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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
No.
No.
No.
code
9232.
PRE?
234.
5 72!
S237:
238:
CONFEENES
BREVIORA
MUSEUM OF COMPARATIVE ZOOLOGY
‘ 5
NUMBERS 231-264
1965
A new anole (Sauria, Iguanidae) from Puerto Rico. By
Ernest E. Williams, Juan A. Rivero and Richard
Thomas. 18 pp. October 29.
Hispaniolan giant anoles (Sauria, [guanidae): new data
and a new subspecies. By Ernest E. Williams. 7 pp.
October 29.
South American Anolis (Sauria, Iguanidae): two new
species of the punctatus group. By Ernest E. Williams.
15 pp. October 29.
Studies on neotropical Pompilidae (Hymenoptera). I.
The genus Agenioideus Ashmead in South America.
By Howard E. Evans. 7 pp. October 29.
A new salamander of the genus Chiropterotriton (Cau-
data: Plethodontidae) from Mexico. By George B.
Rabb. 8 pp. November 19.
. Variation in the number of marginal tooth positions in
three species of iguanid lizards. By Clayton E. Ray.
15 pp. December 10.
A new species of the ant genus Dacetinops from Sarawak.
By Robert W. Taylor. 4 pp. December 15.
1966
An evaluation of Jamaican Dromicus (Serpentes, Colu-
bridae) with the description of a new species. By
Donald W. Buden. 10 pp. February 25.
No.
No.
7S).
240.
241.
. 242.
. 243.
. 244,
. 245.
. 246.
. 247.
. 248.
. 249.
250:
South American anoles: Anolis biporcatus and Anolis
fraseri (Sauria, Iguanidae) compared. By Ernest E.
Williams. 14 pp. February 25.
Gymnothorax galetae, a new moray eel from the Atlantic
Coast of Panama. By Ira Rubinoff. 4 pp. February
ZS,
Avocettinops yanoi, a new nemichthyid eel from the
southern Indian Ocean. By Giles W. Mead and Ira
Rubinoff. 6 pp. February 25.
The supposed “sponge spicules” of Merrill, 1895, from
the Lower Cretaceous (Albian) of Texas. By William
A. S. Sarjeant. 15 pp. February 25.
Quaternary fish fossils from west of Lake Rudolf, Kenya.
By Keith Stewart Thomson. 10 pp. April 29.
A new species of Ashmunella from West Texas (Mol-
lusca: Pulmonata). By W. J. Clench and W. B. Mil-
ler. 6 pp. April 29.
Notes and descriptions of new Urocoptidae from Cuba
and Hispaniola (Mollusca: Pulmonata). By William
J. Clench. 14 pp. April 29.
Pseudanthessius procurrens n.sp., a cyclopoid copepod
associated with a cidarid echinoid in Madagascar. By
Arthur G. Humes. 14 pp. May 3.
The Chanares (Argentina) Triassic reptile fauna. I. In-
troduction. By Alfred Sherwood Romer. 14 pp. May
3h,
A Triassic ammonite from the Hindubagh region, Baluch-
istan, West Pakistan. By Bernhard Kummel. 5 pp.
Sully 29.
Additional notes on the amphisbaenids of greater Puerto
Rico. By Richard Thomas. 23 pp. July 29.
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.
No.
No.
yw 2aily.
B22:
- 253:
. 254.
255):
n250:
H257.
E258:
725) 3).
260.
5261:
Chronological survey of the tetrapod-bearing Triassic of
Argentina. By J. F. Bonaparte. 13 pp. November 4.
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. Jen-
sen. 20 pp. November 4.
A new Hispaniolan gecko. By Richard Thomas. 5 pp.
November 4. |
Preliminary descriptions of new Honey-Eaters (Aves,
Meliphagidae). By Finn Salomonsen. 12 pp. Novem-
ber 4.
A revision of the fossil selenodont artiodactyis from the
Middle Miocene Thomas Farm, Gilchrist County,
Florida. By Vincent Joseph Maglio. 27 pp. Decem-
ber 6.
1967
Anolis chocorum, a new punctatus-like anole from Dari-
én, Panama (Sauria, Iguanidae). By Ernest E. Wil-
liams and William E. Duellman. 12 pp. February 3.
A review of the Clark Fork vertebrate fauna. By Roger
C. Wood. 30 pp. February 3.
Biology of the parthenogenetic fungus beetle Cis fuscipes
Mellié (Coleoptera: Ciidae). By John F. Lawrence.
14 pp. February 3.
Expanding the palpi of male spiders. By William A.
Shear. 27 pp. February 3.
Monograph of the genus Spiroceramus (Mollusca: Pul-
monata: Urocoptidae). By William J. Clench. 10 pp.
February 3.
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 moljuscan 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 Vannuccia 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 Ji es holt enets teats atic ok Sale Gok Lo and eee 251
IB RINCKMANNGV.OSS VANIDAL vis0s sn 5 5 es wad lee eee mee ee 263
BEDE NR IDONATED 2 Watters oh kere as a2 Vosecie chs Cec aan ahaa See ey a 238
CRENCH aWIELIAM Ms 42255 amtod es 2) 6 aa tet ncnee DAE 2455260
OBR aS CAINE Yer eer War ee gis Ohare eee Oe MANE en 250
IDEGENS WE GONPAE SS kate cee lo einen he fp Mere eee 262
DWE IWIENMANFe WEE TANEIES. fates cps. cuss bape aye, © ord Beene ee ee 256
EVANS IOWARD Beers oan ree sarge ene ee oe eee 234
KG TASE TAEN EINE CEVA Mia Decl. Be cars co a At Sayre ar nee ee 262
PERWINDE Sse NREIORG Gi oe Boer tc sek ca ine shavers Were! Stee eer 246
EN SEN: AIMEES Ate aps fache~ ot cv eceis eve Se eae ee ee DIS
OMMET OBE RNETARD (0s 5 hc ahr, oh stand coarse 248
IO ASNIRE GES NOHING Hea. ce urpy ce sees) ty See hen ecw oy eee yeh 258
AGIOS WWINGENT JOSEPH: <4. 56) ors coches eos th oie eee ee DISS)
AE AD) SCRIBES Wier 202 ye uid chtd Sale cer ME Ae Oe 241
IVERICTE BIR TN CPt enact nse okie) 2 su eke Pay Marist, CR cr ty «cae a Baht Bea pws 244
RARKE RA IROBE RSD: gbi.d iG cake ake: ae ele, Gere tater ie vega ets 262
EO NBIBMIGTE OR GIE sites tt a yoo Peat a Aad Se er a ee oes 235
EAs IGAY TONS SO eke 3 SU SON te bn ay honk atoke cet eee 236
FOIE R On WAN PAS trata creche: salts ake Race ots Coe eemoeee oes ee ae 23
ROMER. ALERED: SHERWOOD jis. = oss lake Aco ke DAT. 252, 204
FUBINORE OGURA ES e co sik: & Ghea so ee ree 240, 241
SATOMONSENS@ISINING dope cee ct = epeead eae ees Sie ee 254
SARE AINSI 5 WYLIE LE TAINWAG, “So yio..c. 3% o, o spans eae ast eh regenera 242
SCHWARTZ. OAUEB ERG sks fais chic Ke Sachse «, CROCE eae eee Ren Se 261
SHEAR: BWiDETDANGeAGtE ys. PRY. 088 TRC PEL ce Shee eee meee 259
SPENGERY DEREK Ws Stn eee oe Magis Senter 262
SWAVTORS ROBE RI Wiseds 4\.0 oie <.kvoehs Soho Sie rere 230)
INEONMASIRIGHARD fscsc 30-6 ocr acorrs-e ae < eens oes 231, 2495253: 261
IRHOMSON: UKEITH: STEWART 565) dak as «= uss oe ee 243
NWWHISEeTANS ERNEST I. .osca sex aul 2351, 2232! 2334239256
WOOD ROGER NE@S 5 bo isc dS cuddles os Sie ontos Me Ce DS
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. OcTroBEn 29, 1965 NUMBER 231
A NEW ANOLE (SAURIA, IGUANIDAE) FROM
PUERTO RICO
By ErRNeEst #. WILLIAMS,
JUAN A. RIvERo and RicHARD THOMAS
PART I. DESCRIPTION
By Ernest EK. WIuuiAMs
Museum of Comparative Zoology
and JUAN A. RIVERO
University of Puerto Rico, Mayaguez
On June 16, 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 (Thecodactyllum) sintenisu, which al-
though usually epiphytic, was growing abundantly, together
with Guzmana berteroniana, on the forest floor alongside the
road. Although the cloud forest is fairly heavy in this area,
trees and tree ferns had been eut 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 described 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.
i)
BREVIORA NomZal
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 fly 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 (Drosophila) 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 was never seen to enter a small bromehad 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 (ASE'S) or the Richard
Thomas collection (RT).
In allusion to its long concealment from scientific record we
call the new species by the Latin adjective which means “‘hid-
den’’:
1965
NEW ANOLE FROM PUERTO RICO
Se
<<,
—
&
: Dewlap of MCZ 83666 (9 )
f type, MCZ 80303. Bottom
1ew oO
Dorsal v
Fic. 1. Anolis occultus new species. Top
showing inset (‘‘slotted’’) margin.
(ee)
4 BREVIORA No. 231
ANOLIS OCCULTUS new species
Holotype. MCZ 80503, adult female, on Rd. 143, midway be-
tween Cerro La Punta (1338 m) and Cerro Maravilla (1183 m),
Puerto Rico, J. Rivero ecoll., 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 km N Sabana Grande, 2800 feet (850 meters),
A. Schwartz, Richard Thomas eoll., 25 February, 1965; MCZ
83661-63, same locality, Richard Thomas eoll., 28 February,
1965; ASFS-V6196-97, 10.6 km SSE Villa Pérez, 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 ecoll., 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 ecoll., 2 February 1965; MCZ 83660, same lo-
eality, Richard Thomas coll., 5 February, 1965; ASFS-V5017-19,
same locality, Richard Thomas ecoll., 3 February, 1965. ASFS-
V6662-65, RT 1832, 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 which 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 supracilary 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 seales, 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 ANOLE FROM PUERTO RICO 5
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 from 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 supratemporals. Ten to
eleven supralabials to the center of the eye, the posterior supra-
labials very low and small.
S
Sw
Ses
Fig. 2. Anolis occultus new species. Dorsal, lateral and ventral views of
head of type, MCZ 80303.
6 BREVIORA No. 231
Mentals small, about as deep as wide, in contact with 4 (4-6,
usually 4) seales between the infralabials. Several rows of throat
seales 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
seales small but well developed in well separated rows (2) or
these scales weakly developed, or almost absent ( ¢ ).
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 111 of fourth
toe. Seales of limbs smooth, much smaller than ventrals, supra-
digital scales smooth.
Tail: Round, no dorsal crest. No enlarged postanals in males.
Seales on dorsal surface very small, smooth, imbricate. Seales
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 lght
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,
Fic. 3. Scales around nostril compared in Anolis occultus (right) MCZ
83661 and Anolis evermanni (left) MCZ 61223.
1965 NEW ANOLE FROM PUERTO RICO 7
between the anterior limbs; two 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
Ss
=
<S
Sma
Fig. 4. Tail of A. occultus. Top: MCZ 83666, a verticil showing typical
squamation. Middle: ASFS-V 5490, unregenerated tail tip. Bottom: ASFS-V
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,
evermanm 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 1f in occultus the apparent first infralabials had
segmented off 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. fuscoauwratus, A. chloris, A. maculiwventris,
eve
The Miocene Anolis, 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 ANOLE FROM PUERTO RICO 9
smaller. Other differences are the presence of a small supra-
ciliary, the wide mental, the unicarinate dorsal limb scales, the
multicarinate supradigital scales, the narrower 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 probable that occultus 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 electrwm 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.
LITERATURE CITED
LAZELL, J. D. JR.
1965. An Anolis (Sauria, Iguanidae) in amber. Jour. Paleont.
39: 379-382.
10 BREVIORA No. 231
PART II. FIELD OBSERVATIONS ON
ANOLIS OCCULTUS WILLIAMS AND RIVERO
By Ricuarp THOMAS
10,000 SW 84th Street
Miami, Florida 33143
Having been fortunate enough to collect the first substantial
series of the newly described Anolis occultus, 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 occultus 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 ANOLE FROM PUERTO RICO ikl
not ordinarily allow positive identification of so small 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. When 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
a2 BREVIORA No. 231
eene et
Fic. 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 NNE 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 ANOLE FROM PUERTO RICO 13
Loeality 2: 20.9 km NNE Guayama, 2300 feet (700 rheters),
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
erew what may be ealled 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 below 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.
Loeality 3: 138.7 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 Pérez, 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 lzard 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
1Sinee this manuscript went to press, another visit was made to Puerto
Rieo 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 ANOLE FROM PUERTO RICO 15
‘
A. cristatellus and A. krugi were collected; gundlachi and ever-
mannt were also seen there. A. krugi is an anole of bushes and
grass and is sometimes almost terrestrial in habits. Its sleeping
site preferences are less restricted than those of occultus. 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.
gundlachi was collected in addition to occultus. At locality 4,
cristatellus 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 occultus 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 (seapu-
lar, dorsal, lumbar, and sacral) which may be manifested as
either very hazy, indistinct dark bands 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 yellow 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
daytime and gray or brown at night.
16 BREVIORA No. 231
Inchenate: 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 gray. 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.
LITERATURE CITED
Maerz, A. and M. Rea Pau
1950. A dictionary of color. New York, MeGraw-Hill Book Co.,
pp. i-vii, 1-23, 137-208, 56 pls.
(Received 29 June 1965.)
NEW ANOLE FROM PUERTO RICO 17
1965
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apUudIO
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18 BREVIORA No. 231
TABLE 1
Seale characters in Puerto Rican Anolis
scales across labials
S-V length (mm) lamellae snout at to center
Anolis adult ¢@ 4th toe second canthal loreals of eye
occultus Be 14-18 9-13 2-6 8-13
cuvieri 126 31-33 8-9 6 7-8
evermanni 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
krugi 45-55 21-23 4-7 6-7 6-8
pulchellus 45-50 19-21 4-5 3-5 5-7
poncensis 43-46 18-21 4-6 3-5 5-6
TABLE 2
Seale characters in Puerto Rican Anolis
scales interparietal
between from middorsal rows
Anolis semicircles semicircles enlarged ventrals tail crest
occultus 2-4 2-6 0 sj =
cuviert 3 3 1 si +!
evermanni 0-1 1-4 0 si (+)
stratulus 0-1 0-3 0 si +
cristatellus 0-1 1-4 2 s/k,i + !
gundlachi 1-3 3-8 2 ki +!
krugi 0-2 1-5 4-6 ki (+)
pulchellus 0-2 1-5 ca. 12-18 ki (+)
poncensis 0-1 1-2 ca. 16-20 ki (+)
s = smooth; k = keeled; i = imbricate; j = juxtaposed; ! = strong; ( ) = weak.
TABLE 3
Seale character comparison
maculiventris occultus
scales across snout at second canthal 9-16 9-13
scales bordering rostral posteriorly 6-9 5-9
scales between semicircles 2-4 2-4
loreal rows 6-9 2-6
interparietal from semicircles 5-10 2-6
labials to center of eye 6-9 8-13
scales bordering mental between
infralabials 5-8 4-6
fourth toe lamellae 15-18 14-18
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. OcTOBER 29, 1965 NUMBER 232
HISPANIOLAN GIANT ANOLES (SAURIA, IGUANIDAE):
NEW DATA AND A NEW SUBSPECIES
By ERneEstT E. WILLIAMS
Recent collections of Hispaniolan giant anoles (Anolis ricordiz)
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 RICORDIL RICORDIL Duméril and Bibron
New Records: Haiti. Departement de ’Ouest. Lancerouelle near
Mirebalais, MCZ 69404; Mirebalais, MCZ 68479; Saltrou, MCZ
69405.
Dominican Republic. Monte Cristi Province. Laguna de Salo-
dillo, 7 km SE Pepillo Saleedo, 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 Sae Plain, extends into
the Dominican Republic in the Monte Cristi region. Hence the
suggestion (Wilhams, 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
ricordi ricordii 1s 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 embarrassment remains. No
certain intergrades between the two strikingly different forms
ricordit and baleatus are yet known. In view 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 heavily 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 ricordiz.
Dr. Schwartz (pers. comm.) reports taking the Monte Cristi
series In vines at night in a wooded area about a small cattle pond
1965 HISPANIOLAN GIANT ANOLES 3
between Copey and Pepillo Saleedo — an otherwise arid region.
He states that in general ricordii can be captured with moderate
regularity while sleeping at night in viny 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 RICORDIL 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, ca. 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 Macoris, DWB 271; ca. 4 km NE
Ponton (Rio Cuaba), ASFS V 2987. Samana Province. 6 km E
Sanchez, ASFS V 1904. Peravia Province. 1.1 mi (1.8 km) $8 San
Jose de Ocoa, 1400 feet (425 m), ASFS V 723. El Sezbo Province.
3.5 mi (5.8 km) 8 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 required; 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 ©),
“dark orange” (adult o), ‘‘very bright orange” (adult co), ‘‘dull
grayish orange” (adult @), “dirty orange’ (adult 2), “orange
brown” (adult 9), ‘‘pale grayish orange” (juvenile 07), ‘““dewlap
skin charcoal, scales yellow-green” (juvenile 9). The chin of the
male from southeast of Sosua, 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, varying 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.
ANOLIS RICORDIIT BARAHONAE 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
Enriquillo, 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. Dewlap 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 AMNH 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 ricordiz populations
in the vicinity of Enriquillo consistently show a distinctive pattern
though characteristically barahonae in squamation.
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, AMNH 93713-21, ASFS X
3033-35, 3038-39, 3041-42, same data as type; MCZ 80943-53,
AMNH 93722-36, ASFS X 3182, same data as type except
collected 28 July 1962.
Diagnosis: A subspecies of ricordiz 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 lines more or less broken into and complicated by
light cross-banding. (One or two specimens have this pattern very
much reduced and juveniles are irregularly spotted, without hght
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 other West Indian anoles.
1965
HISPANIOLAN GIANT ANOLES
CJ
#,
Pape me
Gala
Black and white from a water color by D. C. Leber.
Anolis ricordit leberi, new subspecies.
Fig. 1.
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 leberi — MCZ 38277, collected by P. J. Darlington,
Jr., discussed in Wilhams, 1962 — has not been made a paratype
of the new taxon. It is a juvenile which in squamation completely
agrees with leberi. However, while in its spotted, non-lineate
pattern it is roughly like the single topotypic juvenile available
(MCZ 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 Cayes and Jeremie on which Camp Perrin
lies, is largely herpetologically unknown. It cannot be cavalierly
assumed that only one ricordiz race will be native to this area.
Perhaps no surprises await us, but in the West Indies sharp faunal
changes are not unusual and casual generalizations are never
safely made,
REFERENCE CITED
WILiiams, E.E.
1962. Notes on Hispaniolan herpetology. 6. The giant anoles. Breviora,
Mus. Comp. Zool., No. 155: 1-15.
(Received 29 June 1965)
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BREVIORA
Museum 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 punctatus 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 punctatus (and of
its race boulengert O’Shaughnessy) have the rostral swollen, pro-
jecting strongly beyond the lower lip. 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 Orcés. 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 punctatus 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:
2, BREVIORA No. 233
ANOLIS DISSIMILIS new species
Holotype: CNHM! 81369, an adult o’, Itahuania, upper Rio
Madre de Dios, Madre de Dios Province, Peru, Kalinowski ecoll.,
15 October 1954.
Diagnosis: Close to Anolis. proboscis Peters and Orcés, A.
phyllorhinus 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. 14 snout-vent length).
Description. Head: All head scales wrinkled or striate, moderate
to large posteriorly, small anteriorly. Seven scales across head
Fig. 1. Anolis dissimilis new species. Type, CNHM 81369.
1For 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 nine, 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, Machala!, 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, flat, 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 infralabials; 4 gular scales in a
1The 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 probability — more exact.
1965
Fig. 2.
ANOLIS PUNCTATUS GROUP
Anolis nigrolineatus new species. Type, MCZ 38940.
6 BREVIORA No. 233
gentle forward are 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 fan: 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: Seales on limbs unicarinate, the largest on hind limb
larger than ventrals. Supradigital scales multicarinate. Eighteen
lamellae under phalanges ii and 11 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. punctatus (including the sub-
species boulengerz): Specimens from many localities in Brasil,
Bolivia, Peru, Ecuador and the Guianas.
A. 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 8. 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 ANOLIS PUNCTATUS GROUP 7
This situation is well exemplified by the punctatus 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 nzgrolineatus 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 singie prenasal scale
from the rostral. The head scales are flat, pavimentose (even in
boulengert, 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 punciatus. There are no or only a few scales between the
supraorbital semicircles (0-2). The interparietal, larger than the
ear, 1s of moderate size (large in laevis, dissimilis and proboscis) and
separated from the semicircles by 0 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 by naked skin (scales not in
rows in proboscis). Middorsals not or not appreciably larger than
flank seales (a dorsal crest in proboscis). Ventrals smooth (keeled
in boulengeri which, however, intergrades with smooth-sealed
punctatus), squarish, transverse. Tail more or less compressed with
double row of scales dorsally (a single crest in dissizmilis 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,
bluntly pointed structure. A. dissimilis carries the condition of
nigrolineatus to an extreme. In contrast, the bony snout of A.
punctatus 1s 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 similar 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 barkerz, and also Myers
and Carvalho (1945), in deseribing 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-
semblanees between species that on other characters would be
separated. Thus, disstmilis 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 dzssimilis.
More awkward for the desiderate goal of taxonomic clarity are
the species I have described as “fringe species.’’ These are:
transversalis (including buckleyz); 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 A. 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. nzgrolineatus 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) —
boulengert — 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 lcirele of forms which include such
10 BREVIORA No. 233
species as boettgert, chloris, fasciatus, fraseri, frenatus, insignis,
jacare, latifrons, microtus, mirus, nasofrontalis, peraccae, pseudoti-
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 lateral 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 distine-
tive, and geographically quite coherent, occurring in species of
mainland South America (but not those of the West Indies or
Malpelo Island) and also in two or three species — frenatus,
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 strongho!d 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 osteologica!
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 probably 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 achieve their distinctive rostral
structure in different ways. Neighbors to these in the extreme
wing of the alpha series are punctatus and, if I interpret matters
rightly, the two new species that are described in this paper.
Acknowledgments: | am deeply grateful to Dr. Gustavo Orcés-V.
for allowing me to examine his Ecuadorian Anolis — 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 Vanzolini, De-
partamento de Zoologia, Sad 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.
LITERATURE CITED
Corps, 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 morphology. University
Microfilms, Ann Arbor. 236 pp.
Myers, G. 8. anp A. CARVALHO
1945. A strange new leaf-nosed lizard of the genus Anolis from Ama-
zonia. Bol. Mus. Nac., Rio de Janeiro, N. 8. Zool., No. 43: 1-14.
12 BREVIORA No. 233
Prerers, J. A. AND G. Orcks-V.
1956. A third leaf-nosed species of the lizard genus Anolis from South
America. Breviora, No. 62: 1-8.
Scumipt, K. P.
1939. A new lizard from Mexico with a note on the genus Norops.
Zool. Ser. Field Mus. Nat. Hist., 29: 7-10.
RUTHVEN, 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° 70° 60° 50° 40°
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Fig. 3. Map of the distribution of punctatus group anoles except A. punc-
tatus itself. The exact limits of the latter’s very wide range are not known.
13
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1965 ANOLIS PUNCTATUS GROUP 15
Table 2
Character range in South American anoles
alpha anoles beta anoles
number of toe lamellae 14-30 (mode 18-21) 10-27 (mode 14-18)
scales across snout 4-25 7-20
scales between semicircles 0-5 (7 *) 0-4
dorsal scale rows enlarged 0-2 0-12
* Though several alphas range up to five scales between the supraorbital
semicircles, the maximum reported here occurs in two exceptional specimens
of A. princeps Boulenger (= ? frenatus Cope).
ter |
BREVIORA
Museum of Comparative Zoology
Campripez, Mass. OcToBER 29, 1965 NuMBER 234
STUDIES ON NEOTROPICAL POMPILIDAE
(HYMENOPTERA)
I. THE GENUS AGENIOIDEUS ASHMEAD
IN SOUTH AMERICA
By Howarp E. Evans
The rich pompilid 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 Pefia, 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 (Mem. Amer. Ent.
Soc., in press).
2 BREVIORA No. 234
The genus Agenioideus 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 outlined
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 Ageniozdeus. 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 A genizoideus
(which I regard as only two) are closely related and somewhat
divergent from the North American Ageniozdeus, 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, Hnbanksza, 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 a!so 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, witha 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 are except the scutellum and metanotal
dise 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.
Soe., 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 hooklets,
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 > as long as thick, equal to
.35-.45 > 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 xX as long as thick, equal to
.65-.80 the upper interocular distance; wings strongly twice-ban led;
lecsiiuscoussatlleastaimularcespahiene sere mie ee eter ene er 2
2. Pronotum and mesoscutum ferruginous, except the former with some p.le
yellow markings; front relatively narrow, middle interocular distance
HOM LLansiacialadistancen ees: 2a. accoleus accoleus (Banks)
Pronotum and mesoscutum fuscous, except 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... .
OO Oe 2 Ae Oe POD EL Dee eT ote Cente ae ee 2b. accoleus lucanus (Banks)
4 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
Pronotum, mesoscutum, scutellum, metanotum, and upper part of meso-
pleura ferruginous; middle interocular distance .59 X transfacial dis-
GANCC). sce yan a RLS ee aoe re 2a. accoleus accoleus (Banks)
Thorax black; middle interocular distance .62-.66 > transfacial dis-
CANCELs Se”. Ee RARER Ae MN eee) PE Pen 2b. accoleus lucanus (Banks)
i)
1. AGENIOIDEUS (ENBANKSIA) MINUTUS (Banks) new combination
Sericopompilus minutus Banks, 1947, Bull. Mus. Comp. Zool. Harvard,
99: 435. [Type: 2, 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 rufo-testa-
ceous except second and apical few segments usually somewhat
infuscated; legs beyond the trochanters (or at least beyond the
Fig. 1. Male genitalia of Agenioideus (Enbanksia) minutus (Banks), ventral
aspect.
Fig. 2. Same of A.(E.) accoleus lucanus (Banks), left side omitted.
Fig. 3. Tip of subgenital plate of male A.(H#.) minutus (Banks).
Fig. 4. Subgenital plate of male A.(Z.) accoleus lucanus (Banks).
1965 AGENIOIDEUS IN SOUTH AMERICA 5
middle of the femora) bright rufo-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 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 are 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 1.2-1.8 & 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 slightly shorter than metanotum; propodeum in profile
forming a somewhat higher are 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. —8 2 2,30°'c%. BRAZIL: 7 292,14,
Nova Teutonia, Santa Catarina, Nov.—Feb. (F. Plaumann)
[Mus. Comp. Zool., Cornell Univ., Coll. G. R. Ferguson]; 1 9,
Tres Lagoas, Matto Grosso [type, Cornell Univ.]. PARAGUAY:
1 o', Caacupe, Oct. 25, 1955 (F. Schade) [Coll. G. R. Ferguson]
PERU: 1 o&, Avispas, Madre de Dios, 400 meters, Sept. 10- 30°
1962 (L. Pefia) [Mus. Comp. Zool.].
2a. AGENIOIDEUS (ENBANKSIA) ACCOLEUS ACCOLEUs (Banks) new
combination
Sericopompilus accoleus Banks, 1947, Bull. Mus. Comp. Zool. Harvard,
99: 433. [Type: 9, BRAZIL: Maracaji, 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 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 & 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 wideas 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 oceilar triangle. Third antennal seg-
ment rather short although longer than second, measuring about
1.2 < as long as thick. Postnotum nearly as long as metanotum;
slope of propodeum very low and even. Terminaia 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. Mus. Comp. Zool., Harvard,
99: 434. [Type: 9, BRAZIL: Nova Teutonia, Santa Catarina, 25 Jan.
1939 (F. Plaumann) (Mus. Comp. Zool.)|
1965 AGENIOIDEUS IN SOUTH AMERICA i
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 > 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. Middle 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).
Distribution — Southern Brazil (Santa Catarina).
Specimens examined. —13 2 2,4 7%, all from Nova Teutonia,
Santa Catarina, Oct.—March (F. Plaumann) [Mus. Comp. Zool.,
Coll. G. R. Ferguson].
(Received 14 June 1965)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NovEMBER 19, 1965 NUMBER 235
A NEW SALAMANDER OF THE GENUS
CHIROPTEROTRITON (CAUDATA: PLETHODONTIDAE)
FROM MEXICO.
By Grorce B. Rass
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 Potosf, Chiro-
pterotriton mosaueri, is probably restricted to caves. Another
species apparently in this category is described below. For it, I
propose the name:
CHIROPTEROTRITON MAGNIPES Sp. Nov.
Figs. 1, 2
Holotype. — Museum of Comparative Zoology 30607, a young
adult male collected in the Cueva 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.
Paratypes. — MCZ 44033, CNHM 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 type
locality in July 1964 by Kraig Adler; CNHM 142458, UMMZ
125179, and three uncatalogued males in the Instituto Politecnico
collection from Cueva del Madrofio, near Laguna Colorada,
Municipio de Landa de Matamoros, Querétaro, Mexico, 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 Nor 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 2-8, 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 aleohol 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 Madrofio. 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. cit.) also provided data on the Cueva del Madrofio,
which is at an altitude of 1810 m in mixed pine-oak woods con-
taining madrofio (Arbutus) and Carya. 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 (CNHM 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, pl. II).
In proportions, C. magnipes differs from other Chiropterotriton
in having a somewhat wider head (average, 19.1 per cent of sn out-
vent length in males). In relation to head length, C. magnipes has
a large eye (average percentage, 35.5, o'o7; 37.9, 2 2), although
the proportion is matched in C. multidentatus from Rancho del
Cielo, Tamaulipas and El Chico, Hidalgo, and in the short-snouted
species C. bromeliacia. 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. mosauert 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. multidentatus,
C. arboreus).
Acknowledgements. — I am grateful to Dr. E. E. Williams for the
opportunity to deseribe this salamander. Dr. Alejandro Villa-
lobos F. of the Instituto de Biologia, Universidad Auténoma de
Mexico, has kindly supplied data on the type locality. 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. 8. 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 CNHM, 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+96 pp.
Makrtin, P. 8S.
1958. A biogeography of reptiles and amphibians in the Gomez Farias
Region, Tamaulipas, Mexico. Misc. Publ. Mus. Zool. Univ.
Michigan, 101: 1-102, 7 pls.
Rasp, 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 pls.
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.
Taytor, E. H.
1944. The genera of plethodont salamanders in Mexico. Pt. I. Univ.
Kansas Sci. Bull., 30: 189-232.
WakgE, D. B. and A. H. Brame, Jr.
1963. The status of the plethodontid salamander genera Bolitoglossa and
Magnadigita. Copeia, 1963: 382-387.
No. 235
BREVIORA
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1965 NEW CHIROPTEROTRITON 7
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No. 235
BREVIORA
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. DECEMBER 10, 1965 NUMBER 236
VARIATION IN THE NUMBER OF
MARGINAL TOOTH POSITIONS
IN THREE SPECIES OF IGUANID LIZARDS
By
CuayTon 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 implications 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 pectinata with 11 teeth per wave, would be especially
interesting. Hotton (1955, p. 97) in a study of adaptive relations
of dentition to diet in iguanids, including C. s¢milis, 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. pectinata, 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.
bo
BREVIORA No. 236
MATERIAL
Ctenosaura 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 Maya 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 pectinata: 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 Museum, 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
individuals, 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-line 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.
V. = the coefficient of variation of x.
V, = the coefficient of variation of y.
OR, = the observed range of x.
OR, = the observed range of y.
r = the coefficient of correlation, here between x and y.
a = the growth ratio.
co, = the standard error of a.
b = the initial growth index.
Sa = 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.
Da = 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 analysis 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 positions was deter-
mined for this element, as adequate 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 univariate
statistics are presented (Table 1, Figure 1).
Mawillae and dentaries: All of the statistics listed above were
computed for both left and right maxillae and dentaries of the two
samples of Ctenosaura, and for the right side only of Anolis (Tables
2,3, and 4). Asa check on symmetry, the frequency distribution
for length of tooth row and number of tooth positions was in
every case represented by 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 equation:
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
Clenosaura 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 value 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 Clenosaura 7. The fact that premaxillary tooth counts in Anolis
can assume only even values might suggest that the coefficient of
variation (V,) for Anolzs is artificially high. However, if only one
side of the fused element is considered, with tooth counts of 3-6,
V, 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 polymodality, analogous to that demonstrated by Klauber
(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 shert-
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 be of taxonomic
interest only if it lay within the portion of the Ctenosaura dis-
tribution well above the OR of Anolis. 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, sa,
and D4, 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 number of tooth positions in the samples studied
are the relative strength of correlation between the two variates
and the slope of their growth line?. The two samples of Clenosaura
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 = 0 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 growth 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.
2 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.
for)
BREVIORA No. 236
Although so few samples provide 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, 1.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
Epmunp, A. G.
1960. Tooth replacement phenomena in the lower vertebrates. Roy.
Ontario Mus., Life Sci. Div., Contrib. No. 52: 1-190, 58 figs.
Horton, Nicnouas, 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. Bull. Am.
Mus. Nat. Hist., 108: 211-252, 10 figs.
KLAvuBER, L. M.
1937. <A statistical study of the rattlesnakes. IV. The growth of the
rattlesnake. Occ. Pap. San Diego Soc. Nat. Hist., 3: 1—56, 6 figs.
a |
1965 TOOTH VARIATION IN IGUANID LIZARDS
Pouuock, H. E. D., and C. E. Ray
1957. Notes on vertebrate animal remains from Mayapan. Carnegie
Inst. Washington, Dept. Archaeol., Current Repts., 41: 633-656.
Simpson, G. G., ANNE Rog, and R. C. Lewontin
1960. Quantitative zoology. Harcourt, Brace and Co., New York,
vil + 440 pp., rev. ed.
(Received 24 August 1965)
Table 1. Univariate statistical characterization of tooth count (y)
in the fused premaxillae of three iguanid lizards.
Anolis
Statistic Clenosaura similis Clenosaura pectinata carolinensis
N 74 17 58
y 7.05 = .07 (ALS eli 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
No. 236
BREVIORA
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TOOTH VARIATION IN IGUANID LIZARDS
1965
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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 lines.
Maxilla Dentary
C. similis
left - right 22 — .84
C. pectinata
left - right — .317 — .08
C. similis - C. pectinata
left - left el, — 1.28
right - right Ene — 1:02
C’. similis - Anolis
right - right 5.83 5.28
~
. pectinata - Anolis
right - right 5.82 4.70)
11
TOOTH VARIATION IN IGUANID LIZARDS
1965
.
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TOOTH COUNT
ZZ,
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TOOTH COUNT
TOOTH COUNT
Histograms illustrating the frequency distribution of tooth counts
Fig. 1.
in the fused premaxillae of (A) Ctenosaura similis,
pectinata, and
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12
BREVIORA No. 236
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TOOTH COUNT
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LENGTH OF TOOTH ROW (MM_) ‘:
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LENGTH OF TOOTH ROW (MM.)
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TOOTH COUNT
10
6 7 8 9
LENGTH OF TOOTH ROW (MM.)
Fig. 3. Scatter diagrams showing tooth counts (y) and length of tooth
row (x) in the right maxillae of (A) Ctenosaura similis, (B) C. pectinata, and
(C) Anolis carolinensis. The broken line represents the reduced major axis;
the open circle, the joint mean.
1965 TOOTH VARIATION IN IGUANID LIZARDS 13
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(C) Anolis carolinensis. The broken line represents the reduced major axis;
the open circle, the joint mean.
14 BREVIORA No. 236
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1965
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. D&cEMBER 15, 1965 NUMBER 237
A NEW SPECIES OF THE ANT GENUS DACETINOPS
FROM SARAWAK!
By Rosert W. Taytor
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. O. 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 Emerson. This significant and unexpected record
robs Dacetinops of its apparent status as a New Guinean endemic.
In the light 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 Tekalit was a remote field camp established by
the Chicago Natural History Museum, situated about 70 miles
above Kapit on the Batang Balleh—Sungei Mengiong River
system in western Sarawak.
1 Research supported by U. 8. 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. cibdella; 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, strue-
tural details of palopmeres as in czbdella (Brown and Wilson, 1957,
fig. 2). Antennal scrobes as in czbdella; 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 czbdella, bases of teeth
separated by slightly less than twice their length. Propodeal
spiracles directed posterolaterally, placed well back on sides of
propodeum, not as far back as in czbdella. 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 slightly 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
1965 ANT GENUS DACETINOPS 3
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 microsculpture 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 c7zbdella; 4 distinet
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 microsculpture; its basal half with about 10 sharp
4 BREVIORA Now23
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 whitish body hairs extensively developed, proportionately
longer than in czbdella (0.12—-0.65 mm) but similarly distributed;
many on head (particularly its margins), mesosoma and nodes
occupying positions homologous to those of czbdella. 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 czbdella, deep reddish brown, mandi-
bles, antennae, and legs dark yellowish brown.
Discussion. D. concinna is readily distinguished from D. cibdella
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 pomt. 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.
ds let
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
Museum of Comparative Zoology
CAMBRIDGE, Mass. FEBRUARY 25, 1966 NUMBER 238
AN EVALUATION OF JAMAICAN DROMICUS
(SERPENTES, COLUBRIDAE)
WITH THE DESCRIPTION OF A NEW SPECIES
By Donatp W. BupDEN
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 D. 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-
rowed 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 Dowling (1951),
and the terminology in regard to hemipenial descriptions has
been derived mainly from Dowling and Savage (1960). 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 CALLILAEMUS (Gosse, 1851)
Natrix callilaema Gosse, 1851, Naturalist’s Sojourn in Jamaica, p. 384.
Type locality: Bluefields, Jamaica.
Since only two of the three (fide Boulenger, 1894:143) syn-
types of D. callilaemus were examined for the present study, 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
D. 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
subeaudals 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 lone 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 lght 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 subcaudal count, however, correlates with those
of males from other localities.
Distribution: 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
Oo
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 lhghter dorsum; supralabials 7/7; infralabials 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.
4 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 subcaudal; sulcus spermaticus
bifureating at the level of the third subcaudal; 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 infralabials, 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. Right: Dromicus 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, 18887, 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 FUNEREUS (Cope, 1862)
Alsophis funereus Cope, 1862, Proc. Acad. Nat. Sei. 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
subeaudals 90; supralabials 7/7; infralabials 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,
generally 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 +
subeaudals 203-224 in males and 192-218 in females; largest male
479, largest female 462; ratio of tail length to snout-vent length
0.438-0.59 in males, and 0.38-0.47 in females; tail length/snout-
vent length ratio somewhat lower in juveniles than in adults;
—_
Ye
BREVIORA No. 238
dorsum generally dark, unpatterned ; 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; 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.
An account of the hemipenial configuration based upon the
examination of the right extruded organ of ASFS 15172 is as
follows: a single relatively slender, clavate organ extending to
the level of the eighth subeaudal; suleus 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-suleate surface.
Variation: Anomalies in numbers of certain head scales were
occasionally encountered; scales involved were: supralabials,
Fig. 3. Left: Dromicus funereus, hemipenis, view of suleate surface,
ASES 15172, from 1 mi. S Reading, St. James, Jamaica. Right: Dromicus
polylepis, hemipenis, view of suleate surface, ASFS 13536, from Port
Antonio, Portland Parish, Jamaica.
1966 JAMAICAN DROMICUS i
infralabials, prefrontals, preoculars and postoculars. 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 (USNM 108337-38), 5 mi. W Montego Bay, seven (MCZ
44897-900; UMMZ 85941-43), 1 mi. S Reading, 59 (ASFS
147438, 14932, 15060-63, 15066-67, 15146-48, 15159-77, 15264-92) ;
St. Elizabeth Parish, Balaclava, two (USNM 73276-77) ; Trelawny
Parish, Quick Step, one (USNM 117676), Windsor, two (USNM
83719-20) ; Manchester 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.
Paratypes: 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
+ subeaudals 230-245 in males, and 222-237 in females; hemi-
penis single, somewhat bulbous, and heavily spinose.
Description of holotype: An adult male, total length 387, tail
length 150; ventrals 137; anal divided; paired subecaudals 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; infralabials 9/9; prefrontals 2; loreals 1/1;
preoculars 1/1; postoculars 2/2; temporals 1 + 2/1 + 2; dorsals
smooth, dorsal row formula 19-17-17; dorsum dark brown to black
with a pair of ight 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 subeaudals 92-106 in males, 76-93 in females; ventrals +
subeaudals 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; infralabials 9/9; prefrontals 2; tem-
porals 1 + 2/1 + 2; a dorsal row formula of 19-19-17, occasionally
19-17-17; shght 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 sub-
eaudals; sulcus spermaticus bifurecating 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-suleate 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-suleate and suleate 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 fuwnereus 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 subeaudal 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. polylepis 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 Underwood and Williams, 1959), and a hum-
mingbird (Trochilus polytmus scitulus Brewster and Bangs; ef.
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 Richard
Thomas for his assistance in the laboratory as well as for the
10 BREVIORA No. 238
illustrations of the hemipenes of funereus and polylepis; 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 on the herpetology of Jamaica. Bull. Mus. Comp. Zool.,
52(15) :273-301.
BOND, JAMES
1956. Checklist of birds of the West Indies. Acad. Nat. Sci. Philadel-
phia, ed. 4, ix + 214 pp.
BOULENGER, G. E.
1894. Catalogue of the snakes in the British Museum (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.)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. FEBRUARY 25, 1966 NUMBER 239
SOUTH AMERICAN ANOLES:
ANOLIS BIPORCATUS AND ANOLIS FRASERI
(SAURIA, IGUANIDAE) COMPARED
By Ernest E. WILLIAMS
INTRODUCTION
Confusion of the two species Anolis biporcatus Wiegmann 1834
and Anolis frasert 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 biporcatus 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, pl. 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 biporcatus specimen from the syntype series, placing
it in “‘copu’’ (=copei Bocourt 1873, a synonym of biporcatus').
He was, however, apparently quite convineed that ‘‘copii’’ and
fraserv were closely related; he placed them side by side in his
catalogue and a collation of his two descriptions shows 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 biporcatus has suggested that fraseri is the
South American representative of biporcatus.
A contrary view that A. biporcatus and A. fraseri 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 biporcatus or its synonym
copei goes quite unmentioned. Most recently a more explicit
denial of close relationship has been propounded: R. Etheridge,
1 See Stuart (1948) for discussion of the usage of the name biporcatus Wiegmann
rather than copei Bocourt. The question is one of the identity of the Berlin type
ot biporcatus.
2 BREVIORA No. 239
in a doctoral thesis (1960) at the University of Michigan, has
placed A. fraseri and A. biporcatus 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 biporcatus and fraseri has been
the question of the presence of biporcatus 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 biporcatus
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 biporcatus synonym
‘“copu,’’ 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 Gunther syntype in the
svnonymy 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 biporcatus 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 Museum (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 Museum (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 biporcatus Wiegmann does occur in both Colombia
and Eeuador and also in Venezuela.
2. Anolis solifer Ruthven must be considered a synonym of
A. biporcatus Wiegmann.
3. A. biporcatus Wiegmann is at least in a broad sense sym-
patrie with A. fraseri but the sympatric populations appear not
to belong to the nominate race.
4. A. biporcatus 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
a SS =
FE OS
Crees Unsere
sully 9 ano!
Sarcce
a SS
\
Oe Ue S
oy
fA
ea
CZ.
r\
) o>
i Veneers
A
Wiitetrerd
Ss
CSL
i
(
L/
t
Fig. 1. Anolis fraseri, MCZ 70227. Dorsal view of head.
4 BREVIORA
No. 239
TABLE |
South American biporcatus
Head scales, at least anteriorly,
keeled.
2-3 scales between circumnasal
and rostral.
Supraorbital semicircles sepa-
rated by 0-3 scales.
Supraocular seales keeled.
2-3 more or less elongate supra-
ciliary scales followed by a dou-
ble series of scales which 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.
7-12 supralabials to center of
eye.
Dewlap scales strongly keeled,
almost as large as ventrals,
rather uniformly distributed on
the small fan.
Ventral scales strongly keeled,
mucronate.
22-26 lamellae under phalanges
ii and 11 of 4th toe.
fraserv
ITead scales never keeled, usu-
ally flat, pavementose, but some-
times swollen or wrinkled.
1-2 scales between ecircumnasal
and rostral.
Supraorbital senucircles sepa-
rated by 2-4 scales.
Supraocular scales smooth.
Usually no elongate supracili-
ary scales; mstead, the whole
supraciliary margin of round
or squarish scales distinetly
larger than the granules medial
to them.
4-9 loreal rows.
Interparietal larger (larger
than the ear which is moderate
in size).
Subocular scales broadly im con-
tact with supralabials.
6-9 supralabials to center of
eye.
Dewlap scales smooth, small,
almost granular, grouped im
rows which are separated by
naked skin.
Ventral scales smooth or weak-
ly keeled.
20-24 lamellae under phalanges
i1 and iii of 4th toe.
1966 ANOLIS BIPORCATUS AND ANOLIS FRASERI 5
in one species completely overlaps the known range of the other,
I have omitted this character. The most striking differences are
emphasized by 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. The similarities would be even
greater if Mexican and other northern biporcatus were com-
pared with fraserz since the northern animals tend to have 3-4
scales between the supraorbital semicircles (cf. Stuart, 1955).
Overlapping variation in many features, as shown in Table 1,
is characteristic of many sympatric species in Anolis and is one
of the several reasons why this genus is described as ‘‘difficult.’’
As in all such eases, the recognition of the valid species depends
upon recognition of the constant (or almost constant) associa-
tion of characters, however trivial. Smooth or keeled scales, sub-
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 biporcatus will always distin-
euish these two species. (The frasert supraciliaries occur, how-
ever, also and in a more exaggerated degree in latifrons; the
condition is not unique to fraser.)
a a 0) i
ZE PE ES ; Oueeezes
LEELA] BRO CEG Sho eeaess
Ee le AMO EE eee
~ ¢ ic ~ Yr OY =
Cl 7
aya C
BOCES
Fig. 2. Anolis biporcatus parvauritus, new subspecies, MCZ 78935. Dorsal
view of head of type.
6 BREVIORA No. 239
INTRASPECIES VARIATION: A. BIPORCATUS
While I have not endeavored to examine all specimens of A.
biporcatus 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. biporcatus, as I understand it. Thus, the number of
scales between the semicircles, the number of seales 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 precisely those
sympatric with frasers—it seems permissible to regard the
Keuadorian and southern Colombian animals as representing a
distinct subspecies. Some discussion of available names is there-
fore required.
Anolis solifer Ruthven, type locality La Concepcién, 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 biporcatus 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] am indebted to Dr. Allan Leviton of the California Academy of Sciences for
this opportunity.
1966 ANOLIS BIPORCATUS AND ANOLIS FRASERI if
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
Eeuadorian and south Colombian populations. In the critical
characters solifer resembles northern biporcatus.
The other names referable to the species biporcatus are equally
unavailable. None of them refer to South American material.
The nominate race biporcatus Wiegmann 1834 has the type
locality ‘‘Mexico,’’ now restricted to Piedra Parada, Chiapas.
A. copet Bocourt 1873 has the type locality Santa Rosa de Pansas,
Guatemala. Anolis brevirostris Peters 1873, next in date, has the
type locality Chiriqui Prov., Panama. Anolis brevipes Boettger
1893 came from Cairo Plantation, La Junta near Limén, Costa
Rica. Every one of these seems at present best grouped under
the name biporcatus.
The southern race is thus without a name. In reference to the
small ear it may be ealled:
ANOLIS BIPORCATUS PARVAURITUS new subspecies
Holotype: MCZ 78935, an adult ¢, ‘‘banana plantation, woods
and penal colony camp, northern Gorgona Island, Cauea,
Colombia, 5-45 meters altitude,’’ collected by F. Medem and lL.
Salazar G., 1 to 23 February 1961.
Paratypes: Colombia. Cauca: MCZ 78933-34, 78936-41, same
data as type. Narino: MCZ 79142, Rio Mataje. Ecuador. Es-
meraldas: USNM 157105, Cachavi; BM 1901.6.27.1, Carondolet ;
<
awe
sO)
e
Ss
Ss
cn
we
“
Se
i
o
ne,
SS
= IPS Ss PY AW ESI
Se SSS TR
SST YO
SSS Se Sana
Fig. 3. Anolis biporcatus biporcatus, MCZ 56248, from Los Diamantes,
Costa Rica. Dorsal view of head.
8 BREVIORA No. 239
AMNH 6967, Rio Capayas; 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. Imbabura: BM 98.4.28.18,
Paramba. Pichincha: Orces 4596, Santo Domingo de _ los
Colorados. ‘‘ West Ecuador’’: BM 1946.8.13.21 (fraseri syntype).
‘*Heuador’’: USNM 20610, ANSP 7910.
Diagnosis. A subspecies of Anolis biporcatus 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 dorsally), and by 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, ete.) reveal the distinet-
ness of parvauritus, though it is visually apparent immediately
on inspection of the specimens (Figs. 2 and 3). The number of
scales between interparietal and semicircles might be expected
to eorrespond neatly to the very striking difference in the size
of seales surrounding the interparietal in the nominate race and
in parvauritus. In faet, 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 parvauritus an absence
of clear definition of any supraocular disk. This, however, 1s
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 difference. 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 appreciated
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 rather
large. In other regards (scales around interparictal, supraocular
scales) these are northern in type. MCZ 79842 from Sincelejo,
1966 ANOLIS BIPORCATUS AND ANOLIS FRASERI 9
They
EL
ee |
>=
Fig. 4. Ear size in A. biporcatus biporcatus (top) and <A. biporcatus
parvawritus (bottom). Specimens are of the same snout-vent length.
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 (CNHM 43778),
appears, in agreement 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 Colombia.
10 BREVIORA 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 hght 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 seales.’’
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 ‘‘solifer’’ is an available
name, is unknown.
Behavioral notes. F. Medem (field notes) remarks on the
Gorgona Island topotypie series of parvauritus that the gular
sac 1s ‘‘extended laterally’’ and that this species is here found
1 The description by Taylor (1956, p. 136) of the dewlap of Costa Rican speci-
mens as ‘white or slightly greenish” appears on careful reading to result from a
eomplex confusion of notes for A. capito with the text for A. biporcatus, and ap-
parent resulting omission of some of the pertinent data for biporcatus. See the
reference to A. capito, top of page 137.
1966 ANOLIS BIPORCATUS AND ANOLIS FRASERI 141
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. frasert, 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 ight and dark to the vertebral zone. This statement,
of course, apples 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. frasert, 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 Marta Mountains into
Zulia and Tachira in Venezuela. A. frasert—a South American
endemie — is apparently restricted to Colombia and Ecuador
west of the Andes, and is mainly Eeuadorian. I list below the
1 Anolis devillei Boulenger 1880 (type examined) is an unequivocal synonym of
A. fraseri Giinther 1859, and Boulenger (1885) himself synonymized it. The
and indeterminable ; Boulenger utilized it with great hesitation.
12 BREVIORA No. 239
South American A. biporcatus examined (except for the para-
types of parvauritus, which have been listed previously) and also
all specimens of A. fraser.
Anolis biporcatus. Colombia. Antioquia. Villa Arteaga, MCZ
79656-57. Magdalena. La Concepeidn, Santa Marta Mtns., MCZ
6549 (type of solifer Ruthven). Bolwar. Sincelejo, MCZ 78942.
Turbaco (?), Leiden 2807. Norte de Santander. Near Rio Zulia,
32 km N Cuteuta, ILS 19. Choco. Pizarro, CNHM 48778.
Venezuela. Tiichira. La Fria, UMMZ 55994. Zulia. La Kasmera,
Parija 300 m, UCV 8023. ? Brasil. ‘‘Silva, See Saraca,’’ VM
5904.1
Anolis fraserit. Colombia. Cauca. Buenaventura, CNHM
43771; El Tambo, ANSP 25563; Jamundi, CNHM 48772.
Ecuador. Hsmeraldas. San Mateo, CNHM 27681. Imbabura.
Apuela, JAP 4831. Pichincha. Mindo, UMMZ 55525; Nanegal,
BM 83.2.23.11, JAP 925; Quito, BM 72.2.26.16. Gwayas. Rio Pes-
eado, nr. Naranjal, AMNH 23432; Bucay, AMNH 23030. El Oro.
7 km SE Buenavista, JAP 2485. ‘‘ West Eeuador’’: BM 60.6.16.36
(1946.8.8.47) (lectotype of frasert). ‘‘Equateur’’: IRSN 2006
(type of deviller Boulenger ).
REFERENCES CITED
BaRBour, T.
1934. The anoles. II. The mainland species from Mexico southward.
Bull. Mus. Comp. Zool., 77: 121-155.
BoETTGER, O.
1893. Katalog der Reptilien-Sammlung im Museum der Senckenber-
gischen naturforschenden Gesellschaft in Frankfurt am Main.
I Teil. (Rhyneophalen, Schildkréten, Krokodile, Eidechsen,
Chamileons). x + 140 pp. Frankfurt-a-M.
30courT, F.
1873. Etudes sur les reptiles. Mission scientifique au Mexique et dans
1’Amerique Centrale. Pt. 3. Sauriens. Livr. 2: 33-112. Paris.
BOULENGER, G. A.
1880. Reptiles et batraciens recueillis par M. Emile DeVille dans les
Andes de 1’Equateur. Bull. Soc. Zool. France, 1880: 41-48.
1885. Catalogue of the lizards in the British Museum (Natural His-
tory), 2: ix + 497 pp. London.
Burt, C. BE. and M. D. Burt
1931. South American lizards in the collection of the American Museum
of Natural History. Bull. Amer. Mus. Nat. Hist., 61: 227-395.
1{ assume that this is Lago Saraci, Amazonas, but this locality is very remote
from any other record. There are no other specimens of biporcatus from Brasil.
Locality records in the Vienna Museum are so often erroneous that I cannot accept
this without confirmation.
WY
1966 ANOLIS BIPORCATUS AND ANOLIS FRASERI ile
DUELLMAN, W. E.
1963. Amphibians and reptiles of the rain forests of southern El Peten,
Guatemala. Univ. Kansas Publ. Mus. Nat. Hist., 15: 205-249.
DuNN, HE. R.
1937. The giant mainland anoles. Proc. New 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, Ine., Ann Arbor, Michigan.
GUNTHER, A.
1859. Second list of cold-blooded vertebrates collected by Mr. Fraser in
the Andes of western Ecuador. Proc. Zool. Soc. London, 1859:
402-420.
PETERS, W.
1874. Uber neue Saurier (Spoeriodactylus, Anolis, Phrynosoma,
Tropidolepisma, Lygosoma, Ophioscincus) aus Centralamerieca,
Mexico, und Australien. Monatsber. Akad. Wiss. Berlin, 17
November 1873: 738-747.
RUTHVEN, 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 Boquete, Panama, with the
description of a new species of Hydromorphus. Proc. California
Aead. Scei., Ser. 4, 23: 463-480.
SmirH, H. and HE. H. TAYLOR
1950. An annotated checklist and key to the reptiles of Mexico, ex-
elusive 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.
Mise. Publ. Mus. Zool. Univ. Michigan, 91: 1-31.
Tayior, 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.
WILLIAMS, E. E.
1963. Studies on South American anoles. Description of Anolis mirus,
new species, from Rio San Juan, Colombia, with comment on
digital dilation and dewlap as generic and specific characters in
the anoles. Bull. Mus. Comp. Zool., 129: 463-480.
(Received 15 September 1965. )
14 BREVIORA No. 239
80° TO 602
a
El (Qe
* Anolis fraseri
e A. biporcatus parvauritus
200 mi.
/0°
Fig. 5. Map of distributions of A. fraseri and A. b. parvauritus. Inter-
mediates and representatives of the typical race of biporcatus in northern
Colombia and Venezuela omitted.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. FEBRUARY 25, 1966 NuMBER 24(0)
GYMNOTHORAX 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 this 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 GYMNOTHORAX Bloch 1795
+Y MNOTHORAX 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 leneth. 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
‘‘nalatal 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-
eround 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 oceasionally 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: Gymnothorax galetae does not resemble the adult or
immature stages of any other Atlantic species of Gymnothorax.
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. vicinus 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. isingteena and G. stellatus 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 lm-
ited material available.
ACKNOWLEDGMENTS
I wish to acknowledge the aid which the Evolutionary Biology
Committee of Harvard University and the National Science
Foundation (through GF 3450) have extended to the author.
LITERATURE CITED
GINSBURG, I.
1951. The eels of the northern Gulf Coast of the United States and
some related species. Texas Journal of Science, 3 (3): 431-485.
HILDEBRAND, S. F.
1938. The Panama Canal as a passageway for fishes, with lists and
remarks on the fishes and invertebrates observed. Zoologica
CNP YS) 2473) ie) 15-45"
RvusBINorr, 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.)
No. 240
BREVIORA
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. FEBRUARY 25, 1966 NuMBER 241
AVOCETTINOPS YANOI, A NEW NEMICHTHYID EEL
FROM THE SOUTHERN INDIAN OCEAN
By Gites W. Meap 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 Bertin,
1924) ; a second specimen taken off Zanzibar during the ‘‘John
Murray’’ expedition (Norman, 1939; the specimen rather casu-
ally christened Avocettinops norman 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 with 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.
i)
BREVIORA No. 241
Distinctive characters. — Avocettinops yanoi differs from its
congeners 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
(cf. 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. — Meristie 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. Guill slits short (13
in length of head), ventrally directed, and placed below bases of
pectoral fins. Eye cireular, 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. Most 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 suecess 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 fragmentary. 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.
yanor, 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.
4. BREVIORA No. 241
TABLE 1
Proportional dimensions, expressed as per cent of length of head,
of Avocettinops schnudti and A. yanov. 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.
Western North
Type of Atlantic Type of
A. schmidti (SU 47758) A. yanoi
Total length (mm) 510.0 a= 620.0
Leneth of head (mm) 32.0 32.0 50.0
Depth of body at anus
(% of h.) 21.9 24.4
Greatest depth of body 43.8 28.4 39.8
Width of body at anus 9.4 ca. 7.8 15.4
Greatest width of body 10.9 -- 16.4
Greatest depth of head 25.6 ca. 23.4 26.4
Greatest width of head 18.8 17.5 19.8
Leneth of snout 25.0 25.0 24.0
Diameter of orbit 15.6 12.5? 13.4
Postorbital length of head 59.4 ra) 62.4
Length of upper jaw 39.1 33.1 34.8
Length of lower jaw 32.8 24.7 32.8
Leneth of gill slit 13.1 Wie2 7.6
Interorbital width 14.1 — 16.6
Predorsal leneth 90.6 108.8 73.8
Preanal distance 203.1 207.8 127.4
Width of base of pectoral fin 14.1? eT 9.0
Leneth of pectoral fin 28.1 24.7 29.0
Dorsal fin rays 340 = 285
Anal fin rays 315 oe 266
Pectoral fin rays 16-17 5-15 12-13
Total number of vertebrae 194 184
Preanal vertebrae (+1) 20 24 1133
Total number pores in lateral
line 188 = 185
Lateral-line pores, temporal
pore to anus 18? 23 13
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LITERATURE CITED
BERTIN, LEON
1947. Notules ichthyologiques. 1.— Compléments sur le genre Avocet-
tinops. Bull. Soc. Zool. France, 72:54-55.
BOuLKE, JAMES E., and FRANK S. CLIFF
1956. A discussion of the deep-sea eel genus Avocettinops, with notes
on a newly discovered specimen. Copeia, 1956:95-99.
NORMAN, J. R.
1939. Fishes. Sci. Rep. ‘‘ John Murray’’ Expedition, 7(1) :116 pp.
RouLE, Louis, AND LEON BERTIN
1924. Notice préliminaire sur la collection des nemichthydés recueillie
par l’expédition du ‘‘Dana,’’ (1921-1922), suivie de considéra-
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 Nemichthydi-
formes. Rep. ‘‘Dana’’ Exp., 1920-1922, No. 4, 113 pp., 19 pls.
(Received 9 November 1965.)
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‘
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MAss. FEBRUARY 25, 1966 NuMBER 242
THE SUPPOSED ‘‘SPONGE SPICULES” OF
MERRILL, 1895, FROM THE LOWER CRETACEOUS
(ALBIAN) OF TEXAS
By WituiaM A. S§. SARJEANT
Geology Department
University of Nottingham, Nottingham, England
INTRODUCTION
In 1895, a paper was published by J. A. Merrill giving the
results of a study of chert nodules from Texas. In this he de-
seribed 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 which
were doubtfully attributed to the living genus Geodia.
This work eseaped 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 United 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,
pli).
Through the courtesy of Dr. H. B. Whittington, the author
has been permitted to examine Merrill’s type 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
D BREVIORA No. 242
Fredericksburg Formation, Comanchean 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
shdes 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 shde consisted of a square of glass measuring 3” by 2” by
1,” 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 balsam
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 (shde IIT).
Merrill states that ‘‘several slides’? were made from each
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 eases, either that the
holotypes were contained in other slides now lost, or that they
were contained in the missing half of flint section ITI.
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 Ene-
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 hystri-
chospheres were silicified and placed them in the freshwater
desmid genus Xanthidium. 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 in Xanthidium (1845).
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 ‘‘ehosts’’ 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 study, for, as
will be shown in the ensuing section, the microfossils he described
as sponge spicules include spores, pollen, hystrichospheres, wood
and plant fragments, and even carbonate crystals.
Merrill noted the relatively low concentration of microfossils in
the chert (p. 6):‘*...the number 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 Foraminifera 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. ‘‘Monactinelid: Axinella ? sp.’’ The figured spec-
imens were not relocated; however, they appear to consist of
sections through shell fragments which have been 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: Reniera ? sp.’’ 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; acuate 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: Esperites ? 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 replaced shell fragment or a
fragment of organic tissue.
1966 CRETACEOUS CHERT FOSSILS 5
10. ‘*‘Monactinellid: Renicra ? 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: Geodia ? austini n. sp.’’ This species was
tentatively transferred to the genus Systematophora in an earlier
paper (Sarjeant, 1964, p. 175). A specimen closely resembling
the holotype is present in slide I (specimen IA: Pl. 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 (Khrenberg, 1838) O. Wetzel 1933, emend. Davey
and Williams 1965a. The species name austini must therefore be
rejected, under Article 63 of the ‘‘ International Code of Botan-
ical Nomenclature.’’
12. ‘‘Tetractinellid: Geodia ? cretacea n. sp.’’ This species
was transferred to the genus Baltisphaeridium 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 (PI. 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
Xanthidium hirsutum, which is that of a living desmid. Lejeune-
Carpentier (1941) redescribed Ehrenberg’s fossil material, under
the name Hystrichosphaeridium hirsutum, but Deflandre (1946)
correctly pointed out that the trivial name hirsutum could not be
legitimately transferred to another genus and applhed to the fossil
forms. Deflandre had earlier (1937) proposed a new species
H. striolatum 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 cretaceum is
a senior synonym of striolatum, 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 species striolatum
was soundly based, fully described and adequately figured; the
species cretaceum was, in contrast, profoundly misinterpreted
and inadequately figured and described. Moreover, complete cer-
tainty in the recognition of the holotype is not possible; and full
lop)
BREVIORA No. 242
Text-figure A
A reproduction of Merrill’s plate (1895). The specimens are here labelled
according to the interpretation given in the present paper. Fuller discussions
of the assignations are given in the text.
1-4,
Shell fragments sectioned at varying angles.
A fragment of organic material.
Wood fragments.
Shell lining of a unilocular foraminifer (?).
Character not clear.
Fragment of organic material.
Hystrichosphaera ramosa (Ehrenberg, 1838).
Exochosphaeridium cretaceum (Merrill, 1895) =
E. striolatum (Deflandre, 1937).
Hystrichodinium pulchrum Deflandre (?).
Oligosphaeridium complex (White, 1842).
Hystrichosphaera ramosa (Bhrenberg, 1838).
Spore.
Character not clear.
Baltisphaecridium texanum (Merrill, 1895).
Chlamydophorella sp. (2).
Hystrichosphaera ramosa (Ehrenberg, 1838).
Hystrichosphaeridium tubiferum (H. H. White, 1842).
Detached spines of Hystrichosphaera ramosa (Ehrenberg,
1838).
Character not clear.
““Brown bodies’’ of Bryozoa (?).
Carbonate crystals.
Dinoflagellate cyst, probably Microdinium sp. (?).
Character not clear.
Probable sponge spicules.
Fragment of Hystrichosphaera ramosa (Bhrenberg, 1838 )
(2).
Fragment of organic material.
CRETACEOUS CHERT FOSSILS
1966
8 BREVIORA No. 242
and accurate redescription 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-
latwm is in widespread use; the name cretaceum has not been
reused since its original publication, 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
hame cretaceum, im view of its more adequate description. The
correct generic assignation is currently to Hwochosphaeridium
Davey, Downie, Sarjeant and Williams, 1966.
13. **Tetractinellid: Geodia ? spini-curvata n. sp.’’ This species
was transferred to the genus Baltisphaeridium in an earler
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 form with
relatively few, simple spines and havine a precingular archae-
opyle; it is probably referable to the genus Hystrichodinium De-
flandre 1935 emend. Sarjeant 1965, perhaps to H. pulchrum De-
flandre 1935, a species very frequent in Enelsh cherts. How-
ever, since the holotype is lost and since neither description nor
figure permit certain statement of its characteristics, the species
name sprni-curvatum cannot continue in use.
14. ‘‘Tetractinellid: Geodia ? irregularis n. sp.’’ This species
was transferred to the genus Hystrichosphaeridium 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 doubt that this species is a junior
synonym of Oligosphacridium complex (H. H. White, 1842),
Davey and Williams 1966a, a species frequent in the Cenomanian
and known to range down well into the Lower Cretaceous.
15. ‘‘Tetractinellid: Geodia ? tripunctata n. sp.’’? Rejection
of this species on the grounds that it is a junior synonym of
Hystrichosphaera ramosa (Ehrenberg, 1838) O. Wetzel 1933,
emend. Davey and Williams 1966a, has already been proposed
(Sarjeant, 1964, p. 175). A number of representatives of the
species H. ramosa were encountered, including perhaps Merrill’s
holotype (specimen IIIC: Pl. 1, fig. 4); the earlier judgement is
fully supported.
1966 CRETACEOUS CHERT FOSSILS i)
16. ‘*Tetractinellid: Hymeraphia ? sp.’’ The figured specimen
was not found; however, 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 trilete mark) and
WelcAn GRIMS fies 5):
17. **Tetractinellid: Chondrilla ? sp.’’ The figure and deserip-
tion of this form are so indefinite as to render impossible deter-
mination of its true character. One specimen encountered (IIC:
Pl. 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: Geodia? texrana 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 teranwm
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
Chlamydophorella; unfortunately, this probable assignation
cannot be confirmed, since the figured specimen was not relocated.
20. ‘‘Tetractinellid: Geodia? spinipansata n. sp.’’? This
species was transierred to the genus Hystrichosphacridium in an
earlier paper (Sarjeant, 1964, p. 275). It is probable that a speci-
men encountered in slide IIT (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) O. Wetzel
1933, emend. Davey and Williams 1966a; this species shows some
range of variation in morphology (ef. Davey and Willams,
1966a), and also varies in appearance according to orientation.
The spines are not tubular, so that assignation to Hystricho-
sphaeridium is incorrect. The species spinipansatum must be re-
jected, as a junior synonym of ramosa.
10 BREVIORA No. 242
21. ‘‘Tetractinellid: Geodia? hilli n. sp.’’ This species was
transferred to the genus Hystrichosphaeridium (Sarjeant, 1964).
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
apphed to No. 14, ‘* Nothing similar has been found figured,’’ the
specimen he figures is without question attributable to the species
Hystrichosphacridium tubiferum (BKhrenberg, 1838) O. 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 tubiferum.
22. **Tetractinellid: Geodia tripunctata ? n. sp. Fragments
resulting from solution.’’ Unquestionably these are detached
spines of Hystrichosphaera ramosa. A number of such detached
spines were seen, none corresponding precisely to the grouping
shown in the figure, which might in any case 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; Hymeraphia ? sp.’’ Over-simpli-
fied figures and descriptions again render certain recognition
difficult. Pollen grains of comparable morphology were noted,
and also (in slide X) a species of dinoflagellate cyst questionably
attributable to the genus Prolixosphaeridium Davey, Downie,
Sarjeant and Williams 1966, with a similarly elongate shape and
cover of simple spines (PI. 1, fig. 8).
like structures were encountered in several cherts (e.g. VIT B;
Pl. 1, fig. 11). Similar structures occur widely in palynological
material from the Upper Mesozoic; Merrill’s description (p. 17)
is quite accurate. Their interpretation is doubtful; Otto Wetzel
(1961, pl. 1, figs. 8-10) has figured similar forms as ‘‘brown
bodies’ of Bryozoa; it is also possible that they represent remains
of colonial algae, possibly Chlorophyta.
27. ‘*Tetractinellid: Geodia ? sp.’’? Groups of circular, dise-
28. ‘*Tetractinellid: dermal spicules ?’> These are 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 sometimes considered to
1966 CRETACEOUS CHERT FOSSILS 1]
indicate that the chert was formed by replacement of a pre-exist-
ing limestone. Merrill’s misinterpretation of them is surprising
GELS fie),
29. ‘*Tetractinellid: Geodia sp.’’? The figure may correspond
to a specimen in slide III, which resembles it in general features
(specimen IITH). Although full study is difficult because of its
situation deep in the chert, this is certainly a dinoflagellate eyst,
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, possibly
pollen,
31. ‘‘Geodia ?”’ The figure and description are inadequate to
permit recognition of this form.
32. ‘‘Geodia ? Pyramidal or zone spicule.’’ Not relocated ;
most probably correctly interpreted as sponge spicules.
33, 34. ‘‘Hexactinellid. Stauractinellia ? sp.’’> Not relocated ;
most probably correctly interpreted as sponge spicules.
35. ‘‘Frame work of a Hexactinellid?’’ This appears to be a
fragment of a Hystrichosphaera furcata, although the figure is
extremely generalised and precise interpretation is difficult. A
sinilar fragment was encountered in slide III (specimen IIIE).
36. ‘‘Monactinellid: Reniera ? sp.’’ The sausage-shaped ob-
ject in Merrill’s figure is most probably to be interpreted as a
fragment of organic tissue (? plant); many similar fragments
were seen.
CONCLUSIONS
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 dinoflagellate eysts (mainly — hystricho-
spheres) ; these include his eight proposed new species, of which
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 holotype
of the third species does not permit full study.
12 BREVIORA No. 242
The remaining 20 microscopic structures figured 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 foraminiferal shell lining; one is a spore; three
possibly pollen; one represents an association of dise-like strue-
tures, possibly ** brown bodies’’ of Bryozoa; one comprises carbon-
ate crystals; two represent wood fragments; four represent frag-
ments of organic (? plant) tissue of varying shape; and three
are of indeterminate character.
The currently valid species of dinoflagellate cysts recognised
in Merrill’s figures and material are Hystrichosphaera furcata,
Hystrichosphaeridium tubiferum, Exochosphaecridium striolatum,
Oligosphaeridium complex, Hystrichodinium pulchrum (doubt-
ful), and undetermined species of the genera Gonyaulacysta, Pro-
lirosphaeridium, Hystrichodinium, 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 been obtained, the species represented are all ones
which would be likely to occur in Middle to Upper Albian prep-
arations.
REFERENCES
Davey, R. J., C. Downigz, W. A. S. SARJHANT and G. L. WILLIAMS
1966. Fossil dinoflagellate cysts attributable to Baltispraeridium. In
Studies of Mesozoic and Cainozoie dinoflagellate cysts. Bull.
Brit. Mus. (Nat. Hist.) Geol., Suppl. 3 (in press).
Davey, R. J. and G. L. WILLIAMS
1966a. The genera Hystrichosphaera and Achomosphaera. In Studies of
Mesozoic and Cainozoic dinoflagellate cysts. Bull. Brit. Mus.
(Nat. Hist.) Geol., Suppl. 3 (in press).
1966b. The genus Hystrichosphaeridium and its allies. Jn Studies of
Mesozoic and Cainozoic dinoflagellate cysts. Bull. Brit. Mus.
(Nat. Hist.) Geol., Suppl. 3 (in press).
DEFLANDRE, G.
1935. Considérations biologiques sur les micro-organismes d’origine
planectonique conservés dans les silex de la craie. Bull. Biol. Fr.
Belg., vol. 69, pp. 213-44, pls. 5-9.
1937. Microfossiles des silex crétacés II. Ann. Paléont., vol. 26, pp.
51-103, pls. 8-18.
1946. Remarques sur la systématique des hystrichosphaeridés. C. R.
Somm. Soc. Géol. 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, pl. 1.
GLAESSNER, M. F.
1945. Principles of micropalaeontology. Melbourne: University Press,
296 pp., 14 pls.
LEJEUNE-CARPENTIER, M.
1941. L’étude microscopique des silex (9 iéme note). Sur Hystricho-
sphaeridium hirsutum (Ehr.) et quelques formes voisines. Ann.
Soc. Géol. 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. IIT),
vol. 28, no. 1, pp. 1-26, 1 pl.
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 acritarchs. J. Paleont.,
vol. 38, pp. 173-7.
1966. Dinoflagellate cysts with a Gonyaulax type tabulation. In Studies
of Mesozoic and Cainozoie dinoflagellate cysts. Bull. Brit. Mus.
(Nat. Hist.) Geol., Suppl. 3. (in press).
WETZEL, O.
1933. Die in organischer Substanz erhaltenen Mikrofossilien des Bal-
tischen Kreide-Feuersteins. Palaeontographica, vol. 77, pp. 141-
88; vol. 78, pp. 1-110, pls. 1-7.
1961. New miecrofossils from Baltic Cretaceous flintstones. Micro-
palaeontology, vol. 7, no. 3, pp. 337-50, pls. 1-3.
Wuits, H. H.
1842. On fossil Xanthidia. Microsc. J., vol. 11, pp. 35-40, pl. 4.
(Received 25 October 1965)
14 BREVIORA No. 242
Plate 1
Fig. 1. Presumed shell lining of a unilocular Foraminifer (Fig. 7
**“Monactinellid’’ of Merrill). & 500.
Figs. 2, 3. Gonyaulacysta sp. indet., seen in oblique apical view (Fig. 2)
and, by transparency, in oblique antapical view (Fig. 3) —the antapex is
at left. Figure 2 closely resembles ‘‘Tetractinellid; Chondrilla 2? sp.’’ of
Merrill. & 500.
Fig. 4. Hystrichosphaera ramosa (Ehrenberg). The presumed holotype
of Merrill’s invalid species Geodia ? tripunctata. > 500.
Fig. 5. Hystrichosphaera ramosa (Ehrenberg). Possibly the holotype of
Merrill’s invalid species Geodia ? spini-pansata. >< 500.
Fig. 6. Hxochosphaecridium cretaceum (Merrill). The specimen may be
the holotype. & 500.
Fig. 7. Hystrichosphaera ramosa (Ehrenberg). A specimen in terminal
view, illustrating the high degree of variation in appearance given by
orientation. & 500.
Fig. 8. Prolixosphaeridium sp. indet. (Possibly corresponding to Merrill’s
Fig. 26b ‘‘Tetractinellid; Hymeraphia ? sp.’’). ©300.
Fig. 9. Spore, gen. et sp. indet. (Similar to Merrill’s Fig. 16 ‘‘ Tetracti-
nellid; Hymeraphia ? sp.’’?; but lacking a clear trilete mark.) & e300.
Fig. 10. Hystrichosphaera ramosa (Ehrenberg). Possibly the holotype of
Merrill’s invalid species Geodia ? austini. *& 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?’’ > 500.
1966 CRETACEOUS CHERT FOSSILS 15
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. AprRIL 29, 1966 NuMBER 243
QUATERNARY FISH FOSSILS FROM WEST OF LAKE
RUDOLF, KENYA
By KeityH Srewart THOMSON
Department of Zoology, University College London.1
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. 8. 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 voleanies.
Away from the bases of the hills —and here we explored only
eastwards — are flat plains covered with 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 address: Dept. of Biology and Peabody Museum of Natural History,
Yale University, New Haven, Conn.
DZ BREVIORA No. 2438
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 shg¢ht 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 Turkwell River has made a large cut through
some hundred feet of the lake deposits. Patterson and Williams
measured the exposed section as follows:
Upper unit, bed 10 Surface white sands
fSo9 4 feet black clay
OG 3 16 feet white coarse sands with gravel lenses
i SAS feet grey clay
SO 6 10 feet micaceous sand
CG 15s 4 feet black clay
Oe 4: 1% to 5 feet coarse sand and gravel
SG 5 feet black clay
34 foot white sand
2% feet black clay
are 9 feet interbedded black elays and grey
sands
CC eal 12 feet black clays
Lower unit, bed 5 7 feet fine grey sand
oO 4, 346 feet fine grey sand highly indurated with
iron
Cr 3) 5 feet fine white sands, black micaceous
flakes
SG 4 feet medium yellow sands, black micaceous
flakes
co 1) «(12% feet coarse grey sands with large clay
pebbles in upper unit (base con-
cealed )
‘‘Upper unit’’ and ‘‘lower unit”’ are informal, nou-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. Shehtly 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.
1966 QUATERNARY FISHES FROM KENYA 3
Molluse, fish, reptilian and mammalian remains are common in
the upper unit of the exposures, molluse 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.
o Muruangering
Nuees tt -
. Muruadou
3000 v Karsa
Spring
Ferguson :
Gul Gentral
L ksland
ie a6 ° Glendays
Ss E Camp
Chavakono
"2500
° Goritei
Kholotum o
Kangatotha
e
Miles 20 15 I
i
fo) 5
Sketch map of Lake Rudolf 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. Barbus
Two dorsal pterygiophore fragments
Order Percomorphi
Suborder Percoidea
Family Centropomidae |
Lates ef. Lates niloticus Cuvier and Valenciennes |
One incomplete parasphenoid (length 150 mm), —
three vertebrae (diam. 25, 33, 46 mm), four fin
spines (88, 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
ef. Barbus
Tentatively referred to this genus are sections of
two fin spines.
Order Percomorphi
Suborder Perecoidea
Family Centropomidae
Lates ef. Lates niloticus Cuvier and Valenciennes
Four dorsal fin spines (28, 42, 45, 51 mm)
Family Cichlidae
Tilapia sp.
One pterygiophore — first interhaemal (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. Bagrus bayad (Geoffroy )
Posterodorsal portion of a single occiput (estimated
height 28-30 mm)
Family Mochokidae
Synodontis cf. Synodontis schall Cuvier
Humeral process of left cleithrum (49 mm long),
one posttemporal bone, and fragments of four
pectoral spines
Family Claridae
Clarias ef. 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,
12S, ta Ib bs 6. Te oli am)
BREVIORA No. 2438
Suborder Cyprinoidea
Family Cyprinidae
Barbus cf. Barbus bynni 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 ef. Synodontis schall Cuvier.
The humeral process of one cleithrum (length 55
mm), one dorsal fin spine, three pectoral fin
spine fragments
ef. Synodontis
Fragments of three spines
1966 QUATERNARY FISHES FROM KENYA il.
Family Clariidae
Clarias ef. Clarias lazera Cuvier and Valenciennes
Three dermethmoids (2 complete, lengths 34, 45
mm; one incomplete, anterior width 50 mm),
one posttemporal, two sections of supraoccipital
(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
Barbus ef. Barbus bynni Cuvier and Valenciennes
Four pharyngeal bones (approx. length 30, 30, 40,
45 mm), two (?dorsal) pterygiophore frag-
ments
ef. Barbus
Two vertebrae (diam. 14, 15 mm)
Order Percomorphi
Suborder Percoidea
Family Centropomidae
Lates ef. 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 bayad (Geoffroy )
One occiput, incomplete (max. depth 69 mm)
?Family Clariidae
Dermethmoid portion of skull, incomplete
Order Percomorphi
Suborder Percoidea
Family Centropomidae
Lates cf. Lates niloticus 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
Lates ef. 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 bayad, Synodontis schall, Clarias lazera, Lates
niloticus, Barbus bynni, and Tilapia nilotica — only Bagrus
bayad, 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-
seribed 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 Makahan or
Nakuran ‘‘wet phases’’ (see Cooke, 1957, 1963). A radio-carbon
date of 4,800 + 100 years was obtained from a sample of Etheria
elliptica 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
siant forms of Lates, described by Daget, White, 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 1’?Omo 1932-1933. Tome 1. Géologie-
Anthropologie. Fase. II. Mus. Nat. Hist. Natur. (Paris). Pp.
1-74.
Cook, H. B. S.
1957. Observations relating to Quaternary environments in east and
southern Africa. Trans. Geol. Soe. 8S. 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. Bourliére,
eds., African Ecology and Human Evolution. Aldine Co., Chi-
cago.
DAGET, J.
1959. Restes de poissons du Quaternaire saharien. Bull. Inst. Frane.
Afr. Noire, 23: 182-191.
DE HEINZELIN, J.
1957. Les feuilles d’Ishango. Exploration du Pare Albert, 1957: 46-
61.
IMU CHSSaViy LE.
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.
Waits, EH. I.
1926. The geology and paleontology of the Kaiso bone-beds. Part II.
Palaeontology. Fossil Pisces. Oecas. Paper Geol. Surv. Uganda,
2: 45-51.
WORTHINGTON, HE. B. AND C. K. RICARDO
1936. Scientific results of the Cambridge expedition to the East Afri-
ean lakes, 1930-1, no. 15. The fish of Lake Rudolph and Lake
Baringo. J. Linn. Soe. London, Zool., 39: 353-389.
(Received July 26, 1965.)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. AprIL 29, 1966 NUMBER 244
A NEW SPECIES OF ASHMUNELLA FROM WEST TEXAS
(MOLLUSCA: PULMONATA)
By W. J: CLENCH AND W. B. MER!
The first specimens of this new species were collected by our
colleague, Dr. J. C. Bequaert, 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 which
the anatomical drawings were made by the Junior author.
ASHMUNELLA BEQUAERTI new species
Plate 1; text figures 1, 2
Holotype: MCZ 260274, taken from a rockslide about 14 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; Ashmunella bequaerti was found on both sides of the
canyon, at elevations from 4800 ft. to 4900 ft., along with
Humboldtiana palmert Clench and Rehder, in association with
Quercus terana, YQ. hypoleucoides, Acer grandidentata and Rhus
trilobata.
1 University of Arizona, Dept. of Zoology.
2 BREVIORA No. 244
Paratypes: MCZ 260275 from the same loeality.
Greater Lesser
Height Diameter Diameter
mm mm mm
aay) 1 ISIE Holotype
3.8 13.0 Male Paratype
3.7 12.4 ANG) “e
Bie) 12.2 ele =
3.0 UNEZh 105 i
3.4 11.3 10.1 a
3.2 10.7 9:5 28
3.0 10.5 9.3 i
Description: Shell lenticular, shghtly 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 seales 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 appearance to the surface. There are six whorls, the
first half whorl of the embryonice 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 shghtly 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,
1966 NEW SPECIES OF ASHMUNELLA 3
measured from lower suture of body whorl just behind 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
groups of light grey pigment spots.
Jaw with ten ribs.
Genitalia. Penis with lower sac short and wide, upper sac
longer and narrower. In the upper sae 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 which 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 mm, vagina 4.0 mm, free oviduct 2.5 mm, atrium 1.0 mm.
Remarks: Ashmunella bequaerti 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. hebardi 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 A. bequaerti,
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 hebardi, and the upper insertion of the peristome
is only weakly descending, nearly straight, not strongly descend-
ing as in hebardi. 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 trom 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
4. BREVIORA No. 244
5mm.
Fig. 1. Ashmunella bequaerti Clench 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 together. This may be called
the flagellum portion of the epiphallus. There is no free flagellum.
1, Genital orifice; 2, Atrium; 3, Vagina; 4, Oviduct; 5, Prostate; 6,
Uterus; 7, Albumen gland; 8, Talon; 9, Hermaphroditic duct; 10, Sperma-
thecal duct; 11, Lower sac of penis; 12, Upper sae of penis; 13, Penial
retractor; 14, Epiphallus; 15, Vas deferens; 16, Connective tissue; 17,
Flagellum bound with vas deferens; 18, Penis papilla.
1966 NEW SPECIES OF ASHMUNELLA 5
snail on 2 May 1961, obtaining quantities of dead shells but 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.
14
5mm.
Fie. 2. Ashmunella bequaerti Clench and Miller. Lower portion of repro-
ductive system enlarged. (Numbering as in Figure 1.)
REFERENCES
Piussry, H. A.
1940. Land Mollusca of North America. Academy of Natural Sciences,
Philadelphia, Monograph No. 3, vol. 1, pt. 2, pp. 912-977.
1948. The genera Humboldtiana, Sonorella, Oreohelix and Ashmunella.
Proc. Acad. Nat. Sci., Philadelphia, vol. 100, pp. 199-203.
(Received 24 January, 1965. )
6 BREVIORA No. 244
2
Fies. 1-2. Ashmunella bequaerti Clench and Miller, from Goat Cave
Canyon, Black Mt., Davis Mts., Texas. Fig. 1. Paratype, MCZ 260275 (4
X). Fig. 2. Holotype, MCZ 260274 (5.4 X).
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. APRIL 29, 1966 NUMBER 245
NOTES AND DESCRIPTIONS OF NEW UROCOPTIDAE
FROM CUBA AND HISPANIOLA (MOLLUSCA:
PULMONATA)
By WILLIAM J. CLENCH
The following 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-
les 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, Cylindrella crenata Weinland 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 Département 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 BARBOURI Clench
Archegocoptis barbouri Clench, 1935, Proc. Boston Soc. Nat. Hist. 41:5,
pl. 1, figs. B, F (Tardieu, Mt. LaHotte, Haiti, + 3000 feet). [Holotype,
MCZ 108593. ]
BREVIORA No. 245
ARCHEGOCOPTIS CRENATA (Weinland and Martens)
Cylindrella crenata Weinland and Martens, 1859, Malakozoologische Blit-
ter 6:54 (Jérémie, Haiti); Pfeiffer, 1869, Novitates Conchologicae 3:440,
jl, A, ane Boi, acs (alemhn)).
ARCHEGOCOPTIS DARLINGTONI Clench
Archegocoptis darlingtoni Clench, 1935, Proc. Boston Soc. Nat. Hist. 41:6,
pl. 1, figs. C, G (Tardieu, Mt. LaHotte, Haiti, = 3000 feet). [Holotype,
MCZ 108592. }
ARCHEGOCOPTIS DECAPITATA (R6dinge)
oD
Cerion decapitatum Roding, 1798, Museum Boltenianum, p. 90 (no loeality
given [Haiti] ). [See below. ]
ARCHEGOCOPTIS EXIMIA (Pfeiffer)
Cylindrella eximia Pfeiffer, 1857, Malakozoologische Blatter 4:232 (lo-
cality unknown) ; Pfeiffer, 1869, Novitates Conchologicae 3:439, pl. 97, figs.
30-32 (Haiti).
ARCHEGOCOPTIS TIBURONICA Cleneh
Archegocoptis tiburonica Clench, 1935, Proc. Boston Soc. Nat. Hist. 41:6,
pl. 1, fig. I. (Tardieu, Mt. LaHotte, Haiti, between 3-4000 feet). [Holo-
type, MCZ 108591. |
ARCHEGOCOPTIS HAITIENSIS new species
Plate 1, figures 1, 2
Holotype. MCZ 252056, from the top of Morne Rochelois, 25
km WSW of Miragoane, Dépt. du Sud, Haiti, about 3000 feet,
collected by W. J. Eyerdam, July 1927.
Paratypes. A series of 5 paratypes from the same loeality,
MCZ 252057.
Height Width
mm mm
41 10 Holotype (see Remarks )
27 10 Paratype
26.9 10 os
26.5 10 ne
Description. Shell medium in size, reaching 42+ mm in
length (decollated), minutely umbilicate and coarsely sculptured.
1966 NEW SPECIES OF UROCOPTIDAE 3
Color a more or less uniform chocolate-brown and shining on the
coarse, axial riblets and dull in between. Whorls 15+ and flat
sided. Suture well defined. Spire extended, the upper portion
slightly coneave. 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 Archego-
coptis tiburonica 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. tiburonica.
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. havtien-
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. Only the holotype has retained the mid-whorls.
ARCHEGOCOPTIS DECAPITATA (Roding)
Cerion decapitatum Roding, 1798, Museum Boltenianum, p. 90 (refers to
Chemnitz, 1786, Conchylien-Cabinet (1)9: pl. 136, figs. 1256-1257 [fig. 1255,
in error] ).
Cyclostoma fasciata Lamarck, 1816, Eneyelopédie Méthodique, Atlas, p.
12, pl. 461, fig. 7.
Helix truncata Dillwyn, 1817, Descriptive Catalogue of Recent Shells,
London 2:948 (Santo Domingo).
Helix fasciata Lamarck: Férussac, 1822, Tableaux Systématiques, p. 61,
no. 503 (The Antilles).
Cylindrella fasciata ‘Chemnitz’ Pfeiffer, 1862, Malakozoologische Blatter
9:199 (Corail [Haiti] ).
Cylindrella fasciata ‘Chemnitz’ Crosse, 1891, Jour. de Conch. 39:134 (Co-
rail, near Jérémie, SW Haiti).
Urocoptis truncata (Dillwyn): Pilsbry, 1903, Manual of Conchology
(2)15:154, pl. 39, figs. 27-28 (Corail, near Jérémie, 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 644 miles NE of Corail. It is
possible that the specimen found by Weinland may have drifted
from Grande Cayemite. The specimens collected by Eyerdam
are certainly in the genus Archegocoptis and appear to be A.
decapitata Roding, 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) WHITTUMI new species
Plate 2, figure 1
Holotype. MCZ 59580, from Guabairo, near Soledad, Cien-
fuegos, Las Villas, Cuba, collected by Clench, Crozier and Navez,
October 1928.
Paratypes. MCZ 59284, 86289, 105148, all from the above lo-
eality, and collected by various students and visitors to the
Harvard Tropical Gardens.
Height Width
mm mn
11.5 2.7 Paratype
16 2.9 eS
14.8 3 zt
16 3 =
17.3 3.2 Holotype
Ts 3 Paratype
18.5 3
20.5 3.9 at
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 earher
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, cireular, white and reflected.
1966 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.
Urocopris (GONGYLOSTOMA) 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.
Entire specimens Decollated specimens
Height Height
mm mm
14.4 Holotype eT
14.4 1S)
12.9 eg
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 two 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 subgenus Gongylostoma
Albers, near Urocoptis barbourt Torre and Clench. It differs
from U. barbourt, 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 exrquisita. The
free aperture whorl is also much longer than in U. barbourt.
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. livida barbourt and U. livida atkinsi Torre and Clench.
Urocoptis (GONGYLOSTOMA) CANTEROIANA (Arango)
Plate 2, figure 4
Cylindrella canteroiana ‘Gundlach’ Arango, 1875 [1876], Anales Real
Academia Ciencias Medieas, Fisicas y Naturales, Habana 12:284 (La Vigia,
Trinidad [Cuba]); Arango, 1878, Fauna Malacologica Cubana, Habana, p.
117 (environs of Vigia, Trinidad).
Urocoptis canteroiana ‘Gundlach’ Arango: Pilsbry, 1903, Manual of
Conchology (2)15:254.
Height Width
mm mm
1225 2.0 Lectotype
Lectotype. Here selected, MCZ 189252, from La Vigia, Trini-
dad, Cuba, collected by Gundlach and sent to MCZ by Rafael
Arango.
Remarks. 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, Finea Cantero, and Finea La
Pastora, all in the vicinity of Trinidad, Las Villas, Cuba.
Urocoptis (GONGYLOSTOMA) DIAGONALIS new species
Plate 2, figure 3
Holotype. 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.
a
(
1966 NEW SPECIES OF UROCOPTIDAE
Paratype. A single paratype from the same locality is in the
Museum of Zoology, University of Michigan.
Height Width
mm mm
16 3.3 Holotype
15 3 Paratype
Description. Shell reaching 16 mm in height (decollated),
fusiform, imperforate and sculptured. Color a dull gray. Whorls
16 (remaining), very shehtly 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.
Seulpture consisting of numerous fine, axial riblets which are
shghtly 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-
terovana and flattened sigmoid in shape in diagonaiis.
Urocoptis (UROCOPTOLA) CAYEMITENSIS new species
Plate 1, figure 4
Holotype. MCZ 254666, from NE Grande Cayemite, a small
island about 614 miles NE of Corail, Département du Sud, Haiti,
collected by W. J. Eyerdam, July 1927.
Paratypes. MCZ 254667, population 1; and MCZ 254668,
population 2. From the same locality as the holotype.
Height Width
mm mm
aha 6.5 Holotype, population no. 1+
18.5 i Paratype om
17.3 5.8 a 7
15.8 6 ea es
20.5 (ical os population no, 21
20.4 7 a a
20.1 7 i os
Description. Shell oblong, glossy, having the greatest diameter
above the middle, imperforate, sculptured and rather thin in
structure. Color a pale brownish pink. Whorls very shghtly
1 Both populations from the same general area.
BREVIORA No. 245
(© e}
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 cayemitensis differs from U. 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.
Paratypes. MCZ 260872, from the locality of the holotype.
Height Width
mm mm
25 113) Holotype
26 11 Paratype
22.5 10.2 ie
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 rapidly from the
mid-area and produced at an angle of about 55°. Aperture sub-
circular, holostomatous, 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 bencomoi Clench,
but is smaller, proportionally narrower, and has the hp 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 OF 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.
Paratypes. MCZ 260874, from the same locality.
Height Width
mm mm
24.8 12 Holotype
22 11.6 Paratype
22.5 lal: “s
22.5 11.4 se
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 subcireular 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. ekmamni, 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 (IpIoSTEMMA) PERPLICATA (Beck)
Helix (Cochlodina) perplicata Férussac, 1821 [1822], Tableaux Sys-
tématiques Animaux Mollusques, p. 61 or 65, no. 506 (The Antilles) [nomen
nudum).
Brachypodella perplicata Férussac: Beck, 1837, Index Molluscorum, p.
89 [refers to Férussac’s pl. 163, fig. 9].
Clausilia perplicata Férussac: Deshayes [in] Lamarck, 1838, Animaux
sans Vertébres (2)8:216.
Cylindrella perplicata Férussac: Philippi, 1847, Abbildungen Conchylien
2:217, pl. I (Achatina), fig. 9 (The Antilles).
Cylindrella perplicata Férussac, 1851, Histoire Naturelle Générale et
Particuliére 2:229, pl. 163, fig. 9 (The Antilles); non Cylindrella perplicata
Férussac. Pfeiffer 1840 [in] Weigmann, Archiv fiir Naturgeschicte 1:41
(Fundador, Matanzas, Cuba).
10 BREVIORA No. 245
Cylindrella fastigiata ‘Gundlach’ Pfeiffer, 1860, Malakozoologische Blat-
ter 7:20 (Baracoa, Mata, and Yunque, Cuba); Pfeiffer, 1865, Novitates
Conchologicae 2:263, pl. 65, figs. 23-25 [Syntypes, MCZ 26639, 86621].
Urocoptis fastigiata Pfeiffer: Pilsbry, 1903, Manual of Conchology
(2)15:171, pl. 45, figs. 36-41 (Baracoa, El Yunque, Mata, Oriente, Cuba).
Brachypodella perplicata Férussac: Pilsbry, 1903, Manual of Conchology
(2) 16:83, pl. 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 Férussac, as Férussae 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: OrtenTE, 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
Brachypodella (Brevipedella) imitatrix Pilsbry, 1903, Manual of Conchol-
ogy (2)16:47, pl. 8, figs. 54-55 (Port-au-Prince, Sans-Souci, St. Mare, and
La Ferriére, 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. angulifera (Gundlach), occurs in
eastern Cuba.
Specimens examined. Hartt: Bizonton, Lasbaines; Port-au-
Prince and Diquini. SAnto Dominco: Mt. Isabel de Torres,
Puerto Plata, at 1200 to 1600 feet; Sanchez; Penon de Basiles
and Penon de Maria Luisa, both Santa Barbara de Samana, and
Punta Lirio, 2 miles E of Santa Barbara de Samana.
1966 NEW SPECIES OF UROCOPTIDAE 1]
BRACHYPODELLA (GYRAXIS) SAMANA new species
Plate 2, figure 5
Holotype. MCZ 57214, from Penon de Maria Luisa, Santa
Barbara de Samana, Républica 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 Penon de Basiles in the same general area.
Height Width
mim mnt
14.5 2.0 Holotype
12.5 2.4 Paratype
14.5 2.9 i
12.8 2.4 =
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. Whorls 13 to 14 in entire specimens
and moderately convex with the last whorl free. Suture well
defined. Spire acute, aperture holostomatous, subcircular with
the lip flaring 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
shghtly on the suture.
Remarks. This species is closely related to Brachypodella
(Gyravris) sericata Pilsbry. It differs by being larger, having
slightly coarser thread-like costae and having the marbled colora-
tion, sericata being a dull white.
Brachypodella sericata Pilsbry was collected originally by
William M. Gabb 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
Fias. 1-2. Archegocoptis haitiensis n. sp. Morne Rochelois, 25 km WSW
of Miragoane, Dépt. du Sud, Haiti, at 1000 meters. Fig. 1, Paratype, MCZ
252057; fig. 2, Holotype, MCZ 252056 (both 3X).
Fie. 3. Urocoptis (Urocoptola) caribbaea n. sp. 1 mile E of Pte. a Ra-
quette, S central Gonave Island, Haiti. Holotype, MCZ 260873 (3X).
Fie. 4. Urocoptis (Urocoptola) cayemitensis n. sp. NE Grande Cayemite,
Dépt. du Sud, Haiti. Holotype, MCZ 254666 (4X).
Fig. 5. Urocoptis (Urocoptola) ekmani n. sp. La Souree, NW Gonave
Island, Haiti. Holotype, MCZ 260871 (3X).
13
NEW SPECIES OF UROCOPTIDAE
1966
PLATE 1
14 BREVIORA No. 245
Plate 2
Pie. 1. Urocoptis (Gongylostoma) whittumi n. sp. Guabairo, Soledad, Las
Villas, Cuba. Holotype, MCZ 59580 (5.5 X).
Fig. 2. Urocoptis (Gongylostoma) exquisita, n. sp. % mile E of Guabairo,
Soledad, Las Villas, Cuba. Holotype, MCZ 59286 (5.5 X).
Mina Carlota, Sierra
Fig. 3. Urocoptis (Gongylostoma) diagonalis n. sp.
Holotype, MCZ
de San Juan, 8 miles S of Cumanayagua, Las Villas, Cuba.
59292 (5.5 X).
Fia. 4. Urocoptis (Gongylostoma) canteroiana (Arango). (La Vigia,
Trinidad [Las Villas, Cuba]). Lectotype, MCZ 189252 (5.5 X).
Fic. 5. Brachypodella (Gyraxis) samana n. sp. Pefon de Maria Luisa,
Santa Barbara de Samana, Reptblica Dominicana. Holotype, MCZ 57214
(SHY DONe
=a |
roy
e
)
—-
uw =
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. May 3, 1966 NuMBER 246
PSEUDANTHESSIUS PROCURRENS 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 Bé, in northwestern Mada-
gascar, 109 adults and 9 copepodids of the new hchomolgid 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 the 22 species listed by Stock, Humes, and Gooding,
1963, and a new species from a polychaete annelid in Madagascar
whose 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’Océanographie et des Péches at Nosy Bé,
2 BREVIORA No. 246
DESCRIPTION
Family LICHOMOLGIDAE Kossmann, 1877
Genus PSEUDANTHESSIUS Claus, 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-
marek), in 1 meter depth among dead coral (Acropora) at Pte.
Ambarionaomby, Nosy Komba, near Nosy Bé, 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 p.
in length and 133 » 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 » 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 saes
is 107 uw. Behind each attachment area the segment is slightly
1 The specific name procurrens, from Latin procurrere, 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 ».). Each egg sae attachment area (Fig. 4) bears
anteriorly a slender, slightly haired seta (21 » long) and just
posterior to it a short naked seta (6.5 ». 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 v. 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 wu, along the outer side 104 pw, and the width at the
level of the outer seta is 24 vy. The ratio of length to width is
about 4.5:1. The outer seta, inserted 70 ». from the base of the
ramus, is naked and 56 v. in length. The pedicellate dorsal seta is
33 wand slightly haired. The outermost terminal seta (100 yu)
and the innermost terminal seta (72 y.) are haired. The 2 long
median terminal setae are 177 and 250 y 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 ege sacs (Fig. 1) are moderately elongated, often rather
pointed posteriorly, and contain numerous eggs. In one female
the egg sacs measured 385 x 220 wu, with each egg about 57-60 v.
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 » along the anterior margin), 103, 21, 39, 33, 20,
and 17 uw respectively. The formula for the armature is 4, 13, 6,
3, 4 + 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 » along the shorter ventral margin, 93 pv. along
4 BREVIORA No. 246
the dorsal margin, and 19 ». 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 y. 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 » long) and a shorter pointed spine (5 y.) and sub-
terminally a naked hyaline seta (11 »). 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 » 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 selerotization which almost joins the
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) :
Esl a protopod) | .0=1-- 1-0 exp | 0. It; Sih
endie Ot 0-1. 6 Tes
Fo, Sprotopods O-l- A-0) exp > 10.) ik. iio
end 0-1; 0-2; JTIL3
Po protopod. 0-1; 1-0. exp) 4-0- eb Thr
end, 03-9) (0-2, Ieit-2
EA protopod: J0-1-, 91-0) .exp £0. > iets Tis
end II
The inner seta on the coxa of legs 1-3 is long and plumose, but
in leg 4 this seta is minute (7 » 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 exopods 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 wu 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 wu. 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 yu.
long with a minute flagellum, the inner 37 » 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 p), together with a dorsal seta (22 y), 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. AI-
though, as in other species of Pseuwdanthessius, 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 sae (see Fig. 4).
6 BREVIORA No. 246
The color in life in transmitted hght is translucid, the eye red,
the intestine black, the ovary gray, the ege 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
eon aene
The segment of leg 5 is similar to that of the female, and meas-
ures 31 x 94». The genital segment (Fig. 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 pv.
from anterior to posterior.
The caudal ramus resembles that of the female, but is a little
shorter, the inner length being 78 pv, the outer length 73 yu, the
width 23 1, 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, paragnath, 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 wu. 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 y. in length along its
axis, bears a conspicuous terminal lamella. The slightly 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 1
42 y. long with minute lateral spinules, and on its anterior sur-
face there are 2 small naked setules, one 10 yp, the other 4 y 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 v. in length.
The spermatophore, seen only inside the body of the male
(Fig. 24), is oval, about 72 x 45 yp, 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-
euished 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. dubius
G. O. 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.
Hight 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. ob-
scurus A. Scott, 1909, P. sawvagei Canu, 1892, P. spinifer Lind-
berg 1945, P. tenuis Nicholls, 1944, P. thorelli (Brady, 1880),
and P. weberi 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. O. Sars, 1917. The species referred to as P. luber (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
part of the genital segment of the female, has a caudal ramus
about twice as lone 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, I,11,3, instead of I,I1,2 as in the Madagascan species.
P. assimilis G. O. 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. O.
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 Pseudanthessius procurrens is associated with
the cidarid echinoid Phyllacanthus imperialis (iamarck) in the
2 Not Brady and Robertson, 1876. See Stock, Humes, and Gooding, 19638, p. 10,
footnote.
1966 NEW COPEPOD FROM AN ECHINOID 9
region of Nosy Bé, northwestern Madagascar. Within the genus
the copepod 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. liber (Brady,
1880) and P. assimilis G. O. Sars, 1917. Although the host
echinoderm is widely distributed in the Indo-Pacific area, the
new copepod associated with it is known at present only from the
type locality in Madagascar.
REFERENCES CITED
Brapy, G.S.
1880. A monograph of the free and semi-parasitic Copepoda of the
British Islands. Ray Society, London, 3:1-83.
BrabDy, G. S. AND D. ROBERTSON
1876. Report on dredging off the coast of Durham and North York-
shire in 1874. Rept. British Ass. Adv. Sci. (Bristol), 45: 185-
199,
CaNnu, E.
1892. Les Copépodes du Boulonnais, morphologie, embryologie, taxono-
mie. Trav. Lab. Zool. mar. Wimereux-Ambleteuse (Pas-de-
Calais), 6:1-354.
GLAS; C-
1889. Uber neue oder wenig bekannte halbparasitische Copepoden,
insbesondere der Lichomolgiden- und Ascomyzontiden-Gruppe.
Arb. Zool. Inst. Univ. Wien, 8(3) :327-370.
GALLIEN, L.
1935. Pseudanthessius nemertophilus nov. sp., copépode commensal de
Lineus longissimus Sowerby. Bull. Soc. Zool. France, 60:451-459.
GURNEY, 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 espéces de Pseudanthessius (Copepoda, Cyclo-
poida) parasites des oursins @ Madagascar. Mém. Inst. Sci.
Madagasear, sér. F, 1959, 3:67-82.
Humes, A. G. anp J.-S. Ho
In press. New cyclopoid copepods associated with polychaete annelids in
Madagascar. Bull. Mus. Comp. Zool.
Inne, P.
1950. A new copepod, Pseudanthessius latus (Cyclopoida: Lichomolgi-
dae) commensal with a marine flatworm. Jour. Washington
Acad. Sci., 40(4) :129-133.
10 BREVIORA No. 246
LINDBERG, K.
1945. Un nouveau copépode poecilostome de 1’Inde de la famille des
Lichomolgidae; Pseudanthessius spinifer, n. sp. Bull. Soe. Zool.
France, 70:81-84.
NICHOLLS, A. G.
1944. Littoral copepods from South Australia (IL). Calanoida, Cyclo-
poida, Notodelphyoida, Monstrilloida and Caligoida. Ree. So.
Austr. Mus., 8(1) :1-62.
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, Coryecaeidae,
Ergasilidae, Clausiidae, Eunicicolidae, Supplement, pp. 173-225.
Bergen Museum, Bergen.
Scort, A.
1909. The Copepoda of the Siboga Expedition. Part I. Free-swim-
ming, littoral and semi-parasitic Copepoda. Siboga Exped.,
29a)31-323.
SEWELL, R. B. S.
1949. The littoral and semi-parasitic Cyclopoida, the Monstrilloida and
Notodelphyoida. John Murray Exped. 1933-34, Sci. Repts.,
9(2) :17-199.
Stook, 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 Curacao and other Caribbean
Is., 18(77) :1-74.
THOMPSON, I. C. AND A. ScorTr
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 each figure refers to the scale at which the figure
was drawn.
Abbreviations: a; = first antenna, md = mandible, p = paragnath, mx; =
first maxilla, mxg = second maxilla, mxpd = maxilliped, pi = leg 1.
1966 NEW COPEPOD FROM AN ECHINOID 11
scale D. 0.05mm
scale A, 0.5mm
scale E.0.2mm
scale B. 0.2 mm
scale C, 0.05 mm
=
FIGuRES 1-7. Pseudanthessius procurrens u.sp., female. 1, dorsal (A); 2,
lateral (A); 3, urosome, dorsal (B); 4, area of attachment of egg sac, dor-
sal (C); 5, caudal ramus, ventral (D); 6, edge of cephalosome, ventral
(E); 7, rostral area, ventral (F).
12 BREVIORA No. 246
G. 0.03 mm
scale
scale H.0.1mm
+
Figures 8-16. Pseudanthessius procurrens n.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 i133
Ce i) : }
AS ATE\ AN. ///
\ y
Figures 17-21. Pseudanthessius 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 procurrens 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
Museum of Comparative Zoology
CAMBRIDGE, Mass. May 3, 1966 NUMBER 247
THE CHANARES (ARGENTINA) TRIASSIC 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
(ef. Rusconi, 1951, ete., 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. José 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 BREVIORA 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
Nacional 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 earher
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 Mesosaurus of southern Brasil (and
South Africa) and a single labyrinthodont from the Brasilan
Permian (Price, 1948). The earlier Triassic was, before our re-
cent exploration, almost equally unknown as regards reptilian
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 Nacional de Geologia y Mineria,
the Yacimientos Petroliferos Fiscales and the Comisién 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 sery-
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 Félix 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. Héctor 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. Stipanicie, director of geological work for the Comisién
4 BREVIORA No. 247
Nacional de Energia Atomica; and Sr. José F. 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 NSF 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 he in
La Rioja Province; Chilecito was the best available town to visit
periodically for supphes, 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 1% 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 motoreyclette of a type recently
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1966 CHANARES REPTILES
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
foreboding 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, 1951a, 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 by 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 Montana 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 Montana to Puesto
Alojamiento. But as far as our sampling of these vast exposures
went, 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 Peas, between
6 BREVIORA No. 247
Cerro Pena 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, however, 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 ‘‘ Chirothertum’’-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, ete.) had described at length as ‘‘ Paganzo
Il’’ 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 Sanogasta, 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
IL’’; 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 by discussion with Sr.
Bonaparte.
Permo-Triassic beds had been studied in the Sierra de las Qui-
jadas in northwestern San Luis Province by Sr. José Roman
Guinazu of San Luis, and footprints from this region were de-
seribed 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 (Bracaccini, 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 Provinee,
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, Chanhares, and Gualo, geological reconnaissance by Bo-
denbender (1911, ete.) 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 BREVIORA No. 247
Chanares, we camped for the remainder of our stay on the banks
of that stream.
In the following article in this series there will be given a
resumé of the geology of the area. The region is only 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 Chanares-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 would 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 earher expedition) could be readily ae-
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 CHANARES REPTILES 9
a relatively small fraction of ‘‘problematica,’’ 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 Kannemeyeria type. A considerable frac-
tion of the finds were those of small thecodonts, some, at least,
appearing to be forms on the pattern of Euparkeria of the Cynog-
nathus zone of South Africa (recently redescribed by Ewer,
1965). 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.
oe
THE TRIASSIC FAUNAL SEQUENCE
The position of the Chanares 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
part, 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
‘‘Kotriassic’’ Scythian zone. As both Watson (1942) and Lehman
and others (1959) have shown, the Cynognathus and Lystrosau-
rus zones of South Africa are definitely to be regarded as
‘‘Kotriassic’’ and Seythian in age! mainly on the basis of laby-
rinthodonts; the Moenkopi of the early North American Triassic
and the Bunter of Europe are equally well assignable to the
1 Huene (1956) has at times tabulated the Cynognathus zone as middle Triassic.
but has cited no reason for this.
10 BREVIORA No. 247
Seythian on the basis of their content of labyrinthodonts — laby-
rinthodonts of types not too far removed from those of the
‘‘Kotriassic’’ of Greenland and Madagascar. At present it is
perhaps 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 (Lystrosaurus and Cynognathus
zones) of South Africa. Here, therapsids, most notably a variety
of ecynodonts, 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, ete. 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 Muschelkalk 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 Kast 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 af
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-
saurus and Cynognathus zones of South Africa, and the Wuh-
siang beds of Shansi (China) are typical. There are abundant
therapsids, principally cynodonts— 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
ereatly 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 between (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 perhaps 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 rhynchosaurs persist. We
may have here a (B)-(C) intermediate.
Baird and Take (1959) have recently discovered a reptilian
fauna in the Wolfville sandstone of Nova Scotia, generally re-
gvarded 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
Actosaurus of the German Keuper, Walker (1961) believes the
Elgin fauna to be late Triassic in age. There is, however, no rea-
son to belheve 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 Ornithosuchus (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 English equivalent of the Muschelkalk of the continent ; the
early Triassic ‘‘Bunter’’ equivalents are immediately followed
by redbeds 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: 1565-1566.
BODENBENDER, G.
1911. Constitucion geolégica de la parte meridional de La Rioja y
regiones limitrofes (Republica Argentina). Bol. Acad. Nae.
Ciene. Cérdoba, 19: 1-220.
1966 CHANARES REPTILES 13
Borpas, A. F.
1944. Peces tridsicos de la Quebrada de Santa Clara (Mendoza y 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 Rioja). Rev. Soe. Geol. Argent.,
1: 19-61.
BRINK, A. S.
1963. Two eynodonts from the Ntawere Formation in the Luangwa
Valley of Northern Rhodesia. Palaeont. Afr., 8: 77-96.
CASAMIQUELA, R. M.
1960. Noticia preliminar sobre dos nuevos estagonolepoideos argen-
tinos. Ameghiniana, 2: 3-9.
Ewer, R. F.
1965. The anatomy of the thecodont reptile Huparkeria capensis
Broom. Phil. Trans. Roy. Soc. London, B 248: 379-435.
HARRINGTON, H. J.
1941. Investigaciones geolégicas en las Sierras de Villavicencio y Mal
Pais, Provincia de Mendoza. Bol. Direccién Minas y Geologia,
Rep. Argent., 49: 1-31.
1956. Argentina. In: Handbook of South American Geology. Mem.
Geol. Soc. Amer., 65: 131-186.
HUENE, F.
1956. Paldiontologie und Phylogenie der niederen Tetrapoden. Jena,
716 pp.
JAIN, S. L., P. L. ROBINSON, AND T. K. R. CHOWDHURY
1964. A new vertebrate fauna from the Triassic of the Deccan, India.
Quart. Jour. Geol. Soe. London, 120: 115-124.
KITCHING, J. W.
1963. The fossil localities and mammal-like reptiles of the Upper
Luangwa Valley, Northern Rhodesia. South Afr. Jour. Sei.,
59: 259-264.
LEHMAN, J.-P., C. CHATHAU, M. LAURAIN, AND M. NaucHE
1959. Paléontologie de Madagascar. XXVIII. Les poissons de la
Sakamena moyenne. Ann. Paléont., 45: 177-219.
Tot, RS:
1942. Triassic footprints from Argentina. Amer. Jour. Sci., 240: 421-
425.
PRABODY, F. E.
1955. Occurrence of Chirotherium in South America. Bull. Geol. Soc.
Amer., 66: 239-240.
PRICE; Ty: I.
1948. Um anfibio labirintodonte da formaecao Pedra de Fogo, Estado
do Maranhao. Ministerio da Agricultura. Bol. Serv. Geol. Min.
Brasil, 124: 1-32.
14 BREVIORA No. 247
ReiG, O. A.
1961. Acerea de la posicién sistematica de la familia Rauisuchidae y
del género Saurosuchus (Reptilia, Thecodontia). Publ. Mus.
Cienc. Nat. Mar del Plata, 1: 73-114.
Romer, 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.
RuScOoNnI, C.
195la. Laberintodontes tridsicos y pérmicos de Mendoza. Rev. Mus.
Hist. Nat. Mendoza, 5: 33-158.
1951b. Rastros de patas de reptiles pérmicos de Mendoza. Rev. Soe.
Hist. Geog. Cuyo, 3: 1-14.
1956. Mares y organismos extinguidos de Mendoza. Rev. Mus. Hist.
Nat. Mendoza, 9: 2-88.
WALKER, A. D.
1961. Triassic reptiles from the Elgin area; Stagonolepis, Dasygna-
thus and their allies. Phil. Trans. Roy. Soc. London, B 244: 103-
204,
1964. Triassic reptiles from the Elgin area: Ornithosuchus and the
origin of carnosaurs. Phil. Trans. Roy. Soc. London, B 248: 53-
134.
Watson, D. M.S.
1942. On Permian and Triassic tetrapods. Geol. Mag., 79: 81-116.
(Received January 6, 1966.)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. 29 Juty, 1966 NUMBER 248
A TRIASSIC AMMONITE FROM THE HINDUBAGH
REGION, BALUCHISTAN, WEST PAKISTAN
By BERNHARD KUMMEL
Ammonites are extremely rare in the thick development of
Triassic strata between Quetta and Hindubagh 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 specimen of rather good preservation is warranted.
The first record of Triassic ammonites from this region of
Baluchistan was by Mojsisovies (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 Didymites afghanicus Mojsisovies (1896; 1899, pl.
20, fig. 9) and assigned a Norian age. Vredenburg (1904), on
the basis of surveys carried out in the summer of 1901, con-
tributed some regional, 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 ammonite-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. atf. swbcatenato Mojsisovies
Celtites sp. ind. (group of ? acuteplicati)
Paratibetites sp. ind. ex aff. tornquisti Mojsisovies
Dittmarites or Distichites ? sp. ind.
Rhacophyllites vredenburgi 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. subcatenato 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 ammonites.
The fauna as a whole shows very strene affinities 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. Sinee 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 they were Norian in age.
Oo
1966 TRIASSIC AMMONITE
SYSTEMATIC DESCRIPTION
Family TROPICELTITIDAE Spath, 1951
Genus ARIETOCELTITES Diener, 1916
ARIETOCELTITES ARIETITOIDES (Diener)
Figures la, b
Tropiceltites arietitoides Diener, 1906, p. 156, pl. 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 mm for the diameter of the umbilicus. In
degree of involution and general proportion, this specimen is
very much like the type specimen. There are shght 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. With 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 group, between Quetta and Hindubagh, West Pak-
istan.
Repository. — Geological Survey of Pakistan, Quetta, West
Pakistan.
Acknowledgements. —I wish to thank Mr. 8. A. Bilgrami for
the loan of the specimen and for his hospitality during a visit to
Hindubagh. My visit to Pakistan was made possible by the aid
of National Science Foundation Grant No. G19066.
4 BREVIORA No. 248
REFERENCES
ARKELL, W. J. ef al.
1957. ‘Treatise on invertebrate paleontology, Part L, Molusea 4, Ceph-
alopoda, Ammonoidea. Geol. Soc. Am. and Uniy. Kansas Press,
490 pp.
COLOMBO PLAN PROJECT
1961. Reconnaissance geology of part of West Pakistan. Toronto,
590 pp.
DIENER, CARL
1906. Himalayan fossils. Fauna of the Tropites Limestone of Byans.
India Geol. Surv., Palaeont. Indica, (15) 1: 1-201, pls. 1-17.
1906. Notes on an Upper Triassic fauna from the Pishin District,
Baluchistan, collected by E. Vredenburg in the year 1901. Ree.
Geol. Sury. India, 34: 12-21, pls. 3, 4.
1912. The Trias of the Himalayas. Mem. Geol. Surv. India, 36: 1-159.
1916. Kinige Bemerkungen zur Nomenklatur der Trias-cephalopoden.
Centralbl. Mineral. Geol. Palaiont., No. 5: 97-105,
Herron, A. M.
1939. General report of the geological survey of India for the year
1938. Ree. Geol. Surv. India, 74: 1-132.
Mossisovics, E. v.
1896. Beitrige zur Kenntniss der obertriadischen Cephalopoden-
Faunen des Himalayas. Denkschr. Akad. Wiss. Wien, 63: 575-
701, pls. 1-22.
1899. Upper Triassic Cephalopoda faunae of the Himalaya. India
Geol. Surv., Palaeont. Indica, (15) 3: 1-157, pls. 1-22.
SparH, L. F.
1951. Catalogue of the fossil Cephalopoda in the British Museum
(Natural History). Part 5, The Ammonoidea of the Trias.
London, 228 pp.
VREDENBURG, E.
1904. On the oceurrence of a species of Halorites in the Trias of
Baluchistan. Ree. Geol. Sury. India, 31: 162-166, pl. 17.
WILLIAMS, M. DEAN
1959. Stratigraphy of the Lower Indus Basin, West Pakistan. Fifth
World Petroleum Congress, Sec. 1, Paper 19: 1-15.
1966 TRIASSIC AMMONITE 5
Figure 1. Side and ventral view of Arieloceltites arietitoides (Diener)
from Triassic formation near Hindubagh, West Pakistan. * 1.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. 29 JuLy, 1966 NuMBER 249
ADDITIONAL NOTES ON THE AMPHISBAENIDS OF
GREATER PUERTO RICO
By RicHarp 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, Amphisbaena schmidti, from Puerto Rico.
Thorough as it has been, the work 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-Mareh 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 oceupied by an extremely eroded to-
pography of close set, conical mogotes (to above 1500 feet), often
called the Haystack (or Pepino) Hills; but some areas of it are
110,000 SW 84th St., Miami, Florida 33143.
No. 249
BREVIORA
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‘yp jo oSuvt oy} pue ‘sour, [vuoseIp ouy Aq Diaw “Pp JO IsuvI oY} + SOUT] [vuoseIp os1v09 AQ Po}BVOIPUL ST Sp1oded
yuasaid pue snotaoid uo poseq tuaynq ‘py Jo OSuBI UMOUY OY, *(zonsvAvy 1oJ oatpdumsoad ATUO) Spsodar otapedurss
quasordad spoqurdS J[VY OM} YYIM SopoITQ ‘*(SopodITo MOT[OY) wWprWwYyos *P puB ‘(SasSO1o YIM Sopa) Daw *p ‘(SJOp
YIM Sept) rwayng *P ‘(SepPalo pYOS) vIap9 "Pw AOF SpLOdat APTROT MoU SuLMOYS OoTY OJON FO dey ‘LT ‘sty
KAS
~
7
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 12); ‘‘Maricao,’’ MCZ 66513
(= UPR 22); °‘Mayagiiez,’’ UMMZ 55824.
Amphisbaena bakeri 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
4 BREVIORA No. 249
corner of the island, and the only localities from which it was
known ‘‘with’’ bakeri were those which were possibly peripheral
to the range of bakeri (ef. Gans and Alexander, 1962, fig. 4).
A. bakeri 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 bakeri has therefore not been established with cer-
tainty.
One of the recent specimens (ASF'S V5335) represents only a
minor range extension but constitutes the first record from the
northern limestone region; ASFS V6424 is from within the pre-
viously known range. There is no evident explanation for the
small, very restricted, apparent range of bakert. The fact that
a number of specimens of caeca (and no bakerv) were collected
in the vicinity of Utuado indicates that bakert probably does not
occur much farther east than it has been previously reported.
The coloration in life of these two specimens of A. bakeri was
noted as being ‘‘pale pinkish gray’’; ASFS V6424 was 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 (Fig. 2).
\
A B
Fig 2. Hemipenes of A, Amphisbaena schmidti (ASFS V5871), and B,
Amphisbaena caeca (ASFS X4111); view cephalad 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. ASES
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 bakeri 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. schmidti 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
bakeri. Thus, in this area, the three known Puerto Rican species
occur sympatrically, whatever their ecological preferences (see
below for caeca and schmidt).
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’ (835 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,
V0332; 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; 112 km NW Utuado, 1300’ (400 m), ASFS V5922;
1 mi. (1.6 km) NW Dorado, ASFS V5939; 4.1 km NE Villa
Pérez, 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 X7483-34; Mayagiiez,
UPR 84e, 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 Mayagtiez, 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-Adjuntas 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 Afuera. However, the species may be
assumed to occur on Vieques proper. It has long been a matter
of speculation as to which species of Amphisbaena would be
found on Vieques and Culebra, caeca or fenestrata. 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: MCZ 81019, an adult male from 7 km E of Guanica,
Puerto Rico, elevation 600’ (183 m), one of a series collected on
6 Mareh 1965 by Albert Schwartz and Richard Thomas.
Paratypes: PUERTO RICO: UIMNH 56910, ASFS V5646,
USNM 152588, 7.3 km E Guanieca, 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 V5800, 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 V6148, 5 mi. (8 km) ESE San German, south
slope Cerro Algarrobo, 10 July 1965, R. Thomas; ASFS V6423,
1.5 km WNW 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 NW
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 Juana Diaz, 29 July 1965, R.
Thomas ; UMMZ 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, Mayagtiez, 10
8 BREVIORA No. 249
December 1957, J. A. Ramos; UPR 86, Mayagiiez, 8 September
1949, Nestor Nazario; UPR 91, no data.
Diagnosis: A form of Amphisbaena lacking fusion of head
seales, of closest affinities to the species bakeri and caeca, differ-
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.
wera (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 bakert. 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 very similar to those of bakeri and caeca
(hie, 2).
Coloration: The dorsal coloration in life of this form is typi-
cally a pale tan, very reminiscent of the two living specimens of
bakerit 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
11 have objected elsewhere (Thomas, 1965) to the use of the term postmalar ;
however, this was due 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 translucence 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 ight (buffy)
as is the posterior half of the edge of the lower jaw.
Comparisons and discussion: The most pertinent comparisons
of wera are with caeca and bakeri (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 zera,
and a possible exception noted above, the absence of a postmalar
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INTERPREFRONTAL
Fig. 4. Seatter diagram of internasal versus interprefrontal suture
lengths for A. caeca (stippled squares), A. bakeri (solid triangles), and
A, xera (hollow circles). Numbers in the wera 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 vera and caeca represent the values
for the largest specimen of each of these two forms.
1966 AMPHISBAENIDS OF PUERTO RICO 11
row distinguishes zera 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-
gtiez) 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 1s 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 zera 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. rera is strikingly like bakeri 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 bakeri 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
2 BREVIORA No. 249
conditions in bakert and caeca but tending more towards the
former. The 18 xera with but two postgenial rows are even more
suggestive of a trend towards bakerv.
Schmidt (1928) commented on three specimens from Maya-
giiez that appeared to be intermediate between bakeri 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. bakeri. Two specimens of bakert are known from Mayagiiez ;
one (CNHM 12473) has a body annulus 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 Mayaguez 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 xera. More specimens with precise
data from the Mayaguez vicinity may show that bakerv and
xera are Subspecifically related. I am reluctant to so allocate
these forms now because bakeri is so little known.
A. schmidti is distinguished from zxera 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 eaudal 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.
Habitat: 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.
1Heatwole, Torres and Heatwole (Stahlia, 1965, no. 4, p. 1) recorded taking a
specimen of Amphisbaena on Caja de Muertos. I have not examined this speci-
men, but almost certainly it pertains to A. wera, as it was noted to lack the post-
malar row of chin segments and also agreed with vera in other counts given.
te
1966 AMPHISBAENIDS 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 grantt.
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 Amphisbaena 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 loeality 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 Inabén; it too is transitional between the
xeric south and the mesie interior. On the first visit, nine speci-
mens of caeca and one of sera 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 serubby
woods; the spiny palm (Acrocomia), 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.
2.3 km E of Juana Diaz. This locality is a mesic, wooded 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’ (866 m), ASFS V5268; 4 km §
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, 13800’ (400 m), ASFS
75923-24; 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.
Amphisbaena 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. Vélez, 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 Cano.’’ The lo-
cality is in the district of Isabela about six kilometers southeast
of the town of Isabela by the side of Carretera Cano (= 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 present in two
specimens. The organs are relatively slender and deeply bilobed ;
the suleus 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 schmidti
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 snecies
was collected at particular localities within their joint range;
the number of specimens at each locality is indicated in paren-
theses.
schmidti caeca
acacia serub (6) coastal almond (Terminalia)
woods (2)
moderate to well-exposed, scrubby
ravine sides (6)
moderately, exposed hillside (Gl 55. as.
2 od 5 Ee ee ee ee ee coffee grove (3)
2 a an ee ee coffee grove (1)
exposed hillside pasture (2) mesic woods adjacent to
pasture (1)
high, open ravine woods (1) exposed ravine side (1), shady
ravine woods (1)
moderately exposed edge of interior of coffee grove (1)
coffee grove (1)
16 BREVIORA No. 249
schmidti caeca
CANE GANISEIR mie 9 | Nase omadmbo ceed bac
doline pasture (1)
moist, open pasture (2) Ay A OY acta Ee
cleareduslopeyot dolines@l) 9) 9) Paces ae eee
moderately exposed roadside shady coastal woods (1)
ditch (1)
Qoan lnkeiieycarm Gy) eh dba E SS Doo eo
AMPHISBAENA FENESTRATA Cope
Specimens examined: new localities: VIRGIN ISLANDS.
St. John: Lameshur Bay, VINP 1478; Annaberg Road near
Frederickdal, VINP 1479; Caneel 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, ea. 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 postcloacals
Staelhomasiy sane (242-247) .... (2-14)... (14-16) 2. . (8-20)
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 ...... ZAMS) 9. bee MGA ee ee)
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 n7.
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. No other specimens of West
Indian species seen by me (caeca, schmidti, bakert, cubana, imno-
cens, gonavensis, manni) have so prominent a dorsal groove,
although one specimen of baker: has a moderately well indicated
dorsal groove.
Amphisbaena fenestrata 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 off 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 Rieo. Dr. Carl Gans and Mr.
A. Allan Alexander have helpfully read and eriticized the manu-
seript. 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.
Mueh 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. Horacio Mayorga,
University of Puerto Rico at Mayagiiez (UPR). The following
institutions have received paratypes of Amphisbaena xrera: 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.
LITERATURE CITED
GANS, CARL
1964. Amphisbaena 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.
Fow Ler, 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 pls.
THOMAS, RICHARD
1965. Two new subspecies of Amphisbaena (Amphisbaenia, Reptilia)
from the Barahona Peninsula of Hispaniola. Breviora, Mus.
Comp. Zool., No. 215, pp. 1-14.
(Received 2 November, 1965)
iT |
1966 AMPHISBAENIDS OF PUERTO RICO 19
KEY TO GREATER PUERTO RICAN SPECIES OF AMPHISBAENA
le
bo
Nasals separated by posterior extension of rostral which contacts pre-
frontals; body annuli 236-249 ............... ee es ChLestnaua
Nasals in contact with one another ........... Be er. Wee
Number of body annuli low (192-207); high number of caudal annuli
(CL S522) eee ree Menta ks oy ) Wi: arma Ee hye OO Ue. ee ae schmidti
Number of body annuli higher (more than 214); caudal annuli lower
GLROTSICSS)) Lp eee a a ae OO ee ee ee ee ae 3
Internasal suture short (less than one-third interprefrontal suture) ;
bodyaranm ule AS eee aes ac <a ea OO a ee es ae ee caeca
Internasal suture long (one-third or more the length of interprefrontal
SUGUT.O)) Me ee aD ae ners art eee oe nD | or en 4
Number of body annuli higher (239-255); size large ......... bakeri
Number of body annuli lower (225-237), size small ......... _. wera
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.
SPECIMEN NUMBER
A. BAKERI
ASFS V5335
ASFS V6424
MCZ 66512
MCZ 66513
UMMZ 55824
A. CAECA
ASFS V5430
ASFS V5431
UPR 84c
UPR 87
UPR 88
UPR 90
UPR 92
UPR 140
ASFS V5152
ASFS V5329
ASFS V5332
ASFS V5305
ASFS V5267
ASFS V6407
ASFS V5908
ASFS V5909
ASFS V4656
ASFS V4548
ASFS V5939
ASFS V5973
ASFS
ASFS
ASFS
ASFS
ASFS
v6207
V4 3
nnn
Vall 5
V4446
ASFS V4447
ASFS V4448
ASFS V44g
ASFS V4482
ASFS vV4499
ASFS V4500
ASFS V4501
ASFS V4623
ASFS V5860
ASFS V5861
BREVIORA
ANNULI SEGMENTS
LATERALS
CAUDAL
BODY
251+ 4 +12
2554+3/4414
253+ 4 413
245+ 4 413
229+
226+
227+
229+
223+
+14
+14
+15
+16
+15
227+
225+
228+
234+
233+
29 StL
2324+3/44+14
225+ 3 +5*
216+ 3 +16
227+ +5*
+15
+6*
+15
+14
+15
WWWWW WWWWW
221+
221+
229+
227+
232+
+15
+15
+15
+14
+5*
WW EFWW WWW
231+ 3 +14
paane7 acl
259+ 3) lp
2344+2/34+15
232+ 3 +14
235+ 3
230+ 3
232+ 4
3
3
+15
+14
+15
+15
+6*
230+
231+
230+ 3 +15
22942/3+14
23143/3+15
234+ 3 415
235+ 3 +14
DORSAL
VENTRAL
16/16
14/16
16/17
16/18
16/16
16/16
16/16
14/16
17/18
16/17
16/17
16/16
16/18
16/18
16/18
16/16
16/18
17/16
16/18
16/18
16/16
16/17
15/17
15/18
16/17
14/16
16/17
16/16
77,
16/18
14/16
14/16
16/17
14/16
14/16
15/16
15/16
16/18
14/16
WWW
WWWWW WWWWW WWWWW WWWWW WWWWW WWWWW WWWWW
POSTGENIALS
EEE
FREFE FEWHNF BDEFUNW PEE EE FHEEEE NEE EE FEWEE
USSU SD DSO a0 Tet Ue USO Oe rises Oe pine
triwis
*
\O &\O VO BON WO OOWMOO®M DAOWOOM O0M'\'0M DMO OOM Or10.\00.0
POSTMALARS
+10*
v
=)
PRECLOACALS
POSTCLOACALS
PORES
447411
446410
446411
448411
446414
446412
448413
448413
447411
446413
448412
446412
448413
448412
448412
448412
448412
448412
447412
448412
446412
448413
448411
446412
446413
446412
447413
447412
448413
448413
446413
447412
447414
447413
448412
448414
448413
448412
448413
INTERNASAL SUTURE
FUUVEE FLEES FWEWN FEWNW WEEFED BDVUUE FUFEW
No. 249
INTERPREFRONTAL SUTURE
TOTAL LENGTH
TAIL LENGTH
1966
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS .
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
ASFS
V5862
V6620
v6621
V6622
V6623
v6624
V6625
V6626
V6627
V6628
V6631
V6632
V6633
V6634
V6635
V6658
V6659
V6660
V6661
V5755
V5939
v6027
X937
X938
X939
x940
X941
x9h2
X943
X4104
X4105
X4106
X4107
X4108
X4109
X4110
X4111
X4112
X4113
X4114
X4115
xX4116
X4117
x4118
X4119
X4120
X4121
X4122
X4123
x4124
X4125
X7381
X7 382
X7383
X7 384
X7 385
X7 386
X7 387
X7388
X7389
AMPHISBAENIDS OF PUERTO RICO
226+2/34+15
2234+2/3+14
223+ 3 +14
219+ 3 +14
220+ 3 +14
226+ 3 +12
219+2/34+13
221+ 3 +14
221+
224+
3
3
220+ 3
222+ 3
222+ 3
219+ 3
223+2/3+14
220+ 3
220+ 2
223+ 3
222+ 2
225+ 4
223+ 4 +14
2264+3/4412*
224+ 4 +12
222+ 4 +12
226+ 3 +13
223+3/4413
229+ 3 +14
220+ 4 413
22442/34+14
222+ 3 +14
+13
+14
+14
+14
+14
225+ 4
228+ 3
229+ 4 415
220+ 3
223+ 3
218+ 3 +14
230+3/4413
2234+3/4413
218+ 3 +14
22843/4414
228+ 4 +13
221+ & 414
221+ 3 +11
228+ 4 414
2224+3/4413
214+ 4 411
225+ 4 +14
226+ & +14
222+ 3 +13
222+ 3 +13
3
4
226+ 3 +14
225+ +14
227+3/4412
221+ 3 +14
229+ 3 +12
224+ 3 413
224+ 3 414
1
2204+3/4413*
227+ 3 +14
224+ 4 +12
14/16
16/16
16/16
16/16
16/17
17/16
18/18
15/18
16/16
WWWWW WWWWW WWWWWH WWWWW WWWWW WWWWW WWWWW WWWWW WWWWW WWWWW WWWWW WWWWW
FREE FREE FEE E FENDER FENFE FEWEW FREE FREE FWO FOAUFE FEE FN E FEES
© @MMO00 O~A0NO 00000
No.0. ON O.10'0~)0 @yINI®WMO OONIOD DMWOWOODMD OwWOwowodd OO00'0 000
448412
447414
446412
448412
5+84+14
448412
4+7412
448412
446412
447413
446413
447413
447412
448412
448412
448413
448413
446413
447412
448412
447413
448412
446412
446412
447413
446413
446412
446412
448411
446410
447410
446412
446413
446412
446410
446413
446411
446412
446411
446413
446412
446411
446413
+6412
+6412
448410
446411
446411
446411
446412
446412
446411
446410
448410
446413
446411
446410
446413
447410
448412
WADA UNE FENN E FENN UFEVODN UDNED FEE NERF ADDN KWEWFH WEWFEE NAW
ASFS X7390
ASFS X7391
ASFS X7392
ASFS X7393
ASFS X7394
ASFS X7395
ASFS X7396
ASFS X7397
ASFS X7398
ASFS V4798
ASFS V4802
ASFS V6168
A. XERA
UPR 84a
UPR 84d
UPR 86
UPR 91
ASFS V6408
ASFS V6418
ASFS V6423
ASFS V6427
UMMZ 73844
UIMNH 56910
ASFS V5646
USNM 152588
ASFS V5659
ASFS V5661
MCZ 81019
ASFS V5722
ASFS V5723
AMNH 94170
ASFS V6616
ASFS V6617
ASFS V6618
ASFS V6619
ASFS V6636
ASFS V6637
ASFS V6638
ASFS V6639
ASFS V6640
ASFS V6646
ASFS V6647
ASFS V6648
ASFS V6649
ASFS V6650
ASFS V6651
ASFS V6652
ASFS V6653
ASFS V6654
ASFS V6655
ASFS V6656
ASFS V6657
CM 40577
CM 40578
RT 1333
RT 1334
ASFS V6587
UMMZ 73846
ASFS V5800
MCZ 36301
226+ 3 +13
eames
227+ 4 +14
225+ 4 413
222+ 3 +14
229+ 3 +13
227+ 3 +14
224+ 4 +14
2264+4/3+14
219+ 3 415
232+4/2415
228+ 3 +13
233+ 3 +14
229+ 3 +15
236+ 3 415
224+ 4 +13
2214+2/3+14
geie/ 31)
234+ 3 +16
221+
3
220+ 3 +13
220+ 2
233+ 2
226+ 3 +14
Spee TO
234+
231+
225+
232+
232+
+13
+14
226+
231+
229+
233+
2244+
228+
227+
231+
237+
234+
NWNHNW WNWNW NWWNP
+
ru)
i=
232+ 2 +14
229+2/3413
230+ 2 +14
2284+3/2+14
22843/4412
2284+2/34+15
nw
i)
es)
+
FWWWNWNH WNHNHWW NWWW
4
rea
=
BREVIORA
15/18
16/19
16/18
17/18
16/18
16/18
16/17
16/18
16/16
16/16
16/18
17/19
16/18
16/16
14/18
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
WWWWW WWWWWH WWWWW
WWWWWWW WWWWW WWWWWH WWWWW WWWWW WWNWW WWWWW WWwn fw NWWW &
Ferrer FEES FEEEW
FFWWW NERF EN NEWNFU NNER EF HPwWwrwW NFEN EF NEFRN
FWUNNFEW UNF END
Fi nern
Um FPn FHENIF Fiwmit
ine wWiFfit Foil
Pnwiwnm
TE NENE ITN
@M0MM0 OMO000 OMMO0®
446412
446412
446412
446412
446411
446412
448413
446411
446411
449413
448413
448412
446411
448411
448412
446410
448410
448411
446412
448412
446413
447411
446412
446410
44648
446410
447412
446410
446412
446410
446410
446410
447410
448412
448411
448412
448411
448413
447411
446411
448410
448411
447413
448411
447410
448411
447410
448410
448410
448410
448410
448411
448411
44849
2+8410
447411
448410
446412
446411
448412
447412
FANN WWWWEF RDERDWA
=
NON @OOOnN ON@WOOAY) Mo oOo~ MBoOONInN ON@MOO NOVYNND NYNVYOaY NVNADON NNOWOOD
|
ye .
1966 AMPHISBAENIDS OF
A. SCHMIDTI
ASFS V5432 200+ 3 +19 14/16 2 3
ASFS V5433 201+ 3 +20 13/16 2 4
ASFS V5434 199+ 3 +18 14/16 2 3
ASFS V5435 200+ 3 +20 13/16 2 3
ASFS V5436 200+ 3 +20 13/16 2 3
ASFS V5437 196+ 3 +8* 14/16 2 3
ASFS V5334 199+ 3 +21 14/16 3 4
ASFS V5197 200+ 3 +22 14/16 2 4
ASFS V5198 196+ 3 +20 14/16 2 3
ASFS V5199 201+ 3 +20 14/16 2 4
ASFS V5258 202+ 3 +21 14/16 2 3
ASFS V5259 202+ 3 +21 14/16 2 4
ASFS V5260 204+ 3 +21 14/16 2 2
ASFS V5268 200+3/44+20 16/17 3 4
ASFS V5910 197+ 2 +8* 14/16 2 4
ASFS V5923 205+3/2+21 14/16 2 4
ASFS v5924 207+ 3 +21 14/16 2 4&
ASFS V5871 201+ 4 +21 14/16 2 5
ASFS V5999 200+ 3 +21 14/18 2 2
ASFS v6028 201+ 3 +21 14/17 2 4
ASFS V6029 192+ 3 +20 14/16 2 4
ASFS V5959 1964+3/2+20 14/17 2 3
ASFS V5940 195+ 3 +7* 14/16 2 1
A. FPENSSTRATA
ASFS V7504 243+ 2 +413 14/16 3 4
ASFS V7505 250+ 2 +413 14/16 3 4
ASFS V7564 2434+ 2 +12 14/16 3 4
ASFS V8066 248+ 2 +12 14/15 3 4
VINP 1478 247+ 2 +13 14/16
VINP 1479 2434+3/24+13 14/16
VINP 1485 246+ 2 +13 13/14
VINP 1490 24643/2+13 14/16
VINP 1496 251+ 2 +12 14/14
VINP 1497 24142/3+12 14/17
ASFS V7921 247+ 2 414 14/14 3 4
ASFS V7940 2484 2 +13 14/14 3 4
ASFS V7984 243+ 3 414 14/15 3 4
ASFS V7985 248+ 2 +14 14/16 3 4
RT 977 242+ 2 +144 14/15 3 4
ASFS V3805 240+ 2 +12 15/17 3 4
ASFS V3806 245+ 3 +13 15/16 3 4
PUERTO RICO
ADANNAANADNA NAANW BWANYN~ QnN AN~
10
10
\0 10 10 S10 00
446411
44849
448410
448414
446+10
4+749
446412
446411
546412
23
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NoveMBER 4, 1966 NUMBER 250
THE BRAIN OF THE EMU
DROMAEUS NOVAEHOLLANDIAE
Il. ANATOMY OF THE PRINCIPAL NERVE CELL GANGLIA AND
Tracts!
By STANLEY CoBB
INTRODUCTION
This contribution is to supplement a paper published recently
(Cobb and Edinger, 1962) on the gross anatomy of the brain
and pineal body of the emu. Microscopic serial sections! were
made of the whole of one of these brains (#104), embedded
in celloidin, and eut at 25 microns thickness. The sections were
stained with cresyl violet for cell bodies and with Loyez modi-
fication and Weigert’s stain for myelinated nerve fibers. Pieces
of the other brain (#85) 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 with reasonable accuracy.
Such measurements have significance in relation to function
(Cobb, 1964).
1From the Museum of Comparative Zoology, and the Department of Neurology
and Psychiatry, Harvard University, and the Laboratory for Psychiatric Research,
Massachusetts General Hospital.
This investigation was aided by a grant from the National Institute of Neuro-
logical Disease and Blindness, Grant #NB-03429-05.
I wish to thank 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 CEREBRAL VENTRICLES
In order to understand the brain of any bird, one must get
an understanding 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 he 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 Diomedea (albatross) which has big separate
bulbs. In the emu the ventricle is distinct but not large (Figs.
2,3). The ventricle of the olfactory bulb 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 periamyedalar (Figs. 8, 10).
Thus, over approximately the caudal third of the emu’s cerebral
hemispheres, the ventricle 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, or
wulst (Fig. 4) is important. In the emu the wulst is frontally
placed so the ventricles remain largely medial and _ vertical
until 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 base 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
1966 THE BRAIN OF THE EMU 3
merely potential spaces, but at their mesial portions, where
they join the aqueduct, the lumen is distinctly patent. Below
the level of the midbrain the fourth ventricle is seen, wide be-
neath the cerebellum and narrowing down to become covered
with nervous tissue in the medulla near the spinal cord.
ANATOMY OF THE FOREBRAIN
In birds, the main mass of the forebrain is composed of large
eray ganglia (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 ganelia traversed by bands of white
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 ganglha forming axonal connections with each
other. The naming of these ganglha 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 ganegha A, B, C, ete., and Kuhlenbeck (1938),
relying on embryological evidence, developed a nomenclature
for the cerebral gangla of birds that largely disregarded mam-
amalian analogies. His work has been substantiated and _ effee-
tively used by Jones and Levi-Montaleini (1958). Kuhlenbeck’s
investigations show that the ganegla (he ealls them ‘‘nuclei’’)
of the avian cerebral hemisphere arise from four embryonic
zones, dorsomedial and ventromedial (the larger olfactory part )
and dorsolateral and ventrolateral (the striatal nuclei) ; the lat-
ter two form the main masses of the cerebral 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 et 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 brain use a modified Kappers nomenclature with
abbreviations, as shown in my Table I. In mentioning different
parts of the striatum, I use Kuhlenbeck’s names and put
Karten and Hodos’ abbreviations in parentheses; for example:
nucleus epibasalis caudalis (A) is the archistriatum. (See Table
IT and Fig. 1, and the list of abbreviations. )
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IVIORA
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1966 THE BRAIN OF THE EMU 5
Figure 1 is drawn from a reconstruction of the forebrain
of the emu, the main ganglia or nuclei being shown in seven
sections at the levels indicated by the vertical lines drawn on
the lateral view of the whole brain (insert). The wulst or
nucleus diffusus (HA, 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.
Other main nuclei can be similarly traced to show their antero-
posterior relationships. The relation of one nucleus to another
in any one section can be seen in the photographie reproduction
CORTICOID
» N. DIFFUSUS DORSALIS AREAS
LAMINA FRONTALIS SUPREMA—————* OR
\ HYPERSTRIATUM
ACCESSORIUM
LAMINA FRONTALIS SUPERIOR———»~
w
N. EPIBASALIS DORSALIS a
LAMINA OR z
oO
HYPERSTRIAT HYPERSTRIATUM VENTRALE °
a
x=
N. EPIBASALIS CENTRALIS
OR
NEOSTRIATUM J ~-— ~ VENTRICLE
aN
man EPI. CENT. ACCESS.
OR ECTOSTRIATUM
N ENTOPEDUNCULARIS
vt
Sees \or
PALEOSTRIATUM PRIMITIVUM
a)
PERIAMYGDALAR
CORTICOID AREA
N. BASALIS
OR
PALEOSTRIATUM
AUGMENTATUM
ny Olf LAMINA MEDULLARIS DORSALIS
SEPTAL NUCLEI
Fig. 1A. A diagrammatic representation of the dorsoventral relations of
the principal forebrain ganglia, but no one section could show all of these.
6 BREVIORA No. 250
of the sections that represent various levels (Figs. 2 to 11). The
nucleus intercalatus 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. +). 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 diffusus) (HA, HD)
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
MiNi
——— !
“a |
LE: |
Zz a;
4 DORSAL
(Ge ae
|
Ty: CENTRAL Se
La
POU
Is
CENTRALIS )
Fig. 1B. 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 drawn 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 many cells lving in
the interstices. Some of these are small and might be said to
form a group which could be called nucleus interealatus (Figs.
6, 7). This is not clear in sections farther forward (Fig. 4).
In facet, no boundaries would seem to be definite enough to
justify deseribing 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 (HV) 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 year 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 usine 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 distinet (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 caudalis (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 surely not homologous
8 BREVIORA No. 250
with mammahan 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 with
what is called here ‘‘parahippocampal ecorticoid,’’ and called
‘‘oeneral cortex’’ by Kappers, Huber and Crosby (1936). 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 the embryo, 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 corticoid areas
are homologous with the temporal mesocortex of mammals, and
that the praepyriform corticoid areas are homologous with the
entorhinal areas of mammals. One thing 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 (1966).
COMPARATIVE MEASUREMENTS OF GANGLIA IN THE
FOREBRAIN
Believing that increase of any function in an animal is posi-
tively correlated with increase in size of the parts of the brain
significantly involved, it is of interest to compare the sizes of
the different parts of the brains of birds of various famihes 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
eanelia of the forebrain (Fritz, 1949). Since we know so little
about the function of the different ganglia of the forebrain, it
is difficult to choose which ganglia to measure. Fritz showed
differences in size of the ganglia he measured, but no correlations
with any functions. He chose to measure the volume of ‘‘hyper-
striatum’’ (n. diffusus plus n. epibasalis dorsalis), ‘‘neostria-
tum’ (n. epibasalis centralis), ‘‘archistriatum’’ (n. epibasalis
eaudalis), 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 parts plus n.
interealatus (i.e. all the ganglionic mass above the lamina
frontalis superior) ; 2) the n. epibasalis dorsalis (HV); 3) the
n. epibasalis centralis (N, E). The first was chosen because
it seemed, grossly, to vary so much from species to species, the
1966 THE BRAIN OF THE EMU 7)
other two because they are large forebrain ganelia and are
possibly associative in function and hence related to ‘‘higher
integrations’? and ‘‘intelligent’’ behavior (Table IT).
In the brainstem the n. isthmi, n. mesencephalicus, and tectum
opticum were chosen because of their relation to hearing and
vision, as described in a previous paper (Cobb, 1964), and by
Karten (1966). The method of estimating the volume of each
of these ganglia and of the whole hemisphere, with which they
are compared, has been deseribed previously (Cobb, 1964).
3riefly, it is to project and draw the structures to scale, and,
by knowing the thickness of the sections and their distances
apart and number, to compute the volume. Absolute values for
volumes of different structures are not reliable because of dif-
ferences in shrinkage from one brain to another, but relative
measurements in any one series of slides made from one brain
are significant.
An attempt was made to measure the areas of the hippocampus
and the corticoid structures, but because of their lone and nar-
row shapes, it was difficult to use the planimeter to measure
their areas. Photographs were therefore 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. diffusus
(HA and HD) had a volume 28 per cent as large as the hemis-
phere, and that the hippocampus plus the corticoid areas had a
volume 13 per cent of the size of the hemisphere. By the projee-
tion planimeter method (Cobb, 1964) the n. diffusus (HA and
HD) was also found to be 28 per cent of the hemisphere. For
these quantitative data on the emu brain see Table IT.
Table Il
VOLUMES OF CERTAIN Parts oF Emu BRAIN
Volume of one cerebral hemisphere (before fixation) approximately 6.3 ¢¢
percentage of hemisphere
Nucleus Diffusus (HA & HD) 28
N. Epibasalis Dorsalis (HV) 11
N. Epibasalis Centralis (N & E)? 32
Volume of one optic lobe (before fixation) approximately 0.5 ce
percentage of optic lobe
N. Mesencephaleus Lateralis 5.4
N. Isthmi Magnocellularis 1.9
N. Isthmi Parvoeellularis 2.5
Teetum Opticum 81.0
1 As defined in section on abbreviations and nomenclature and Fig. 1.
10 BREVIORA Nos 250
As to the compactness of the emu’s brain, it is obvious by
microscopic inspection that there are fewer nerve cell bodies
(perikarya) im 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 (Selasphorus platycercus).
Knowing the thickness of the section, the number of cells per
eubic millimeter was computed for stratum E of the optic tee-
tum of the midbrain, and for the midposterior part of the n.
epibasalis centralis (N). In each of these eases 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 two 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 figured. 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
eelloidin, there is much shrinkage of the brain tissue, and the
amount of shrinkage in two separately prepared series of brain
sections cannot be 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 III
ESTIMATES OF COMPACTNESS OF EMU BRAIN
COMPARED TO HUMMINGBIRD
Dromaeus Selasphorus
novaehollandiae platycercus
Number of cells per mm® of n. epi- 42,800 125,000
basalis centralis (N).
Number of cells per mm® of stratum 40,000 100,000
E of tectum
Purkinje cells per 100 linear miera 2.2 4.8
Average diameter of Purkinje cells 26.0 miera 12.0 micra
1966 THE BRAIN OF THE EMU 11
GRAY NUCLEI AND LAMINAE OF MIDBRAIN AND
HINDBRAIN
Figures 8 to 12 show the main gray masses of the midbrain
and hindbrain; these masses are either nuclei, large and small,
or laminated structures such as the nerve cell layers of the tee-
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 aeme of such de-
velopment is seen in the mammalhan 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, but
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, but were stained only
with 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 lke 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 parahippocampal corticoid
areas. The size of these areas is probably related to the medium
size of the olfactory bulbs. 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
iy BREVIORA No. 250
aceessorium ) which is relatively larger and is bounded below by
a loosely arranged lamina frontahs. This cannot be divided into
two parts, superior and suprema. The nucleus intercalatus is
represented by a group of small cells in the lateral part of the
area erossed by 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 emu’s eye sug-
gests a reason for the big wulst.
The midbrain is remarkable for its large optic teetum 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 birds with skill
in flying.
REFERENCES
Bores, J., EH. FirKkova and J. MARSALA
1960. Leao’s spread in depression in pigeons. J. Comp. Neurol., 114:1.
COBB, 8S.
1960. A note on the size of the avian olfactory bulb. Epilepsia, 1:394-
402.
1964. A comparison of the size of an auditory nucleus (n. mesence-
phalieus lateralis, pars dorsalis) with the size of the optic lobe
in 27 species of birds. J. Comp. Neurol., 122: 271-280.
Coss, 8S. and T’. EDINGER
1962. The brain of the emu (Dromaeus novaehoilandiae, Lath) 1.
Gross anatomy of the brain and pineal body. Breviora, Mus.
Comp. Zool. No. 170: 1-18.
Craiciz, EH. H.
1935a. The hippocampal and parahippocampal cortex of the emu. J.
Comp. Neurol., 61: 563.
1935b. The cerebral hemispheres of the kiwi and emu. J. Anat. 69:
380-393.
FRitz, W.
1949. Vergleichende Studien tiber den Antiel von Striatumteilen am
Hemisphiirenvolumen des Vogelhirns. Rev. Suisse Zool., 56: 461-
491.
GoupBy, IF’. and H. J. GAMBLE
1957. The reptilian cerebral hemispheres. Biol. Rev., 32: 383-420.
Huser, C. G. and E. C. Crosby
1929, The avian diencephalon. J. Comp. Neurol., 48: 1-177.
1966 THE BRAIN OF THE EMU 13
JONES, A. E. and R. LEvI-MONTALCINI
1958. Patterns of differentiation of the nerve centers and fiber tracts
in the avian cerebral hemispheres. Arch. Ital. Biol., 96: 231-281.
Kapperrs, C. U. A., G. C. HUBER and E. C. CrosBy
1936. The comparative anatomy of the nervous system of vertebrates,
including man. New York, MaeMillan, 2 vols.
KaArRTEN, H. J. and W. Hopos
1966. Stereotaxic atlas of the pigeon’s brain. (In press).
IKUHLENBECK, H.
1938. The ontogenetic development and phylogenetic significance of
the cortex telencephali in the chick. J. Comp. Neurol., 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 Research, 10: 49-86.
Roskg, M.
1914. Ueber die cytoarchitektonische Gliederung des Vorderhirns der
Vogel. J. Psychol. Neurol., 21: 278-352.
SMITH, G. H.
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. Vergleichende qualitative und quantitative Untersuchungen an
der Horbahn von Végeln. 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.
ABBREVIATIONS 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. VEL. 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. Purkinje cells
3. granular
4. medullary
CENTR. GRAY Central gray matter around ventricle
14
CHORIOID
DORS. LAT. CORTI-
COID OR DORSOLAT.
CORTICOID
FASC. LONG. POST.
HABENULA
HEM.
HIPPOCAMP.
INNER GRAN.
LAM. FRONT.
LAM. HYPERSTRIAT.
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.
a
eS
J. BASALIS
+
N. CEREBEL. LAT.
N. DIFF. DORS.
. DORSOMED.
N. DORSOLAT.
N. ENTOPED.
N. EPI. CAUDALIS
N. EPI. CENTRALIS
N. EPI. DORSALIS
N. GENIC.
INE ESE OFMEVAGE:
BREVIORA No. 250
Plexus chorioideus 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 epithalamus
Cerebral hemisphere
Hippocampus or cortex ammonalis
Inner granular layer of cells.
Lamina frontalis. Sometimes separated into su-
perioris and suprema
Lamina hyperstriatica
Lamina medullaris dorsalis
Lateral forebrain bundle, contains tracts to thala-
mus, midbrain, and hindbrain
Lateral lemniscus, lies in fibrous capsule of n.
mesen. lat.
Dorsal pedunele
Ventral peduncle
Lateral ventricle of the cerebral hemisphere
Lateral out-poeketing of the midbrain ventricle
beneath the tectum
Medulla oblongata
Paleostriatum augmentatum, the nucleus separa-
ted from neostriatum above and laterally by 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 be-
low the lamina frontalis; HV (Karten); Field D
(Rose)
Nucleus geniculatus lateralis
Nucleus hypothalamicus
1966 THE BRAIN OF THE EMU 15
N. INTERCAL.
N. INTERPED.
N. ISTHMI
N. MESEN. LAT.
N. MESEN. PROF.
N. NERV. ABDUC.
. OCULOMOT.
SOLMMVE SUR:
- OVOED:
. PARAVENT.
. PRETECT.
. ROTUNDUS
. RUBER
. SPIRIFORM.
. SUBPRETECT.
2 VES. IeAu:
VESTIB. VENTRO-
LAT.
OLF.
OLE. VENT.
OR:
OP. CHIASM
OUTER GRAN.
PARAHIP. CORTICOID
PERIAMYG. CORTI-
COID
r b b b i b b b b b
4 ZA 4 A 4 4 Z 4 4 4
Z
PONT GRAY.
POST. COM. OR P.
COM.
PREPYR. CORTICOID
STRAT. FIBR.
SUPRAOPTIC.
TECTUM OP.
TR. FRONTO-
ARCHISTRIAT.
Nucleus intercalatus hyperstriaticus; HIS (Kar-
ten); Field A (Rose), of nucleus diffusus dorso-
lateralis
Nucleus interpeduncularis
Nueleus isthmi, pars magnocellularis
Nucleus mesencephalicus lateralis, pars dorsalis,
also called torus semicireularis
Nucleus mesencephalicus profundus or tegmento-
pedunculopontinus
Nucleus nervus abducentis
Nucleus oculomotorius
Nucleus olivarius superior
Nucleus ovoidalis
Nucleus paraventricularis magnocellularis
Nucleus pretectalis
Nucleus rotundus
Nucleus ruber or red nucleus
Nucleus spiriformis
Nueleus subpretectalis
Nueleus vestibularis lateralis, Deiter’s nucleus
Nueleus vestibularis ventrolateralis
Olfactory bulb
Ventricle of olfactory bulb
Optic lobe
Optie chiasm
Outer granular layer of cells
Parahippocampal corticoid areas
Periamygdalar corticoid area, contiguous with
the parahippocampal areas; not outside the ven-
tricle
Pontine gray nuclei
Posterior commissure, from tectum to opposite
tectum and also connects spiriform and dorsal
mesencephalic nuclei
Praepyriform corticoid area
Stratum fibrosum
Dorsal supraoptic decussation
Teetum 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 periventriculare
F. Stratum fibrosum periventriculare
Tractus fronto-archistriaticus
16
TR.
TR.
TR.
TR.
OCCIP.-MESEN.
OP. MARG.
SEPTOMESEN.
THAL-STRIAT.
V. OR VAL.
VENT. M.B.
Ww.
BREVIORA No. 250
Tractus occipito-mesencephalicus
Tractus opticus marginalis, lamina A of the op-
tie tectum. Input from optic nerve to tectum
Tractus septomesencephalicus
Tractus thalamo-striatalis
Vallecula
Midbrain ventricle or aqueduct of Sylvius
Wulst
Ly
EMU
OF THE
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BREVIORA
Museum of Comparative Zoolog
rA
CAMBRIDGE, Mass. NoveMBER 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 Técnicas. 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 Provinee, and, in addi-
tion, from a new locality in the Puesto Viejo Formation south
of San Rafael, Mendoza Province, discovered by the former
Direccién Nacional de Geologia y Mineria. 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, Universidad Nacional de Tucuman; and Consejo Nacional
de Investigaciones Cientificas y Técnicas, Buenos Aires, Republica Argentina.
iS)
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 obtained
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 Stipanicie,
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 Neuquén provinees. The correlation proposed by these
authors, between the Argentinian Choiyoilitense and the Porfidic-
queratofirie series of the Chilean coast south of Los Vilos, is also
accepted by Bonetti. She concludes that the Argentinian Trias-
sie series with Dicroidium flora are essentially of Keuper age
(Carnian at the lower levels, Norian at the upper). This conelu-
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 lutitie level intercalated in
the upper part of the Chilean effusive complex with the Argen-
tinian Triassic flora from the Potrerillos, Cacheuta, Barreal, Cor-
taderita, ete., formations. In her clear and brief analysis, how-
ever, Bonetti poimts out the need for re-examination of the
determinations of most of the marine fauna below the Los Vilos
effusives. Quite apparently, the chronology of the Chilean effu-
Sive series 1s 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 — Koanisian. Romer (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 ‘‘ Kotriasico
Superior’’ (Scythian) than to any later time. As to the age of
the Sierra de las Higueras Formation as given by Stipanicic,
1966 TETRAPOD-BEARING TRIASSIC OF ARGENTINA a
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 Bonetti Reig Romer
1957 1963 1961 , 1963 1960 , 1962
Los Colorados
Los Colorados
NORIAN
A
CARNIAN Ischichuca
re
| PES es Ischigualasto
Table 1. Recent chronological interpretations of some Triassic forma-
tions of Argentina.
LADINIAN
ANALYSIS OF VERTEBRATE FAUNAS
Puesto Viejo Formation (Mendoza Province). — The fauna of
these red sandstones comprises, so far as is now known, only
therapsids of the infraorders Cynodontia and Dicynodontia.
The eynodont Pascualgnathus polanskii Bonaparte (in press.b),
known from very complete remains, is a diademodontid. It has
affinities of great significance with Trirachodon, 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 palate, 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, which immediately follows the Cynognathus 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,¢).
The dicynodont of this formation has been referred to the
African genus Kannemeyerta, 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
posteranial 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-
meyeria is cited by Kitching (1963) from the Ntawere Forma-
tion, and a Kannemeyeria from the Manda Formation of the
Rubhuhu 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 Seythian or Anisian in age. The presence of
Pascualgnathus, however, leads us to consider the therapsids of
the Puesto Viejo Formation as Anisian.
Potrerillos Formation (Mendoza Province). — Fragmentary
remains of a cynodont jaw constitute the only available tetrapod
material from this formation. Minoprio (1954) studied these
remains, and erected a new genus, Colbertosaurus, which he
placed in the Ictidosauria. However, the principal anatomical
feature used by Minoprio in his identification of Colbertosaurus
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 Middle Triassic and the South American
Middle or Upper Triassic; Colbertosaurus is thus more reason-
ably considered a gomphodont eynodont than an ictidosaurian
(see Bonaparte, 1962). Colbertosaurus can best be compared
with Pascualgnathus, from the Puesto Viejo Formation. The
two have many features in common: the type of postcanine 1m-
plantation, the number and size of the incisors, the large section
of canines, the larger size of the posteanines in the middle of
the tooth row, and the type of jaw constriction behind the
canines. Pascualgnathus has eleven posteanines and Colberto-
saurus nine.
1966 TETRAPOD-BEARING TRIASSIC OF ARGENTINA By
It is evident that there is a significant, although limited, re-
semblance between the two 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.
Cacheuta Formation (Mendoza Province). — From this for-
mation we know two brachyopid labyrinthodonts, apparently
representing two separate genera — Pelorocephalus Cabrera
(1944) and Chigutisaurus Rusconi (1948). We also have the
postcranial skeleton of a proterosuchian, Cuyosuchus Reig (1961),
which was described by Rusconi in 1951 as a brachyopid.
Watson (1956) agreed that Pelorocephalus 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 Seythian.
It would, of course, be out of the question to call them Keuper;
tentatively we may consider them to be of Anisian age. Cuyo-
suchus, as mentioned above, is considered by Reig (1961) to be
a proterosuchian; Hughes (1963) and Tatarinoy (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 Province). — 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. barthi from the Moenkopi Formation of the United
6 BREVIORA No: il
States. Reig (1961) and Casamiquela (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. higuerensis and the
Rigalites-like genus together, rather than to examine C. higueren-
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 C.
higuerensis. 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 Sauwro-
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 concordantly by the Ischi-
eualasto Formation, the age of which is fairly well defined. On
this basis, we conelude 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 Frenguell. 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 el
1965), Romer (1962a), and Bonaparte (1962, 1963a-e, in press.a,
b). The fauna of the Ischigualasto Formation is composed of six
different groups; these will be discussed individually.
(1) Labyrinthodontia. Promastodonsaurus bellmanni 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 eynodonts are Exaereto-
don Cabrera (1944), Proexaeretodon Bonaparte (1963d), and
Ischignathus 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
Ischigualastia (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 Aetosauroides and Argentinosuchus (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-
suchus from Brasil or Stagonosuchus from Africa. We can as-
sume that these characters represent anatomical advances over
the Brasilian and African rauisuchids, comparable to the posi-
tion noted in the cynodonts. 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 Aetosauroides may be slightly more primi-
tive than Aetosaurus (Casamiquela, pers. com.). Aetosaurus
and the related genus Stagonolepis (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 posteranial
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 posteranial material.
Reig reports some affinities with Coelophysis 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 Jschisaurus, 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 Triassie forms, we have no valid
arguments at present to consider them older than the Keuper;
we may, then, assign them to the Carnian.
(6) Rhynehosauridae. Complete remains of these curious
forms have been found at different levels of the Ischigualasto
Formation. Until the present, 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 Ischignathus, and the three saurischians appear as forms
1966 TETRAPOD-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 supposing that ecological conditions permitted a later
expansion of these forms. The advanced features of the cyno-
dont Ischignathus, which has departed far from the known
structure of the African Middle Triassic eynodonts, would sup-
port this view. The presence of both rhynchosaurs and cynodonts
has been considered by Romer (1960, 1962b) as strong evidence
that this is an upper Middle Triassic fauna —as noted above,
‘surely pre-Norian 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 with the supposedly
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 les concordantly
over the Ischigualasto Formation, and is about 800 m thick.
At its base has been found a skull and Jaws comparable to Ischi-
gualastia, indicating the possibility of the survival of some
genera common 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 Seythian 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
humerous 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
Los Colorados
UPPER TRIASSIC
Ischigualasto
Sierra
de las
Los Rastros
Higueras
Ischichuca
LADINIAN
MIDDLE TRIASSIC
Potrerillos
Puesto Famatinense
Viejo
Paganzo I11
Las Cabras
Table 2. A new attempt at chronology of some Triassic formations in
Argentina.
LITERATURE CITED
BONAPARTE, J. F.
1962. Deseripcién del craneo y mandibula de Huaeretodon frenguellii
Cabrera y su comparacién. ... Publ. Mus. Cienc. Nat., Mar
del Plata, 1: 135-202.
1963a. Promastodonsaurus bellmanni n.g. et n.sp., capitosdurido del
Tridsico Medio de Argentina. Ameghiniana, 3: 67-78.
1963b. Deseripeién del esqueleto posteraneano de Hxaeretodon. Acta
Geol. Lilloana, 4: 5-52.
1963¢. Deseripeiédn de Ischignathus sudamericanus n.gen, et n.sp.,
nuevo cinodonte gonfodonte del Tridsico Medio superior de San
Juan, Argentina. Acta Geol. Lilloana, 4: 111-128.
1966 TETRAPOD-BEARING TRIASSIC OF ARGENTINA ie
1963d. Un nuevo cinodonte gonfodonte del Tridsico Medio superior de
San Juan, Proexaeretodon vincei. Acta Geol. Lilloana, 4: 129-
1833}.
1963e. La familia Traversodontidae. Acta Geol. Lilloana, 4: 163-194.
(In press.a) Chiniquodon Huene, en el Triasico de Ischigualasto, Argen-
tina. Consideraciones sobre su asignacién familiar. Acta Geol.
Lilloana, 8:
(In press.b) Sobre nuevos terapsidos Triasicos hallados en el centro de
la Provincia de Mendoza, Argentina. Acta Geol. Lilloana, 8:
(In press.c) Una nueva ‘‘fauna’’ Trifsica de Argentina. Consideraciones
filogenéticas y paleobiogeograficas. Ameghiniana.
Bonetti, M. I. R.
1963. Contribucién al conocimiento de la flora fésil de Barreal, Dpto.
de Calingasta (San Juan). Thesis, University of Buenos Aires.
BRINK, A. S.
1963. Two cynodonts from the 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 (Paleont. No. 55), 8: 317-331.
1944. Sobre un estegocéfalo 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. Ameghiniana, 2: 3-9.
1962. Dos nuevos estagonolepoideos argentinos. Rey. Asoc. Geol.
Argentina, 16 (3-4): 143-203.
1964. Estudios icnolégicos. Gobierno Prov. Rio Negro. Min. Asuntos
Sociales. Buenos Aires, 229 pp.
Cox, (CB:
1962. Preliminary diagnosis of Ischigualastia, a new genus of dicyno-
dont from Argentina. Breviora, Mus. Comp. Zool., No. 156: 8-9.
1965. New Triassic dicynodonts from South America, their origins
and relationships. Phil. Trans. Roy. Soe. 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 tritylodonts. Ann. S. Afr. Mus., 44: 1-14.
GROEBER, P. F. C. AND P. N. STIPANICIC
1953. Tridsico. In: Geografia de la Reptblica Argentina. Gaea,
74 (okie yeilehle
HuENE, F. v.
1931. Die fossilen Fihrten im Rhit 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. Ropinson, AND T. K. R. CHOWDHURY
1964. A new vertebrate fauna from the Triassic of the Deccan, India.
Quart. J. Geol. Soe. London, 120: 115-124.
KXITCHING, J. W.
1968. The fossil localities and mammal-like reptiles of the Upper
Luangwa Valley, Northern Rhodesia. S. Afr. J. Sei., 59: 259-
264.
MrInoprio, J. L.
1954. Theriodonte en el Triasico de Mendoza. An. Soe. Cien. Argen-
tina, 157: 31-37.
PrEapopy, F. L.
1955. Occurrence of Chirotherium in South America. Bull. Geol. Soe.
Amer., 66: 239-240.
Reig, O. A.
1959. Primeros datos descriptivos sobre nuevos reptiles arcosaurios del
Trifsico de Ischigualasto (San Juan, Argentina). Rev. Asoe.
Geol. Argentina, 13: 257-270.
1961. Acerea de la posicién sistematica de la familia Rauisuchidae y
del género Saurosuchus (Reptilia, Thecodontia). Publ. Mus.
Cien. Nat. Mar del Plata, 1: 73-114.
1963. La presencia de dinosaurios saurisquios en los ‘‘Estratos de
Ischigualasto’’ (Mesotridsico superior) de las provincias de
San Juan y La Rioja (Reptblica Argentina). Ameghiniana,
3: 3-20.
Romer, A. 8.
1960. Vertebrate-bearing continental Triassic strata in Mendoza Re-
gion, Argentina. Bull. Geol. Soe. 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.
Rusconl, C.
1948. Nuevos laberintodontes del Tridsico de Mendoza. Rev. Mus.
Hist. Nat. Mendoza, 2: 225-229.
195la. Laberintodontes Tridsicos y Pérmicos de Mendoza. Rev. Mus.
Hist. Nat. Mendoza, 5: 33-158.
1951b. Rastros de patas de reptiles Pérmicos de Mendoza. Rev. Soe.
Hist. Geogr. Cuyo, 3: 1-14.
STIPANICIC, P. N.
1957. El sistema Tridsico en la Argentina. Internat. Geol. Congr.,
20th, Mexico, Tr. See. 2: 73-111.
TATARINOV, L. P.
1961. Materiales de pseudosuchios de la URSS. Paleont. Zhurn.
URSS, No. 1: 117-132. [In Russian. ]
1966 TETRAPOD-BEARING TRIASSIC OF ARGENTINA 15
WALKER, A. D.
1961. Triassic reptiles from the Elgin area: Stagonolepis, Dasygna-
thus and their allies. Phil. Trans. Roy. Soe. London, B 244: 103-
204.
Watson, D. M.S.
1948. Dicynodon and its allies. Proce. Zool. Soe. London, 118: 823-877.
1956. The brachyopid labyrinthodonts. Bull. Brit. Mus. (Nat. Hist.),
Geol., 2: 315-392.
(Received 6 May, 1966.)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NovemBeEr 4, 1966 NUMBER 252
~
THE CHANARES (ARGENTINA) TRIASSIC REPTILE
FAUNA
If. SKETCH OF THE GEOLOGY OF THE RIO
CHANARES-RIO GUALO REGION
By ALFRED SHERWOOD ROMER AND JAMES A. JENSEN
TALAMPAYA BASIN
In 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
ean be placed in its proper setting. Since our expedition was
aimed solely at fossil collecting, proper instruments for geologic
work 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
Mesozoic continental sediments located mainly in western La
Rioja Province south of Villa Unidn. No name has been applied
to this region; we shall here term it the Talampaya Basin, be-
eause 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 Sanogasta, a southern
1Frenguelli (1948) speaks of the ‘“‘Cuenea de Ischigualasto-Ischichuca,” but
this properly applies only to the southwestern portion of the basin.
9 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 Rio Guandacol (or de la Troya).
The sediments of the basin extend northward to the Villa Unién
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 Sanhogasta 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 Huaco 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 Morada, Cerro Rajado,
Cerro Bola, and Cerro Villa Uni6én; 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 Sanogasta. East of the
village of Paganecillo is the Loma de los Loros, composed of
Tertiary deposits (de Alba, 1954, pp. 54-56), farther south the
striking Sierra de los Tarjados, formed by Permo-Triassie sedi-
ments. South of the region of the Rio Talampaya, the west flank
of the Sierra de Sahogasta descends abruptly (due, presumably,
to major faulting [Fidalgo, 1963]) to a sand-covered plain,
averaging 10 to 15 kilometers in breadth, which slopes gently
1966 RIO CHANARES-RIO GUALO GEOLOGY 3
westward toward the Campo de Talampaya. But for the last few
kilometers before the campo is reached, erosion in the valleys
of the Chafares 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 Rioja 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.
68°
672
2
<
Dp 3
" ie)
S z
6 z
A
°
ws
°
Guandacol
s :
——_—
aN
\
ENS
t= Jachal
your"t 38
ies
66°
67°
Fig. 1. Sketch map of parts of La Rioja and San Juan provinces, to
show the position of the area studied. Rivers and principal roads indi-
cated; provincial boundary in broken line. Scale approximately 1/1,200,000.
4 BREVIORA No. 9252
During the preparation of the present paper we have further
corresponded, to our advantage, with Dr. Pedro N. Stipanicie of
the Comisién 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 em] per annum) ; railroads are entirely lack-
ing, and roads few. Irrigation from the Rio Vinchina has per-
mitted the growth of population to about 3500 in the Villa
Unién 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 Unién is connected by national roads with Guan-
daecol 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 Unién 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 Sanogasta. In general, the sandy
‘‘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 Bodenbender’s emphatie (and italicized) warning (1911, p. 96):
“NOTA —A los futuros explorades hago presente que no se puede contar —
excepto anos de mucha lluvia—con sufficiente pasto para los animales,” etc.
There is, however, a mule track, of considerable antiquity and still used, from
Jachal to Patquia, which crosses the Rio Bermejo at Paso Ferreyro, traverses the
Ischigualasto Valley, and continues on to the southeast.
é
1966 RIO CHANARES-RIO GUALO GEOLOGY 5
the area in 1873, and later by Bodenbender, 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
Provinee and adjacent regions of San Juan. It is to Boden-
bender that we owe the invention of the term ‘‘ Paganzo,’’ com-
monly used by 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, ¢,
1945, and especially 1948), in which he appears to have relied
greatly on the work of de la Mota and Ramaccioni. An excellent
résumé of this work, as regards the Triassic beds, is given by
Groeber and Stipanicie (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
superficial 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
6 BREVIORA INo.4252
“| Sand Dunes
Undifferentiated
ertiary
Los Colorados
Ichigualasto
i-—J Los Rastros
4 Chanares
PROVISIONAL GEOLOGIC MAP <<
OF THE
GUALO REGION,
LA RIOJA,
ARGENTINA
de Agua
Escondida
Approximate Scale, i
Hotes
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 Chanares drainage, of which the lower part
is notable for a considerable development of sand dunes which
1966 RIO CHANARES-RIO GUALO GEOLOGY tf
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 Plano del Gualo. Here it receives
several short tributaries. One from the northeast has along its
course the only 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
Plano 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 Plano del Gualo is a
prominent peak of voleanic rock, the Mogote de Agua Escondida.
Arising in this region is a stream whose channel may be termed
the Arroyo de Agua Escondida. 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; only 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 Escondida, as seen
in Figure 2) the highest beds of the series are vertically placed
and even somewhat overturned.
KL 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. dlo, etc.) called these redbeds the Estratos de Patquia; 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 Safogasta; some have been
carried by diastrophism to high elevations in the range. West of
the Sierra de Sanogasta, El Chacho redbeds are abundantly 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
Unién 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 Sanogasta 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.
TALAMPAYA 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 RIO CHANARES-RIO GUALO GEOLOGY 9
the overlying Tarjados Formation, the results are striking; the
soft nature of the Talampaya 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 cliffs 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 Talampaya 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 Chanares
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 Plano 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, light 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.’’ Stipanicie (in litteris) suggests
that the Talampaya, as well as the overlying Tarjados Forma-
tion, can be considered as part of ‘‘Paganzo III’ or ‘‘Famati-
rh»)
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 by him
at Cerro Bola (the remainder of this set of beds include erup-
tives and porphyritic conglomerates).
TARJADOS FORMATION
Above the Talampaya Formation he 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 mainly of red-colored sandstones, the upper
predominantly white. However, the contrast between 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
Plano del Gualo; scraps of skull and posteranial 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
Plano 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-
elomerate, are a dull red sandstone, 40 to 45 meters thick, with
1966 RIO CHANARES-RIO GUALO GEOLOGY ala
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 cliffs of the underlying Talampaya Formation.
Whenever the entire thickness of the Tarjados 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 systems of plants which grew on the un-
eroded surface of the Tarjados before the beginning of the
deposition of the overlying Chanares beds.
Bodenbender’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’’ (ef. Bodenbender, 1924; Turner, 1960). At all
events, it seems probable 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.’’ Stipanicie (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 I1’’ 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 BREVIORA Now2a2
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, which 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 which are in-
eluded 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 tuffs, 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 Chanares were from the lowest 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 tuffs, 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 Chanares 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 cliffs.
At their top, the Chanares tuffs are followed without any un-
conformity by the beds 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 the Chanares beds, somewhat
interrupted by faults, follows the base of the Los Rastros For-
mation from the northwest part of the Rio Chanares drainage,
eastward and southward to the Rio Gualo just beyond the point
where it leaves the Plano del Gualo; faulting has caused the
series to appear again to the eastward of this line, well up the
courses of both the Rio Chanares and Rio Gualo.
Because of lack of unconformity, it would be possible to con-
sider the Chanares 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 Chanares. 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 Chanares beds
do not, however, compare with the basal part of the Ischichuca
in the Rio de la Pena region as deseribed by Frenguelli (1948,
p. 191 and fig. 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 Mota’s unpublished
work.
Future study of the fauna will presumably give evidence as to
the age of the Chanares Formation. As seen in the field, the
material suggested a somewhat later evolutionary stage than that
seen in the Cynognathus Zone of South Africa. This zone seems
certainly to be of early Triassic, Scythian, date (Watson, 1942,
Lehman, et al., 1959). Hence the Chanares 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
Chanares 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 BREVIORA No. 252
The conformable contact between the Los Rastros and the under-
lying Chanares Formation is exposed over very considerable
areas; on the other hand, a Los Rastros-Ischigualasto 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-Ischichuca 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 continuous 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; tt 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 Chanares — 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-
eretions 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 Chanares canyon, near its mouth, there is a notable display
of a eycheal type of deposition which appears to be to some
degree characteristic of the formation as a whole. Here four
eyeles 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 Ischichueca 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
ia
1966 RIO CHANARES-RIO GUALO GEOLOGY 15
Los Rastros formations. This is also done by Frenguelli (1948)
who, it seems, was 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 Pefa 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 Ischichuea.
ISCHIGUALASTO FORMATION
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 ineludes some
poorly 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. Above (as in the Ischigualasto Valley)
16 BREVIORA No e252,
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
30odenbender (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 western 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 ease, 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 hmited 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 immediately 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 RIO CHANARES-RIO GUALO GEOLOGY ay
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 Rhaetie in 1953. No fossils have been described
from these beds; however, Sr. Bonaparte of the Instituo Lillo
of Tucuman has recently discovered dinosaur remains of Upper
Triassic type in the 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 Norian,
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, ete.) 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, hke 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 these 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 the Permo-Triassice basin, voleanic
rocks are prominent. De la Mota, for example, in his unpub-
lished works on Cerro Bola describes thick intrusions of mela-
phyrite; Frenguelli (1944a) likewise 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 the Cerro Morado 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 Mogote 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 Plano del Gualo. The dyke is
composed of green crystalline material which is less resistant to
erosion than the metamorphosed sediments immediately adjacent.
The Agua Escondida is not a true spring, but a eateh-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 Chanares
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.
LITERATURE CITED
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1966
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1916.
1924.
FIpALeo, F.
1963.
RIO CHANARES-RIO GUALO GEOLOGY 19
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Nae. Ciene. Cérdoba, 15: 201-252.
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de Mendoza, y de las Sierras centrales de la Reptblica Argen-
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El Calehaquefio y los estratos de la Puna de Penck. Bol. Acad.
2
Nae. Ciene. Cordoba, 27: 405-468.
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1963.
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Excursiones al territorio del Neuquén y a las provinecias de
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20 BREVIORA No. 252
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NOVEMBER 4, 1966 NuMBER 253
A NEW HISPANIOLAN GECKO
By Ricuarp THOMAS
10,000 S.W. 84th St., Miami, Florida 33143
Recent collecting in Haiti yielded a single specimen of an un-
described species of Sphaerodactylus 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, who 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 sahent distinguishing character, this
new gecko may be known as
SPHAERODACTYLUS ELASMORHYNCHUS! new species
Holotype: MCZ 81119, a female, taken ca. 5 km (airline) SSE
Marché Léon, Dept. du Sud, Haiti, at an elevation of 2600 feet
(790 meters), on 15 March 1966 by Richard Thomas. Original
number ASFS V9353.
Diagnosis: A species of Sphacrodactylus distinguished by: 1)
much enlarged supranasals 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 elasmorhynchus from all other
known species of Sphacrodactylus.
Description of holotype (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 From the Greek, elasmos, plate, and rhynchos, snout.
2 Although I could not see a definite nasal seale, even under high magnifica-
tion, I have called the large scale behind the naris the postnasal because of its
position (see Smith, 1946, fig. 27).
2 BREVIORA No. 253
Fig. 1. Holotype of Sphaerodactylus elasmorhynchus: 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. Seales of snout be-
hind platelike covering swollen, subimbricate to imbricate, and
keeled. Seales 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. Seales on anterodorsal parts of limbs flat-
tened and imbricate. Gular scales granular; ventral scales large,
rounded, flattened, smooth and imbrieate; transition between the
two on posterior part of neck abrupt. Dorsal seales 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, midventral 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 Sphaerodactylus. Also
distinctive are the size difference between the dorsal and ventral
scales and the relatively abrupt 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 Sphaerodac-
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 S. 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 elasmorhynchus is an adult or subadult, not a juve-
nile, and represents a small-sized species.
The distinctness of S. elasmorhynchus prevents an assessment
of its relationships at this time; it is not obviously a member of
any of the recognized groups of the genus. Although elas-
morhynchus has granular dorsal scales, it differs strongly from
all members of the predominantly granular scaled decoratus
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 probably a reflection of the small size of the
species.
4 BREVIORA No. 253
MM
HEAD LENGTH
10 20 30
SNOUT-VENT MM
Fig. 2. Scatter diagram of head length (anterior border of ear to tip of
snout) versus snout-vent length for six species of Sphaerodactylus: solid
rhombs, S. cinereus; hollow triangles, S. difficilis; crosses, S. roosevelti;
hollow circles, S. n. nicholsi; solid circles, S. parthenopion; solid hexagon,
type of S. elasmorhynchus. For each species, aside from elasmorhynchus,
the range of values plotted includes near-hatchling juveniles through mature
adults.
Grant (1957) has shown that the eniematie S. mgropunctatus
Gray, known from a single type of unknown provenance, is a
eranular scaled form. However, references to the snout seala-
tion of nigropunctatus in Grant (1957), and a drawing of its
snout (Boulenger, 1885, pl. 18), indicate that it has the normal
Sphaerodactylus snout scale configuration ; the type is also large
in size.
1966 NEW HISPANIOLAN GECKO 5
Remarks: The type of S. elasmorhynchus was found among
the roots of a standing but rotten tree on the side of a ravine
filled with a jumble of limestone roeks and boulders. Although
the surrounding countryside was much cut over, the ravine was
overgrown with mesic vegetation. Also taken at this locality
were specimens of the relatively little known Anolis monticola
Shreve. No other sphaerodactyls were found at this locality. Of
the named Hispaniolan species, S. copet Steindachner and S.
cinereus are the only ones occurring in this general region of
Haiti.
LITERATURE CITED
BOULENGER, GEORGE ALBERT
1885. Catalogue of the lizards in the British Museum (Natural His-
tory). London. 2d ed., vol. 1, pp. i-xil, 1-436.
GRANT, CHAPMAN
1957. The status of Sphaerodactylus nigropunctatus, a neotropical
gecko. Nat. Hist. Misc., Chicago Acad. Sci., No. 158, pp. 1-4.
SmirH, Hospart M.
1946. Handbook of lizards. Comstock Pub. Co., Ithaca, New York, pp.
INCRE) Ofihe
THOMAS, RICHARD, AND ALBERT SCHWARTZ
1966. The Sphaerodactylus decoratus complex in the West Indies.
Brigham Young Univ. Sci. Bull., Biol. Ser. (In press.)
(Received 23 May, 1966.)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. NovemBeEr 4, 1966 NUMBER 254
PRELIMINARY DESCRIPTIONS OF NEW
HONEY-EATERS
(AVES, MELIPHAGIDAE)
By Finn SALOMONSEN 1
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, Volume 12, which is now in press.
MELILESTES MEGARHYNCHUS BRUNNEUS hew subspecies
Type: AMNH No. 294369, adult male; Siwi, Arfak Moun-
tains, New Guinea; May 2, 1928; Ernst Mayr collector.
Diagnosis: Darker and more brownish than nominate mega-
rhynchus (type locality: Aru Islands). The upper parts are
warm brown without the olive tinge of megarhynchus, the under
parts are darker brownish grey than megarhynchus, 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 Salawati in the western Papuan Islands,
Vogelkop and Onin Peninsula in northwestern New Guinea, in
the Weyland Mountains tending towards nominate megarhyn-
chus.
1 Zoological Museuin, Copenhagen.
bo
BREVIORA No. 254
OEDISTOMA PYGMAEUM WAIGEUENSE new subspecies
Type: AMNH No. 301021, adult male; Waigeu Island; June
5, 1931; G. Stein collector.
Diagnosis: Differs from nominate pygmaeunm (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 pygmaeum. Pro-
portions smaller. Wing length of adult males 45-46 mm, in
pygmaeum 47-51 mm; bill 11.8-12.8 mm, in pygmaewm 13-14 mm.
Range: Waigeu 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 waigewense. 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 Watut 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 Huon Peninsula.
LICHMERA INCANA MAREENSIS new subspecies
Type: AMNH No. 337641, adult male; Maré Island, Loyalty
Islands; Nov. 8, 1937; L. Maemillan collector.
Diagnosis: Very similar to L. 7. 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: Maré Island, Loyalty Islands.
1966 NEW MELIPHAGIDAE 3
LICHMERA INCANA GRISEOVIRIDIS new subspecies
Type: AMNH No. 212924, adult male; Lopevi Island, New
Hebrides; Aug. 9, 1926; J. G. Correia (The Whitney South Sea
Expedition) collector.
Diagnosis: Very similar to L. 7. flavotincta (type locality:
Erromanga, New Hebrides), but paler throughout; crown paler
grey, upper parts paler and less brownish green, and, particu-
larly, under parts paler with less yellowish olive on lower breast
and abdomen. Differs from mareensis 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-
nouneed.
Range: Central New Hebrides (Efate, Makura, Mai, Tongoa,
Epi, Lopevi, Pauuma, Ambrym, and Malekula).
MyYzoMELA NIGRITA NIGERRIMA new subspecies
Type: AMNH No. 422677, adult male; Long Island, off the
northeastern coast of New Guinea; Dee. 2, 1933; W. F. Coultas
(The Whitney South Sea Expedition) collector.
Diagnosis: Similar to M. n. ernstmayri (type locality: Manu
(= 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 mm, in ernstmayri 75-77; adult fe-
males 71 mm, in ernstmayri 66-69 mm.
Range: ong Island, off the northeastern coast of New
Guinea.
MyzOMELA CARDINALIS ASUNCIONIS new subspecies
Type: AMNH 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) exeept 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 parts 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 saffordt.
MELIPHAGA MIMIKAE RARA hew subspecies
Type: AMNH 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, granti (type locality: Mafulu, mountains of
southeastern New Guinea) and nominate minuikae (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 mimilkae 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. minukae in its general diagnostic
characters (dark coloration, especially on upper parts, ochra-
ceous under wing-coverts and pale buffish 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
Type: AMNH No. 422342, adult male; Wuroi, Oriomo River,
3ritish Papua (western division), southern New Guinea; Jan.
28, 1934; The Archbold Expedition collection.
Diagnosis: Very similar to nominate analoga (type 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-coverts 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 ANALOGA 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 flavida (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 slightly 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 dusky 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 flavida and stevensi. Wing length of adult males of
topotypical flavida 82-85 mm, of connectens 79-83 mm, of
stevenst T7-80.
Range: 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’Entrecasteaux 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 Western 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
Type: AMNH No. 348828, adult male; Ward River, south of
Charleville, southwestern Queensland, Australia; March 1, 1940;
L. Maemillan 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 vellow, cheeks and ear-coverts yellow, without
the olive greenish tinge found in nominate penicillata; black ear-
stripe reduced or absent. Also slightly smaller. Wing length of
adult males of topotypical penicillata 85-92 mm, of tnterioris
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: AMNH No. 428247, adult male; Tarara, Wassi Kussa
River, 90 miles west of Daru, British Papua (western division),
southern New Guinea; Dee. 8, 1936; The Archbold Expedtition
collection.
1966 NEW MELIPHAGIDAE 7
~
Diagnosis: Much paler throughout than saturatior (type lo-
eality: Aru Islands) and giulianettw (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 giulianettu 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 distinetly paler earth-brown than in saturatior
and giulianettu, 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 VIRIDIFRONS hew subspecies
Type: Acad. Nat. Sci. Philadelphia No. 132851, adult female
(labelled male) ; Bamoskaboe, Karoon, Tamrau Mountains, Vo-
gelkop Peninsula, 2300 feet altitude, northwestern New Guinea ;
March 1, 1938; D. Ripley (Denison-Crockett South Pacifie Ex-
pedition) collector.
Diagnosis: Differs from nominate obscura (type locality: Mt.
Seratchley, southeastern New Guinea) in having front and crown
green, like nape and upper parts, while in obscura front and
crown are grey, contrasting with nape.
Range: Mountains of Vogelkop Peninsula, northwestern New
Guinea. Specimens from the Weyland Mountains are interme-
diate between viridifrons and nominate obscura.
MELITHREPTUS LAETIOR NORMANTONIENSIS new subspecies
Type: AMNH No. 691532, adult male; Normanton, northwest-
ern Queensland, Australia; Jan. 31, 1914; R. Kemp collector.
Diagnosis: Differing 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, shghtly 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 june-
tion of Black and Palmer rivers, 100 meters altitude, British
Papua (western division), southern New Guinea; July 13, 1936;
The Archbold Expedition collection.
Diagnosis: Ditters 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 txoides 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 ¢inna-
mon throat contrasting with greyish brown breast and abdomen
(in troides 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 wing-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 «wroides, although slightly tending towards
cineretfrons.
PHILEMON CITREOGULARIS CARPENTARIAE new subspecies
Type: AMNH No. 696823, adult male; Normanton, north-
western Queensland, Austraha; May 1914; R. Kemp collector.
Diagnosis: In coloration, hardly differing from nominate
citreogularis (type locality: interior of New South Wales) and
johnstont (type locality: Johnston River, northern Queensland),
only very slightly paler generally, but differg from nominate
citreogularis in having a much larger bill, and from johnstoni in
having much greater wing length. Wing length of adult males:
1966 NEW MELIPHAGIDAE 9
citreogularis 136-140 mm, carpentariae 135-141 mm, johnstoni
127-131 mm; bill length: citreogularis 30.5-31.5 mm, carpen-
tariae 33-34 mm, johnston 33-34 mm.
Range: Southern coast of Gulf of Carpentaria, northwestern
Queensland.
Remarks: It is possible that the name pseudonymus Mathews
and Neumann, 1939 (type locality: Cloneurry, 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
Cloncurry. 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-
eality: Triton Bay, southwestern New Guinea), brevipennis
(type locality: Utakwa River, southern New Guinea) and arwen-
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 brevipennis in being
much larger. Wing length of adult males: topotypical brevi-
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.
Meek collector.
Diagnosis: Same general coloration as subtuberosus (type lo-
cality: Fergusson Island, D’Entrecasteaux Archipelago), but
slightly lighter (the subtuberosus from the Trobriand Islands
are the darkest), with distinetly longer bill and, especially, with
larger casque on bill. Length of bill in adult males of trivialis
47.5, 50.2 mm, in subtuberosus 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 Kumusi 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
elliott (type locality: Mt. Elhot, northern Queensland). Wing
leneth of adult males: corniculatus 159-166 mm, clamans 154-
156 mm, elliote 137-147 mm.
Range: Southeastern Queensland.
MELIDECTES FUSCUS GILLIARDI new subspecies
Type: AMNH No. 705793, adult male; Mt. Wilhelm, Bismarck
Range, 11,000 feet altitude, east-central New Guinea; June 7,
1950; KE. T. Gillard collector.
Diagnosis: Same coloration as occidentalis (type locality:
Wichmann Mountains, central New Guinea) and nominate fws-
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 fuscus 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
gilliardy 114-116 ole D-35.0 32.9-39.0
fuscus 113-117 34.0-37.0 33.0-35.0
Range: Bismarek Mountains, eastern New Guinea.
Remarks: Named after the collector, the late Dr. E. Thomas
Gilhard.
MyzA CELEBENSIS PARVIROSTRIS new subspecies
Type: AMNH No. 3002382, adult male; Mt. Tanke Salokko,
2000 meters altitude, Mengkoka Mts., southeastern Celebes; July
22, 1931; G. Heinrich collector.
1966 NEW MELIPHAGIDAE ial
Diagnosis: Differs from nominate celebensis (type locality:
Bone Mts., Gorontalo, northern Celebes) in its much smaller
proportions, especially its smaller bill. Wing length of adult
males: nominate celebensis (from Latimodjong Mts., central
Celebes) 85-87 mm, parvirostris 81-83 mm; bill length: nominate
celebensis 32-84 mm, parvirostris 28-30 mm. No material from
the type locality of celebensis has been examined, but a series
from Ile-Ile, northern Celebes, is similar to the Latimodjong
birds in measurements, although not quite so big.
Range: Mengkoka Mountains, southeastern Celebes.
PHYLIDONYRIS 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 with 75-83
mm in all other subspecies.
Range: King Island and Flinders Island in Bass Strait,
Australia.
ACANTECRHYNCHUS TENUIROSTRIS TROCHILOIDES new subspecies
Type: AMNH No. 703283, adult male; Bunya Mountains,
2000 feet altitude, southeastern Queensland, Australia; Oct. 3,
1940; L. Maemillan collector.
Diagnosis: Differs from nominate tenwrostris (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: Restricted to the isolated pockets of rainforest in
southeastern Queensland.
ACANTHORHYNCHUS TENUIROSTRIS REGIUS new subspecies
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 tenwirostris, with bill
24.5 mm, tarsus 18.2 mm, wing 68 mm. The largest bill of adult
birds of any subspecies is 31 mm, reached in nominate tenutros-
tris; the bill 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.
Range: King Island in Bass Strait; probably also on Flinders
Tsland.
(Received 23 May 1966.)
BREVIORA
Museum of Comparative 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 floridanus, 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!
FLORIDATRAGULUS DOLICHANTHEREUS White
Floridatragulus dolichanthereus White, 1940, p. 35, pl. VII.
Syndyoceras australis White, 1941 (in part), p. 97, pl. XV, figs. la, 1b.
Hypermekops olseni White, 1942, pp. 11-12, pl. 8.
Floridatragulus barbouri White, 1947, pp. 505-507, fig. 4.
Hypodigm: MCZ 3635, type, incomplete mandible with M, to
Ms and alveoli for P; to Ps, anterior crescent of right M, and
hypoconulid of right M; missing, both ascending rami and the
symphysis anterior to the C-P, diastema missing; MCZ 4086
(type of F. barbouri), right mandible with partial symphysis and
complete ascending ramus, left canine, right P. to Ms and the
alveoli for P;; MCZ 3711 (type of Hypermekops olseni), incom-
plete skull with right It and I°, P+ to M3, left I?, P* to M3, 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 P; to Ms and alveoli for P; and Py»;
MCZ 7780, 30 astragali; MCZ 7781, 11 calcanea; MCZ 7782, 15
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, pl. 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 I! 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,
flat, 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 Nom255
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 P,; 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 P,, 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 Pj, as in Myotylopus bathygnathus.
The dental formula is 13, C1, P{, M3. In MCZ 3711 I 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 I! and I?; this is the longest spacing
within the dental row. I? 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.
I° is separated from I? by a diastema equal to somewhat less
than haif 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 I?. 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 P!. 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 I! and I?
separates P! 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.
P® 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 M1! to M3,
as in Paratylopus primaevus, and are as brachyodont as in that
species. M1 is wider than it is long resembling the nearly square
first molar of P. primaevus and M. bathygnathus, more so than the
elongated M! 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
“JT am not convinced that cameloides belongs in this genus.
6 BREVIORA IN@: 0255:
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.
MCZ 37Il | MCZ 3642 MCZ 37Il
MCZ 4086 | MCZ 3635
Figure 1. Floridatragulus dolichanthereus. a, Diagram of upper denti-
tion, X 4/5. b, c, Composite crown view of left cheek teeth; b, upper P?
to M2, c, lower Ps to Ms, X1.
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 SELENCDONT 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.
P, is separated from the canine by a long diastema somewhat
exceeding the combined length of M, and Mb in the type, but
variable in referred specimens. P, is not preserved, but the alve-
oli indicate a small, one-rooted tooth probably similar to P! and
roughly equivalent to it in size.
After another long diastema the second premolar follows (Fig.
1c) 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 compietely unworn condition.
The diastema between P, and Ps is quite variable; that of MCZ
4086 is 52 per cent smaller than that of the type. Ps is similar to
P. 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.
P, follows immediately behind Ps; 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 Ps;, but
does not form the prominent right angie 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 INO: 2255
early to the entoconid and protoconid. The hypoconulid 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 P,-Ps and P.-P., diastemata are
quite variable, but this may be due to the functionless condition
of P,. 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 P, 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
3G. M. Allen, in a personal communication to T. Barbour, expressed
this view more than 20 years ago (correspondence in the Harvard Archives).
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 I® 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 Mz; 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 Dorcatherium, 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 M; divided.
TABLE 1
Comparative measurements of Floridatragulus
dolichanthereus in mm
Floridatragulus Paratylopus
dolichanthereus primaevus
MCZ 3711 MCZ 3642 AMNH 9806
Skull length to condyles *385 — *200
Width of palate at P’ 45 — 21
Width of palate at M? *716 _- 52
Ratio: snout (Pmx to orbit)
to skull length *0.70 — 0.51
Tip of Pmx to 23 — =
I1 length 2) — —
I! width 7 — =
I? length a — —
I? width 4.5 — —
I! to I? diastema S)/55) — -—-
I? to canine 9 —- 3
Canine to P! diastema 9 —
P! to P? diastema 28 — 75
P! length 11.5 — =
P! width 4 _- “=
P® length — 10 =
P® width _- 5 —
P*4 length 9 9 8
P+ width 8.5 8.5 “8
M!? length PZ 13:5 “10
M! width Pils 115) mil
M? length 16 17, "13
M? width 17 17 *113
M® length 18.5 — 16
M2 width 18.5 — 14.5
1966 | SELENODONT ARTIODACTYLS FROM THOMAS FARM 11
Floridatragulus dolichanthereus
MCZ 3635 MCZ 4086 MCZ 4291
Jaw length — *300 —
Length of symphysis behind P; 13 14 3
Width of jaw at P; SE, 19:5 —
Width at Mz — *70 ~-
Depth of jaw under P; DCS: 21 14
Depth under P, “32 32 23
Depth under M;, *38 am 32)
Canine to P; diastema — 28.5 --
P; to Po diastema 33 37 24
P2 length —_ 10.5 —
Ps width _- Se, —
Ps to Pz diastema 21 10.5 19:5
P; length — 10.5 SLOSS
P; width —: 4.5 “4.5
P, length -— 10 “10
P, width ~- 6 6
M; length =13 10 10.5
M, width 2 9 8.5
Mp2 length LSy55) 15 15#5
Mp» width 11 11 (ld
M3; length Zed 13 13
M3; width 12-5 13 13
“estimate based on incomplete specimen.
» worn.
ALTICAMELINAE
NOTHOKEMAS FLORIDANUS (Simpson) 1932
Oxydactylus floridanus Simpson, 1932, pp. 35-37, figs. 20, 21.
Paratylopus grandis White, 1940, p. 33, pl. V.
Nothokemas grandis White, 1947, pp. 508-515, figs. 5, 6.
Hypodigm: FSGS V-5247, type, incomplete right maxilla with
P= to. Maz ESGS, VY-5238; paratype, right lower jaw with Ps; to
M;; MCZ 3636 (type of N. grandis), incomplete right mandible
with P; to Mz and alveolus for Pj; MCZ 4322, right maxilla with
P, to Ms; MCZ 4323, left mandible with P, to Ms and anterior
crescent pair of M;; MCZ 4324, partial left mandible with P. to
12 BREVIORA No. 255
P4; MCZ 4325, left mandible with M, to Ms and alveoli for P.
to P,; MCZ 4326, left mandible with dP, to Ms; MCZ 4329,
crushed skull with poorly preserved P? to M®, rostrum anterior
to P1-P? diastema missing; MCZ 4541, crushed skull with dP? to
M3, anterior snout missing; MCZ 7297, left mandible with dP,
to Mz; MCZ 7592, incomplete left mandible with My to M3;
AMNH 22672, left mandible with Ps to Ms; MCZ 7767, 22
astragali; 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 P!. 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 P? is a long diastema approximately equal to the combined
lengths of P+, M1! and M?.
2 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
P 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 M8. In this character Nothokemas
resembles “P.” cameloides and contrasts with the pillarless molars
of Oxydactylus.
M! 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
I, to P;, is unknown. MCZ 4325 has a broken symphyseal area
which contains what may be the socket for P,. 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 P,
14 BREVIORA INOPZ55
and P, 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. P. 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.
P. has the anterior cusp very strongly inflected internally, more
so than P,. The posterior intermediate crest is somewhat more
developed than in P., enclosing with the posterior crest a deeper
but still medially open valley. P4 is similar to P;, but with the
posterior intermediate crest long and completely enclosing a deep
valley, 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 My. 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 My.
The tarsal bones and phalanges that may be referred to Notho-
kemas on the basis of size and relative abundance are very Oxy-
dactylus-like, 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, P; 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
Oxydactylus “Paratylopus”
Nothokemas floridanus longipes cameloides
MCZ 4329 MCZ 4322 MCZ 4541 CM 918 YPM 10917
Skull length “340 — — 340 »278
P1-P2 diastema *53. — — 18 —
P? length 13 12.5 — ili! —
P2 width 6 Sed) — 6 —
P® length 11833) 13.4 — 14 —
P? width 8.5 VS — 8 —
P+ length =113} 13 — 14 11
P+ width 11 2 — 10 10
M! length — 18.6 19.7 19 13
M! width — 18.5 18.5 16 —
M? length — “Bp 22 2B es
M? width — DES 20.5 20 16.5
M® length 23 23 23 nS 17.4
M® width 19 IS) 18.8 20 17.3
Oxydactylus “Paratylopus”
Nothokemas floridanus longipes cameloides
MCZ 3636 MCZ 4325 MCZ 4324 CM 918 AMNH 8197
P,-Ps diastema — 45.5 — 19 aN
Ps length — = 10.8 10 10
Ps width — — 4.5 5 —
P; length 13.8 — 12.8 12 2S
Ps width 6.4 — 5)5) 5 4.3
P, length IS oa 14.5 12 13
Py width tS — W383 7 5)
M; length “18 19.2 — 18 14
M, width 10.7 10.7 — 10 14
M2 length DLS 23.6 — 23 eS
Me» width 14 13.2 — 14 ital
M3 length 35 33 — 31 DS
Ms: width 16.5 14.1 — 13 122
*“ estimated from incomplete specimen.
» estimated from figure, Dougherty, 1940.
16 BREVIORA No. 255
crests. P, is marked by a very strongly inflected parastylid in
Hypisodus and Hypertragulus, 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-like 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
N. 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. INDET.
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).
P® 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 7
PROTOCERATIDAE
SYNTHETOCERAS (PROSYNTHETOCERAS) RILEYI AUSTRALIS
(White), 1941 new rank as subspecies
Syndyoceras australis White, 1941 (in part), p. 97, pl. XV, figs. 2, 2a.
Synthetoceras (Prosynthetoceras) douglasi White, 1947, pp. 504-505, fig. 3a.
Hypodigm: MCZ 3654, type, right mandible with P,, P, to Ms,
and alveoli for P, and P:, coronoid process and symphysis an-
terior to P, missing; MCZ 3708, right mandible with P,; to Ms;
and alveoli for I, to Pp; MCZ 3655, right mandible with P, to M;
and partial alveoli for I, to the canine; MCZ 3656, right mandible
with P, to Msg and partial alveoli for I, to the canine; MCZ 3666,
partial right mandible with P; to M3; MCZ 4065 (type of Synthe-
toceras douglasi), palate with right and left P? to M*; MCZ 7635,
right mandible with P, to Ms and alveoli for I, to P;; MCZ 7636,
incomplete left mandible with P, to M3; MCZ 7637, left mandible
with P; to Mg and alveoli for P; and P,;; MCZ 7774, 13 astragali;
MCZ iid>;. 3. cuboids: MCZ, 77.76, 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 P; is 13 per cent smaller, and median cusp of Py 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 P., the jaw begins
to flare outward so that P, 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 I,, suggesting that
it was a functional member of the incisor series, as in Protoceras.
P, follows after a short diastema equal to M, in length. In the
+ Frick’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 INOs255
type and MCZ 3656 and 7637, P, 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 P, and P.. The latter is a two-rooted,
simple, robust tooth equal to Ps in length (Fig. 2). Ps 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.
P, is similar to P; 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. My,
and M, have narrow crescents, strong parastylids, and weak
metastylids. The median ridges are well developed, especially on
Ms. Mg 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.
g SSS SS =, —S
MCZ 3654 cna MCZ 3660
EIS
MCZ 3656
Figure 2. Synthetoceras (Prosynthetoceras) rileyi australis, composite
crown view of right lower cheek teeth, X1.
In the upper dentition, the referred palate most closely resem-
bles that of Synthetoceras francisi 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
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.
P* 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 S. 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 ot
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
TABLE 3
No. 255
Comparative measurements of Synthetoceras
(Prosynthetoceras ) rileyi australis in mm
Width of palate between M!
P® to M® length
P® length
P? width
P*# length
P* width
M! length
M! width
M? length
M2 width
M® length
M® width
Depth of jaw
under Po
Depth under P4
Depth under M;
P2 to Mz length
P; to Py diastema
Ps length*
Ps width*
P; length
Ps width
Py length
P, width
M; length
My, width
Mo length
Mo» width
M3 length
Ms width
MCZ 3654 MCZ 3708
PIMeg2 DESI
23.6 a
29M DY
85.4 76.6
42.0 46.0
— 8.5
— 4.0
10.5 10.5
6.5 6.3
13.0 12.2
10.0 OD
18.0 16.1
11.0 10.5
YES) 25.0
11.4 IN dleal
Synthetoceras (P.) rileyi australis
MCZ 4065
Synthetoceras (P.)
rileyi australis
64.5
64.8
9.0
6.1
9.0
3:6
13.6
14.8
V7/e2
15.9
17.0
16.8
MCZ 7635
Pp bcs)
23.0
28.4
76.7
10.1
Soi!
14.4
9.4
15.4
21.6
21.6
10.5
Synthetoceras (P.)
rileyi rileyi
AMNH 34181,
type
1933
20.5
77.4
8.2
Siell
9.7
4.3
10.4
6.4
12.4
9.4
16.7
22.4
22.4
10.1
“ Known only from MCZ 3656; length, 8.2 mm, width, 4.0 mm.
1966 | SELENODONT ARTIODACTYLS FROM THOMAS FARM P|
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 Merychyus 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 anteroposteriorly, 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), X1%.
There can be little doubt of the oreodont affinities of the tarsal
elements, which probably belong 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 INO: 7255
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
BLASTOMERY™*. (PARABLASTOMERYX ) FLORIDANUS White, 1940
Hypodigr.:: MCZ 3626, type, partial left mandible with P4,
M,, the ante: or external crescent of M., and the alevoli for Ps
and P;; MCZ 3627, paratype, upper left molar; MCZ 3706, right
mandible with dP, to M2; MCZ 3707, left mandible with dP. to
Ms; MCZ 3912, left mandible with dP, to Ms; MCZ 4220, left
mandible with P,, dP; to M3; MCZ 4221, right mandible with
P; to M3; MCZ 4222, right mandibie with P., M, 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: Middie Miocene, Hawthorne Formation,
Thomas Farm, Gilchrist County, Florida.
Diagnosis: White (1940, p. 34) diagnosed this species as fol-
lows: “A large Parablastomeryx, paraconid of Py 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.
MCZ 7763 MCZ 4349
Figure 4. Blastomeryx floridanus. a, Composite crown view of left lower
cheek teeth, X12. b, MCZ 3706, right dP4, X3.
The hypoconulid of M; has a prominent anterointernal cusp,
and a strong palaeomeryx fold is present on the protoconid. The
internal stylid of P, which White described as being “very small”
on the type varies from strong on MCZ 7763 to absent on MCZ
4221. In Ps 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 enciosed 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 mediaily, 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, dP, to dP, are known from two speci-
mens, MCZ 3706 and 3707. DP, (Fig. 4b) is five cusped with
an elongated pillar between the anterior inner cusps and a conical
pillar between the posterior inner cusps. DP. is very similar to
Ps in structure and length, but is much narrower. There is little
difference between dP» and Po».
24 BREVIORA INo#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.) floridanus in mm
MCZ MCZ MCZ MCZ
7763 4221 4349 3626
Jaw and lower dentition:
Jaw length — plis2 — —
Ps to M3 — 60.8 66.9 —
Ps length 5.6 — — —
Ps width 2.6 -- — —
Ps length 7.3 8.0 8.2 —-
P3 width 4.3 4.3 4.6 —
P, length 8.9 8.4 8.7 8.2
Py width 5.4 5.0 5.0 4.9
M; length — 10.1 10.3 9.0
M, width — ES 7.0 6.3
Mp length — 11.8 silo z —
Me width — 8.2 TS) —-
M3 length — 16.2 eS —
Ms width — 8.2 Vel =H
“estimated from incomplete specimen.
1966 SELENODONT ARTIODACTYLS FROM THOMAS FARM 25
MACHAEROMERYX GILCHRISTENSIS White, 1941
Hypodigm: MCZ 3651, type, partial right mandible with P, to
Mz, and alveoli for P2, P; and M3; MCZ 3652, right mandibular
fragment with M,, and a left upper molar; MCZ 3709, partial
right mandible with P; to Mz, and part of M;; MCZ 3710, partial
right palate with M! and M2; 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,
My 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
since 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 floridanus and Machaero-
meryXx 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 Nossa»)
REFERENCES
BADER, R. S.
1956. A quantitative study of the Equidae of the Thomas Farm
Miocene. Bull. Mus. Comp. Zool., 115: 49-78.
Cook; CW.
1945. Geology of Florida. Bull. Florida Geol. Surv., No. 29: 1-339.
DouGHERTY, J. F.
1940. Skull and skeletal remains of the camel Paratylopus cameloides
from the John Day, Oregon. Carnegie Inst. Washington Publ.
No. 514: 49-58.
Fisk, HN:
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: 1-11.
PATTERSON, B.
1949. Rates of evolution in taeniodonts. Pp. 243-278. In G. L.
Jepsen, G. G. Simpson, and E. Mayr eds., Genetics, Paleon-
tology, and Evolution, Princeton Univ. Press.
Purl, 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.
RAv, CE:
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 af
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. la, 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 Pz to Mz read P?
to M?.
Page 15, Table 2, M, width, AMNH 8197, for 14 read 9.5.
Page 17, lines 9, 10. MCZ 3655, for P, to Mz and partial alveoli
for I, to canine, read M,-Mz and alveoli for P,-P4,.
Page 17, line 11, for MCZ 3666 read 3660.
Page 20, Table 3. M2 width, MCZ 7635, for 21.6 read 10.3;
AMNH 34181, for 22.4 read 10.4.
Page 22, Blastomeryx, Hypodigm. Add MCZ 7763, partial left
jaw with P,-P,, and MCZ 4349, left jaw with P;-Ms.
qh Tobe
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BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MAss. 3 FEBRUARY, 1967 NUMBER 256
ANOLIS CHOCORUM, A NEW PUNCTATUS-LIKE ANOLE
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, Darién, 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, Darién, 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, Darién, Panama, collected by a native for Charles W.
Myers, 26 July 1965.
Paratypes. Panama: Darién. KU 76027, young ¢é, 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. Duméril 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 supranasals.
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 3
SS
SOQ5c
See RES
S = IS
Fig. 1. Anolis chocorum, 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 ventrals. 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.
4 BREVIORA No. 256
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7,
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Fig. 2. Anolis chocorum, new species. Side view of head of type,
KU 96934. Actual snout-vent length 79 mm.
Size (snout/vent length): 4 (Type) 79 mm; longest @ 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 Darién localities from which A. chocorum 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 chocorum 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 Darién; these include two large arboreal
species (A. biporcatus and A. frenatus), the moderate-sized arbo-
real A. capito, the smaller bush-dwelling A. chloris, limifrons,
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 Darién 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 Darién with A. chocorum, is
likewise a South American autochton, well known in Chocoan,
Colombia (there known under the names A. palmeri and A. gor-
gonae), and in trans-Andean Ecuador. The distribution of 4.
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 Darién 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 Darién, and in the Colombian Choco.
Of the endemic South American group, Anolis chocorum is very
clearly most similar to Anolis punctatus Daudin of Brazil, the
6 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 Darién-Choc6 form thus far known is
N. end Serraniae
de Pirré
5S
<
=~
Q
P ry
e
Cerro Cituro Tacarcuna
Village
Quebrada fla UTE
20 30 40
Kilometers
Fig. 3. Map of the distribution of Anolis chocorum, 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. punctatus 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 | 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.
8 BREVIORA
TABLE 1
Scale Counts
Anolis chocorum and Relatives
No. 256
punctatus
(including
boulengeri) chocorum transversalis
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
punctatus
(including
boulengert) chocorum transversalis
Snout Swollen in 2 Not swollen in Not swollen in
either sex either sex
Ventrals Smooth (punc- Smooth Smooth
tatus) or keeled
(boulengeri)
Mental At least as deep Wider than deep Deeper than wide
as wide
Sublabials | = + (large)
Dewlap (4 ) 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 (2 ) Absent Large as in male Large as in male
but differently but differently
colored colored
Dorsal pattern Not sexually Not sexually Strong sexual
dichromatic dichromatic
dichromatism
1967 NEW ANOLIS FROM PANAMA 9
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
As in 6, but center of belly not
light in any specimen
As in 4
As in 4, 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 punctatus group. Breviora, No. 233:1-15.
(Received 5 April, 1966.)
No: 256
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Museum of Comparative Zoology
CAMBRIDGE, MaASss. 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 ef 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 MacIntyre,
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-
cerning 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.
CLARK FORK VERTEBRATE FAUNA
1967
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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” (bid.,
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 Champsosaurus 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 Clark Fork and Sand Couiee 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.
No. 257
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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 McCulloch 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 paleontological 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 McCulloch 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 McCulloch Peak] described above and although
fossils were scarce some SO individuals were obtained. The small
phenacodonts constituted about 12 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 McCulloch 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
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12 BREVIORA INO 257
Footnotes for Table I (continued)
Denison (1938, p. 175) states: “Amer. Mus. No. 16068 from the Clark
Fork horizon, described by Matthew .. . as ‘?Oxyaena 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.
3 AMNH No. 18668 has been tentatively referred to this genus, but specific
identification is difficult because of the scrappy nature of the specimen.
4Simpson (1937) recognizes three, or possibly four, species of Esthonyx
from the Clark Fork beds: E. bisulcatus, E. grangeri, E. latidens, and per-
haps E. spatularius. AMNH No. 16065 is labelled as “E. cf. spatularius
(type),” but is figured in Simpson (1937, p. 6, fig. 1) as E. ?bisulcatus. 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. spatularius being AMNH No. 4809 and that
for E. bisulcatus being USNM No. 1103 (Gazin, 1953, pp. 17, 21). None
of the specimens in the collection are presently labelled as representing E.
bisulcatus.
>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. priscum 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 Palae-
oryctes punctatus by Van Valen (1966, p. 56).
SMany 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. intermedius, and P. hemiconus, 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 Littlke 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’Hara 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 £5
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,
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1967 CLARK FORK VERTEBRATE FAUNA Ly
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 Phenacodus
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-
maevus 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 Probathyopsis 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 Probathyopsis
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 different 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.
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1967 CLARK FORK VERTEBRATE FAUNA Das
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-
nensis 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, 3257
reason for referring this specimen to a Paleocene horizon is that it
was found in rocks mapped, presumably on non-paleontological
grounds, as being of Paleocene age. Whether C. proterus is of
Paleocene or Eocene age cannot be determined on the basis of
available evidence.
Simuarly, 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 Paleocene
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 Little 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 Paleocene 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 Paleocene 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 P,4, M:2; PU No. 13311 is an M,.)
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. 1
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 2S
sp. is more similar to P. praecursor 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. 0. 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) sthat
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 P,
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 P, of P. dubius.” 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
collection belongs to a late Paleocene or an early Eocene fauna.
A wide ranging stratigraphic distribution or a lack of adequate
knowiedge regarding phylogenetic 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 INO257
matthewi) may be unique to what has been described as the Clark
Fork fauna. Carpolestes dubius 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 praecursor 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.
Core, ED:
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, IlI-VI, pp. 1-72, 1 map.
Gazin, C. L.
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1967 CLARK FORK VERTEBRATE FAUNA 29
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| plate.
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MaASss. 3 FEBRUARY, 1967 NUMBER 258
BIOLOGY OF THE PARTHENOGENETIC FUNGUS BEETLE
CIS FUSCIPES MELLIE (COLEOPTERA: CIIDAE)
By John 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 beeties in the family Ciidae,
I discovered an apparent case of thelytoky in the species Cis
fuscipes Mellié, 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 fuscipes Mellié
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. fuscipes. Females of the two species are apparently
indistinguishable (Casey, 1898; Dury, 1917).
nN
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 F. larvae in the dishes. By July 19,
adult F. 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 Fs generation by October 24, and one of these was
retained to produce F,’s by the end of the year.
Because of intermittent field work, the exact generation time
could not be recorded, but the F, 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 Veshima
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 100th 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 usuaily cause white rot in dead or dying angio-
sperms. It is especially common in Polyporus versicolor, which
may be 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, 7 from P. hirsutus, 6 from Lenzites betulina, 2 from P. con-
chifer, and a single record each from P. squamosus, Trametes
suaveolens, Fomes fraxineus, 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 californicum Casey, and
Octotemnus laevis Casey. In California, extensive coliecting 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 (Oliviér). 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. pargamenus, 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 fuscipes 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 CF 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
PARTHENOGENETIC FUNGUS BEETLE
1967
‘SOLOS OY} UI B[VU OUO svoT 3v JO DOUaSOId OY) O}LdIPUT S1v]s Spop1OdI1 Ud9q DALY STB
-o} A[Uo YoIyM WoT sarRdo] JUdsaidar sapIIIO Youtg “sPEW Sadiosn{f s19 Jo uoNNgLYSIP POM MON oy} Jo dey
aun!
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 (81.0%) 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 subparailel 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; dua! and subseriate elytrai 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 simpie aedeagus. The species group extends from eastern
1967 PARTHENOGENETIC FUNGUS BEETLE |
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. fuscipes occur mainly in Siberia
and Japan. These include Cis seriatopilosus Motschulsky (Siberia:
Amur, Sakhalin; Japan); Cis seriatulus 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 thelytoky in
animais have been demonstrated to be glacial phenomena; this is
discussed by Suomalainen (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
rapidiy 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: 25's
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 fuscipes 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 Doli-
chocis indistinctus Hatch extend southward only in montane
regions and may occur in the southern Appalachians or in the
mountains of Arizona. Sulcacis curtulus (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. fuscipes, and both are absent from
the Southern Coastal Plain and Gulf Coast. Two other wide-
ranging species, Ennearthron californicum Casey and Ennearthron
thoracicorne (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. fuscipes 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. fuscipes belongs to a group restricted to
the eastern Palaearctic and Oriental regions and having some
relationships with palaeotropical forms. It may be that C. fuscipes
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. fuscipes 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 comptetely
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.
Mellié’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 Mellié’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
Commonweaith Institute of Entomology. Aithough 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 coilection 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 established 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. Mellié
(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
Mellié in the Salié and Pic collections at the Muséum 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 preter
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 Mellié
Cis fuscipes Mellié, 1848: 271, pl. 2, fig. 23; Wollaston, 1854: 281; Wollaston,
1865: 234; Cas2y, 1898: 78; Blatchley, 1910: 898; Dalla Torre, 1911: 11;
Dury, 1917: 11; Leng, 1920: 246; Weiss, 1920: 138 (host); Weiss and
West, 1920: 8 (host); Weiss and West, 1921b: 169 (host); B6ving and
Craighead, 1931: 270-71, pl. 92, figs. K-R (larva); Arnett, 1962: 827,
fig. 1.98; Hatch, 1962: 231; Lawrence, 1965: 279. Type locality: “Bos-
ton.” Lectotype, female, Pic collection, Mus. Natl. Hist. Nat., Paris.
Cis atripennis Mellié, 1848: 258, pl. 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 chevrolatii Mellié, 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 dubius Mellié, 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: 3.) Dury, 1917 11 keng:
1920: 246. Type locality: “North Carolina (Asheville).” Holotype,
female, Casey collection, U.S. Nat. Mus., Washington, D.C.
Cis impressa Casey, 1898: 79; Pallawome slo = 12. Dany oly. 1; 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 pallens Casey, 1898: 78; Dalla Torre, 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 Xj, 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. Limnsley, 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 Naturai 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, IIl.; Cin-
cinnati Museum of Natural History, Cincinnati, Ohio; Cornell
University, Ithaca, N. Y.; Hlinois Natural History Survey, Urbana,
Ill.; Museum of Comparative Zoology, Cambridge, Mass.; Muséum
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.
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1962. The beetles of the United States (A manual for identification).
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BAKER, H. G.
1955. Self-compatibility and establishment after “long-distance” dis-
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BLATCHLEY, W. S.
1910. An illustrated descriptive catalogue of the Coleoptera or beetles
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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
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86-125.
(GASEY- alles
1898. Studies in the Ptinidae, Cioidae, and Sphindidae of America.
New York Ent. Soc., 6(2): 61-93.
CHUJO, M.
1939. Onthe Japanese Ciidae (Coleoptera). Mushi, 12(1): 1-10.
DALLA Torre, K. W. VON
1911. Cioidae. Pars 30. Jn W. Junk and S. Schenkling, edit., Coleop-
terorum Catalogus. Junk, Berlin. 32 pp.
DarEwWSKI, I. S. and W. N. KULIKOWA
1961. Natiirliche Parthenogenese in der polymorphen Gruppe der
kaukasischen Felseidechse (Lacerta saxicola Eversmann). Zool.
Jahrbuch., 89(1): 119-176, 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-
terms Je Ecole sii: ail-23:
Fukupa, A.
1940. A study of the Japanese Ciidae, Coleoptera (II). [In Japanese}
Trans. Nat. Hist. Soc. Formosa, 30: 48-53, 5 figs.
Hatcu, M. H.
1962. The beetles of the Pacific Northwest. Part III: Pselaphidae and
Diversicornia I. Univ. Washington Press, Seattle. ix + 503 pp.,
66 pls.
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.
Louse, 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.
MELLIE, J.
1848. Monographie de l’ancien genre Cis des auteurs. Ann. Soc. Ent.
France, Ser. 2, 6: 205-274,313-396, pls. 9-12.
MIYATAKE, M.
1954. Studies on the Japanese Ciidae, I (Coleoptera). Sci. Rep.
Matsuyama Agr. Coll., 14: 40-67, 11 pls.
PAVIOUR-SMITH, KITTY
1960. The fruiting-bodies of macrofungi as habitats for beetles of the
family Ciidae (Coleoptera). Oikos, 11(1): 43-71, 1 pl.
REITTER, E.
1902. Analytische Uebersicht der palaearctischen Gattungen und
Arten der Coleopteren-Familien: Byrrhidae (Anobiidae) und
Cioidae. Verh. 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 wiih Polyporus 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 pl.
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 pl.
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 +
634 pp., 13 pls.
1865. Coleoptera Atlantidum, being an enumeration of the coleopter-
ous insects of the Madeiras, Salvages, and Canaries. Van
Voorst, London. xlvii + 526 pp., appendix, 140 pp.
(Received 5 July, 1966).
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MaAss. 3 Fepruary, 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
tN
BREVIORA No. 259
I wish to express my indebtedness to Dr. C. Clayton Hoff 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. 111) 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 sclerites 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 A gelenopsis
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 (7), an inappropri-
ately named ring of heavily sclerotized tissue connected to the
subtegulum by the middle hematodocha (mh). 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 (1) 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 (/sa). The terminal apophysis (fa) is located at the
very tip of the organ.
OBSERVATIONS
Order ARANEAE
Family THERIDITDAE
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)
Pigss 12S
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 (7), 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. 3).
Family LINYPHITDAE
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 alteranda Chamberlin
Figs, 10514
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 (/sa) 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 (ft), 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 (/sa) is more satisfactorily observed.
1967 PALPI OF MALE SPIDERS 7
Linyphia marginata C. L. Koch
Pips. 23
In this species, studied by Blauvelt (1936), the lateral sub-
terminal apophysis (/sa), 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 (7) 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 (/sa) 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 Merreit’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 (¢), and embolus (e). The shape
of the tibia (f7) 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 formosa (Banks)
Figs l6;pal7
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.
Family 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 20521
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 (ft), terminal apophysis (fa), 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 Dr
the hook-like lateral subterminal apophysis (/sa) 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.
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 (bh) appears as the most basal part. The tegulum
(t), embolus (e), and well-developed conductor (c) are all read-
ily visible. The median apophysis (ma) 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 Ivie (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 (ft) 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 (ft) and the median apophysis.
Singa pratensis Emerton
Figs; 26,27
The members of the genus Singa are structurally close to the
theridiids (Kaston, 1948), but the structure of the palpus is
closer to Araneus 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 (¢), median apophysis
(ma), and terminal apophysis (7a) 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 compiex 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 (ft) 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, 30593), 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 (sf).
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.
A gelenopsis 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 (ft) 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: 355/36
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 (¢), 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 (bf) 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 (ti) 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 (ft) 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 Jenta 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 agrestis 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 (¢), and embolus (e) are subequal.
The expanded right palpus (Fig. 40) shows the ectad rotation of
the embolus (e¢), exposing its fingerlike tip.
Aysha 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 (ft) 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 (sf) is now visible. The track of the internal sperm
duct through the tegulum is prominent.
Family SALTICIDAE
The Saiticidae 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 (7), which
is enlarged and greatly extended proximally into a hollow on the
1967 PALPI OF MALE SPIDERS 15
ventral surface of the palpal tibia (ti), and the fused embolus-
conductor (e, c). In the ectodorsal view of an expanded paip
(Fig. 44), the basal hematodocha (bh) 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 (st) 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 (t) 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) 51
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 (ft) 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 compiex palpi, and those with simple
palpi.
Drapetisca 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 Singa, 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 frondeum, Steatoda
borealis, Argiope trifasciata, Araniella displicata, Wadotes calcara-
tus, Agelenopsis utahana, Coras lamellosus, Cicurina 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 usualiy visible be-
fore expansion.
The third group, species with simple male palpi, includes:
Soulgas corticarius, Ceratinopsidis formosa, 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.
Ui tees Mikes il 7/e
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.
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BisHoPp, S. C., and C. R. Crossy.
1930. Studies in American spiders: genera Ceratinopsis, Ceratinop-
sidis, and Tutaibo. J. New York Entomol. Soc. 38: 15-34.
BLAUVELT, H. H.
1936. The comparative morphology of the secondary sexual organs
of Linyphia. 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 Cicurina. Bull. Univ. Utah,
Biol. Ser. 5(9): 1-108.
1942. A hundred new species of American spiders. Bull. Univ. Utah,
Biol; Ser. -7@1) 2 1-187.
1943. New genera and species of North American linyphiid spiders.
Bull. Univ. Utah, Biol. Ser. 7(6): 1-39.
ComsTOock, J. H.
1940. The spider book, revised and edited by W. J. Gertsch. Com-
stock Publishing Company, New York. 721 pp.
Crospy. C. R., and S. C. BISHOP
1936. Studies in American spiders: miscellaneous genera of Erigo-
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Epwarps, 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, VY.
1910. Beitrage zur Kenntnis der weiblichen Copulationsorgane eini-
ger Spinnen. Z. Wiss. Zool. 96: 32-117.
EXLINE, H.
1936. Nearctic spiders of the genus Cicurina Menge. Amer. Mus.
Novit. No. 850: 1-26.
GERING, R. L.
1953. Structure and function of the genitalia in some American
agelenid spiders. Smithson. Misc. Collec. 121(4): 1-84.
GertTscH, W. J.
1941. New American spiders of the family Clubionidae. II. 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.
KasTOoN, B. J.
1948. Spiders of Connecticut. Connecticut State Geol. Natur. Hist.
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KasTON, B. J.
1952. How to know the spiders. Wm. C. Brown Company, Dubuque,
Iowa. 220 pp.
Levi, H. W.
1957a. The spider genera Crustulina and Steatoda in North America,
Central America, and the West Indies (Araneae, Theridiidae).
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1957b. The spider genera Enoplognatha, Theridion, and Paidisca 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.
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1946. North American Agelenidae of the genus Coras Simon. Amer.
Mus. Novit. No. 1329: 1-20.
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1964. The spider family Uloboridae in North America north of
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1925. External reproductive organs of the common grass spider,
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1928. The biology of spiders. Sidgwick and Jackson, London. 376 pp.
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1967
PALPI OF MALE SPIDERS
ABBREVIATIONS USED IN FIGURES
basal hematodocha
conductor
cymbium
distal hematodocha
embolus
lateral subterminal apophysis
median apophysis
middle hematodocha
paracymbium
radix
stipes
subtegulum
tegulum
terminal apophysis
tibia
i
i)
BREVIORA No. 259
Fics. 1-3. Steatoda borealis. 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. Fics. 4, 5. Enoplognatha tecta. 4, Ectal view of an
unexpanded left palpus. 5, Ectal view of an expanded left palpus. Fics.
6, 7. Theridion differens. 6, Ventral view of an unexpanded right palpus.
7, Ventral view of an expanded right palpus. Fics. 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 23
Fics. 10, 11. Drapetisca alteranda. 10, Mesal view of an unexpanded
right palpus. 11, Mesal view of an expanded right palpus. Fics. 12, 13.
Linyphia marginata. 12, Ectal view of an unexpanded right palpus. 13,
Ectal view of an expanded right palpus. Fics. 14, 15. Soulgas corticarius.
14, Ectal view of an unexpanded right palpus. 15, Mesal view of an ex-
panded right palpus. Fics. 16, 17. Ceratinopsidis formosa. 16, Ectal view
of an unexpanded right palpus. 17, Dorsoectal view of an expanded right
palpus.
24 BREVIORA No. 259
Fics. 18-21. Araneus sericatus. 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.
Fics. 22, 23. Argiope trifasciata. 22, Mesal view of an unexpanded right
palpus. 23, Dorsomesal view of an expanded right palpus. Fics. 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 DS
SB)
Fics. 26, 27. Singa pratensis. 26, Mesal view of an unexpanded right
palpus. 27, Ectal view of an expanded right palpus. Fics. 28, 29. Mangora
gibberosa. 28, Ectal view of an unexpanded right palpus. 29, Ventroectal
view of an expanded right palpus. Fics. 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. Fics. 33, 34.
Agelenopsis utahana. 33, Ectal view of an unexpanded right palpus. 34,
Mesal view of an expanded right palpus.
26 BREVIORA No. 259
Fics. 35, 36. Tegenaria domestica. 35, Ventral view of an unexpanded
right palpus. 36, Ectoventral view of an expanded right palpus. Fics. 37.
38. Cicurina robusta. 37, Ectal view of an unexpanded right palpus. 38,
Ectal view of an expanded right palpus. Fics. 39, 40. Clubiona bryantae.
39, Ectal view of an unexpanded right palpus. 40, Ectal view of an ex-
panded right palpus. Fics. 41, 42. Aysha gracilis. 41, Ventral view of an
expanded right palpus. 42, Mesal view of an unexpanded right palpus. Fic.
43. Metacyrba undata, ventral view of an unexpanded right palpus.
1967 PALPI OF MALE SPIDERS 27
Fic. 44. Metacyrba undata, ectodorsal view of an expanded right palpus.
Fics. 45, 46. Tutelina elegans. 45, Ventral view of an unexpanded right
palpus. 46, Ectodorsal view of an expanded right palpus. Fics. 47-49.
Hyptiotes cavatus. 47, Ectal view of an unexpanded right palpus. 48, Mesal
view of an unexpanded right palpus. 49, Ectoventral view of an expanded
right palpus. Fics. 50, 51. Dictyna sublata. 50, Mesal view of an unex-
panded right palpus. 51, Ventral view of an expanded right palpus.
a ‘ ve ¥ :
arti f ; ade te) by,
Any 4 J
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. 3 Fesruary, 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 Spiroceramus. This species was S. amplus
(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 Camagiiey 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
Spiroceramus Pilsbry and Vanatta, 1898, Proc. Acad. Nat. Sci. Philadel-
phia, 50: 281 (type species, Macroceramus amplus 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. amplus, 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 hay-
ing the oblique sculpture of fine striae similar on all whorls. The
suture with or without crenulations.
SPIROCERAMUS (SPIROCERAMUS) AMPLUS (Pfeiffer)
Plate 1, figures 6, 7
Macroceramus amplus Gundlach, 1856, [in] F. Poey, Memorias Sobre
la Historia Natural de la Isla de Cuba, Habana 2: 8, no. 378 (Cuba).
[Nomen nudum. |
Macroceramus 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, pl. 89, figs. 12-14; Arango, 1878, Contribucion a la
Fauna Malacologica Cubana, Habana, p. 84 (Guisa and San Andres,
Bayamo, Cuba). [Syntypes, MCZ 39318.]
Microceramus (Spiroceramus) amplus Pfeiffer. Pilsbry, 1904, Man. of
Conch: (2) 16: 172," pl: 26, figs. 24°" 29:
Measurements
Height Width
mm mm
10.5 3:9 Syntype. Guisa, Bayamo, Cuba
it 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. Whoris 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 5
angle of about 35°. Aperture subcircular, with the outer lip
slightly refiected. 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 112
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 amplus barbouri Aguayo, 1935, Mem. Soc. Cubana Hist.
Nat., 9: 126, pl. 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
55) 2: Holotype
4.5 1.8 Paratype
4.75 2, Paratype
DES D 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 1%2 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 amplus
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 Tuabagiiey, 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
Sail 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: §. 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,
Camagiiey 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
Tuabagiey, near the Cueva del Indio (MCZ 188844), all Sierra
de Cubitas, Camagiiey Province, Cuba.
Measurements !
Height Width
mm mm
9.4 4.6 Holotype
10 4.5 Paratype
8.6 4.5 Paratype
9 4.2 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
1 All specimens measured are from the type locality.
6 BREVIORA No. 260
whorls finely lirate. 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) robertsoni Clench, 1963, Bull. Mus.
Comp. Zool. 128: 406, pl. 3, fig. 1 (Delectable Bay Settlement, Acklins
Island, Bahama Islands). [Holotype, MCZ 225313.]
Measurements
Height Width
mm mm
6.7 120 Holotype
Description: Shell reaching 6.7 mm in height, thin, imperforate,
dull and sculptured. Color probably gray, whorls 11 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
little smaller.
SPIROCERAMUS (INSULACERAMUS) GREENWAYI (Clench)
Plate 1, figure 1
Microceramus (Spiroceramus) greenwayi Clench, 1938, Mem. Soc. Cu-
bana Hist. Nat., 12: 329, pl. 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, Camagiiey, Cuba (5.3 X). Holotype, MCZ 188845.
Fig. 4. Spiroceramus (Spiroceramus) vanattai n. sp., Los Cangilones, Sierra
de Cubitas, Camagiiey, 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 237877.
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 9
10 BREVIORA No. 260
Plate 2
Map showing distribution of the species. 1. Spiroceramus (Spiroceramus)
amplus (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, Camagitiey, Cuba.
4. Spiroceramus (Spiroceramus) vanattai n. sp., Los Cangilones, Sierra de
Cubitas, Camagiiey, Cuba. 5. Spiroceramus (Insulaceramus) robertsoni
(Clench), Delectable Bay, Acklins Island, Bahama Islands. 6. Spiroceramus
(Insulaceramus) greenwayi (Clench), Fox Town, Little Abaco Island,
Bahama Islands.
me
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, MASss. 31 Marcu, 1967 NUMBER 261
THE MONTICOLA GROUP OF THE LIZARD GENUS
ANOLIS 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 Ile 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
Laferri¢re 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. christophei. 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 3
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 CHRISTOPHE! Williams
The apparently most primitive member of the christophet-
etheridgei-monticola trio of Hispaniolan anoles, A. christophei, was
described from two female specimens from the Citadelle Laferriere,
Dépt. 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-l’Eveque 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 5
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
SZ LE
—~ Df Giee yA
ZG QZ a d CATY
Fae ae
We : = — = > x
EY IAS KS OSS GSS
ZEEE OCLs ae
= = SS = S
eee
Figure 2. Lateral view of head of Anolis christophei, ASFS X9193.
Snout-vent length 45 mm.
contrast to that of such larger anoles as A. cybotes, A. distichus,
and 4. 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 Cordiliera 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 (Pl. 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 effect 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 Th
length; ASFS X8835) than females (45 mm snout-vent length;
ASFS V1957). 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 Anolis christophei, ASFS X9193.
Snout-vent length 45 mm.
Specimens examined: HAITI, Dépt. du Nord, Citadelle Lafer-
riere, 20 (MCZ 66900-19); Dépt. de l’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, V1866-67); 4 km SW El Rio, 4000 feet
(1220 m), 5 (ASFS X8552-55, X8737); 4 km SW EI Rio, 3500
feet (1068 m), 3 (ASFS X8886-88); 6 km E El Rio, 3600 feet
(1098 mi); 7 CASES -X9205-11); 7.1 mi: (1-4 kmoee, El Rio;
3500 feet (1068 mm) CASES X8113))223- km, E. El Rio, 3050
feet (930 m), 3 (ASFS X9193-95); Espaillat Province, 2 km N
Puesto Grande, 2200 feet (671 m), 6 (ASFS V1956-61); San
Cristébal 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).
s BREVIORA No. 261
ANOLIS ETHERIDGE! Williams
Anolis etheridgei Williams (= Anolis darlingtoni Cochran,
1939) was described from four localities in the Cordillera Central
in the Republica 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
= 5
ie”.
yA Me Ee
ws — SIE oT k
= SARE
SSS SSE
mecaccor
Figure 4. Dorsal view of head of Anolis etheridgei, ASFS X9146. Snout-
vent length 42 mm.
1967 ANOLIS MONTICOLA GROUP 9
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
facility 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.
<>
>=
SSS
=
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 (Pl. 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 i
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 O (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 km SW EI Rio, 3500 feet (1068 m), 3 (ASFS X8889-91 );
6 km E El Rio, 3600 feet (1098 m), 1 (ASFS X9212); 23 km
EEE Rio; 3050 feet (930) m), 14 (ASES 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 Bayjito, 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 Géodésie et Cartog-
raphie 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.”
ft 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 Jérémie 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, 115 miles south of Beaumont on the
Les Cayes-Jérémie road, and with AMNH 49845 from 32 miles
from Les Cayes on the Jérémie 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 Sphaerodactylus copei Steindachner was noted as being indis-
tinguishable from S. copei picturatus Garman 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 Laferricre 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 monticola quadrisartus, 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 IT and II 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
Anolis, as follows: upper 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., Reptblica Dominicana, painted from living
spec:'men; center left, Anolis monticola monticola, ASFS V9624, ca. 5 km
SSE Marché Léon, Dépt. du Sud, Haiti, painted from Thomas’ field notes
and color transparencies of living specimens; center right, Anolis monticola
quadrisartus, MCZ 63004, paratype, Tombeau Cheval, Dépt. du Sud, Haiti,
painted from color noted by collectors, as quoted by Williams (1962:3-4);
lower, Anolis rimarum, AMNH 96469, paratype, 8 to 9 km W Marmelade,
Dept. de l’Artibonite, Haiti, 3500 feet, painted from Thomas’ field notes
on living specimens.
1967 ANOLIS MONTICOLA GROUP 15
ANOLIS MONTICOLA MONTICOLA Shreve
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.
Sens
:
I
()
'
Ka
nan
fi
0,
CH
ith
Ve
f
\)
te
i
’
}
i
My
()
;
t
SSeS so.
4s (ee = TRS
SS S= 2S Se SS OHS
SS SSIES
— <= Ss, Soe,
SS OSES SSS
<a SSS SQSS5 _, CIS TIO
SSS SRF ESD
SSeS, SSS
CO —_
SX Se
=<
Figure 8. Lateral view of head of Anolis monticola quadrisartus, 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
aste! biue to blue-green stripe. The saddies 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 (Pl. 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
Marché Léon, 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 Marché Léon, 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, Dépt. du Sud: ASFS V9196-211,
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 Marché Léon, 2600 feet (793 m); MCZ 74866-
70, Trou Bois, about 1% km S of Beaumont, on Jérémie 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 Jérémie road 32 miles from Les
Cayes, 2000-3000 feet (610-915 m) approx.; MCZ 65026, “Grande
Cayemite.”
Ss SSS [ae
Ss sasS° Boas
( == Bees seeses B= soos
SS A oe
x Sew Sema ase sese
SSS SSS RG
SSS wee RE Smee SSS
SS SS SS SSS EES
WSS SSE ae
=
SS
Figure 9. Mental region of head of Anolis monticola quadrisartus, 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, Dépt. 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.”
18 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.
GRANDE
CAYEMITE
JEREMIE
¢ eee ae
Go
BEAUMONT
; Le Moises
eg
PIC MACAYA’ Yj: TOMBE AU
A CHEVAL
Z Yj); Vj CAMP QO
SS ae Ri
—
LES CAYES
Figure 10. Tip of the Tiburon 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. quadrisartus. Line
connecting Les Cayes and Jérémie 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, Dépt. de l’Artibonite, 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 81129, 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
Cl6=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 pl. 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.
PORE
Rt TAN geen
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
Secas
eee:
sien
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 rimarum, 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. Rimarum 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 lizards 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
dewiap 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
CocuRaN, 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 pls.
SCHWARTZ, ALBERT, and RICHARD THOMAS
1964. Subspeciation in Sphaerodactylus 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 Williams. 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 christophei
ite
i)
Snout most elongate, most con-
cave.
Canthus
edged.
pronouncedly — sharp-
Scales 6-10 (mode 8) across
snout at level of second canthal.
Loreal rows 4-8 (mode 6).
Scales in frontal depression nu-
merous, polygonal to rounded,
smooth or slightly rugose.
Supraoculars moderate to small
in size, somewhat elongate,
weakly keeled, ca. 9-11 in num-
ber.
Supraorbital semicircles keeled,
separated by 1-2 scales (Fig.
ie
Lower eyelid covered with gran-
ules, without “window” of
squarish scales.
Anterior canthal scales abruptly
smaller.
. Ventrals rounded to squarish,
smooth, slightly imbricate and
arranged in transverse rows.
Dewlap very large, extending
from mid-gular region to mid-
venter; scales at edge of dewlap
smooth, imbricate, larger than
ventrals.
Anterior femoral scales keeled,
imbricate, not tranversely en-
larged.
Interparietal small, ca. 2 or
less size of ear opening.
Scales between interparietal and
supraorbital semicircles 3-7.
ANOLIS MONTICOLA GROUP 25
Anolis etheridgei
ile
tN
10.
Hil.
14.
Snout bluntest, not pronounc-
edly concave.
Canthus distinct but not prom-
inently sharp-edged.
Scales 8-15 (mode 10) across
snout at level of second can-
thal.
Loreal rows 5-10 (mode 6).
Scales in frontal depression nu-
merous, polygonal, keeled.
Supraoculars small to moder-
ate, slightly elongate, keeled, ca.
11 in number.
Supraorbital semicircles keeled,
separated by 0-4 scales (Fig.
4).
Lower eyelid covered with gran-
ules, without “window” of
squarish scales.
Anterior canthal scales not ab-
ruptly smaller.,
Ventrals rounded to squarish,
subimbricate, smooth and in
transverse rows.
Dewlap, small, extending from
mid-gular region onto chest,
scales keeled.
Anterior femoral scales keeled,
imbricate, only slightly enlarged
distally.
Interparietal small, less than 2
size of ear opening.
Scales between interparietal and
supraorbital semicircles 3-6.
26
BREVIORA
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-
thal.
4. Loreal rows 6-9 (mode 7).
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.
Y2 size of ear opening but may
be equal to ear opening in size.
14. Scales between interparietal and
supraorbital semicircles 3-6.
Mablen tl:
Hispaniolan anoles.
No. 261
Anolis rimarum
If
1s
14.
Snout elongate, somewhat con-
cave.
Canthus sharp-edged.
Scales 7-10 (mode 8) across
snout at level of second can-
thal.
Loreal rows 4-6 (mode 5).
Scales in frontal depression rel-
tively few, polygonal, pavimen-
tous.
Supraoculars large, not elon-
gate, smooth or only faintly
keeled, ca. 5-7 in number.
Supraorbital semicircles smooth
or feebly keeled, in contact or
separated by one scale only
(Bios Wilh):
Lower eyelid with “window” of
enlarged squarish scales.
Anterior canthal scales abruptly
smaller.
Ventrals squarish, juxtaposed to
subimbricate, smooth and in
transverse rows.
Dewlap very small, occupying
only the region between the
angle of the jaw and the chest,
scales smooth, imbricate.
Anterior femoral scales trans-
versely enlarged, feebly multi-
carinate.
Interparietal large, about size
of ear opening.
Scales between interparietal and
supraorbital semicircles 2-3.
Comparison of the four species of the monticola group of
27
ANOLIS MONTICOLA GROUP
1967
‘uosexoy
»
ov of oz tO
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‘py ismopjoy se ‘dnosd yjoouuow syoup ay) Jo saioeds anoy oy} 10F spsodse1 AyPEdo] SuIMoys ‘gjotuedsip, ‘pl ons
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. 31 Marcu, 1967 NUMBER 262
A PHYLOGENETIC SURVEY OF MOLLUSCAN SHELL
MATRIX PROTEINS
By
Michael T. Ghiselin
Systematics-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.
D 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 al., 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-G18727.
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). “ihe sinner
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
(Boggild, 1930) and protein configurations (Grégoire, 1961) are
quite intricate, suggesting a highly organized system. In mollusks,
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 effective 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
1% at the 10° molar level for identical runs. The overall varia-
tion of two identical samples from the same collection site is less
than 3%; 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%
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 different 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 HI 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 differences 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 ef al., 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; simultaneous!y, 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 mollusks 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
(Chaetopleura), 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, Spirula 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, Spirula, 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 difference 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 fal
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 al., 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 Ptenoglossa 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. A ply-
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
Umbraculum (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. Douvilleé’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.
16 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 Hi 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
a
1967 MOLLUSCAN SHELL PROTEINS i07/
the primitive state; the freshwater forms change to a greater de-
gree 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
phenolics. 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. 45% 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 moliusks 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; Chaeto-
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 90% 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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BaBA, K.
1961. On the identification and the affinity of Tamanovalva limax,
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CaIn, A. J. (ed.)
1959. Function and taxonomic importance. Systematics Association,
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1961. Structure of the conchiolin cases of the prisms in Mytilus edulis
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1962. The amino acid composition of the organic matrix of some
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22 BREVIORA No. 262
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1967 MOLLUSCAN SHELL PROTEINS 23
RUDALL, K. M.
1963. The chitin/protein complexes of insect cuticles. Advances
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(Received 13 September, 1966.)
BREVIORA
1967 MOLLUSCAN SHELL PROTEINS Fe)
PLATE |
GASTROPODA
Figure 1. Achatinella lorata Pfeiffer, 1848, Oahu, Hawaiian Islands,
Pease collection, MCZ (Mollusk Department) — aperture view. X 1.
Figure 2. Acmaea pustulata (Helbling, 1799), Puerto Sosya, Dominican
Republic, MCZ, coll. 1937 — top view. X 2/3.
Figure 3. Akera soluta (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 regium (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 (Linné, 1767), Hadley Harbor, Elizabeth
Islands, Massachusetts, Sta. P-2211-65, 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; 10B, exterior view. X DY Bs
Figure 11. Colus 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 (Linné, 1758), Miami, Florida, MCZ (uncat.)
— aperture view. X 1/3.
Figure 13. Dolabella callosa (Lamarck, 1801) = 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).
X23:
26 BREVIORA No. 262
PLATE | (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. Janthina janthina (Linné, 1767), Cape Florida Key, Biscayne,
Florida, MCZ No. 155173 — aperture view. X 5/6.
Figure 19, Littorina littorea (Linné, 1758), Hadley Harbor, Elizabeth-
Islands, Massachusetts, Sta. P-0610-65, Systematics-Ecology Program, Ma-
rine Laboratory — aperture view. X 1.
Figure 20. Lunatia 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. Nassarius trivittatus (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 duplicatus (Say, 1822), Provincetown, Massachu-
setts, MCZ (uncat.) aperture view. X 1 1/3.
Figure 26. Siphonaria alternata (Say, 1826), Bermuda, coll. Bryant,
1903, MCZ No. 24213 — top view (exterior). X 1 1/3.
Figure 27. Succinea ovalis (Say, 1817), bank of Grand River, W. Bridge
Street Ferry, Ottawa County, Michigan, MCZ No. 166697 — aperture view.
XG.
Figure 28. Turritella terebra (Lamarck, 1822), Manila Bay, near
Cavite, Luzon, Philippine Islands, MCZ No. 138070—aperture view. X 1/3.
Figure 29. Umbraculum indicum (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-0612-65, Systematics-Ecology Program,
Marine Biological Laboratory — aperture view. X 4 2/3.
1967 MOLLUSCAN SHELL PROTEINS 749
PLATE | (Continued)
Figure 31. Viviparus georgianus (I. Lea, 1834), Lake Woodruff near
Fick Island, Volusa County, Florida, MCZ No. 186800 — aperture view.
Ke2 3.
AMPHINEURA
Figure 32. Chaetopleura apiculata (Say, 1830), Hadley Harbor, Eliza-
beth Islands, Sta. P-7-63, Systematics-Ecology Program, Marine Biological
Laboratory — top view (exterior). X 2.
No. 262
BREVIORA
28
1967 MOLLUSCAN SHELL PROTEINS 29
PLATE Il
CEPHALOPODA
Figure 33. Argonauta hians (Dillwyn, 1817), southern Tropical Atlan-
tic, MCZ (uncat.) — side view of egg case. X 1/3.
Figure 34. Nautilus pompilius (Linné, 1758), Southwest Pacific Ocean,
MCZ (uncat.) — side view. X 1/3.
Figure 35. Spirula spirula (Linné, 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 (Linné, 1758), Georges Bank, Massachu-
setts) MCZ No. 14325 — interior view, right valve. X iW/ 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-141-64, 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.
e273:
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 BREVIORA No. 262
PLATE Il (Continued)
Figure 46. Mercenaria mercenaria (Linné, 1758), Holding Tank, Supply
Department, Marine Biological Laboratory, Woods Hole, Massachusetts —
exterior view, left valve. X 2 1/3.
Figure 47. Mulinia 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 (Linné, 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,
Australia, 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 leanum (Conrad, 1831), Martha’s Vineyard, Chap-
paquidick Island, Massachusetts, MCZ No. 192934 —§interior view, left
valve. X 1/6.
Figure 53. Petricola pholadiformis (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. Pitar cordata (Schwengle, 1951), off Port Aransas, Texas, 80
m, MCZ No. 194372 — interior view, right valve. X 2/3.
Figure 55. Pitar morrhuana (Linsley, 1845), Hadley Harbor, Elizabeth
Islands, Massachusetts, Sta. P-189-64, Systematics-Ecology Program, Ma-
rine Biological Laboratory — exterior view, left valve. X 2 1/3.
Figure 56. Saxidomus nuttalli (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, Elizabeth 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
TABLE |
DISTANCE BETWEEN GENERA WHEN CONSIDERED AS POINTS
IN FACTOR SCORE SPACE
JONAXO
SNTYWSSVN
VILWNN1
VINd1d3u9
XNId1wSOuN
WNIdOLLT1
wantivHy | Sl lolol Alle
MOLLUSCAN SHELL PROTEINS
(SNOISN3WIO 8) vaOdO4LSY9
ARCHITECTONICA
ACHATINELLA
ASTRAEA
LITTORINA
UROSALPINX
CREPIDULA
SIPHONARIA
CAVOLINA
FISSURELLA
NASSARIUS
SUCCINEA
POLINICES
APLYSIA
UMB RACULUM
JANTHINA
DOLABELLA
EPITONIUM
NERITA
MELANELLA
HELISOMA
MUREX
HALIOTIS
VIVIPARUS
VINOSIYLOIN
SN139VL
SNWOGIXVS
o
yadLsOssvad
Cal
VI1LI8v
BERREEEE
BB“BBEE
LAEVICARDIUM
AEQUIPECTEN
PETRICOLA
MERCENARIA
LIMOPSIS
MALLETIA
CRASSOSTREA
SAXIDOMUS
CORBICULA
(SNOISN3WIG 6) WO0dAD3913d
PERIPLOMA
NEOTRIGONIA
2 {3
fells] __vwwanem
z(ele
:
E
2 iz (2| N3193d INDI
<
BE WOIGHWIIA3WT
Pe
e[ sua
a
VISNOAT
SISdOWI1
v1091ulad
VINITOW
. VavaVNy
TURRITELLA
AKERA
31
No. 262
BREVIORA
*sueaw OLJawyylue aULWeSOXeYy/ULa}Odd
Ob° Scr O9L 6l2 ble el oe @
Ge 08 0s v2 Le 68 82 8
92 ve S S LL if b°0 8
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L8 20l 9 2ll 69 b9l €8 bl
juawnbaquy | yuawnbazuy wndeuzsoluad (uad)
ParplLopej |] parseopeouny YFLM LLeys LLeusS} LLeUs LLEUS LLeys}] = LLe4s
*sqeuy plunz4od PULL LGOON | SNL LIAW] ELNINN | SLZOLLEH I snLLznen OBL [07
"PpalsLo_eo g SpatyLo_eoun satoads ¢ - sueaWw ILuzawoan y
(auoq
W10L OOOL Y3d SOILVY GIDV ONIWV
ZT AAV;
2 2
22 G9
6b v
BE le
ne ral
l2 L2
09 G2
1g eb
02 G2
Ol ol
Ob 6€
82 le
90L 99
o2k e0L
LOL LL
el aa
86 pol
eb Lg
9LL O€l
2133N2)
LLey4s LLey4s
eidas | epnards
86°0
LE
L€
as)
0€
él
St
bl
88
LEL
£6
86
89
09
82l
LLE4S
eunald
-07yaeYD
96°St
0S
aque
eunald
-dJaeUD
SauLWweSOXa}
uLayOdd
auluLbay
SULPLASLH
auLsAy
auLsh]-HO
auLuele|Auaud
DULSOUAL
auLonay
duLoNaLoOs]
BULUOLYZOW
SULLEA
[$Ley] aurysé9
auLue Ly
BuLoAl[9
BULLOUd
Ploy otweyn[o
auLuas
duLuoadyy
ploy otquedsy
aul LO4d-HO
1967 MOLLUSCAN SHELL PROTEINS 3)3)
TABLE 3
FACTOR SCORES. GASTROPODA.
(Shell Tissues)
] 2 3 4 5 6
7 8
Phylogenetic Positive
Loading OHPRO
Relationships | Varimax
Factors
Negative
Loading
2] SIPHONARIA
HELISOMA
PLANORBIS
ACHATINELLA
SUCCINEA
CAVOLINA
UMBRACULUM
ARCHITECTONICA
AKERA
OXYNOE
APLYSIA
BULLA
DOLABELLA
— mNNnN W OOO mM OW
Cony ony Ups" Cony ins) (=)
own rw OT ON W
oOo fF Oo Mw W PM
Somos 2 Sw &
—- ON NN — OC —
58) 2
.6 4
2 ]
al 6
4 4
-6 1
0 0
4 3
EPITONIUM
JANTHINA
TURRITELLA
MUREX
UROSALPINX
NASSARIUS
COLUS
MELANELLA
POLINICES
LUNATIA
LITTORINA
CYPRAEA
CREPIDULA PLANA
CREPIDULA FORNICATA
FISSURELLA
ACMAEA
VIVIPARUS
NERITA
ASTRAEA
HALIOTIS
SS
———
—)
Dwnn o1w nn FN —] FP S|] Wm
MD OWWnN DWH HM CO ("NH WHO MY W WO OO
ON WKN OWN DY FON OO MH W ©
OM —]| WHaanant WN? NY —| NY W OO
M~aMnDO DAO 1H oO fF fF ON
34 BREVIORA No. 262
TABLE 4
FACTOR SCORES. PELECYPODA.
(Shell Tissues)
] 2 3 4 5 6 7 8 9
Phylogenetic Positive ISOLEU
A : Loadin LEU
Relationships 9 VAL
Varimax HIS
Factors
LAEVICARDIUM
MACOMA
MULINIA
TAGELUS
CORBICULA
ARCTICA
AEQUIPECTEN
CRASSOSTREA
MYTILUS
ANADARA
LIMOPSIS
NEOTRIGONIA
PERIPLOMA
LYONSIA
NUCULA
1967
OH-Proline
Aspartic Acid
Threonine
Serine
Glutamic Acid
Proline
Glycine
Alanine
Cystine [half]
Valine
Methionine
Isoleucine
Leucine
Tyrosine
Phenylalanine
OH-Lysine
Lysine
Histidine
Arginine
Protein
Hexosamine
Nerita
Marine
66
MOLLUSCAN SHELL PROTEINS
AMINO ACID RATIOS PER 1000 TOTAL
Viviparus | Archi-
Fresh
Water
1060
tectonica
Marine
22
TABLE 5
Siphonaria
Marine
23
Helisoma|Planorbis
Fresh
Water
1]
Fresh
Water
114
Succinea
Land
77
35
Achatinella
Land
82
BREVIORA
Museum of Comparative 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, Vannuccia, 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 Vannuccia, 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 0. gen.
Type species. Vannuccia forbesii (Mayer) 1894: 236, pl. 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, pl. 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 perisare 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 whor]
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 3
PHAN AR GE 1 WAH AN wan
DURES: Wieletecsct/,
bs ere
ava
1mm
ee!
Figure 1. Vannuccia forbesii, 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.
4 BREVIORA No. 263
©:2 5mm
Fig. 2. Vannuccia 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, pl. IIT). 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.
Sunod jo sasvys eyuswdojaaap da.1y) ‘“Msaqsof PloonuUuDA *¢ dINBIA
HYDROID OF VANNUCCIA FORBESII
6 BREVIORA No. 263
Reproduction and development. All of my medusae of V. for-
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 Euphysa aurata
and Vannuccia forbesii. 1, Euphysa aurata; Wl, 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 Euphysa 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.
forbesti. 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. forbesii 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. for-
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
| I
Figure 5. Oral tentacles of young Vannuccia forbesii (1), and young
Corymorpha nutans (11).
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 forbesii, 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 of Vannuccia forbesii 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. forbesii and strongly developed in
C. nutans. Similarly, the anchoring filaments are feebly developed
1967 HYDROID OF VANNUCCIA FORBESII 9
in Hypolytus obvoluta, become more apparent in Vannuccia for-
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.
[2854
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 pls.
KRAMBP, P. L.
1933. Coelenterata, Ctenophora and Chaetognatha. The Scoresby
Sound Comm. 2nd East Greenland Exped. in 1932. Meddel.
om Gronland., 104(11): 1-20.
1949. Origin of the hydroid family Corymorphidae. Vidensk. Medd.
fra Dansk Natur. Foren., 111: 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 pls.
1910. Medusae of the world. Part I, Hydromedusae. Carnegie Inst.
Washington, Publ. 109(1): 1-230, 29 pls.
MursacH, L.
1899. Hydroids from Wood’s Holl, Mass. Hypolytus 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.
Ucnipa, TH.
1927. Studies on Japanese Hydromedusae. I, Anthomedusae. J. Imp.
Univ. Tokyo, Sect. 4, Zool., 1: 145-241, 47 figs., 2 pls.
(Received 26 September, 1966.)
BREVIORA
Museum of Comparative Zoology
CAMBRIDGE, Mass. 6 ApriL, 1967 NUMBER 264
THE CHANARES (ARGENTINA) TRIASSIC REPTILE FAUNA
Ill. 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 Chafares-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 Chanares
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 inclined to believe 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-
fares Formation about two miles north of the point where the
Chanares 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 11 lower cheek teeth in available specimens.
Description. Massetognathus pascuali appears to be an exceed-
ingly common member of the Chanares 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 Massetognathus 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 5
Figure 2. Dorsal and ventral views of the skulls of Massetognathus pas-
cuali. 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 Massetognathus 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 11 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.
8 BREVIORA No. 264
Figure 4. Left (above) and right (below) upper cheek tooth series of a
specimen of Massetognathus pascuali. 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 9
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 pascuali. A, Crown
view. 8B, 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,
mediaily, 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 Massetognathus ter-
ugegil. 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 Massetognathus teruggii. 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 BREVIORA No. 264
Figure 8. Laterial view of the skull of Massetognathus teruggii. X %4.
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
ither 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 113)
Figure 9. Dorsal and ventral views of the skull of Massetognathus ter-
uggil. X %.
14 BREVIORA No. 264
Figure 10. Posterior view of the skull of Massetognathus teruggii. X %.
Schréder (1935), and most recently by Brink (11955; etc*):
Trirachodon, 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 Placerias. 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 Chanares 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
“Traité” also included all gomphodonts (including Gomphodonto-
suchus) in the Diademodontidae, although effecting a subdivision
into subfamilies.1 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
(melior Diademodontoidea?) of advanced cynodonts, in contrast
to the carnivores — superfamily Cynognathoidea.
1 His term “Eudiademodontinae” is, of course, inadmissable, not being
based on the name of a genus.
1967 TWO NEW GOMPHODONTS Ly.
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 Ellenberger
(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 Chanares 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, 11 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
Chanares 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 [schignathus 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.
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1967 TWO NEW GOMPHODONTS 25
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(Received 26 September, 1966.)
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