by considering each species 
as a point within the 8 or 9-dimensional factor space. The factor 
scores are the coordinates of the points. Species that have similar 
scores will be separated by only a short distance, while those 
with greatly different scores will be well separated. Table 1 pre- 
sents the linear distances between each of the species in the factor 
space. This kind of comparison, although suggestive of relation- 
ships, was ambiguous and misleading. It only told how "similar" 
the material was, in terms of the new factors. On the basis of other 
evidence, it was obvious that certain clearly unrelated forms were 
grouped close together. This is readily understood, for the dif- 
ferent amino acid variations are not strictly commeasurable, and 



1967 MOLLUSCAN SHELL PROTEINS 5 

the values give only sequences. That is, the ordering is an inten- 
sive, not an extensive one (Ghiselin, 1966a). It is not surprising, 
therefore, that the summation of similarities grouped together: 1) 
unrelated forms which had changed little from the ancestral state; 
2) unrelated forms which had undergone the same change to an 
extreme degree; and 3) "advanced" forms which had attained 
the same grade through parallelism. 

Yet by taking the series of factor scores as independent evidence, 
and arranging the species in series which show changes for each 
factor in a particular direction, it was possible to arrange the 
various forms in series corresponding to divergent genealogical 
lineages. The forms previously misplaced then fell into more 
reasonable positions, and it could be demonstrated that the earlier 
misplacement had been due, say, to great alteration, in diverse 
lineages, of one particular factor. It was possible to verify these 
inferred relationships by means of comparative anatomical evi- 
dence. Our data are here presented (Tables 3 and 4) in the form 
of series showing progressive changes in factor scores. The series 
correspond to a system of inferred phylogenetic relationships, dia- 
grammed as a phylogenetic tree shown on the left, with the direc- 
tion of trends shown by arrows. The phylogenetic trees are strictly 
genealogical; distances mean nothing. The diagrammed arrange- 
ments are those which are most strongly supported by our amino 
acid evidence, and reasonably consistent with comparative anat- 
omy. As a number of alternative interpretations are possible, the 
reader may wish to evaluate them. This can easily be done by 
writing out the factor scores on paper, and cutting out strips giving 
the scores; these may then be placed in any desired sequence. 

SYSTEMATIC METHODS 

Although we have stressed supra-generic relationships, future 
work will probably depend on precise species identifications. 
Therefore we have taken the precaution of figuring (Plates 1, 2) 
representatives of most of our samples, and have drawn upon 
materials deposited in the Museum of Comparative Zoology. 

Future work may have to incorporate a larger and more rigor- 
ously selected sample than does this preliminary study, in order to 
account for variation. We have selected several specimens from 
widely separated localities and different ecological conditions, and 
find that there is considerable temperature-dependent variation 
(Degens et al, 1966). The particular type of variation that oc- 
curs, however, appears to be different for each species. Some 



6 BREVIORA No. 262 

evidence that convergences occur when there is a change from 
marine to freshwater or terrestrial habitats will be discussed 
below. The environmental effects can be considered "noise" in 
evaluating genealogical relationships. If we could remove some of 
the environmental effects, perhaps some of the anomalous rela- 
tionships could be resolved. The removal of these effects would 
require accurate environmental data for each species, selected 
for a wide range of conditions, and, preferably, experimental 
studies. 

To some extent, this study is based upon traditional comparative 
anatomical methods. The material studied consists of highly-struc- 
tured proteins, the parts of which have relational properties such 
as allow one to abstract a common type from which all may be 
derived. To be sure, we do not know the details of structure, but 
our techniques do allow us to infer that diiferent changes have 
taken place, and to correlate these with each other. However, our 
argument is not limited to such formal comparison, as our hypo- 
thetico-deductive system includes premises about chemistry and 
evolution which affect the results. 

The hypothetical biochemical explanations outlined above imply 
that there will be a gradual diminution in the quantity of carbo- 
hydrate and protein in correlation with evolutionary development. 
At the same time, there should be a general trend toward increase 
in chemical groupings useful for nucleation (e.g., aspartic acid) 
and for cross-linkages (e.g., phenolic compounds). Further, 
forms which retain a high amount of protein should be those with 
a structure which may readily serve as a precursor for more modi- 
fied forms. The primitive condition may therefore be inferred on 
the basis of physiological criteria, providing independent evi- 
dence for inferences based on divergent specialized types. We do 
not, however, presuppose an overall "primitiveness" for any par- 
ticular organism. Different structures may evolve at different rates, 
and the ancestral form of a group may be reconstructed with 
rigor only when the hypothesized properties of each system are 
supported by empirical evidence. This point is crucial in the pres- 
ent study, which shows that there is no strict correspondence be- 
tween the degree of evolutionary advance in the shell and in other 
structures. 

Perhaps the major difficulty for any phylogenetic study is paral- 
lelism, and the present study is no exception. Evolutionary changes 
may tend in the same direction, and occur repeatedly. When 
this happens, a grouping together of similar forms on the basis of 
such changes leads to a series of grades rather than clades. It is 



1967 MOLLUSCAN SHELL PROTEINS 7 

abundantly clear in bivalves, for example, that some of the struc- 
tural changes (in hinges, gills, stomachs, etc.) used as phylogen- 
etic evidence have evolved a number of times. Classifications 
erected on only one such line of evidence take it as a priori that 
some character can only have evolved once (e.g., Purchon, 1959). 
Such premises are not only without support, but are flatly contra- 
dicted by analogy with such polyphyletic structures as the mam- 
malian ear-bones. If a structure evolves once, it can evolve re- 
peatedly. The problem of parallelism has been de-emphasized by 
Remane (1956), and Sokal and Sneath (1963), on the grounds 
that sometimes there is divergence rather than parallelism. This 
misses the point; there is no way to estimate the probability for a 
structure to evolve in more than one direction. 

A solution to the problem of parallelism has been proposed by 
Bock (1959, 1960) and has found some application by one of us 
(Ghiselin, 1966b) in gastropod phylogeny. To understand the 
technique, one must realize that it entails a shift in emphasis from 
observed characters to a model of divergent genetic and anatomical 
potentialities. It is assumed that however many times a character 
develops, closely related lineages may evolve in different direc- 
tions; this follows from similarities and differences inherent in the 
genetic potentialities within each lineage. If the forms can be 
placed in at least two sequences corresponding to divergent ten- 
dencies (rather than parallel stages), then the arrangement sup- 
ports the view that each member of one divergent group is not 
related to any member of any other such group. From placing 
them in sequences which do not diverge, but proceed to only one 
derived state, no systematic inferences can be drawn, as any num- 
ber of changes would explain the facts; founding groups on lack 
of divergence in a derived trait is therefore the logical fallacy of 
denying the antecedent in a conditional statement. In other words, 
the development of a character is not evidence as such for rela- 
tionship to a form also having that character, but only that both of 
the two forms having it are not related, for the same reason, to a 
form having some different alternative. For example: Purchon 
(1958, 1959) has distinguished five types of stomachs in bivalves. 
Three of these occur in the "higher" bivalves studied here: types 
III, IV and V. Both III and V may be derived from IV, and 
the advanced types overcome certain functional inefficiencies in the 
ancestral form. If the bivalves are divided into groups, including 
one for forms with type V stomachs, and one for those with type 
III, reference to other kinds of evidence demonstrates that forms 
within each grouping are most closely related to forms with type 



8 BREVIORA No. 262 

IV. Therefore, the changes from IV to V, and from IV to III have 
occurred more than once. Nonetheless, a system of relationships 
which abstracts one group with type IV and only type V, and 
another with type IV and only type III, fits in quite well with other 
evidence (cf. Newell, 1965). The biochemical evidence given 
here likewise supports this division as indicative of relationships. 
A valid phylogenetic method, then, admits any number of uni- 
directional changes, as IV to III or IV to V; a contradiction arises 
only when the divergent, derived forms (III and V) both occur 
within more than one lineage. Such a shift in emphasis may seem 
minor, especially when the logic of conditional statements is over- 
looked, but it seems to resolve much of the confusion that has 
resulted through efforts to erect systems of classes differentiated 
by visible, intrinsic properties, rather than corresponding to the 
more fundamental genetic and evolutionary order. Classification, 
in other words, involves, not simply grouping organisms on the 
basis of resemblances, but sorting them out into groups which 
differ in their programmed genetic information, and therefore have 
divergent potentialities and tendencies, irrespective of whether 
these at any moment are realized or not. 

PHYLOGENETIC SURVEY 

Ancestral models, Amphineura, Cephalopoda, Monoplacophora. 
By analogy with other studies on protein evolution, one might as- 
sume that difterences in the matrix protein have resulted from re- 
placement of particular amino acids owing to corresponding chro- 
mosome mutations affecting the template. Such modifications may 
underlie some of the diversity, but we feel that the major reason 
is to be sought elsewhere. Chemical and electron microscope 
studies (for references cf. Degens et ai, 1966) indicate that the 
protein is heterogeneous. Solubility tests (Degens and Spencer. 
1966) suggest that the protein of Mercenaria resembles collagen in 
solubility and molecular weight, although our quantitative data 
show differences in amino acid ratios; for example, the molluscan 
shell matrix differs from its analogue in vertebrate bone, in lack- 
ing the hydroxyproline characteristic of collagen. 

Some amino sugar is present in both shell and mantle of mol- 
lusks. The amount of carbohydrate in the shell varies consider- 
ably throughout the phylum. The relative proportion of carbo- 
hydrate to protein seems to have decreased progressively with 
functional improvement in shell structure; simultaneously, the per- 
centage of protein has decreased, but at a different rate. These 



1967 MOLLUSCAN SHELL PROTEINS 9 

trends may be explained if one hypothesizes that the shell was 
produced from a protein-carbohydrate complex which has been 
modified so as to improve nucleation. Such a change has occurred 
in calcified arthropod integuments (Rudall, 1963; Carey et al., 
MS), which show a decrease in the proportion of protein and an 
increase in the amount of hexosamine with calcification. In gastro- 
pods and bivalves, on the other hand, amino sugars decrease 
greatly. The integument of arthropods and moUusks may be de- 
rived from a common ancestral precursor (presumably originat- 
ing at an annelidan stage of evolution) in which a non-calcified 
protein was linked to carbohydrate. In arthropods, the amino 
sugars have evidently been retained, while protein has been so 
altered as to expose the acidic and basic side-chains responsible, 
respectively, for concentrating calcium and carbonate ions. Mol- 
lusks differ in having lost much of the amino sugar, while elaborat- 
ing the protein. 

Our data suggest that the molluscan shell has been evolved by 
modification of proteins which occurred in the mantle. Reference 
to Table 2 shows the type of relationships and evolutionary changes 
that seem to characterize the phylum. Here comparisons are given 
of the amino acid ratios in both mantle and shell of a chiton 
(Cliaetopleura), the shells of several cephalopods, representative 
"primitive" gastropods and pelecypods, a monoplacophoran (Neo- 
pilina), and the calcified and non-calcified integuments of portunid 
crabs. The relationships shown by this table are, at first glance, 
ambiguous. It may be seen that certain amino acids are more (or 
less) abundant in the shell than in the mantle of Chaetopleura. 
Often, these particular amino acids occur in about the same con- 
centration, as in Chaetopleura, Spirilla and Sepia, whereas in 
Loligo, Nautilus and other mollusks, the range is distinctly dif- 
ferent (e.g., aspartic and glutamic acids, perhaps glycine and ala- 
nine, and cystine). This relationship is explicable in terms of a 
high degree of calcification, without loss of carbohydrate, in 
Chaetopleura, Spirula and Sepia. Perhaps Loligo somewhat re- 
sembles Nautilus because although it is rich in carbohydrate, it is 
not highly calcified. In a number of other amino acids, however, 
Loligo is (in correlation with the high ratio of carbohydrate to 
protein, but independent of calcification) close to Chaetopleura, 
Spirula and Sepia, and most distinct from Nautilus, Haliotis, Nu- 
cula and Mytilus (threonine, proline, perhaps glycine and alanine, 
valine, methionine, isoleucine, leucine, tyrosine, phenylalanine and 
perhaps hydroxylysine). Of these, only four (threonine, methio- 
nine, tyrosine and phenylalanine) would seem to correlate with 



10 BREVIORA No. 262 

the high level of carbohydrate in Chaetopleura, Spirilla, Sepia and 
Loligo. Any attempt to give phylogenetic interpretations to these 
relationships without weighting would be foredoomed to failure. 
Nautilus, which retains many primitive traits, more closely resem- 
bles members of other classes than its closest relatives. The only 
cephalopod much like it is Loligo. But Loligo is, in a way, inter- 
mediate between Nautilus and the other cephalopods in just those 
characters in which the latter resemble the Amphineura. There 
must have been convergences. 

The ambiguity may be resolved if one supposes that there 
have been three, largely parallel, stages in the evolution of the 
shell. First, there would be the development of a calcified, but 
carbohydrate-rich shell. Chaetopleura, Spirula and Sepia would 
represent this stage. Loligo would retain much of the character of 
the first stage, but have lost (or conceivably never have developed) 
some of the functional units related to calcification, while retaining 
others significant in carbohydrate physiology. Second, there would 
be a loss of carbohydrate. This would explain the diiTerence be- 
tween Nautilus and other cephalopods, and, as a convergence, the 
resemblances between Nautilus and Haliotis, etc. Third, there 
would be an elimination of that part of the protein which is 
superfluous as a structural element or in calcification, and the de- 
velopment of more efficient means of cross-linkage. These three 
stages would occur more or less in parallel fashion, perhaps si- 
multaneously, but a stepwise sequence would explain such evident 
gaps as that between Nautilus and other cephalopods. 

Within the Cephalopoda, the shell of Nautilus is the most ad- 
vanced in its chemical evolution, in sharp contrast to its many 
primitive anatomical features. The other cephalopods would ap- 
pear to have survived in spite of a poorly-developed shell, largely 
by converting it to an endoskeleton. The Chaetopleura shell is of 
equally low chemical grade; there is no reason to think that this 
has occurred by degeneration. Neopilina belongs to the class 
Monoplacophora, thought to be ancestral to many other classes 
of mollusks, but it does not show any close agreement in amino 
acid ratios to any other mollusk. However, as we were unable 
to separate all the periostracum from our sample, the data are hard 
to interpret. On the other hand, the proportions of carbohydrate, 
protein and mineral show that the shell of Neopilina has undergone 
considerable modification. 

It appears that the major, early steps in shell evolution (calci- 
fication, loss of carbohydrate) are parallel and unidirectional. There- 
fore, while of physiological interest, they are uninformative for 



1967 MOLLUSCAN SHELL PROTEINS 11 

cladal phylogeny. In later stages, however, many divergences oc- 
cur, and these are most useful in establishing genealogical rela- 
tionships. Such changes may be largely due to different kinds of 
cross-linkages. These are primarily covalent linkages, such as: 
1) various linkages to carbohydrate moieties; 2) disulfide connec- 
tions; 3) unusual amino acids (e.g., desmosine and isodesmosine); 
4) phenolic compounds and quinones (for details, see Degens 
et ai, 1966). The matrix seems to be secreted as soluble pre- 
cursors which, at about the time of secretion, are acted upon by 
enzymes and other substances which produce the cross-linkages. 
Differences in composition may result from variations in the rates 
and kinds of such processes of secretion and modification, as well 
as from differences in the raw materials. The subsequent discus- 
sions of gastropod and bivalve shell evolution trace out progressive 
changes in various groups; these changes evidently are related to 
different kinds of secretory processes and materials. 

Gastropoda. Table 2 shows the raw data for Haliotis, a form 
which approximates a hypothetical ancestral gastropod. Changes 
within the class are shown in Table 3. As would be expected 
from the usual classification of the order Archaeogastropoda, Hali- 
otis (superfamily Zeugobranchia) and Astraea (superfamily Tro- 
chacea) are quite similar. But Fissurella, another zeugobranch, 
is quite distinct from Haliotis; as it shows no clear relationship to 
any other group, the difference is probably due to divergence. The 
superfamily Patellacea, as shown by Acmaea, is likewise distinct 
and isolated. The representative of the other archaeogastropod 
superfamily, Neritacea (Nerita), resembles Viviparus, a form usu- 
ally placed in the order Mesogastropoda. Both Nerita and Vivi- 
parus are in some ways transitional between the Archaeogastro- 
poda and Mesogastropoda; the resemblance may be convergent, 
but there is no compelling reason to think that it is. 

The rest of the gastropods studied break down into three major 
groupings which agree fairly well with generally recognized natu- 
ral groups. The less-modified members of each group are quite 
similar, indicating a close relationship and common origin. 

The first of these groups (Table 3, Crepidula to Murex) in- 
cludes the order Neogastropoda as a distinct sub-group and most 
of the Mesogastropoda (excepting only some forms known to 
have opisthobranch affinities). The Neogastropoda (Colus, Nas- 
sarius, Urosalpinx and Murex) show distinctive tendencies in the 
modification of several factors, although the original pattern is not 
far removed from that of mesogastropods. This fits in with other 
biological data; the neogastropods are distinct but not greatly 



12 BREVIORA No. 262 

modified in terms of morphology, except for the more specialized 
members of the group. But neogastropods are distinct in chromo- 
some number (Nishikawa, 1962) and evidently in many physi- 
ological characters also. The superfamily Naticacea (here, Lunatia 
and Polinices) has a pattern of change clearly distinct from other 
mesogastropods, and proceeding in a direction opposite to that of, 
say, Littorina and Cypraea. One of the unexpected results of this 
study was that Melanella has the same tendencies as occur in the 
Naticacea. Melanella has been placed in the superfamily Aglossa, 
which may be an artificial assemblage of parasitic forms with tall 
shells. An affinity of Melanella to the Pyramidellidae (now 
grouped as opisthobranchs), as suggested by Thiele (1935), is 
not consistent with our findings; similarly, Melanella lacks the 
characteristic reproductive system and larval shell of opistho- 
branchs. 

Littorina, which, along with other members of the superfamily 
Littorinacea, is often thought to stand near the base of the meso- 
gastropod series, has a pattern of change which tends in the direc- 
tion characteristic of Cypraea and Crepidula, fitting in well with 
the concepts of Thiele (1935). An affinity between the Cypraea- 
cea {Cypraea) and Calyptraeacea {Crepidula) follows from in- 
termediate forms (Lamellariacea) (cf. Fretter and Graham, 
1962). The type of variation that occurs within a genus may be 
seen in Crepidula. The range is fairly narrow and in conformity 
with overall trends, but some of the factors appear to have changed 
more than others. Such variation clearly indicates that a larger 
sample will be necessary in future work, but the utility of the 
method stands. 

Another distinct grouping of mesogastropods consists of Tur- 
ritella (Cerithiacea) and Epitonium and Janthina (Ptenoglossa). 
Both superfamilies differ from all other gastropods in possessing a 
curious, spermatophore-like structure, the spermatozeugma. The 
Cerithiacea are of disputed taxonomic position, often looked upon 
as transitional between various major lineages of mesogastropods 
and opisthobranchs; our data suggest a closer affinity to opistho- 
branchs. However, Cerithiacea and Ptenoolossa also have much 
in common with Neogastropoda, and the basal members of each 
of our three main groupings {Architectonica, Turritella, Littorina, 
etc.) are quite similar. Therefore, the precise cladistic relationships 
of each group remain uncertain. 

The third group corresponds to the Euthyneura, or the sub- 
classes Pulmonata and Opisthobranchia. The Euthyneura are char- 
acterized by hermaphroditism, a tendency to loss of the effects of 



1967 MOLLUSCAN SHELL PROTEINS 13 

torsion, a distinctive type of spermatozoon, a peculiar ctenidium, 
and a heterostrophic larval shell (except where secondarily mod- 
ified). Fretter and Graham (1949) have placed the family Pyra- 
midellidae in this group. As Architectonica has a heterostrophic 
larval shell, Robertson (1963; also, Robertson and Merrill, 1963) 
and others have argued for its close affinity to the Euthyneura, and 
this placement is adopted here. 

Some workers have advocated abandonment of the subclasses 
Pulmonata and Opisthobranchia (cf. Taylor and Sohl, 1962). 
However, this has been done solely on the basis of highly specu- 
lative assertions that Pulmonata is a polyphyletic assemblage; there 
is no real evidence for this view (cf. Ghiselin, 1965, 1965b). 

In its low protein content, and in the amino-acid ratios, Archi- 
tectonica is an excellent precursor for the opisthobranchs and 
pulmonates, the members of each of which can readily be derived 
by modification of different factors. This fits in with the larval 
shell types. Chemically, Architectonica differs but slightly from 
such Cerithiacea as Turritella. Indeed, Thiele (1929) put Archi- 
tectonica in the Cerithiacea. On the basis of fossil evidence. 
Knight, Batten and Yochelson (1954) relate the superfamily Pte- 
noglossa to the Pyramidellidae and group both with the opistho- 
branchs. Clench and Turner (1951) likewise stress the opistho- 
branch affinities of Ptenoglossa. Thus the close relationships be- 
tween these groups, on the basis of other evidence, supports the 
inference from shell biochemistry. 

The general conclusions reached by one of us on the basis of 
various lines of evidence on the relationships of opisthobranchs 
(Ghiselin, 1966b) are fairly well borne out by the present study. 
Bulla shows a beginning in the development of trends character- 
istic of the Anaspidea (Akera, Aplysia); gizzard morphology and 
anatomy generally support this relationship. Both Akera and 
Oxynoe are distinguished from all other mollusks by the presence 
of hydroxyproline in the shell. This amino acid is distinctive of 
collagen; its adaptive advantage is probably one of making the 
shell flexible, and the same explanation suffices for the rather high 
protein content. Perrier and Fischer (1911) describe a muscle 
inserted near the rim of the shell in Akera which bends the shell 
and closes the mantle cavity. An evidently homologous muscle pro- 
vides a respiratory current in Cylindrobulla, a form transitional 
between Anaspidea and Sacoglossa (including Oxynoe) (cf. Mar- 
cus and Marcus, 1956); from it may be derived the shell adductor 
muscle in the bivalved gastropod (Baba, 1961). These homologies. 



14 BREVIORA No. 262 

both morphological and chemical, strongly support the relation- 
ship previously inferred (Ghiselin, 1966b) on the basis of repro- 
ductive anatomy and spermatozoon morphology. Although both 
Aplysia and Dolabella are universally looked upon as closely 
related to Akera, neither contains hydroxyproline in its shell. Aply- 
sia is closest to Akera, but deviant, and Dolabella is so distinct 
that no particular relationship for it follows from our data. Al- 
though Table 3 shows Akera and Oxynoe more closely related to 
each other than to Aplysia, which is a possibility, it seems more 
reasonable to infer that Aplysia and Dolabella have lost their 
former resemblances to Akera; their shells are vestigial and no 
longer function in the adults. Cavolina (order Thecosomata) and 
Umbraciilum (order Notaspidea) are here grouped together on 
the basis of a few similarities, especially factor 4; however, to 
place Cavolina with the Akera-Bulla group, as previously sug- 
gested on the basis of admittedly weak anatomical evidence (Ghi- 
selin, 1966b), would be fairly consistent with our data. 

Insufficient information was available to establish relationships 
for the basommatophorous Pulmonata (Siphonaria, Helisoma and 
Planorbis), beyond relating them to other Euthyneura. In the Sty- 
lommatophora, on the other hand, the relationships seem clear: 
Achatinella (suborder Orthurethra) and Succinea (suborder Hete- 
rurethra) are very close. Admitting the possibility of convergence 
due to a terrestrial habitat, our data contradict the view of Rigby 
(1965) that Succinea is an opisthobranch with affinities to the 
Anaspidea {Akera, Aplysia, and Dolabella) . The pulmonate na- 
ture of Succinea is further supported by its stylommatophoran 
chromosome morphology (Inaba, 1959), gizzard type, nervous 
system, and reproductive morphology and cytology. 

Bivalvia. The relationships of bivalves have long been contro- 
versial, owing to parallelism, convergence, and an insufficient num- 
ber and variety of characters. (For a useful summary of the data 
which have been used, see Newell, 1965.) Hinge-teeth (Dall, 1894, 
and others) have been very useful, especially with fossils, and are 
particularly useful in that some types are divergent, but some 
convergences have taken place. The attempts of Pelseneer (e.g., 
1911) to divide the bivalves on the basis of gill elaboration result 
in partially artificial groupings, because of unidirectional, parallel 
evolution. There are analogous difficulties with palp types (Stasek, 
1963), kidneys (Odhner, 1912) and the pallial sinus. Douville's 
(1912) division into normal sedentary and burrowing branches 
does show fair correspondence to other groupings, and this is to 



1967 MOLLUSCAN SHELL PROTEINS 15 

be expected, as the branches are divergent. Again, certain spe- 
cializations for which there seems to be no unidirectional trend 
give groupings which correlate with other types of evidence: cer- 
tain types of cilia on the gill (Atkins, 1938), and the anisomyarian 
state. We have already alluded to the value of some modified and 
divergent stomach types (Purchon, 1958, 1959). When the 
available evidence is properly weighted — emphasizing divergences 
and recognizing possible effects of parallelism and convergence — 
the bivalves fall quite readily into natural groupings which may be 
supported on the basis of various kinds of evidence. The same 
is true of the new information summarized here. Although the pre- 
cise sequence of genealogical relationships remains uncertain, the 
general pattern obtained correlates quite rigorously with other 
types of evidence. Indeed, our groupings correspond almost per- 
fectly with the classification proposed by Newell (1965), except 
that our data (and some other evidence) suggest removing one 
group to another, closely related one. 

Our data are presented as factor scores in Table 4. Nucula may 
be considered as having properties closest to the common ancestor 
of the class. We give no starting point for the tree because the 
data are consistent with a variety of interpretations, although the 
common stem presumably is near to Nucula. 

Nucula, Solemya, Malletia and Yoldia are members of the 
order Protobranchia, a group generally held to be representative 
of the ancestral bivalve stock. The placement of Malletia and Yol- 
dia close together, distinct from Nucula is in agreement with the 
usual classifications (McAlester, 1964). Solemya is similar to 
Nucula but somewhat modified, and its relationship to the Nucul- 
anidae {Malletia, Yoldia) is only weakly supported by our data. 
Newell's (1965) removal of Solemya to a taxon of rank equal to 
that of other protobranchs is perhaps based on an overemphasis of 
shell morphology. 

Periploma and Lyonsia are not greatly altered from the Nucula 
stage. The pattern of modification agrees well with their usual 
classification as Pandoracea, and with the view that they are but 
distantly related to the other major groups of higher bivalves in- 
cluded in this study. Neotrigonia is of uncertain relationship. The 
position shown is only weakly supported by our evidence. Indeed, 
it would in many ways be more satisfactory to relate it to the 
Arcidae (Anadara and Limopsis) as suggested by Odhner (1912) 
and others. It shows a pattern of change which is intermediate be- 
tween Mytilus and Anadara in factors 3, 4, 5, and 9; only factor 1 
is out of place, and this could be a divergent or ancestral state. 



1 6 BREVIORA No. 262 

The connections shown on the diagram for Anadara and Limop- 
sis serve only to suggest a possible relationship of other bivalves 
to one lineage: the two genera are closely related. A close re- 
lationship of the Arcidae to the Anisomyaria (Mytilus, Crassostrea 
and Aequipecten) is supported by the presence, in the Arcidae, of 
micro-laterofrontal cilia, which are common in Anisomyaria (At- 
kins, 1938), and also by the type III stomach (Purchon, 1957) 
which occurs only in Pteriomorpha {Mytilus and Crassostrea, but 
not Aequipecten) . 

The remaining bivalves are Heterodonta (sensu Newell, 1965). 
Pitar, Mercenaria, Saxidomus and Petricola are all members of 
the Veneracea, and clearly form a natural group, as do Corbicula 
and Arctica (Corbiculacea). The placement of Macoma and 
Tagelus (Tellinacea) with Mulinia and Laevicardium is in agree- 
ment with Newell's (1965) classification, but inclusion of the 
Corbiculacea in this grouping is not. However, Mulinia and some 
of its allies share with the Corbiculacea a desmodont hinge, unlike 
the Veneracea; the hinge of the Tellinacea is likewise aberrant, 
although Laevicardium (here placed as an early offshoot of the 
Tellinacea-Corbiculacea line) is heterodont. 

NOTES ON ECOLOGY 

Invasion of land and fresh water. In the present study we have, 
on the whole, attempted to hold the environment constant by 
selecting our material from similar habitats. An exception is a 
few freshwater and terrestrial forms. The effects of such a drastic 
change in habitat are striking. In Table 5 are compared two fairly 
closely related prosobranchs, the marine Nerita and freshwater 
Viviparus, and a similar series of two marine Euthyneura (Archi- 
tectonica and Siphonaria), two freshwater pulmonates (Helisoma 
and Planorbis) and two land pulmonates (Succinea and Acha- 
tinella). In general, the same amino acids change in the same 
direction with shifts to both land and fresh water: aspartic acid 
(factor 8); threonine, glutamic acid, glycine, methionine (factor 
1); perhaps histidine (factor 3); proline, arginine (factor 4); 
serine, alanine, phenylalanine (factor 6). It holds true as a general 
rule, that the change in concentration for these amino acids is in 
the direction of the concentrations that prevail in more primitive 
mollusks such as Haliotis and Nucula (Table 2). Other amino 
acids either remain constant or change in the opposite direction 
in terrestrial forms: valine, isoleucine and leucine, i.e., factor 2. 
Tyrosine (factor 3) seems to change in the direction away from 



1967 MOLLUSCAN SHELL PROTEINS 17 

the primitive state; the freshwater forms change to a greater de- 
cree than the terrestrial ones. The other members of factor 3 
(hydroxylysine, lysine and histidine) show no clear-cut pattern, 
but tend to approach the primitive level. Tyrosine and phenylala- 
nine seem anomalous, but this may be due to their both being 
phenoiics. Although we do not have enough data to obtain con- 
clusive results (especially since we do not have several lineages of 
terrestrial forms), the evidence suggests that convergent changes 
occur in freshwater and terrestrial environments. It is probable 
that the shift to a new habitat makes it advantageous to concentrate 
the same functional units which had been decreased, in proportion, 
among advanced marine forms. Such a pattern of change is con- 
sistent with the presumed heterogeneity of the protein. When the 
structure and function of the substances underlying the variation 
are known, it should be possible to explain just why particular 
changes take place. 

Effects of salinity and temperature. The sample given here is 
large enough only to be suggestive, but there is reason to think 
that within a species, salinity and temperature may have some 
effect on amino acid ratios. However, the effect seems to be char- 
acteristic for each species, and there is little evidence that particu- 
lar amino acids vary consistently in correlation with salinity or 
temperature for larger taxa as a whole. Hare (1962) measured 
the amino acid content for Mytilus californianus over a wide range 
of temperatures, and found no correlated change in protein compo- 
sition. Comparison of other species (M. edulis and M. viridis) 
from widely different habitats supports the same view (Degens 
and Spencer, 1966). However, the periostracum of Tagelus divisus 
is distinctly different in forms from Bermuda, on the one hand, and 
Nantucket and Long Island, on the other (Degens et al, 1966). A 
factor analysis of the shell matrix proteins in Polinices duplicatus, 
Mulinia lateralis and Anadara transversa (Degens et al., 1966) 
shows that there is a distinct correlation between mean temperature 
and salinity and certain of the factors used in establishing genea- 
logical relationships. Mulinia and Anadara both showed the same 
direction of change in factors 1 and 5, but only Mulinia changed 
progressively in factor 8, while Anadara changed in factors 2, 6 
and 9. Polinices changed in the opposite direction in the threonine- 
glutamic acid-glycine factor. For Mulinia, a multiple regression 
analysis was run to evaluate the degree of predictibility of each 
amino acid from the following environmental parameters: median 
temperature, range of temperature, median salinity, range of salin- 
ity, and depth. These analyses showed that only in the case of iso- 
leucine, leucine, valine and cystine, was a statistically significant 



18 BREVIORA No. 262 

linear regression obtained. For example, 82% of the variance of 
isoleucine is explained in terms of the environmental parameters. 
Of these, the median temperature and range of salinity were by 
far the most important, accounting for ca. 457© and ca. 30% of 
the variance, respectively. Such temperature dependent variation 
as occurs may reasonably be attributed to differential effects of 
temperature on the various chemical processes involved in shell 
deposition. Some of the patterns of variation may ultimately be 
related to genetic differences and therefore cast light on the evo- 
lution of the underlying physiological mechanisms. The precise 
significance of such variation must await the accumulation of a suf- 
ficient body of relevant data, but the problem is under investiga- 
tion. 

CONCLUSIONS 

Success in grouping mollusks on the basis of shell matrix 
proteins suggests that the method may prove useful in the future. 
However, the precise placements given by this study must re- 
main speculative until more adequate samples and better tech- 
niques are available for dealing with ecological variation and 
other problems. Preliminary results (Degens et al., 1966) indi- 
cate that periostracum also is useful for phylogenetic inference, 
and other tissues show promise. 

In any such study as this, it is essential that rigorous methodol- 
ogy be employed. This is clear-cut evidence that various lineages 
have undergone the same changes repeatedly and independently. 
Because of mosaic evolution, the rates at which changes occur in 
a single lineage are not necessarily the same for different structures. 
It is both bad biology and fallacious logic to place whole organisms 
in a series from "primitive" to "advanced" forms and argue that 
one has a real series of genealogical relationships. The term 
"primitive" is correctly used in a descriptive sense, in referring 
to the earlier stages of an evolutionary sequence. In such compari- 
sons it is perfectly valid to assert, for example, that the noncalcified 
molluscan shell is more primitive than the calcified. But such com- 
parisons give only intensive sequences: it is nonsense to meas- 
ure "primitiveness" by summing up the degree of advancement 
in several different systems, as the various comparisons are not in 
commeasurable units. The present study illustrates this point very 
well. In terms of the degree to which metamerism has been lost, 
one could rank some mollusks thus: Neopilina; Nautilus; C/iaeto- 
pleura; Sepia; Haliotis and Nucula. In terms of eye anagenesis, the 



1967 MOLLUSCAN SHELL PROTEINS 19 

order would be: Neopilina (?); Chaetopleura and Nucula; Halio- 
tis; Nautilus; Sepia. For shells it would be: Chaetopleura and 
Sepia; Nautilus, Haliotis and Nucula; Neopilina. The absurdity of 
trying to elaborate a calculus of evolutionary advance should be 
obvious. Nonetheless, one often reads assertions that a particular 
relationship should be rejected because modern members of some 
group are not "primitive" enough for some of their characters to 
represent the ancestral state. The present confusion in molluscan 
taxonomy may be expected to continue as long as such fallacy is 
perpetuated. 

The approach has obvious use in developing a comparative 
biochemistry and physiology of calcification. The matrix is some- 
times preserved in fossils, and study of its paleontology has al- 
ready begun (Degens and Love, 1965; Degens and Schmidt, 
1966). The effects of environmental factors should be particularly 
interesting. A change to fresh water from the sea would affect the 
pH and various ion concentrations, as is suggested by the rather 
high protein levels in freshwater forms. Where the calcification 
mechanism is inefficient, and where the medium in which calcifica- 
tion occurs is not regulated, slight changes in the environment — 
ionic ratios, pH, temperature, etc. — might have great effect on 
biota. Conceivably, physical and chemical changes in the environ- 
ment might help to explain the greater abundance of fossils in 
Cambrian and younger rocks than in older ones, and mass ex- 
tinction of ammonites and nautiloids. It is well known that the 
formation of coral reefs is dependent on temperature, and it seems 
a reasonable analogy that the success or failure of many organisms 
may be intimately connected with the process of calcification. 

SUMMARY 

1. Measurements of the amino acids in the shells of selected 
mollusks show a wide diversity. Some differences may be related 
to progressive evolutionary development of the shell. 

2. Factor analysis shows that 907^ of the variance may be 
explained in terms of a few factors. A technique which overcomes 
problems of parallelism and convergence has been developed for 
inferring phylogenies on the basis of differences in the factor struc- 
ture. 

3. Various phylogenetic hypotheses and classification schemes 
are evaluated on the basis of the evidence obtained in the study. 
The results are in close agreement with conventional classification 
systems, and cast some light on the positions of forms of disputed 
relationship. 



20 BREVIORA No. 262 

4. Preliminary results show that there are convergences with 
shifts from marine to freshwater habitats, and that temperature and 
salinity dependent variation poses a considerable, but not insol- 
uble, problem of interpretation. 

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1935. Jena. 2 vols. 
Wilbur, K. M. 

1964. Shell formation and regeneration. In: K. M. Wilbur and C. M. 
Yonge (eds.). Physiology of Mollusca. Academic press. New 
York, Ch. 8: xiii -f 473 pp. 

(Received 13 September, 1966.) 



24 



BREVIORA 



No. 262 




1967 MOLLUSCAN SHELL PROTEINS 25 

PLATE I 
GASTROPODA 

Figure 1. Achatinella lorata Pfeiflfer, 1848, Oahu, Hawaiian Islands, 
Pease collection, MCZ (Mollusk Department) — aperture view. X 1. 

Figure 2. Acniaea piistiilata (Helbling, 1799), Puerto Sosya, Dominican 
Republic, MCZ, coll. 1937 — top view. X 2/3. 

Figure 3. Akera soliita (Gmelin, 1791), Zanzibar, MCZ No. 4444 — 
aperture view. X 1/3. 

Figure 4. Architectonica nobilis (Roding, 1798), Middle Atlantic Coast, 
2-10 m, MCZ No. 90867 — aperture view. X 2. 

Figure 5. Astraea caelata (Gmelin, 1798), Pelican Shoals, Florida, 
MCZ, coll. 1939 — aperture view. X 2/3. 

Figure 6. Bulla striata (Bruguiere, 1792), Puerto Vieja, Dominican Re- 
public, MCZ, coll. R. H. Parker, April 1965 — aperture view. X 2/3. 

Figure 7. Cavolina tridentata (Forskal, 1776), pelagic. West Indies, 
Alexander Agassiz coll. 1879, MCZ (uncat.) — aperture view. X 2. 

Figure 8. Cerion regiiim (Benson, 1849), northeast end of Castle Island, 
Crooked Island Group, Bahamas, B.W.I. — coll. Robertson and Scott, 
1958. MCZ (uncat.) — aperture view. X 2/3. 

Figure 9. Crepidula fornicata (Linne, 1767), Hadley Harbor, Elizabeth 
Islands, Massachusetts, Sta. P-22 11-65, Systematics-Ecology Program, Ma- 
rine Biological Laboratory — interior view; 9A, exterior view, X 1 1/3. 

Figure 10. Crepidula plana (Say, 1822), Hadley Harbor, Elizabeth Is- 
lands, Massachusetts, Sta. P-275-65, Systematics-Ecology Program, Marine 
Biological Laboratory — interior view; lOB, exterior view. X 2/3. 

Figure 11. Coins trophius (Dall, 1919), off San Francisco, California, 
1874-1929 m, coll. R. H. Parker, MCZ (uncat.) — aperture view. X 1 1/3. 

Figure 12. Cypraea zebra (Linne, 1758), Miami, Florida, MCZ (uncat.) 
— aperture view. X 1/3. 

Figure 13. Dolabella callosa (Lamarck, ISOl) = scapula (Martyn), 
Calapan, Mindoro, Philippine Islands, MCZ No. 96107 — side view. X 2/3. 

Figure 14. Epitonium angulatum (Say, 1831), near mouth of New 
Brazos River, Freeport, Texas, MCZ No. 230893 — aperture view. X 2. 

Figure 15. Fissurella barbadensis (Gmelin, 1791), Romey Point, Bonn- 
guen Air Force Base. Puerto Rico, MCZ No. 1956 — top view (exterior). 

X2/3. 



26 BREVIORA No. 262 

PLATE I (Continued) 

Figure 16. Haliotis cracherodi (Leach, 1814), San Diego, California, 
MCZ No. 58659 — exterior view. X 1/3. 

Figure 17. Helisoma trivalvis (Say, 1817), La Porte, Indiana, MCZ No. 
63234 — side view; 17B, aperture view. X 1 1/3. 

Figure 18. Jantliina jantliina (Linne, 1767), Cape Florida Key, Biscayne, 
Florida, MCZ No. 155173 — aperture view. X 5/6. 

Figure 19, Littorina Uttorea (Linne, 1758), Hadley Harbor, Elizabeth- 
Islands, Massachusetts, Sta. P-0610-65, Systematics-Ecology Program, Ma- 
rine Laboratory — aperture view. X 1. 

Figure 20. Liinatia triseriata (Say, 1826), Hadley Harbor, Elizabeth 
Islands, Massachusetts, Sta. P-247-65. Systematics-Ecology Program, Marine 
Biological Laboratory — aperture view. X 2. 

Figure 21. Murex brevifrons (Lamarck, 1822), Mayaguez, Puerto Rico, 
coll. M. R. Carriker, Systematics-Ecology Program, Marine Biological 
Laboratory — aperture view. X 1/3. 

Figure 22. Nassariiis trivittatiis (Say, 1822), Hadley Harbor, Elizabeth 
Islands, Massachusetts, Sta. P-0605-65, Systematics-Ecology Program, Ma- 
rine Biological Laboratory — aperture view. X 2. 

Figure 23. Nerita plexa (Dillwyn. 1817), Mauritius, MCZ No. 139024 
■ — -aperture view. X 1/3. 

Figure 24. Oxynoe viridis (Pease, 1864), from Pease coll., MCZ (uncat.) 
aperture view. X 2 1/3. 

Figure 25. Polinices diiplicatus (Say, 1822), Provincetown, Massachu- 
setts, MCZ (uncat.) — aperture view. X 1 1/3. 

Figure 26. Siphonaria cdternata (Say, 1826), Bermuda, coll. Bryant. 
1903, MCZ No. 24213 — top view (exterior). X 1 1/3. 

Figure 27. Siiccinea oralis (Say, 1817), bank of Grand River, W. Bridge 
Street Ferry, Ottawa County, Michigan, MCZ No. 166697 — aperture view. 
X 2. 

Figure 28. Tiirritella terebra (Lamarck, 1822), Manila Bay. near 
Cavite, Luzon, Philippine Islands, MCZ No. 138070 — aperture view. X 1/3. 

Figure 29. Umbracuhim indicitm (Lamarck, 1819), Hawaii, C. B. Adams 
coll., MCZ No. 1173 — top view (exterior); 29A, side view. X 1/3. 

Figure 30. Urosalpinx cinerea (Say, 1822), Hadley Harbor, Elizabeth 
Islands, Massachusetts, Sta. P-06 12-65, Systematics-Ecology Program, 
Marine Biological Laboratory — aperture view. X 4 2/3. 



1967 MOLLUSCAN SHELL PROTEINS 27 

PLATE I (Continued) 

Figure 31. Vivipanis georgianus (1. Lea, 1834), Lake Woodruff near 
Pick Island, Volusa County, Florida, MCZ No. 186800 — aperture view. 

X2/3. 

AMPHINEURA 

Figure 32. Chaetopleitra apiculata (Say, 1830), Hadley Harbor, Eliza- 
beth Islands, Sta. P-7-63, Systematics-Ecology Program, Marine Biological 
Laboratory — top view (exterior). X 2. 



28 



BREVIORA 



No. 262 




1967 MOLLUSCAN SHELL PROTEINS 29 

PLATE II 

CEPHALOPODA 

Figure 33. Argonaiita hians (Dillwyn, 1817), southern Tropical Atlan- 
tic. MCZ (uncat.) — side view of egg case. X 1/3. 

Figure 34. Nautilus pompilius (Linne, 1758), Southwest Pacific Ocean, 
MCZ (uncat.) — side view. X 1/3. 

Figure 35. Spimla spirula (Linne, 1758), Saint Kitts Island, B.W.L, 
MCZ (uncat.) — side view. X 1 1/3. 

PELECYPODA 

Figure 36. Aequipecten irradians (Lamarck, 1819), Hadley Harbor, Eliz- 
abeth Islands, Massachusetts, Sta. P-65-64, Systematics-Ecology Program, 
Marine Biological Laboratory — exterior view, right valve. X 1. 

Figure 37. Anadara transversa (Say, 1822), Hadley Harbor, Elizabeth 
Islands, Massachusetts, Sta. P-77-64, Systematics-Ecology Program, Marine 
Biological Laboratory — exterior view, left valve. X 9 1/3. 

Figure 38. Arctica islandica (Linne, 1758), Georges Bank, Massachu- 
setts^ MCZ No. 14325 — interior view, right valve. X 1/3. 

Figure 39. Corbicula consobrina (Cailliand, 1827), Nile River, Egypt, 
MCZ No. 14676 — interior view, right valve. X 1 1/3. 

Figure 40. Crassostrea virginica (Gmelin, 1791), Holding Tank, Supply 
Department. Marine Biological Laboratory, Woods Hole. Massachusetts — 
interior view, right valve. X 1/3. 

Figure 41. Laevicardium mortoni (Conrad. 1831), Hadley Harbor, Eliz- 
abeth Islands. Massachusetts. Sta. P- 14 1-64, Systematics-Ecology Program. 
Marine Biological Laboratory — exterior view, left valve. X 8 1/3. 

Figure 42. Limopsis compressus (Dall, 1908), off Salina Cruz. Mexico, 
1020-1240 m. coll. R. H. Parker, MCZ (uncat.) — exterior view, left valve. 
X 2/3. 

Figure 43. Lyonsia hyalina (Conrad, 1831), Hadley Harbor, Elizabeth 
Islands. Massachusetts, Sta. P-200-64, Systematics-Ecology Program, Ma- 
rine Biological Laboratory — exterior view, left valve. X 5. 

Figure 44. Macoma tenta (Say, 1834), Hadley Harbor, Elizabeth Is- 
lands, Massachusetts, Sta. P-91-64, Systematics-Ecology Program. Marine 
Biological Laboratory — exterior view, left valve. X 2 1/3. 

Figure 45. Malletia, species "M." Atlantic Abyssal Plain, 4970 m. coll. 
H. L. Sanders, in coll. at Woods Hole Oceanographic Institution — exterior 
view, left valve. X 2 1/3. 



30 BREVIOKA No. 262 

PLATE II (Continued) 

Figure 46. Mercenaria mercenaria (Linne, 1758), Holding Tank, Supply 
Department, Marine Biological Laboratory, Woods Hole, Massachusetts — 
exterior view, left valve. X 2 1/3. 

Figure 47. Midinia lateralis (Say, 1822), Hadley Harbor, Elizabeth Is- 
lands, Massachusetts, Sta. P-0902-65, Systematics-Ecology Program, Marine 
Biological Laboratory — exterior view, left valve. X 2 1/3. 

Figure 48. Mytilus edulis (Linne, 1758), Hadley Harbor, Elizabeth Is- 
lands, Massachusetts, Sta. P- 148-64, Systematics-Ecology Program, Marine 
Biological Laboratory — exterior view, left valve. X 2 2/3. 

Figure 49. Neotrigonia margaritacea (Lamarck, 1803), New South Wales, 
AustraHa. from coll. of A. L. McAlester. Yale University — exterior view, 
right valve. X 2/3. 

Figure 50. Nucula proxima (Say, 1822), Hadley Harbor, Elizabeth Is- 
lands, Massachusetts, Sta. P-92-64, Systematics-Ecology Program, Marine 
Biological Laboratory — exterior view, left valve. X 2 1/3. 

Figure 51. Nucula proxima variety truncula (Dall, 1898), Buzzards Bay, 
Massachusetts, Sta. "R," coll. H. L. Sanders, Woods Hole Oceanographic 
Institution — exterior view, left valve. X 4 2/3. 

Figure 52. Periploma leanuni (Conrad, 1831), Martha's Vineyard, Chap- 
paquidick Island. Massachusetts, MCZ No. 192934 — interior view, left 
valve. X 1/6. 

Figure 53. Petricola pholadiforinis (Lamarck, 1818), Black Beach, West 
Falmouth, SEP-949, George M. Gray Museum, Systematics-Ecology Pro- 
gram, Marine Biological Laboratory — exterior view, left valve. X 2 1/3. 

Figure 54. Pilar cordata (Schwengie, 1951). off Port Aransas, Texas, 80 
m, MCZ No. 194372 — interior view, right valve. X 2/3. 

Figure 55. Pitar morrhuatia (Linsley, 1845), Hadley Harbor, Elizabeth 
Islands, Massachusetts, Sta. P-1 89-64, Systematics-Ecology Program, Ma- 
rine Biological Laboratory — exterior view, left valve. X 2 1/3. 

Figure 56. Saxidomiis mittalli (Conrad, 1837), Gulf of Georgia, British 
Columbia, Canada, MCZ No. 5235 — interior view, right valve. X 1/3. 

Figure 57. Solemya velum (Say, 1822), Hadley Harbor, Ehzabeth Is- 
lands, Massachusetts, Sta. P-225-65, Systematics-Ecology Program, Marine 
Biological Laboratory — exterior view, left valve. X 2 1/3. 

Figure 58. Tagelus divisus (Spengler, 1794), Hadley Harbor, Elizabeth 
Islands, Massachusetts, Sta. P-21-63, Systematics-Ecology Program, Marine 
Biological Laboratory — exterior view, left valve. X 2 1/3. 

Figure 59. Yoldia limatula (Say, 1831), Hadley Harbor, Elizabeth Is- 
lands, Massachusetts, Sta. P-267-65, Systematics-Ecology Program, Marine 
Biological Laboratory — exterior view, left valve. X 2 1/3. 



1967 



MOLLUSCAN SHELL PROTEINS 



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32 



BREVIORA 



No. 262 



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o 




o 












O 


o 








re 




^ 


' ^ 


CVJ 




uo 


^3- 


o 


2; 




CVJ 




^ 


CM 


n 


ro 




CvJ 




CO 


=^ 




i/i 






































^ 










































, 


































<^ 




-^ 
























































o 


cr. 


OD 


00 


















o 




O 


CO 


5 




^ 


1 CO 


^ 


IT 


'=^ 




en 


f3 




■" 




'~ 


CvJ 


^ 


LO 








CM 


^ 




1/0 






































o 




















CM 






















C71 
















































1 -sr 


o 










CO 


o 




O 


■3- 




CO 








CO 


CO 






OJ OJ 


















VD 






^D 


la 












O 




-C CL 










































oo — 










































,__ 










































OJ 










































c 










































o 










































-Q 










































OJ 


1 vn 


0-) 


CO 


ro 


r^ 


o 


UD 


00 


O 


O 


O 


,— 


o 


,— 


1 


CO 


cr. 


CM 










■st- 


cr. 




O 




<3 












U3 








■=r 








»— 4-> 






































•^ 




"oi 3 






































OJ 














































Ln~^ 






































<n 




































































































































1 o 






r^ 




ro 


^ 




CTi 


o 


































O 


























a. 




S 


*" 




*" 


*~~ 




^ 


























+-> 


Z 








































QJ 


3 




1 CO 


O 


CO 


CO 






00 








O 


















QJ 


OJ 








01 






CO 




»r 














■^ 




^o 












































(_) 


"o. 


00 






































o 


ro 


OJ 


























VO 








^ 














































OJ 


5 


■^ 


00 o 


C^ 


•o 


Oi 


Oi 


ii> 


IC 


n 


CO 


'^ 


CO 


s 


O 


cn 


■^^ 


^ 


o 


ID 












































LJ 


Q. 


s 







































c 
•.- u 

O -*-» C OJ E c 
o c ~ 

OJ ■— 



Ol OJ OJ OJ 
_ OJ P= 

s- o 

ro OJ 

O. S- S- 3 O >, 
_ S- .— 



C QJ O 



.— >» fO OJ 



(U c TC) -^ 

u o c: _i 
at >i -c: ic 



<U OJ 



1967 



MOLLUSCAN SHELL PROTEINS 



33 



Table 3 

FACTOR SCORES. GASTROPODA. 
(Shell Tissues) 
12 3 4 



Phylogenetic 
Relationships 



n 



n 



Varimax 
Factors 



Positive 
Loading 



Negative 
Loading 



SIPHONARIA 

HELISOMA 

PLANORBIS 

ACHATINELLA 
SUCCINEA 

CAVOLINA 

UMBRACULUM 

ARCHITECTONICA 

AKERA 

OXYNOE 

APLYSIA 

BULLA 

DOLABELLA 

EPITONIUM 

JANTHINA 

TURRITELLA 

MUREX 

UROSALPINX 

NASSARIUS 

COLUS 

MELANELLA 

POLIMICES 

LUNATIA 

LITTORINA 

CYPRAEA 

CREPIDULA PLANA 

CREPIDULA FORNICATA 

FISSURELLA 

ACMAEA 

VIVIPARUS 

NERITA 

ASTRAEA 

HALIOTIS 



GLY 



GLU 
THR 
MET 



0.8 

2.2 

0.6 

1.8 
3.0 

•0.3 
■0.6 
•0.2 
•0.1 
•0.4 
•0.6 
■0.0 
0.4 

■1.0 
-1.0 
-0.4 

-0.8^ 

-0.8 

-0.3 

0.2 

0.9 

0.2 

-0.1 

-1.0 

-1.2 

-0.61^ 

-0.6 

-0.8 

-0.7 

2.3 

-0.8 

1.0 

1.2 



VAL 

ISOLEU 

LEU 



-0.2 

-0.4 

0.9 

2.0 
2.0 

-0.2 

-0.4 

-0.1 

0.6 

0.4 

1.1 

1.0 

-1.3 

-0.5 
-0.1 
-0.7 

-0.7 

0.7 

1.4 

0.8 

-1.3 

-0.7 

0.8 

0.4 

0.4 

0.8 

1.6 

-0.5 

0.9 

-1.6 

-1.0 

-1.6 

-1.8 



OHLYS 
LYS 
HIS 
TYR 



-0.6 

0.3 

5.2 

-0.4 
-0.4 

0.2 
-0.3 

0.2 
-0.5 
-0.5 
-0.4 

0.0 

0.0 

1.5 

0.0 
0.2 

1.8 
-0.3 
-0.2 
-0.9 
-0.5 
-0.5 
-0.4 
-0.2 
-0.6 
-0.7 
-0.3 

0.5 
-0.2 
-0.1 

0.6 
-0.7 

0.0 



PRO 



ARC 



-0.5 

0.7 

0.9 

-0.7 
0.8 

2.5' 

1.7 

1.0 

-0.0 

0.3 

-1.2 

-0.2 

-0.1 

-0.7 

-0.3 

0.3 

-2.1 

-0.9 

0.3 

1.0 

-0.6 

0.7 

-1.3 

-0.2 

-0.1 

0.2 

-1.1 

0.2 

0.4 

0.4 

-0.4 

-1.8 

-1.5 



CYS 



-0.3 

0.9 

-0.2 

0.6 
0.2 

0.1' 

-0.2 

0.1 

0.4 

0.4 

-0.1 

-0.1 

0.4 

0.5 
0.4 
0.1 

0.9 

0.1 

0.1 

-0.6 

-0.1 

0.2 

0.4 

0.3 

-0.1 

-0.1 

-0.8 

-0.1 

0.1 

0.5 

0.3 

0.0 

0.1 



SER 
ALA 



PHE 



-2.1 

0.2 

0.8 

-1.7 
-0.1 

-0.1 
0.0 
0.1 
0.2 

-0.2 

-0.2 
0.9 

-1.1 

0.3 

-0.5 

0.3 

-0.8 

-0.5 

-0.3 

0.5 

2.2 

1.2 

0.2 

-0.5 

-0.3 

0.2 

0.1 

-0.7 

0.2 

-2.9 

-1 

0.9 

0.0 



::f 



OHPRO 



0.1 

-0.3 

0.5 

-0.3 
-0.7 

0.0 
-0.4 
-0.2 

1.4 

5.6 ' 
-0.2 
-0.6 
-0.6 

-0.3 
-0.3 
-0.3 

-0.6 
-0.3 
-0.2 
-0.1 

0.2 

0.1 
-0.2 
-0.3 
-0.1 
-0.6 
-0.6 
-0.1 
-0.3 

0.1 

0.2 

0.5 

0.2 



ASP 



0.1 

-1.7 

-0.3 

1.2 
1.4 

-0.3 
-0.5 
-0.5 
-0.2 
-0.1 
-0.6 
-1.3 
-3.9 

1.4 , 
0.4 
-0.1 

0.7 

0.3 

0.1 

-1.4 

i-U 

0.7 
-0.4 
-0.2 

0.5 
-0.3 
-0.5 

0.2 
-0.2 

1.3i 

0.6l 
-0.2 
-1.3 



34 



BREVIORA 



No. 262 



Table 4 

FACTOR SCORES. PELECYPODA. 

(Shell Tissues) 

12 3 4 5 



Phylogenetic 
Relationships 



TZ 



JI 



JZ 



n 



Varimax 
Factors 



Positive 

Loading 



Negative 
Loading 



PI TAR. 
MERCENARIA 
SAXIDOMUS 
PETRICOLA 

LAEVICARDIUM 

MACOMA 

MULINIA 

TAGELUS 

CORBICULA 

ARCTICA 

AEQUIPECTEN 

CRASSOSTREA 

MYTILUS 

ANADARA 

LIMOPSIS 

NEOTRIGONIA 
PERIPLOMA 
LYONSIA 
NUCULA 

SOLEMYA 

MALLETIA 
YOLDIA 



THR 
GLU 
PRO 



0.3 
2.4 
1.3 
0.5 

2.2 
-0.6 
0.1 
0.8 
0.7 
2.9 

-0.6 
-0.9 

-0.8 
-0.5 
0.1 

-1.1 
-1.2 
-1.1 
-0.6 

-0.9 

0.5 
0.9 



VAL 



0.3 
-1.0 
-1.1 
-0.6 

0.5 
0.1 
0.7 
-0.6 
-0.3 
0.3 

-3.1 
-2.1 
-0.3 
1.3 
-1.2 

-0.8 

-0.6 

-0.6 

0.1 



LYS 
HIS 
ARG 



0.2 

1.8 

-0.4 

-1.0 

-1.0 
-0.4 
-0.5 
-0.5 
-0.3 
-0.5 

-0.5 
0.3 

-0.6 
1.7 
2.0 

0.1 
0.6 
0.2 
0.2 

-0.8 

0.9 
0.2 



ALA 



MET 



-0.5 
0.2 
0.5 

-1.1 

-0.8 

-1.8 

-0.2 

0.1 

0.4 
1.1 

-0.6 

-0.5 

3.0 

0.1 

-0.8 

1.5 

-0.8 

1.1 

0.7 

-1.1 

0.2 
0.6 



ISOLEU 
LEU 
VAL 
HIS 



-0.7 

0.5 
1.5 

0.4|^ 
-0.4 
-0.5 
-1.0 
-0.8 
-0.1 



-0.6 
-1.2 
0.5 
-0.2 

-o.sr 

0.0 
-0.4 

0.0 
-0.7 

0.2 

-1.5 
-0.6 



TYR 



1.8 
0.3 
1.9 

-1.2 
-0.9 
0.1 
0.0 
1.6 
1.1 

-0.7 
1.2 
0.0 
1.4 

-1.8 

-1.0 
2.6 
0.3 

-0.7 

-0.7 
-0.3 



OHLYS 



-0.4 
-1.3 
-0.2 
-0.2 

-1.5 
-0.1 
-0.1 

0.0 
-0.3 

4.8 

0.1 
-0.2 
-0.4 

0.1 
-0.3 

1.2 
-0.4 
0.3 
0.0 

0.2 

-0.5 



-0.1 0.2 



ASP 



-1.9 
1.4 

-0.9 
1.0- 

1.6^ 

1.6 

-1.0 

0.3 

-0.7 

-0.4 

-1.5|k 
-0.6 

0.4 
-0.4 

0.8 

0.7 
0.9 
1.2 
1.0 

0.8 

0.6 
-1.2 



PHE 



-0.9 

0.4 

-1.4 

-0.9 

2.01 
0.7 
0.3 
-0.4 
0.9 
0.0 

1-n 

1.2 

1.0 

-0.1 

-2.3r 

0.2 
-1.8 
-1.4 
-0.9 

-0.6 

0.4 
-0.2 



1967 



molluscan shell proteins 
Table 5 

AMINO ACID RATIOS PER 1000 TOTAL 



35 



OH-Proline 

Aspartic Acid 

Threonine 

Serine 

Glutamic Acid 

Pr9l ine 

Glycine 

Alanine 

Cystine [half] 

Valine 

Methionine 

Isoleucine 

Leucine 

Tyrosine 

Phenyl alaniae 

OH-Lysine 

Lysine 

Histidine 

Arginine 



Nerita 


Viviparus 


Archi- 


Siphonaria 


Helisoma 


Planorbis 


Succinea 


Achatinella 


tectonica 






Marine 


Fresh 

Water 


Marine 


Marine 


Fresh 
Water 


Fresh 
Water 


Land 


Land 


103 


48 


132 


133 


143 


98 


11 


47 


50 


11 


57 


46 


23 


56 


11 


16 


70 


34 


88 


58 


74 


61 


37 


25 


114 


26 


114 


103 


69 


91 


9 


29 


85 


54 


90 


82 


39 


59 


57 


50 


116 


519 


159 


116 


380 


114 


526 


411 


108 


15 


81 


74 


62 


88 


19 


30 


2 


8 


15 


16 


3 


21 


4 


4 


48 


17 


40 


63 


38 


61 


83 


78 


26 


11 


19 


30 


3 


10 


1 


2 


23 


26 


36 


35 


15 


37 


50 


64 


68 


59 


76 


76 


50 


56 


121 


110 


23 


46 


14 


15 


42 


67 


28 


11 


44 


95 


19 


74 


17 


-- 


16 


74 


1 


-- 


3 


— 


1 


16 


-- 


0.3 


36 


15 


26 


34 


25 


82 


12 


39 


16 


9 


6 


1 


1 


34 


0.2 


0.2 


67 


6 


25 


45 


15 


48 


6 


9 



Protein 



66 



1060 



22 



BREVIORA 

MiiseiLiioi of CoHiparative Zoology 

Cambridge, Mass. 31 March, 1967 Number 263 

THE HYDROID OF VANNUCCIA FORBESII 
(ANTHOMEDUSAE, TUBULARIIDAE) 

By Anita Brinckmann-Voss 

Stazione Zoologica, Napoli, Italy, and Institute of Animal 
Science, University of Manitoba, Winnipeg, Manitoba, Canada 



Abstract: This paper deals with a hydroid of the order Anthomedusae 
(Athecatae), family Tubulariidae, formerly described as Hybocodon forbesii 
Mayer 1894. The structure of the hydroid proves that the species does not 
belong to the genus Hybocodon: a new genus, Vanniiccia, is therefore intro- 
duced. The hydroid, the budding medusae, the asexual reproduction and 
the young medusae are described in this paper. 

INTRODUCTION 

During the preparation of a monograph of the Anthomedusae 
and athecate hydroids from the Gulf of Naples a tubularian hy- 
droid was found on mud in the Bay of Naples. This hydroid 
liberated medusae, in the laboratory, which appeared to belong 
to the species Hybocodon forbesii Mayer 1894. 

But since the hydroid does not have the structure of other 
Hybocodon hydroids and does not resemble any other hydroid of 
the Tubulariidae, a new genus had to be established for the former 
Hybocodon forbesii. The new genus is named Vanniiccia, in rec- 
ognition of Dr. Martha Vannucci, the head of the plankton section 
of the Oceanographic Institute of Sao Paulo, who has done ex- 
tensive research on the plankton around the Brazilian coast. 

The find reported here is the first and only case in which Van- 
nuccia forbesii (Mayer) has been recorded from the Mediter- 
ranean. Only one specimen of the hydroid stage could be found 
and no free medusae were collected from the sea. However, the 
species proved easy to keep in the laboratory, and numerous 
medusae as well as hydroids were liberated from the one specimen. 

The research at Naples was supported by a grant from the U.S. 



2 BREVIORA No. 263 

Office of Naval Research, No. 2100(00). Completion of the re- 
search was made possible by a grant from the National Science 
Foundation, No. GB2358 under the sponsorship of the Museum 
of Comparative Zoology, Harvard University, Cambridge. 

Vannuccia n. gen. 

Type species. Vannuccia forbesii (Mayer) 1894: 236, pi. I, 
fig. 1 ; the gender of the new generic name is feminine. 

Generic characters. Tubulariidae with solitary hydroids with 
one whorl of moniliform oral tentacles and an aboral whorl of 
filiform tentacles. With endodermal stem canals and basal tuft of 
rooting filaments. Medusa buds borne on blastostyles just above the 
aboral tentacle whorl. Free medusae with one tentacle and no 
exumbrellar cnidocyst rows. 

Vannuccia forbesii (Mayer) 

Hybocodon forbesii Mayer, 1894: 236, pi. I, fig. 1. 

Specific characters. Hydroid with 12 to 14 oral tentacles and 
16 to 20 aboral tentacles. Asexual reproduction by transverse 
fission of the stem. Free medusae with slight asymmetrical bell, one 
fully developed tentacle with a large terminal cnidocyst swelling. 

Distribution. Tortugas, Bahama Islands, and Florida (Mayer, 
1894; 1910); Trivandrum and Madras, India (Nair, 1951); south- 
ern Japan (Uchida, 1927); Bay of Naples, Mediterranean. 

Description of the species. The hydroids of V. forbesii are very 
similar to those of Corymorpha nutans. The hydranth of V. forbesi 
consists of a stem or hydrocaulus and the hydranth body itself. 
The base of the stem or the "foot" (Fig. 6) shows a similar ar- 
rangement of rooting filaments and endodermal canals to that seen 
in Corymorpha nutans. The whole stem is surrounded by a 
flexible perisarc which extends slightly below the hydranth body. 
There the perisarc is usually marked by a slight constriction of 
the stem (Fig. 1). 

The length of hydranth and stem measures 2-3 cm for speci- 
mens with medusae buds, but there is great variability in the 
length of the stem. 

The oral tentacle whorl consists of 12 to 14 tentacles each of 
them carrying 4 to 6 cnidocyst swellings. The aboral tentacle whorl 
consists of 16 to 20 tentacles. Each aboral tentacle is very long 
and has a small swelling at its tip which is more or less pronounced 
depending on whether the tentacles are contracted or expanded. 



1967 



HYDROID OF VANNUCCIA FORBESII 




Figure 1. Vannuccia jorbesii, adult hydroid with medusae buds. Drawn 
by Ilona Richter. 



The medusa buds develop in clusters on short blastostyles. As 
the buds are naked, the one tentacle of each bud can be seen in 
the more advanced stages of the medusa formation. The liberated 
medusae are 1.8-2.0 mm high. The exumbrella is apple shaped 
and somewhat asymmetrical. Young medusae show all the features 
of the specific characters of the adult medusae with the exception 
of the gonads, which develop later. This is typical of the Cory- 
morpha-Tubularia group. 



BREVIORA 



No. 263 




0.25mm 



Fig. 2. Vanniiccia forbesii, male medusa two days old. Drawn by Ilona 
Richter. 



Medusae of V. forbesii carry one fully developed tentacle and 
three tentacle bulbs. The tentacle bulb which is opposite the 
fully developed tentacle is larger than the other two (Fig. 2). 
This bulb starts to grow out on the second day after liberation to 
form a small rudimentary tentacle. Mayer, who gives a descrip- 
tion of a fairly young and an adult medusa of V. forbesii (Mayer, 
1894; 1910), shows that this tentacle may grow longer in fully 
mature specimens (Mayer, 1910, pi. III). The one fully developed 
tentacle has a large swelling at its tip, which is provided with 
numerous cnidocysts. The manubrium is club shaped without a 
peduncle and extends to the velum. Usually the manubrium does 
not stay in the center of the subumbrellar cavity, as is generally 
expected of anthomedusan species, but is located slightly nearer 
the tentacle bulb opposite the fully developed tentacle. The gonads 
start to develop soon after the liberation of the medusa. The 
stomach is completely encircled by the gonads. As the medusae 
are difficult to feed, I did not manage to get specimens with fully 
mature gonads. The velum is thin and narrow. 



1967 



HYDROID OF VANNUCCIA FORBESII 




BREVIORA 



No. 263 



Reproduction and development. All of my medusae of V. jor- 
besii were males. Therefore, the sexual reproduction could not 
be observed. However, there is a very interesting type of asexual 
reproduction of the hydroid: the base of the foot of the hydroid 
thickens slightly and is then constricted off from the remainder of 
the stem (Fig. 3), and is freed from the perisarcal sheath. It 
develops four oral and six aboral tentacle buds after it is freed 
(Fig. 4). Sometimes, however, the tentacle buds are already de- 
veloped inside the perisarc of the old hydranth. A similar type of 




Figure 4. Comparison between the asexual budding of Eiiphysa aiirata 
and Vanniiccia forbesii. I, Euphysa aiirata; II, Vannuccia forbesii. a, 
Hydranth stem; b, young buds which are constricted off from the foot of 
the hydranth; c, developing young hydranths; d, perisarc. For the clarity 
of the figure the rooting filaments are omitted in II. 



1967 HYDROID OF VANNUCCIA FORBESII 7 

budding is known from EupJiysa aurata (Forbes) and Hypolytus 
peregrinus Murbach. But in Euphysa aurata the oral tentacles of 
the bud always develop distal to the aboral ones, in relation to the 
"mother" stem (Fig. 3). 

The liberated hydranths do not settle immediately but move 
around slowly until they affix themselves after some hours or even 
one or two days. This asexual reproduction may become so 
prolific that a whole tuft of hydroids is formed out of one speci- 
men in about a week. 

Ecological observations. When found in May 1962, the young 
hydroid of V. forbesii had only six oral and ten aboral tentacles. 
During its development the number of tentacles increased. The 
young one was found in mud at 30 m depth. In the same region 
I found Euphysa aurata, but no other hydroids or medusae of V. 
forbesii. The hydroid was kept at 18°C. The asexual reproduction 
started at once, after the hydroid had been brought to the lab- 
oratory. Medusae production took place in autumn and winter. 
The asexual reproduction did not cease while the development of 
medusa buds was taking place. 

SYSTEMATIC DISCUSSION 

The medusae which were liberated from the above described 
hydroid match Mayer's description of Hybocodon forbesii (Mayer, 
1894), with one exception: There is a velum in our specimens. 
But then, while Mayer says in his original description "without 
velum" in 1894, his redescription, based on more abundant ma- 
terial, in his "Medusae of the World" (Mayer, 1910:42) men- 
tions: "The velum is narrow." 

Mayer placed his medusa species in the genus Hybocodon on 
account of the slightly asymmetrical bell and the one tentacle. 
Kramp is the first one to question the placement of Hybocodon 
forbesii in the genus Hybocodon, suggesting however that it should 
provisionally remain in that genus (Kramp, 1959, 1961). The 
medusa stage is somewhat reminiscent of Euphysa and Corymor- 
pha as well. 

Regarding the hydroid stage, difficulties arise about placing 
the species in the proper systematic position. The morphology of 
the hydroid demands that the species (up to now known from its 
medusa stage only) not remain in the genus Hybocodon any 
more, because it is certainly not a Hybocodon hydroid. The estab- 
lishment of the new genus Vannuccia for the former Hybocodon 
forbesii is suggested therefore. 



8 



BREVIORA 



No. 263 



The combination of moniliform oral and filiform aboral tenta- 
cles in V. jorbesii hydroids is unique among the Tubulariidae. Yet, 
upon checking the different solitary tubulariid hydroids, either from 
literature or preserved material, I find that the hydroid of V. jor- 
besii bears some relationship to Hypolytus obvoluta (Kramp) 
1933; this was first described as Corymorpha obvoluta, and later 
placed in the genus Hypolytus by Rees, being cogeneric with H. 
peregrinus Murbach 1899, on account of the moniliform tentacles 
in both sets (Rees, 1957). However, if one compares the figures 
of H. peregrinus Murbach and H. obvoluta (Kramp), it seems that 
the aboral tentacles of H. obvoluta are much less moniliform 
than the aboral tentacles of H. peregrinus. This points to a certain 




Figures. Oral tentacles of young Vaiiniiccia jorbesii (I), and young 
Corymorpha nutans (II). 



order insofar as there are: (1) moniliform tentacle whorls in 
both sets in H. peregrinus Murbach, (2) "reduced" aboral monili- 
form tentacles in H. obvoluta (Kramp), (3) filiform aboral and 
moniliform oral tentacles in Vannuccia jorbesii, and (4) filiform 
tentacles in both whorls, slightly moniliform in the oral set when 
young, in Corymorpha nutans. In this comparison, Vannuccia is 
placed between Hypolytus and Corymorpha. There are also other 
characters indicating the relationship o/ Vannuccia jorbesii with 
H. obvoluta, on the one hand, and with Corymorpha nutans, on 
the other: Hypolytus obvoluta shows faint endodermal channels; 
they are more apparent in V. jorbesii and strongly developed in 
C. nutans. Similarly, the anchoring filaments are feebly developed 



1967 



HYDROID OF VANNUCCIA FORBESII 



9 



in Hypolytiis obvoluta, become more apparent in Vannuccia jor- 
besii, and are strongly developed in Corymorpha nutans. 

In summary, the genus Vannuccia is related to the genera 
Hypolytus and Corymorpha, representing an intermediate stage be- 
tween them. Medusa and hydroid of Vannuccia forbesii are de- 
cidedly different from those of either genus, so that the new genus, 
Vannuccia may be justified. 



255 m 




Figure 6. Vannuccia forbesii, base of hydranth stem. The rooting fila- 
ments form a dense tuft, but in order to make the figure clearer only a few 
of them are drawn in this figure. A, Gelatinous perisarc; B. ectoderm; C, 
endodermal ridge; D, developing rooting filament; E, rooting filament. 



The genus Vannuccia is closer to Corymorpha than to Tubularia. 
If the family Tubulariidae is divided into two families, the Cory- 
morphidae and Tubulariidae (Rees, 1957; Brinckmann-Voss, MS 
in preparation), the genus Vannuccia would have to be placed in 
the Corymorphidae. 



10 BREVIORA No. 263 

ACKNOWLEDGMENTS 

I wish to thank Mrs. Sofia Giaquinto-Boag for her most efficient 
help in maintaining the polyps, and the director of the Naples 
Zoological Station, Dr. P. Dohrn, for his generous support. 



REFERENCES CITED 

Allman, G. J. 

1872. A monograph of the gymnoblastic or tubularian hydroids. Ray 
Society, London: 1-450, 84 figs., 32 pis. 

Kramp, p. L. 

1933. Coelenterata, Ctenophora and Chaetognatha. The Scoresby 

Sound Comm. 2nd East Greenland Exped. in 1932. Meddel. 

omGronland., 104(11): 1-20. 
1949. Origin of the hydroid family Corymorphidae. Vidensk. Medd. 

fra Dansk Natur. Foren., Ill: 183-215. 
1959. The Hydromedusae of the Atlantic Ocean and adjacent waters. 

Copenhagen. Dana Report No. 46: 1-283. 
1961. Synopsis of the medusae of the world. J. Mar. Biol. Assoc. 

United Kingdom, 40: 1-469. 

Mayer, A. G. 

1894. An account of some medusae obtained in the Bahamas. Bull. 

Mus. Comp. Zool., Harvard College, 25(11): 235-242, 3 pis. 
1910. Medusae of the world. Part I, Hydromedusae. Carnegie Inst. 

Washington, Publ. 109(1): 1-230, 29 pis. 

MURBACH, L. 

1899. Hydroids from Wood's HoU, Mass. Hypolytiis peregrinus, a 
new unattached marine hydroid. Quart. J. Microscop. Sci., 
(N.S.) 42: 341-360. 

Nair, K. K. 

1951. Medusae of the Trivandrum Coast, Part I, Systematics. Bull. 
Centr. Res. Inst. Univ. Travancore (Trivandrum), 2 C (Nat. 
Sci.) No. 1: 47-75. 

Rees, W. J. 

1957. Evolutionary trends in the classification of capitate hydroids 
and medusae. Bull. Brit. Mus. (Nat. Hist.), Zool., 4(9): 
453-534. 

UCHIDA, TH. 

1927. Studies on Japanese Hydromedusae. I, Anthomedusae. J. Imp. 
Univ. Tokyo, Sect. 4. Zool., 1: 145-241, 47 figs., 2 pis. 



(Received 26 September, 1966.) 



BREVIORA 

Mmseiiim of Comparsitive Zoology 

Cambridge, Mass. 6 April, 1967 Number 264 

THE CHANARES (ARGENTINA) TRIASSIC REPTILE FAUNA 

III. TWO NEW GOMPHODONTS, 
MASSETOGNATHUS PASCUALI AND M. TERUGGII 

By Alfred Sherwood Romer 



In previous numbers of this series an account was given of the 
1964-1965 expedition of the Museo de la Plata and the Museum 
of Comparative Zoology to the Permo-Triassic of western Ar- 
gentina, and the Permo-Triassic geology of the Chanares-Gualo 
region was briefly described (Romer, 1966; Romer and Jensen, 
1966). This and succeeding papers will be devoted to descrip- 
tions of the new genera and species contained in the Chaiiares 
fauna. 

Owing to a sequence of political incidents which need not be 
detailed here, the greater part of the collection has only recently 
(December 1966) reached the Museum of Comparative Zoology 
after the lapse of nearly two years. The one box of specimens 
which reached Cambridge earlier and is now partially prepared, 
contains, however, a considerable variety of reptilian remains, par- 
ticularly skulls and jaws. Represented, in addition to indeterminate 
fragmentary materials, are a small pseudosuchian, a moderately 
large dicynodont with powerful tusks, two carnivorous cynodonts, 
and at least four (possibly more) gomphodont cynodonts. For 
most of these forms, the materials at hand are none too complete, 
even as regards crania and dentitions, and it hence seems wise to 
defer description until the arrival of the main body of the collec- 
tion, in which further specimens are almost surely present. Two of 
the gomphodonts, however, are represented by a number of well- 
preserved skulls and jaws, and will be described here as Masse- 
tognathus pascuali and M. teruggii. As will be seen from the 
descriptions and illustrations, the two are comparable in many 
ways, and are obviously closely allied. The skulls assigned to 
M. teruggii are on the average about 43 per cent larger than 



2 BREVIORA No. 264 

those of M. pascuali, and the differences between the two are in 
great measure differences which might reasonably be associated 
with the contrast in size. I was at first inchned to beheve that we 
were possibly dealing with growth stages of a single form. How- 
ever, the skulls sort clearly into two size groups, rather than form- 
ing the graded series expected had they represented growth stages 
of a single species. The difference in size between the two is con- 
siderably greater than that expected in sex differences, and it is 
hence reasonable to believe that we are dealing with two species 
of a single genus. 

Massetognathus gen. nov. 
Type species: M. pascuali sp. nov. 

Diagnosis. Traversodontid gomphodonts of modest size, known 
skull lengths (snout to condyles) ranging from 82 to 138 mm. 
The skull is short and broad, the width about seven-tenths the 
length to condyles. Muzzle expanded laterally so that the maxilla 
is broadly exposed ventrally lateral to the cheek tooth row. As 
in gomphodonts generally, the zygomatic arch is broad but with 
a constricted posterior base; the arches widely expanded for 
their full length in a line parallel to the long axis of the skull; the 
epipterygoid-pterygoid bar extending broadly back to the quadrate 
region; the angular process of the dentary well developed and 
thickened at its ventral border; posterior to this, the angular-sur- 
angular well exposed laterally. No suborbital process of the jugal; 
external exposure of the squamosal on the zygomatic arch re- 
stricted to upper and posterior borders; no posterior projection of 
the angular process of the dentary. Four upper, three lower in- 
cisors; canines modestly developed; 12-15 upper, 11-13 lower 
cheek teeth; anterior teeth smaller and simpler in structure than 
main series, but no distinct "premolar" series. Teeth of typical 
traversodont pattern; upper "molars" with two lateral cusps, the 
more posterior one the more prominent; a high posterior transverse 
ridge with cusps close to and at the posteromedial corner. Teeth 
subquadrate. but extending farther laterally toward the posterior 
corner; anterior and posterior margins essentially transverse and 
parallel, but slightly convex in outline anteriorly toward the ex- 
ternal margin and similarly concave posteriorly, so that each tooth 
"shoulders" to a slight degree into the area of its next anterior 
neighbor. The generic name (masculine) refers to the obviously 
excellent chewing powers of the jaws. 



1967 TWO NEW GOMPHODONTS 3 

Massetognathus pascuali sp. nov. 

Holotype. No. 65-XI-14-1, Museo de la Plata, skull and jaws, 
collected by the 1964-1965 expedition of the Museo de la Plata 
and the Museum of Comparative Zoology. 

The specific name is in honor of Dr. Rosendo Pascual, Pro- 
fessor of Paleontology in the Universidad de la Plata, who accom- 
panied the expedition during much of its stay in western Argentina, 
and was exceedingly helpful to us before, during, and after our 
field work. 

Horizon and locality. From an exposure in the Triassic Cha- 
nares Formation about two miles north of the point where the 
Chafiares River debouches into the Campo de Talampaya, in west- 
ern La Rioja Province, Argentina. 

Diagnosis. A small form, with a mean length, snout to con- 
dyles, of about 87 mm in available specimens. Posterior portion of 
frontals and anterior portion of parietals form a flat triangular dor- 
sal area, the posterior end of the triangle extending back to a 
parietal foramen of modest size; the postorbitals extend back on 
either side to a point about opposite the parietal foramen. The 
angular process of the dentary descends to form approximately a 
right angle with the ventral margin of the bone. Twelve upper 
and 1 1 lower cheek teeth in available specimens. 

Description. Massetognathus pascuali appears to be an exceed- 
ingly common member of the Chafiares fauna; even in the re- 
stricted material currently available from our collection there are 
some seven fairly good skulls, as well as less complete skull and 
jaw specimens and considerable postcranial material. This is a 
relatively small animal, the mean length of available specimens, 
from premaxilla to occipital condyle (allowing for minor anterior 
or posterior deficiencies), being 87.3 mm, with a range from 82 
to 95 mm. As noted above, the first thought that these specimens 
might be growth stages leading to M. teruggii appears to be negated 
by the fact that the series here assigned to M. pasuali seems to clus- 
ter closely about the size mean, with an average deviation of but 
3.6 mm, or about 4 per cent. 

The general nature of the skull is obvious from Figures 1-3. Its 
structure is for the most part that familiar in advanced cynodonts 
generally and gomphodonts in particular, and in many regards does 
not call for detailed description. In its general proportions it is 
relatively broad and short. The seeming shortness is exaggerated by 
the most distinctive feature of the genus — the "swollen" muzzle. 
As in all typical gomphodonts, the two tooth rows of the cheek 



4 BREVIORA No. 264 

battery are close together on the ventral surface of the skull. In 
most gomphodonts there is little lateral extension of the muzzle 
beyond the line of the tooth row, and (as may be seen, for ex- 
ample, in Diademodon or Exaeretodon) the snout region is slen- 
der. Here, however, the maxillae extend far out dorsally (with a 
downward slope) to a point about opposite the lower margin of the 
orbits, then, curving downward and inward, present a broad ven- 
tral surface lateral to the tooth rows. 



Figure 1. Lateral view of the skull of Mossetognathits pascuali. Natural 
size. 



The skull is low, and the orbits face somewhat more dorsally 
than laterally and slightly anteriorly. The prefrontal and post- 
orbital meet broadly above the orbits. The nasals and frontals 
form an essentially flat dorsal platform atop the skull. On either 
side, the prefrontals and postorbitals are somewhat raised above 
the flat surface of the frontals and anterior part of the parietals. 
Posterior to the orbits, the ridges formed by the postorbitals con- 
verge to leave a triangular wedge of flat dorsal surface formed 
by the back portion of the frontals and the anterior portion of the 
parietals. This narrows posteriorly, with the rather small parietal 
foramen at the apex of the triangle. Posterior to this, the con- 
joined parietals form a sharply developed parietal crest. The post- 
orbitals on either side extend backward, sheathing the parietals 
laterally, to about the level of the parietal foramen. 

The lateral surfaces of the braincase appear to be constricted in 
typical cynodont fashion; I shall postpone description of this region 
and the interior of the orbit until further material is available for 
"dissection" and sectioning. The occipital surface of the skull is of 
normal cynodont construction. Laterally, as in gomphodonts gen- 
erally, but in contrast to other cynodonts, the squamosal descends 



1967 



TWO NEW GOMPHODONTS 





Figure 2. Dorsal and ventral views of the skulls of Massetognathiis pas- 
citali. Natural size. 



6 BREVIORA No. 264 

ventrally so that the connection between the portion of the bone 
forming the lateral wing of the occipital region and that associated 
with the zygoma is only by a narrow neck ventrally; in posterior 
view, the dorsal margins of the two portions form a V, widely 
open above. The pterygoid-epipterygoid bar reaches broadly back 
to the quadrate region above a pterygo-paroccipital fenestra. The 
squamosal forms the boundaries, medial and lateral, of a distinct 




Figure 3. Posterior view of the skull of Massetognuthus pascuali. Natu- 
ral size. 



external auditory meatus. The postorbital bar is slender; the zygo- 
matic arch, however, is deep dorsoventrally. In contrast to cynog- 
nathids, in which the arch slants diagonally outward and backward 
from the orbital region to attain its greatest length posteriorly, the 
arch, as in gomphodonts generally, attains its full breadth at the 
posterior margin of the orbit, and runs back from this point parallel 
to the main axis of the skull, allowing a high development of the 
temporal musculature. The external exposure of the squamosal on 
the arch is restricted to a narrow area at the top of the arch, 
and, curving downward posteriorly, along the anterior margin of 
the external meatus. 

Ventrally, the two rows of cheek teeth are close together an- 
teriorly, and diverge gently posteriorly. The secondary palate ex- 
tends back most of the length of the tooth row and (although the 
sutures are none too certain) the palatines appear to form the most 
posterior portion of the secondary palate. Back of the secondary 
palate, little of sutures can be made out in the material available. 
On either side, the lateral margins of the pterygoids extend back- 
ward as raised ridges which terminate in sharply-pointed flanges di- 
rected downward and backward. From the median bar formed 
posteriorly by the conjoined pterygoids and basicranial axis, a pair 
of ridges run diagonally forward and laterally toward the posterior 
ends of the tooth rows, and a median ridge runs directly forward. 



1967 TWO NEW GOMPHODONTS 7 

The posterior ventral structures are of the type commonly seen in 
cynodonts, with an excavated basioccipital-sphenoid area bounded 
by a pair of ridges; on either side, there is a well-developed vagal 
foramen, and farther forward, the fenestra ovalis. A V-shaped 
channel, bounded medially by the braincase wall and laterally by 
the pterygoid-epipterygoid bar, represents the primitive cranio- 
quadrate passage; anteriorly it is roofed below by a median ex- 
tension of the pterygoid. In most specimens the quadrate (plus 
quadratojugal, loosely attached to the skull) has dropped out, 
leaving a pair of notches to represent its area of articulation. In 
the jaw the dentary is highly developed, with a high and broad 
coronoid process, a posterior process which reaches back nearly 
to the articular area, and a pronounced angular process of the 
dentary which is thickened and rounded ventrally. Posteriorly, in 
contrast to non-gomphodonts, the angular and surangular are well 
exposed on the outer surface. The elements of the jaw apart from 
the dentary are incompletely preserved in the available material, 
and I shall reserve description of the inner surface of the mandible 
for a future occasion. 

There are four upper and three lower incisors of modest size, 
somewhat chisel-shaped and with longitudinal striations. The 
canines are relatively little developed. The cheek teeth are sep- 
arated from the canines by a short diastema; both above and be- 
low, the members of each cheek series (Figs. 4, 6) are crowded 
close to one another, and the two rows, rather close together in 
front, curve gently outward posteriorly. As is now well known in 
other gomphodonts (Crompton, 1955, etc.), the composition of a 
gomphodont cheek battery may change during "adult" life by the 
eruption of additional teeth posteriorly and resorption of anterior 
ones, so that no precise tooth count is possible. In our material 
there is no positive evidence of anterior resorption, but in most 
specimens there is evidence of posterior addition in the presence, at 
the back end of the battery, of either a newly erupted but unworn 
tooth, one well formed by not yet erupted, or a partially developed 
tooth still in its alveolus. In the specimens currently available to 
us there are generally 12 maxillary teeth, and 1 1 in the only two 
mandibles now in our possession. Although, in contrast to dia- 
demodonts, there is no sharp contrast between small "premolars" 
and a "molar" series, the more anterior cheek teeth are smaller 
and with a simpler pattern. The wear on the teeth is obviously 
great, so that little detail of the crown pattern can be seen except 
in teeth at the back end of the series which are as yet unerupted or 
very freshly erupted. 



BREVIORA 



No. 264 





Figure 4. Left (above) and right (below) upper cheek tooth series of a 
specimen of Mussetognathiis pasciioli. X 3. 



Such a tooth from the maxillary series is shown in Figure 5. In 
surface view the crown is subquadrate, nearly twice as broad as 
long anteroposteriorly. The inner margin is curved; anterior and 
posterior margins are nearly straight and parallel to one another, 
except that towards the outer border the anterior margin is some- 
what convex, the posterior border opposite somewhat concave. Each 
tooth thus "shoulders" somewhat into the proper area of its next 
anterior neighbor — a feature which is much more pronounced in 
the more advanced traversodonts of the later Ischigualasto horizon. 
On the outer border, the base of the cone forming the postero- 
external cusp extends considerably outward and backward, so that 
the external boundary is a diagonal line. The outer portion of the 
tooth is a high anteroposterior ridge, on which a major cusp de- 
velops posteriorly; a minor cusp, only slightly separated from it. 



1967 



TWO NEW GOMPHODONTS 



is developed along the downward anterior slope of the ridge. Me- 
dial to the two cusps, the external ridge descends abruptly into 
a basin which occupies most of the crown area. This basin, apart 
from its straight lateral border, is an essentially oval concave 
area, without internal markings. It is surrounded on posterior, 
medial, and anterior borders by a curved ridge, considerably 
higher along the posterior margin than medially and anteriorly. An 
elongate cusp is present about two-thirds the way in along the pos- 
terior ridge, a second cusp near the posteromedial corner, and a 
lower cusp part way along the anterior ridge. In a freshly erupted 
tooth a series of tiny bead-like denticulations may be seen along 




Figure 5. A right upper "molar" of Massetognathus pasciiali. A, Crown 
view. B, Posterior and somewhat ventral. C, Medial. D, Anterior and 
somewhat dorsal. E, Lateral. X 4. 



most of the extent of the ridges surrounding the basin posteriorly, 
medially, and anteriorly, and a few may even be present along 
the external ridge. With wear, the denticulations rapidly disap- 
pear; with somewhat further wear, the anterior ridge is ground 
down, leaving the posterior ridge projecting, but with distinction 
between its two cusps lost, and the internal ridge still prominent. 
With still further wear, the posterior ridge, too, becomes worn 
away, and the basins of successive teeth become a continuous sur- 
face, flat anteroposteriorly but concave as seen in end view, 
rising medially to a persistent medial ridge, and rising still higher 
laterally to the outer longitudinal ridge. 

The general position of the lower cheek series corresponds to 
that above. (Because of imperfections in available specimens of 
M. pascuali, I have shown instead, in Figure 6, that of M. teruggii.) 
As in the upper jaw, the most anterior cheek teeth are smaller and 
simpler in pattern than the main series, although there is no sharp 
division into "premolars" and "molars." Just as in the case of 



10 



BREVIORA 



No. 264 




Figure 6. Right cheek tooth series of a specimen of Massetognathiis ter- 
iiggii. X 2. 

the upper "molars," wear tends to obliterate rapidly most of the 
details of the crown pattern. A freshly erupted tooth is shown in 
Figure 7. The lower "molars" are of the diagnostic traversodont 
pattern. This pattern, as regards position in the jaw, gives them 
a build contrasting sharply with those of the upper jaw. Curi- 
ously, however, if one imagines a lower tooth rotated 90°, so that 
the anterior border faces, instead, to the outside, the pattern be- 
comes highly comparable to that of the upper tooth. The lower 
cheek teeth are subquadrate in shape. The anterior portion forms 
a high transverse ridge, partially subdivided into two cusps, of 
which the more external is the larger and higher. Just as the ex- 
ternal ridge of the upper tooth descends steeply on its inner sur- 
face to the basin of the tooth, so here the posterior face of the 
anterior transverse ridge descends abruptly into the subquadrate, 
smoothly concave tooth basin. As in the case of the upper tooth, 
the basin is surrounded by a continuous ridge on the other three 




Figure 7. A right lower "molar" of Massetognathiis teriiggii. A , Crown 
view. B, Lateral. C, Anterior. D, Medial. E, Posterior. X 4. 



1967 TWO NEW GOMPHODONTS 11 

sides; the external ridge is much the highest, the internal the low- 
est. As in the margins of the upper basin, so here a nearly continu- 
ous series of bead-like cuspules is present in the unworn tooth, 
but in contrast to the upper teeth, no conspicuous cusps are present, 
even in the unworn tooth. With wear, the denticulations soon dis- 
appear, and except for the strong anterior cross-ridge, the tooth 
is reduced to a somewhat concave basin with a somewhat raised 
rim externally. 

Massetognathus teruggii sp. nov. 

Holotype. No. 65-XI-14-2, Museo de la Plata, skull and jaws, 
collected by the 1964-1965 expedition of the Museo de la Plata 
and the Museum of Comparative Zoology. 

The species is named for Professor Mario Teruggi, who was 
most helpful in the carrying out of our field work and in our post- 
expeditionary difficulties. 

Horizon and locality. From the same horizon and locality as 
the type of M. pascuali. 

Diagnosis. A larger species than M. pascuali, with a mean 
skull length of about 125 mm in available specimens. The flat- 
tened dorsal area formed by the frontals and anterior portions of 
the parietals is more restricted than in M. pascuali, the roof con- 
stricting to a sagittal crest in front of the parietal foramen, which 
is reduced to a tiny slit. The postorbitals extend backward on 
either side of the crest to a point well back of the foramen. The 
angular process is not as sharply developed as in M. pascuali, the 
posterior and ventral borders of the process meeting at an obtuse 
angle. Thirteen to 15 maxillary, and 11 to 13 mandibular cheek 
teeth in available specimens. 

Description. M. teruggii seems, next to M. pascuali, to be the 
most common of Chanares reptiles; our present restricted materials 
contain some five skulls of this form. As noted above, this species 
is considerably larger (more than 40 per cent) than M. pascuali. 
This is, of course, a considerably greater difference than would be 
expected if we were dealing with sex differences. The skulls at- 
tributed to M. teruggii, like those of M. pascuali, seem to form a 
compact series, ranging from 115 to 138 mm in length, with an 
average deviation from the mean of about 9 mm, or about 7 per 
cent only. 

The skull of this species (Figs. 8-10) is very similar in most 
regards to that of its associate, M. pascuali. Since we have de- 
scribed, above, the skull of this smaller form in some detail, we 



12 



BREVICRA 



No. 264 




Figure 8. Laterial view of the skull of Massetognathus teruggii. X %. 



can here avoid unnecessary and monotonous repetition of descrip- 
tion of their numerous similar features, and call attention only to 
the few points of difference. These are in part associated with size. 
As is, I think, well known (and should be obvious), the chewing 
apparatus of a large form must be disproportionately large as com- 
pared with that of a relative of smaller size. Here the differences in 
this feature have to do with a somewhat greater development of 
a sagittal crest. This is accomplished by a greater forward develop- 
ment of the crest, so that the flattened area of the top of the skull 
is "pinched in" to a greater degree than in M. pascuali, and a 
median ridge is, in contrast, formed anterior to the area of the 
parietal foramen, which is reduced almost to the vanishing point. 
The posterior flanges of the postorbitals which clamp in upon 
either side of the parietals here reach farther back than is the case 
in M. pascuali, to extend to a point well back of the parietal 
foramen. As may be seen from Figure 8, the angle of the dentary 
is less sharp than in M. pascuali, the posterior border of the dentary 
sloping downward and forward at an angle, rather than descending 
vertically to the apex. As noted, the tooth count of the cheek bat- 
tery is somewhat higher than in M. pascuali. I find no essential dif- 
ference in the pattern of the "molars" in unworn teeth of the 
two species. 



DISCUSSION 

"Gomphodont" cynodonts, with a masticatory series of cheek 
teeth, were early discovered in the Cynognathus Zone of the early 
Triassic of South Africa. Diademodon (of which Gomphognathus 
and several other generic names are synonyms) is the best known, 
and has been well described by Watson (1911, 1913), Broili and 



1967 



TWO NEW GOMPHODONTS 



13 





Figure 9. Dorsal and ventral views of the skull of Massetognathiis ter- 
uggii. X % . 



14 



BREVIORA 



No. 264 




Figure 10. Posterior view of the skull of Massetognathus teruggii. X %. 



Schroder (1935), and most recently by Brink (1955, etc.); 
Thrachodon, Protacmon, Trirachodontoides, Gomphodontoides, 
and Inusitatodon are less well known forms from the Cynognathus 
Zone; a recent discovery is Cragievarus (Brink, 1965). 

Until recent decades, no remains of gomphodonts were known 
in other areas or in later Triassic horizons, and it was assumed 
that the gomphodonts were a short-lived and unimportant branch 
of the cynodont stock. This, however, was due to our lack of 
adequate knowledge of Middle Triassic faunas. Explorations from 
the nineteen-thirties onward, of beds of this age in South Amer- 
ica and Africa, are now revealing the fact that the gomphodonts 
long remained a flourishing group which formed a major com- 
ponent of the Middle Triassic faunas — at least those of the south- 
ern continents. 

From the Santa Maria beds of southern Brazil, von Huene 
(1928, 1942) described Gomphodontosuchus and Traversodon. 
From collections made in the Manda beds of East Africa by Par- 
rington and for von Huene have been described Theropsodon 
(Huene, 1950), and Aleodon, Cricodon, and Scalenodon (Cromp- 
ton, 1955). In South Africa, the Middle Triassic Molteno beds, 
long considered barren, have begun to yield fossils to Crompton, 
including a gomphodont Scalenodontoides (Crompton and Ellen- 
berger, 1957). Brink ( 1963) has recently described a Diademodon 
specimen from Northern Rhodesia at a level which he considers 
equivalent to the Cynognathus Zone, and a new gomphodont. 
Luangwa, from a horizon in the Ntawere Formation which is prob- 
ably a Molteno equivalent. Large gomphodonts, not yet described, 
are said to persist into the lower part of the Upper Triassic Red- 
beds of Basutoland (Crompton, 1964). A recent British expedi- 
tion resulted in the discovery of further gomphodont material in 
Northern Rhodesia and Tanganyika. In Argentina a visit by Fren- 
guelli to the Middle Triassic deposits of Ischigualasto produced a 



1967 TWO NEW GOMPHODONTS 15 

few scraps of gomphodont skull to which Cabrera (1943) gave 
the name Exaeretodon and Theropsis. Exploration of this region 
by a Harvard-Buenos Aires Museum expedition in 1958 revealed 
the presence of a rich deposit of Middle Triassic vertebrates. As a 
result of further extensive collections made by the Instituto Lillo 
of Tucuman, Bonaparte (1962, 1963c) has been able to give a 
comprehensive description of Exaeretodon (with which Theropsis 
proves to be synonymous), and has described two further genera of 
gomphodonts, Proexaeretodon and Ischignathus (Bonaparte, 
1963d, 1963a). Sr. Bonaparte informs me that he has under de- 
scription a small diademodont gomphodont from the early Triassic 
Puesto Viejo Formation of southern Mendoza Province. 

According to Dr. Donald Baird, a gomphodont, as yet unde- 
scribed, is present in the lower beds of the Newark series of Nova 
Scotia — the only gomphodont as yet reported from the Northern 
Hemisphere. 

Most recent of gomphodont-bearing faunas to be discovered is 
that of the Chanares Formation. Forms of this nature are here 
exceedingly abundant, and, as seen in the field, appear to make 
up a very large proportion (? one-half) of the total number of 
specimens recovered. In the small fraction of our collection now 
available for study there appear to be at least two further distinct 
genera present, in addition to the forms described above. 

The gomphodonts are obviously an herbivorous side branch of 
the advanced cynodont stock. The circumstances which led to this 
radical departure from the typically carnivorous mode of life of 
other cynodonts and, indeed, of all other theriodonts generally, are 
far from clear. In the late Permian a host of dicynodonts, large 
and small, were the typical herbivores of the times; as anyone 
who has collected in the Endothiodon or Cistecephalus zones of 
South Africa knows, 90 per cent or so of all specimens found in 
these beds are members of the dicynodont group. By the time we 
reach the Cynognathus Zone, the dicynodonts are much reduced in 
numbers and variety, and survive then and later in the Triassic 
only in the form of such large types as Kannemeyeria, Ischigual- 
astia, and Plocerias. In such typical Middle Triassic faunas as 
those of Ischigualasto and Santa Maria, the common herbivores 
are, instead, of two types in about equal numbers — rhynchosaurs 
and gomphodonts. In the Chafiares fauna we are dealing with 
an earlier Meso-Triassic stage. Rhynchosaurs must have been in 
existence (a few ancestral types are found in the Cynognathus Zone 
fauna), but presumably were little developed. (We have found no 
trace of a rhynchosaur in the sample of the collection currently 



16 BREVIORA No. 264 

available to us, and saw no sure rhynchosaur material while col- 
lecting.) Gomphodonts are extremely abundant and apparently 
varied and, in the way of percentages, occupy much the same posi- 
tion in the Chanares that the dicynodonts did in the Upper Per- 
mian. Seemingly, it was only later in the Middle Triassic that 
rhynchosaurs became fully evolved and came to rival the gompho- 
donts in numbers. 

Judging by the very considerable number of new forms and 
the amount of new data unearthed during the last few years, we 
are currently undergoing, so to speak, an explosive period in our 
acquisition of knowledge concerning gomphodonts. For this rea- 
son, and because our knowledge of many of the forms concerned 
is very incomplete, it is inadvisable at this time to attempt any 
comprehensive study of gomphodont classification and phylogeny. 
Pertinent facts, however, may be discussed. 

In 1954 Haughton and Brink included all African forms then 
known in the Diademodontidae. In our 1956 classification of 
therapsids, Watson and I included all known gomphodonts except 
Gomphodontosuchus in the single family Diademodontidae, and 
I followed the same pattern in my "Osteology of the Reptiles," 
published the same year. Lehman, in 1961, in the Piveteau 
"Traite" also included all gomphodonts (including Gomphodonto- 
suchus) in the Diademodontidae, although effecting a subdivision 
into subfamilies.^ Von Huene, however, had erected a second fam- 
ily, Traversodontidae, for the Brazilian genera Traversodon and 
Gomphodontosuchus, and Crompton and Ellenberger advocated a 
division of the gomphodonts into two major groups, Diademodon- 
tidae and Traversodontidae. 

Study of South American gomphodonts brought me in 1961, in 
contrast to my earlier opinion, to the conclusion that diademodonts 
and traversodonts (to which Exaeretodon obviously belonged) 
were distinct groups, but also to feel that the Trirachodon-Cricodon 
tooth type indicated the presence of a third group. Bonaparte 
(1963b) advocated two families, but {contra Crompton) would 
include Gomphodontosuchus, as well as the newly discovered 
Ischigualasto genera, in the Traversodontidae. Brink (1963) 
would unite these families into a superfamily Gomphodontoidea 
{meUor Diademodontoidea?) of advanced cynodonts, in contrast 
to the carnivores — superfamily Cynognathoidea. 



1 His term "Eudiademodontinae" is, of course, inadmissabie, not being 
based on the name of a genus. 



1967 TWO NEW GOMPHODONTS 17 

Actually, it would seem, on the basis of present knowledge, that 
three, rather than two, groups of gomphodonts are present in the 
Lower and Middle Triassic. 

DiADEMODONTIDAE 

Here I would place a series of Cynognathus Zone forms typified 
by Diademodon, and a series of named genera identical or closely 
related, including: Cyclogomphodon, ?Cynochampsa, Diastemo- 
don, Gomphodontoides, Gomphognathus, Microhelodon, Octagom- 
phus, and Protacmon. The entire anatomy is, as mentioned above, 
adequately known only in Diademodon. 

The skull is relatively long, as in typical cynognathids, with a 
rather long and slender facial region, and with the greatest width 
toward the posterior end. The squamosal is deeply incised medial 
to the external auditory meatus, giving a V-shaped dorsal outline 
to the bone in posterior view; this is quite in contrast to the condi- 
tion in primitive cynodonts and even cynognathids, in which the 
two portions of the squamosal are broadly connected at the pos- 
terior margin of the temporal fenestra. In cynognathids the skull 
shape, as seen from above, is that of an isosceles triangle, the 
breadth gradually increasing backward to its maximum at the 
posterior end of the zygoma, whereas in diademodonts there is 
a major increase in width at the orbits, so that the zygomatic 
arches lie nearly parallel to the main axis of the skull for their 
entire lengths. The arches are deep, and are mainly formed by the 
jugals; the squamosals have but a narrow external exposure pos- 
teriorly and dorsally. A peculiar feature is a prominent ventral 
flange below the orbit formed by the jugal. In contrast to some 
cynognathids there persists in Diademodon (but not in Protacmon) 
a primitive condition in that the pterygoid-epipterygoid bar extends 
back to the quadrate. Certain of the features here listed for 
Diademodon appear to be present in gomphodonts generally; how- 
ever, the proportions of the "muzzle'' are variable, the ventral 
jugal flange may be absent in some groups, and a broader external 
exposure of the squamosal is reported in some cases. 

Four incisors and a stout canine are present above and below; 
beginning after a short diastema, the diademodont dentition in- 
cludes a long row of cheek teeth on either side, the two tooth rows 
diverging with an outward curvature posteriorly. Thirteen or more 
cheek teeth may be present; here and in at least certain gom- 
phodonts, teeth may be added posteriorly to the cheek battery 
during "maturity," and in Diademodon small anterior teeth may 



18 BREVIORA No. 264 

be lost and their alveoli closed. The most anterior cheek teeth are 
peg-like, the most posterior retain a primitive, laterally compressed 
shape like that of carnivorous cynodonts. The main portion of 
the dental battery consists of gomphodont "grinders." Here, as 
in other gomphodonts, there is a strong tendency for the de- 
velopment of rows of small bead-like denticulations along the 
ridges and cusps of the "molars"; however, these rapidly disappear 
with wear and are seldom seen, except in teeth in process of erup- 
tion. The evolution of the typical "molars" appears to have taken 
place by an inward growth, from the original compressed carnivore 
tooth type, of a "heel" overlapping the lower teeth. The original 
longitudinal tooth line is represented by two stout cusps along 
the outer margin; variable cusps may be developed medially. There 
may be some development of small cusps along anterior and pos- 
terior margins. A low ridge develops crossing the center of the 
"heel" from side to side; this, when unworn, shows a row of tiny 
denticles, but never a developed cusp. The lower "molars" are 
circular to subquadrate in shape, with a ring of cusps and cuspules 
surrounding a central basin. There is some suggestion of compari- 
son with the upper "molars" in that the marginal cusps tend to be 
more highly developed on the medial and lateral margins, and there 
is some development of a transverse ridge crossing the basin. 

The diademodont tooth type is, as far as known, mainly con- 
fined to forms from the Cynognathus Zone of South Africa. How- 
ever, Crompton (1955: 656-659, fig. 14 A-D) has described frag- 
mentary jaws from the Manda beds with mandibular "molars" of 
diademodont pattern. Further, the imperfect skull and jaws from 
these same beds which form the type of Aleodon brachyrhamphus 
(Crompton, 1955) are suggestive, as Crompton notes, of diade- 
modont relationships, but the teeth are so worn that almost noth- 
ing can be made of their pattern. 

Trirachodontidae 

Trirachodon and Trirachodontoides of the Cynognathus Zone 
and Cricodon of the Manda beds are forms with a dentition of 
very distinctive type. There are ten to eleven cheek teeth in Cri- 
codon, but only six reported in Trirachodon. The cheek tooth 
rows are nearly straight, but with a slight outward curvature pos- 
teriorly in Cricodon. In this genus the most anterior cheek teeth 
are small subcircular "premolars" and, at least in the lower jaw, 
the most posterior teeth, as in Diademodon, "revert" to the narrow 
cynognathid type. In the Cynognathus Zone forms the lower 



1967 TWO NEW GOMPHODONTS 19 

"molars" are unknown. The upper cheek teeth of Trirachodon and 
both uppers and lowers of Cricodon are transversely broadened, 
but narrow anteroposteriorly. There is a single internal cusp and 
a single external one, and connecting the two a transverse ridge 
which with wear develops a prominent longitudinal cusp. 

It seems clear that this tooth type has developed independently 
of that of the diademodonts, by medial lateral development from 
an ancestral type with but a single major cusp; the diademodont 
type, on the other hand, expanded laterally from a tooth with 
greater anteroposterior length. Both Crompton and EUenberger 
(1957) and Bonaparte (1963d) would include these forms in the 
Traversodontidae. But while the upper "molars" of typical traver- 
sodonts might perhaps be derivable from the type seen in the pres- 
ent group, I think it is impossible to conceive of the lower "molars" 
of the traversodonts having been derived from the radically differ- 
ent type seen in Cricodon and presumably present in Trirachodon. 

A minor but perhaps significant feature suggesting at least a 
distant trirachodont-diademodont relationship is the presence in 
Trirachodon of a suborbital process of the jugal comparable to that 
of Diademodon and Protacmon. 

Probably of no significance is the fact that the fragment of skull 
from the Upper Triassic Redbeds named Tritheledon by Broom 
(1912) shows a row of transversely widened multicuspidate teeth, 
somewhat suggestive of a trirachodont. 

Traversodontidae 

At present this family (as here restricted) definitely includes two 
African forms — Scalenodon of the Manda beds and Scalenodon- 
toides of the Molteno — and a series of South American forms — 
Traversodon from Santa Maria, Exaeretodon, Proexaeretodon, and 
Ischignathus from Ischigualasto, and from the Chafiares the two 
species of Massetognathus described above, as well as other forms 
to be described later. Many of these are incompletely known — 
Scalenodontoides from a lower jaw alone. Scalenodon from numer- 
ous but fragmentary remains, Traversodon from isolated elements 
from which von Huene has restored a skull; Proexaeretodon and 
Ischignathus are represented by incompletely preserved skulls. As 
yet Exaeretodon and Massetognathus are the only traversodontids 
in which adequate skull material is known. Considerable variation 
in skull structure appears to have been present in the group. All 
appear to have had, as seemingly in gomphodonts generally, zygo- 
matic arches placed far laterally in a plane parallel to the long 



20 BREVIORA No. 264 

axis of the skull. In general the muzzle is narrow; only in Masse- 
tognathus, as far as known, do we find a "swollen" maxillary 
region. In all forms where data are available, the angular process 
of the dentary is more pronounced than in other cynodonts (includ- 
ing Diademodon) and the angular-surangular more exposed lat- 
erally. The number of cheek teeth is variable — 8 maxillary teeth, 
for example, are figured in Scalenodon and Traversodon, 9 in 
Exaeretodon, 1 1 in Proexaeretodon, 7 in Ischignathus, 11 to 15, 
as noted, in Massetognathus. 

The "molars," and most especially those of the mandible, with 
a pair of prominent crests across the anterior border (behind which 
is a broad "heel"), are the most clearly diagnostic feature of the 
group. The mandibular teeth are not at all comparable with those 
of Cricodon (or those expected in Trirachodon), and offer a major 
reason for separating those genera from the Traversodontidae. 

As far as they are known in members of the South American 
fauna, the upper "molars" are nearly equally distinctive. There are 
here two external cusps, the posterior the major one; a prominent 
transverse ridge along the posterior margin, with a typical develop- 
ment of two cusps toward and at the internal edge; anterior to the 
posterior crest a concave basin, bounded in front and laterally by a 
ridge much lower than that behind. The cheek teeth are set closely 
in series, and even in Massetognathus each tooth "shoulders" 
slightly into the one ahead somewhat medial to the lateral border. 
This "shouldering" is much more pronounced in Proexaeretodon 
and Exaeretodon, so that in the latter genus the tooth appears to 
consist of two portions, external and internal, set at a considerable 
angle to one another. 

Of the two African forms, Scalenodon and Scalenodontoides, 
definitely assignable to the traversodonts, both exhibit lower 
"molar" patterns characteristic of the family. The upper dentition 
is unknown in the latter genus, but it is present and well described 
by Crompton in the former. The maxillary "molars" are basically 
comparable to those of South American forms, but differ in that 
there is but a single external cusp, without development of the 
accessory anterior one, and, further, in that the prominent trans- 
verse ridge, bearing two cusps toward and at the inner margin, lies 
not at the posterior border of the tooth but some distance anterior 
to it. These differences suggest that Scalenodon is rather more 
primitive than the described South American forms. Possibly we 
may see here a stage in the evolution of the traversodont upper 
"molar" from one resembling that of Trirachodon, but as remarked 



1967 TWO NEW GOMPHODONTS 21 

above, such an ancestry seems out of the question as regards the 
lower cheek teeth. 

However, there is evidence in the Manda beds of a form with 
more advanced upper "molars" — this is represented by a maxilla 
which Crompton (1955: 659-660, fig. 14 E) compared tentatively 
with the Brazilian Gomphodontosuchus, but which is obviously 
distinct (there are alveoli for at least ten cheek teeth, whereas the 
short-jawed Gomphodontosuchus has but six). The two "molars" 
preserved are somewhat worn, but show definitely the presence of 
two external cusps, a central basin, a low, crenulated, anterior 
transverse ridge, and a high posterior one. Further, there is a 
slight posterior marginal concavity on each tooth, indicating a be- 
ginning of the "shouldering" which became more prominent in 
some of the Argentinian genera. It appears probable that we are 
dealing with a form closely related to, and not impossibly belonging 
to the genus Massetognathus. Significant in this connection is the 
fact that in this specimen there is, as Crompton notes, a pro- 
nounced outward "bulge" of the maxilla lateral to the tooth row, 
as in Massetognathus but not reported in any other described 
gomphodont. 

Forms of doubtful assignment 

Apart from genera which seem clearly to belong to one or an- 
other of the three groups distinguished above, there are a number 
of forms which cannot be assigned, mainly because of lack of 
knowledge of molar pattern. 

Inusitatodon (Brink and Kitching, 1953) is a small Cynognathus 
Zone form represented by an incomplete skull. The jaws are 
tightly locked and the crown pattern of the cheek dental battery 

— which includes two "premolars" and apparently eight "molars" 

— is unknown. Brink and Kitching compare the skull with that 
of Diademodon in various regards, but, without giving reasons 
for this, state that Trirachodon is the closest relative. 

In the newly described Cragievarus (Brink, 1965) from the 
Cynognathus Zone, the cheek tooth pattern is, unfortunately, not 
described. The "molars" are subquadrate in outline, suggesting 
reference to the Diademodontidae, and Brink notes various points 
of agreement with Diademodon. However, the cheek tooth row 
is short (? nine teeth above, seven below), the skull lacks the 
sharp expansion in width at the orbits usual in gomphodonts, the 
zygomatic arch is weak, the jugal lacks the ventral process seen in 
Diademodon and Protacmon, and the angular process of the 
dentary is less developed than in typical gomphodonts. Possibly 



22 BREVIORA No. 264 

this form is a primitive but somewhat aberrant diademodontid. 

Luangwa (Brink, 1963: 89-93), from the Ntawere Formation of 
Northern Rhodesia, is obviously, from its expanded cheek teeth, a 
gomphodont, but the crown pattern is unknown. The shortness of 
its tooth row (only five "molars" and a miniscule "premolar") is 
unmatched, except in Trirachodon and Gomphodontosuchus , but 
the shape of its subquadrate "molars" is quite different from those 
of the former (narrow anteroposteriorly ) and from the triangular 
"molars" of the latter genus. The general skull pattern and the 
"incipient" ventral jugal process suggest the Diademodontidae; pos- 
sibly we are dealing with a juvenile. Brink states that its closest 
ally is Scalenodon, but gives no reasons. 

Aleodon from the Manda (Crompton, 1955) is, again, surely 
a gomphodont, but, again, we have almost no data on tooth cusps. 
The skull is little known. The rather "fat" shape of the "molars" 
suggests the Diademodontidae. 

Theropsodon of the Manda beds was described by von Huene 
(1950) on the basis of a nearly complete but poorly preserved 
skull. It is definitely a gomphodont, according to von Huene, but 
the tooth pattern cannot be determined. Possibly it may be identi- 
cal with either Cricodon or Scalenodon, from these same beds, or 
may represent a true diademodont, of which individual teeth are 
known to be present there. 

Gomphodontosuchus from the Brasilian beds is apparently a 
somewhat aberrant form, with a short face, short tooth row, and 
massive jaws. The crown pattern of the "molars" is imperfectly 
preserved. Watson and I, in 1956, assigned this form to a separate 
family; Crompton and Ellenberger (1957) assigned it to the diade- 
modonts; Bonaparte advocates a position in the Traversodontidae. 
The last is perhaps the best suggestion, but better material is 
needed. 

Bonaparte (1963b) has suggested that Belesodon of the Santa 
Maria beds might prove to be a traversodont. However, the 
Chafiares fauna includes a form, as yet undescribed, which is obvi- 
ously related and probably antecedent to Belesodon, in which teeth 
of cynognathid type are preserved. 

Watson, Kiihne, Crompton, and Bonaparte have all successively 
suggested that the tritylodonts were of gomphodont derivation. 
This may well be the case. In no other therapsid group do we 
find any appreciable trend for the development of multicuspidate 
molars. In forms such as Ischignathus we find a shortened, com- 
pact, "molar" series and a considerable postcanine diastema — 
features to be sought in a tritylodont ancestor. But although 



1967 TWO NEW GOMPHODONTS 23 

Crompton has hopefully pointed out that it would be possible to 
derive tritylodont molars from those of traversodonts, there is still 
a very considerable structural gap to be bridged. It is to be hoped 
that further exploration of Middle Triassic deposits will yield 
forms bridging this gap. 

A further possibility is that the gomphodonts may be in some 
way related to the ancestry of the monotremes. Their ancestry is a 
complete blank (despite a suggestion at one time of docodont rela- 
tionships ) and it is generally felt that they probably evolved from 
therapsids of some sort quite separately from other mammals. The 
curiously multicuspidate "molars" of the forms here described are 
suggestive in general (although not in detail) of the likewise multi- 
cuspidate molars of Ornithorhynchus. It may well be that mono- 
tremes are descended from gomphodonts of some sort. 

LITERATURE CITED 

Attridge, J.. H. W. Ball, A. J. Charig, and C. B. Cox 

1964. The British Museum (Natural History) — University of London 
joint paleontological expedition to Northern Rhodesia and Tan- 
ganyika, 1963. Nature, 201: 445-449. 

Bonaparte, J. F. 

1962. Descripcion del craneo y mandibula de Exaeretodon frenguellii, 
Cabrera y su comparacion con Diademodontidae, Tritylodonti- 
dae y los cinodontes sudamericanos. Publ. Mus. Munic. Cienc. 
Nat. Tradic. Mar del Plata, 1: 135-202. 

1963a. Descripcion de Ischignathus siidamericanus n. gen. n. sp., nuevo 
cinodonte gonfodonte del Triasico Medio superior de San Juan, 
Argentina. Acta Geol. Lilloana, 4: 111-118. 

1963b. La familia Traversodontidae. Acta Geol. Lilloana, 4: 163-194. 

1963c. Descripcion del esqueleto postcraneano de Exaeretodon. Acta 
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1963d. Un nuevo cinodonte gonfodonte del Triasico Medio superior de 
San Juan, Proexaeretodon vincei n. gen., n. sp. Acta Geol. Lil- 
loana, 4: 129-133. 
Brink, A. S. 

1955. A study on the skeleton of Diademodon. Palaeont. Afr., 3: 
3-39. 

1963. Two cynodonts from the Ntawere Formation in the Luangwa 
Valley of Northern Rhodesia. Palaeont. Afr., 8: 77-96. 

1965. A new gomphodont from the Cynognathiis zone of South 
Africa. P'alaeont, Afr., 9: 97-105. 

Brink, A. S. and J. W. Kitching 

1953. On some new Cynognathiis zone specimens. Palaeont. Afr., 1: 
29-48. 



24 BREVIORA No. 264 

Broili, F. and J. Schroeder 

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1935. Beobachtungen an Wirbeltieren der Karrooformation. IX. Uber 
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Broom, R. 

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1955. On some Triassic cynodonts from Tanganyika. Proc. Zool. Soc. 
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1964. A preliminary description of a new mammal from the Upper 

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1957. On a new cynodont from the Molteno beds and the origin of 

the tritylodonts. Ann. S. Afr. Mus., 44: 1-14. 
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1954. A bibliographical list of Reptilia from the Karroo beds of Africa. 

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1967 TWO NEW GOMPHODONTS 25 

RoMER, A. S. and J. A. Jensen 

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(Received 26 September, 1966.) 



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