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HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



B R E V I R A 

Museum of Comparative Zoology 



HARVARD UNIVERSITY 



Numbers 437-463 
1976-1980 



MUS. COMP. 200L 
LIBRARY 

FEB - 1^ 1983 

HARVARD 
UNIVERSITY 



CAMBRIDGE, MASSACHUSETTS 02138 U.S.A. 

1980 



BREVIORA 

Museum of Comparative Zoology 

CONTENTS 

Numbers 437-463 

1976 

No. 437. Anolis Alumina, New Species of Grass Anole from 
the Barahona Peninsula of Hispaniola. By Paul E. 
Hertz. 19 pp. July 30. 

No. 438. Crocodilians from the Late Tertiary of Northwestern 
Venezuela: Melanosuchus Fisheri Sp. Nov. By 
Carmen Julia Medina. 14 pp. July 30. 

No. 439. On a New Specimen of the Lower Cretaceous Ther- 
opod Dinosaur Deinonychus Autirrhopus. By 
John H. Ostrom. 21 pp. July 30. 

No. 440. West Indian Anoles: A Taxonomic and Evolutionary 
Summary. 1. Introduction and a Species List. By 
Ernest E. Williams. 21 pp. July 30. 

No. 441. Lizard Karotypes from the Galapagos Islands: 
Chromosomes in Phylogeny and Evolution. By D. 
Paull, E. E. Williams, and W. P. Hall. 31 pp. 
December 27. 

1977 

No. 442. A Laboratory Study of the Turkish Hamster Meso- 
cricetus Brandti. By Charles P. Lyman and Regina 
C. O'Brien. 27 pp. May 27. 

No. 443. Epigonus Trewavasae Poll, A Junior Synonym of 
Epigonus Constanciae (Giglioli) (Perciformes, 
Apogonidae). By Garry F. Mayer and Enrico Tor- 
tonese. 13 pp. May 27. 



No. 444. Stations of the Thayer Expedition to Brazil 1865- 
1866. By Myvanwy M. Dick. 37 pp. May 27. 

No. 445. Natural History of Cerion. VII. Geographic Varia- 
tion of Cerion (Mollusca: Pulmonata) from the 
Eastern End of Its Range (Hispaniola to the Virgin 
Islands): Coherent Patterns and Taxonomic Sim- 
plification. By Stephen Jay Gould and Charles 
Paull. 24 pp. November 30. 

1978 

No. 446. A New Interpretation of the Mammalian Teeth of 
Tribosphenic Pattern from the Albian of Texas. By 
Percy M. Butler. 27 pp. December 20. 

No. 447. Food Selection by Beavers: Sampling Behavior. By 
Stephen H. Jenkins. 6 pp. December 20. 

No. 448. Systematic Notes on the Loons (Gaviidae: Aves). By 
Robert W. Storer. 8 pp. December 20. 

1979 

No. 449. South American Anoles: The Species Groups. 2. The 
Proboscis Anoles {Anolis Laevis Group). By 
Ernest E. Williams. 19 pp. February 21. 

No. 450. A New Species of Moenkhausia from the Mato 
Grosso Region of Brazil (Pisces: Characidae). By 
William L. Fink. 12 pp. February 21. 

No. 451. A New Species of Cybotoid Anole (Sauria, Iguani- 
dae) from Hispaniola. By Albert Schwartz. 27 pp. 
February 21. 

No. 452. Lizards of the Sceloporus Orcutti Complex of the 
Cape Region of Baja California. By William P. 
Hall and Hobart M. Smith. 26 pp. February 21. 



No. 453. The Large Palaeotragine Giraffid, Palaeotragus 
Germaini, from Late Miocene Deposits of Lotha- 
gam Hill, Kenya. By C. S. Churcher. 8 pp. Febru- 
ary 21. 

No. 454. Evolution of Life Histories: A Comparison of Anolis 
- Lizards from Matched Island and Mainland Habi- 
tats. By Robin M. Andrews. 51 pp. July 31. 

No. 455. A New Paleocene Palaeanodont and the Origin of 
the Metacheiromyidae (Mammalia). By Kenneth 
D. Rose. 14 pp. July 31. 

No. 456. Description of a New Hawaiian Gobid Fish of the 
Genus Trimma. By Phillip S. Lobel. 1 5 pp. July 3 1 . 

1980 

No. 457. Two new Species of Eleutherodactylus (Amphibia: 
Leptodactylidae) from the Lowlands and Lower 
Cloud Forests of Western Ecuador. By John D. 
Lynch and Kenneth Miyata. 12 pp. January 31. 

No. 458. A New Species oiAtelopus ( Anura: Bufonidae) from 
the Cloud Forests of Northwestern Ecuador. By 
Kenneth Miyata. 10 pp. January 31. 

No. 459. A New Species of Dendrobates (Anura: Dendrobati- 
dae) from the Lowland Rain Forests of Western 
Ecuador. By Gregory O. Vigle and Kenneth 
Miyata. 7 pp. January 31. 

No. 460. On the Evolution of the Jaw Adductor Musculature 
in Primitive Gnathostome Fishes. By George V. 
Lauder, Jr. 10 pp. June 30. 

No. 461 . Geographic Variation in Anolis Brevirosths (Sauria: 
Iguanidae) in Hispaniola. By Douglas L. Arnold. 
31 pp. June 30. 



No. 462. Eleutherodactylus Eremitus, a New Trans-Andean 
Species of the Lacrimosus Assembly from Ecuador 
(Amphibia: LeptodactyUdae). By John D. Lynch. 
7 pp. June 30. 

No. 463. Jaw Musculature of the West Indian Sn2i\iQ Alsophis 
Canthehgerus Brooksi (Colubridae, Reptilia). By 
Kenneth V. Kardong. 26 pp. August 15. 



BREVIORA 

Museum of Comparative Zoology 

Index of Authors 

> Numbers 437-463 

1976-1980 

Andrews, Robin M 454 

Arnold, Douglas L 461 

Butler, Percy M 446 

Churcher, C. S 453 

Dick, Myvanwy M 444 

Fink, William L 450 

Gould, Stephen Jay 445 

Hall, William P 441, 452 

Hertz, Paul E 437 

Jenkins, Stephen H 447 

Kardong, Kenneth V 463 

Lauder, George V., Jr 460 

LoBEL, Phillip S 456 

Lyman, Charles P 442 

Lynch, John D 457, 462 



Mayer, Garry F 443 

Medina, Carmen Julia 438 

MiYATA, Kenneth 457, 458, 459 

O'Brien, Regina C 442 

OsTROM, John H 439 

Paull, Charles 445 

Paull, D 441 

Rose, Kenneth D 455 

Schwartz, Albert 451 

Smith, Hobart M 452 

Storer, Robert W 448 

ToRTONESE, Enrico 443 

ViGLE, Gregory O 459 

Williams, Ernest E 440, 441, 449 



^^S. CCMP. ZOOL 
LlDr?ARY 

MAR 1 fi m^ 

B R E V I Q,R...A 

Museum of Comparative Zoology 

_US ISSN_0006-9698_ 

Cambridge, Mass. 30 July 1976 Number 437 

ANOLIS ALUMINA, NEW SPECIES OF 

GRASS ANGLE FRGM THE 

BARAHGNA PENINSULA GF HISPANIGLA 

Paul E. Hertz^ 

Abstract. A new species, Anolis alumina, from the Baiahona Peninsula 
of Hispaniola is described on the basis of two scale characters and dewlap 
color in males. Electrophoretic and morphological analyses show that 
(iliiini)ia is more closely related to A. semilmentus than to A. olssoni. 
Limited distributional data suggest that the new species is more eiuytopic 
than either of the other two species of Hispaniolan grass anoles, occurring 
in habitats ranging from lowland desert scrub to high elevation pine 
savannah. 

Durins^ 1973 and the winter of 1974, I initiated an electro- 
phoretic study of the gras.s anoles of Hispaniola, using material 
that had been collected by various researchers in the Harvard 
Anolis group; the results, though incomplete, suggested that 
animals found south of the Massif de la Selle — Sierra de 
Baoruco mountain chain are taxonomically differentiated from 
the vvhite-dewlapped A. semilineatus Cope of the northern part 
of Hispaniola. 

While collecting in the Dominican Republic during the 
summer of 1974, Raymond B. Huey and I journeyed to the 
south slopes of the Sierra de Baoruco to look for grass anoles. 
We were fortunate to enjoy the hospitalit)- of the Alcoa Ex- 
ploration Company at their bauxite mining operation at Cabo 
Rojo, Pedernales Province, while we explored the mine area 
and surrounding countr\'side. 

As we drove up to the mine one morning, Huey spotted a 
small animal dart across the road in front of our vehicle. After 



iMuseum of Comparative Zoology, Harvard University. Cambridge, 
Massachusetts 02138. 



2 BREVIORA No. 437 

perhaps ten minutes of frantic searching and chasing the elusive 
lizard in the pine savannah into which it fled, I succeeded in 
grabbing the animal which promptly exhibited his displeasure 
by erecting his pale greenish-yellow gular fan. I suspected at 
once that we had captured a previously undescribed species of 
Anolis which, in honor of our hosts, I name 

Anolis alnmino, new species 

Holotype. 31.5 km north of Cabo Rojo, Pcdeinales Province, Dominican 
Republic (elevation 1150 m) , MCZ 143824, P. E. Hertz and R. B. Huey 
collectors, 8 July 1974. 

Paratypes. Dominican Republic: Pedernales Pro\'ince: Peder- 
nalcs: ASFS V2816, R. Thomas collector, 3 Julv 1964; 5 km 
north of Pedernales: ASFS V2544-V2545, R. Thomas collec- 
tor, 25 June 1964; 9 km north of Pedernales: ASFS \' 2 1496, 
R. Thomas collector, 27 Julv 1969; 6 miles north of Peder- 
nales: ASFS V30118-V30127, D. C. Fowler and A. Schwartz 
collectors, 23 August 1971; 21 km north of Cabo Rojo: ASFS 
V30058, D. C. Fowler collector, 21 August 1970;" 23.5 km 
north of Cabo Rojo: MCZ 143849-143851, P. E. Hertz and 
R. B. Huey collectors, 8 Julv 1974; 28 km north of Cabo Rojo: 
MCZ 143822-143823, P. E. Hertz and R. B. Huev collectors, 
7 July 1974; 30 km north of Cabo Rojo: MCZ 146632-146633, 
W. E. Haas collector, 20 Julv 1975; 31.5 km north of Cabo 
Rojo: MCZ 143825-143827, P. E. Hertz and R. B. Huev col- 
lectors, 8 July 1974: MCZ 146627-146631, W. E. Haas and 
E. E. Williams collectors, 19 Julv 1975; 7 km north, 17.6 km 
southeast of Cabo Rojo: ASFS V30079-V30083, D. C. Fowler 
and A. Schwartz collectors, 22 August 1971; 5 miles northeast 
of Oviedo: ASFS V289, R. Thomas collector, 7 August 1963; 
13.1 miles southv/est of Enriquillo: x^SFS X9966, A. Schwartz 
collector, 30 Julv 1963. Barahona Province: southern outskirts 
of Barahona:" ASFS V30980, B. R. Shenlan collector, 12 Sep- 
tember 1971; MCZ 106995, E. E. Williams, A. S. Rand, and 
E. Marcano collectors, 28 Julv 1968; 5 km south of Barahona: 
ASFS V20552, R. Thomas 'collector, 22 June 1969; 7 km 
southwest of Barahona: ASFS V23423, A. Schwartz collector, 
4 January 1971; 4.1 miles southwest of Barahona: ASFS 
V30407-V30415, D. C. Fowler, A. Schwartz, and B. R. Shep- 
lan collectors, 9 December 1971; 4 km northwest of Naranjal: 



1976 



ANOLIS ALUMINA 



ASFS V20954, R. Thomaa collector, 4 July 1969; 1 km south 
of Cabral: MCZ 140011-140012, T. P. Webster collector, 
4 November 1973; 4.9 miles northwest, 0.3 miles west of 
Cabral: ASFS V30815, D. C. Fowler collector, 8 September 
1971; Polo: AMNH 50320, W. G. Hassler collector, no col- 
lectins: date. Haiti: Departement de I'Ouest: Belle-Anse: MCZ 
140104-140111, T. P. Webster collector, September 1973. 

Diagnosis' An Anolis closely related to A. semilineatus and 
A. olssoni Schmidt, distinguished from both by smooth scales in 
the frontal depression ( Fig. 1 ) , enlarged middorsal scales grad- 
ing into the granular scales of the flank (rather than abruptly 
distinct) (Fig. 2), and a pale greenish-yellow gular fan in males 
( rather than white or orange ) . 




Figure 1. Dorsal and lateral views of the head of holotype of Anolis 
alumijia (MCZ 143824). 



BREVIORA 



No. 437 



m^Wimi^ 





Figure 2. Enlarged middorsal scales of (left) Anolis ahanhia (MCZ 
143824) and (right) A. semilineatus (ASFS V8093) . 



Head. Most head scales keeled. Six to eight scales across 
head between second and third canthals. Frontal depression 
extremely shallow (except in one specimen), its scales smooth 
and varying in size but never larger than the anterior supra- 
orbital. 

Supraorbital semicircles usually separated by one scale row 
(separated by two scales in one specimen, in contact in two 
specimens) and separated from supraocular disc by one row of 
small scales or in contact with supraocular disc (two speci- 
mens). Supraocular disc consists of two to five large keeled 
scales and zero to three smaller keeled scales, usually surrounded 
by granular scales. Supraocular disc separated from elongate 
supraciliary by one to three rows of granular scales and/or a 
small elongate scale at anterior side of contact. 

Canthus distinct, canthal scales four to five, second largest, 
diminishing in size anteriorly. Naris anterior to canthal ridge. 
Anterior nasal scale small, in contact with rostral scale. Four 
to five vertical loreal rows. 

Temporal scales subgranular. Two to four rows of granular 
supratemporals ( larger than temporals ) , grading into subgranu- 
lar scales which grade dorsally into the larger scales surrounding 
the interparietal. Interparietal about as large as the ear, sepa- 
rated from supraorbital semicircles by two to three scales (four 
in one specimen). Suboculars keeled (sometimes markedly so) 
and in contact with supralabials. Four to six supralabials from 
rostral to center of the eye. Rostral scale markedly rounded on 
dorsal surface. 



1976 ANOLIS ALUMINA 5 

Mentals wider than long, two small elongate scales inserted 
between posterior tips. One to five (usually three) sublabials 
on each side contact infralabials. Central throat scales keeled, 
elongate. Gular fan large in males, absent in females. Scales of 
gular fan singly keeled, elongate or oval, about the same size 
as ventrals, not clearly arranged in rows. 

Trunk. Unicarinate middorsal scales, arranged in longitudinal 
rows, about' as broad as long, grading laterally into small keeled 
or granular flank scales. Seventeen to 24 middorsals in standard 
distance (tip of snout to center of eye) in males, 15 to 19 in 
females. Ventrals in longitudinal rows, unicarinate, imbricate, 
and in some specimens slightly mucronate. Fifteen to 24 ven- 
trals in standard distance in males, 11 to 18 in females. En- 
larged postanal scales present in males. 

Limbs and digits. Hand and foot scales multicarinate. Larg- 
est arm and leg scales unicarinate, about as large as enlarged 
middorsals. Fifteen to 18 lamellae under phalanges two and 
three of fourth toe. 

Tail. Oval to circular in cross section two and one-half to 
three times snout-\'ent length. 

Size. Largest male 40 mm snout-vent length. Largest female 
37 mm snout-vent length. 

Color in life. Dorsal surface of head tan to chestnut. Flanks 
chestnut. Middorsal stripe (or series of elongate blotches which 
meet medially) a peppered ofT-white, bordered on either side 
by a narrow tan stripe. \^enter and mandible of both sexes and 
throat of females white with moderate brown peppering. Con- 
spicuous white lateral stripe on maxilla from tip of snout along 
flank to hind leg, extending in some specimens to base of tail. 
Scales of gular fan (in males only) white with brown peppering. 
Skin of gular fan pale greenish-yellow. Iris blue. 

Distribution. Anolis alumina appears to be widely distributed 
on the Barahona Peninsula and on the south slopes of the Sierra 
de Baoruco-Massif de la Selle mountain range (Fig. 3). Most 
specimens from this area which had been previously assigned to 
seynilineatus are now designated as paratypes of alumina on the 
basis of the two diagnostic scale characters. 

Preliminary observations suggest that the habitat preferences 
of Anolis alumina are similar to those of other Hispaniolan 
grass anoles. The type series was collected in the undergrowth 
of a pine savannah at middle ele\'ations; the animals were 



6 BREVIORA No. 437 

spotted while perching below two feet in the vegetation, and 
usually fled by dropping to the ground and scurrying into dense 
clumps of grass or between rocks. Other collectors have re- 
ported the animals sleeping on living and dead desert shrubs, 
on dead Acacia in and along the margins of cleared fields, and 
on dead grass and twigs in or near meadows. Albert Schwartz 
(personal communication) has collected three specimens at night 
from under rocks in hammock woods, a sleeping habit previously 
unreported for members of the sernilineatus group. D. C. Fowler 
collected a single specimen asleep at a height of four feet on the 
leaf of a tree. 

COMPARISONS WITH OTHER SOUTHERN ISLAND 
POPULATIONS IN THE SEMILINEATUS COMPLEX 

Comparison of alumina with neighboring populations of grass 
anoles on the south island of Hispaniola supports the distinction 
of alumina as a full species. A. alumina and semilineatus are 
easily distinguished from olssoni on the basis of the size of the 
throat scales in females and the gular fan scales in males: in the 
latter species, throat scales are greatly enlarged and are much 
larger than the ventrals. We can then refer to alumina and 
semilineatus as members of the more primitive semilineatus com- 
plex within the semilineatus species group, and consider the 
enlarged throat scales of olssoni a derived character (Williams, 
1961). A. olssoni can then be ignored (as unambiguously dis- 
tinct) in morphological comparisons of southern island grass 
anoles because the only confusion of alumina and olssoni appears 
to have arisen from the fact that both species have pigmented 
(as opposed to "white," as in semilineatus) dewlaps. 

Cope (1864) lists the type locaHty of semilineatus as "Hayti." 
The type specimen (BMNH 1946.8.5.85) resembles the Petion- 
ville semilineatus series in its keeled head scales and its dorsal 
standard distance counts. Since the Port-au-Prince area was, 
and still is, the most travelled part of Haiti, one is tempted to 
suggest that the type locality of semilineatus may plausibly be 
restricted to Petionville. 

A series of semilineatus (ASFS V16698-V16707) geographi- 
cally close to alumina was collected at the top of the road to the 
Alcoa mine near Cabo Rojo at an elevation of 4400 feet, ap- 
proximately eight road kilometers above the type locality of 
alumina. The two diagnostic scale characters indicate that the 



1976 ANOLIS ALUMINA 7 

ASFS series is unquestionably semilineatus. In addition, their 
maxillary-lateral stripes are considerably darker than those on 
the nearest alimiina, though this may be the result of different 
preservation techniques. These animals are smaller than their 
geographically closest Haitian conspecifics, but several males are 
larger than any of the specimens of alumina that we have at 
hand. It is- possible that semilineatus replaces alumina at high 
elexations in the Sierra de Baoruco and Massif de la Selle, but 
much more material is needed to establish this point. 

Because of inadequate collecting, our knowledge of the fauna 
on the dry north slopes of the Sierra de Baoruco is scant. Semi- 
lineatus is unknown in the \^alle de Neiba and Plaine du Cul- 
de-Sac except at its westernmost edge near Port-au-Prince. A. 
olssoni is somewhat better represented in the valley itself, but 
there are few records of its presence on the north slopes of the 
Sierra de Baoruco. Based upon our limited knowledge of this 
area, it appears that alumina is restricted to the lowlands in the 
southeastern corner of the Valle de Neiba near Barahona and 
Cabral. 

To the west of the range of alumina, in the central section of 
the southern island, there are a series of populations here re- 
garded as semilineatus, characterized by large body sizes (snout- 
vent length of largest adult male is 46 mm), ver\- distinct mid- 
dorsal strioes, markedly keeled head scales (including those of 
the frontal depression), and distinctly enlarged middorsal scales 
which do not grade into the granular flank scales. The follow- 
ing are the south island specimens so characterized: 

HAITI: 5 km south of Dufort, MCZ 63046-63047; ^a-Ira, 
MCZ 64841, 64844-64846, 64850, 64852, 64857, 64858; 4 
miles southwest of Gressier, ASFS V8335; 10 miles north of 
Jacmel, ASFS V37867; 4 miles northwest of Jacmel, ASFS 
V9804; 3 miles east of Jacmel, ASFS V9759-V9760; Bascap 
Rouge, MCZ 65024; 3 miles east of Caves Jacmel, ASFS 
V9723-V9725; 2 miles west of Marigot, ASFS V9770; 9.7 miles 
southwest of Seguin, ASFS V38159-V38160; 3.8 miles south- 
west of Seguin, ASFS V38187; 10 miles north-northeast of 
Marigot, ASFS V9732-V9735; mountains south of Gasseleine 
River at Marbial, AMNH 69036; Peneau, ASFS XI 354, 
X1550; Obleon, MCZ 60015-60016; Basin Bleu, MCZ 60017- 
60020; Furcv, MCZ 58011, 59553, 60025-60027, 63417- 
63426, ASFS'X1596, X1905, V8357-V8366; 4.8 km south of 
PetionviUe, ASFS V8083-V8116; Boutilliers Road, MCZ 



8 



BREVIORA 



No. 437 



60021-60024, 60028-60029, 63038-63039, 112099-112100, 
ASFS V36199, V36231-V36232, V36551; 6 miles west of 
Petionville, ASFS V8347; Petionville, ASFS X3340-X3345. 

In addition, we have a single specimen of semilineatus (MCZ 
131150), which had been mistakenly included in a series of 
olssoni, from Savanne Zombi, a high elevation (1500 meters) 
locality in eastern Haiti. Since olssoni is otherwise known to 
occur only in relatively xeric lowlands (and along some middle 
elevation roadsides on the north island), I consider the locality 
data for these specimens doubtful and have omitted them from 
the range maps (Fig. 3) for these species. 



^=^ 




Figure 3. Locality records for (top) AnoUs alumina (solid circles) , 
A. semilineatus (open circles) , and (bottom) A. olssoni. Locality records 
from collections in American Museum of Natural History, Museum of 
Comparative Zoology, United States National Museum of Natural History, 
and Albert Schwartz Field Series. Dotted lines indicate the Massif de la 
Selle and Sierra de Baoruco. 



1976 ANOLIS ALUMINA 9 

A. semilineatus specimens from localities at the western end of 
the Tiburon Peninsula (near Duchity, Carrefour Zaboka, Jere- 
mie, and Les Platons) are similar to, though smaller than, those 
just described. The most variable characters in these lizards are 
dorsal scale size and body size which, as in populations on the 
north island, appear to vary clinally with elevation (Hertz, in 
preparation). It seems reasonable to regard these as serniUne- 
atus, at least provisionally. 

ELECTROPHORETIC STUDIES 

As a check on the taxonomic status of alumina, I have ana- 
lyzed eleven samples of grass anoles using starch gel electropho- 
resis; sample sizes varied between six and 42 animals (Fig. 4). 
Five of these populations are white-dewlapped animals from the 
north island of Hispaniola. Constanza (population A) is the 
type locality of A. cochranae, which Williams and Rand (1961) 
described as distinct from semilineatus, but which I consider an 
altitudinal variant of semilineatus. In addition, I have grouped 
the Petionville sample with those from the north island because 
the animals there are morphologically and electrophoretically 
indistinguishable from north island seiyiilineatus. [Williams 
(1965) has reported that Anolis aliniger, a north island green 
anole, has been found near Petionville, and Schwartz (1974) 
noted that several other northern island lizard species are known 
from southern island localities in the area. Thus, the western 
side of the Cul-de-Sac Plain appears to be an area of "faunal 
leakage" from the north island to the south island, and Petion- 
\ille may, at least for my purposes in this description, be con- 
sidered a north island locality.] 

I used material from two alumina populations in the electro- 
phoretic analysis: animals from the type locality (28-31.5 km 
north of Cabo Rojo) and those from Belle-Anse, Haiti. The 
latter population exists on the south coast of Haiti at the western 
boundary of the Barahona Peninsula, cut off on the north from 
the rest of the island by the Massif de la Selle, the Haitian 
analogue of the Sierra de Baoruco. 

Three populations from the north island were available to 
represent olssoni. Since both olssoni and semilineatus are with- 
out question good species, electrophoretic comparison of them 
provides a means of assessing the significance of various degrees 
of genie differentiation within the species group. 



BREVIORA 



No. 437 




1976 AXOLIS ALUMINA 11 

The methods of sample preparation and horizontal starch gel 
electrophoresis are adapted from techniques described by Selan- 
der et al. (1971). All proteins were extracted from tissue and 
water homogenates prepared from material which was frozen in 
the field. (The less-than-ideal freezing conditions in the field 
resulted in some protein denaturation; two proteins, Indophenol 
oxidase and an Isocitrate dehydrogenase, were eliminated from 
consideration because of their lability.) Buffer systems and as- 
says used in this study are similar to those used in previous 
Anolis studies (e.g., Webster 1975) ; details of the technique and 
assay formulas are available from the author upon request. 

TABLE 1 

Proteins assayed in the electrophoretic sur\ey 

Albumin Alcohol dehydrogenase 

Protein A a-Glycerophosphate 
Protein B dehydrogenase 

Protein C Glutamic oxaloacetic 
Leucine aminopeptidase transaminase 

Phosphoglucose isomerase 6-Phosphogluconate 
Lactate dehydrogenase- dehydrogenase 

1 and 2 Phosphoglucomutase-1 and 2 

Isocitrate dehydrogenase-2 Peptidase 
Malate dehydrogenase- 1 and 2 Fumarase 

Nineteen proteins produced consistently scorable bands in 
all individuals (Table 1); allele frequencies for these proteins 
are used for the calculation of indices of electrophoretic similar- 
ity of pairs of populations. Webster (1975) has argued that 
Nei's (1972) normalized identity of genes, 7, is the preferred 
index of genetic similarity of two populations. W^hen 7 = 0, 
the populations share no alleles ; when 7 = 1 , all gene frequen- 
cies in the two populations are identical. I use Nei's 7 as an 
index of genetic similarity here and assume in the calculation 
of 7 N'alues that each polypeptide is the product of one gene. 

I present 7 values for paired localities in Table 2. It is readily 
apparent that the populations fall into three distinct groups 
which correspond to the three species recognized on the basis of 
scale characters. The average similarity \-alue for paired com- 
parisons of six semilineatus populations from the north island 



12 BREVIORA No. 437 



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1976 ANOLIS ALUMINA 13 

is 0.96 (range 0.90-0.99), for the three olssoni populations 
7^0.96 (range 0.95-0.97), and for two alu?nina populations 
I = 0.99. 

Average similarity values between paired populations of the 
different species are much lower: semiUneatus to alumina, I ^= 
0.80 (range 0.75-0.85) ; semiUneatus to olssoni, I = 0.84 (range 
0.81-0.87); and alumina to olssoni, 7 = 0.70 (range 0.64- 
0.73). 

Only one of the 19 proteins assayed is completely differen- 
tiated in animals from the topotypic sample of Anolis alumina, 
a fast variant (i.e., one that migrates further from the origin) 
of albumin which is also the dominant albumin variant in the 
animals from Belle-Anse. However, the topotypic almnina pop- 
ulation is fixed for an allele of Isocitrate dehydrogenase-2 which 
appears in only two other populations: 94 per cent at Belle- 
Anse (also alumina) and 7 per cent at Petionville [semiUnea- 
tus) . Similarly, a slow variant of 6-Phosphogluconate dehydro- 
genase is the only 6-PGD allele in topotypic alurriina and the 
predominant (89 per cent) 6-PGD allele at Belle-Anse, but 
only occurs at a maximum frequency of 20 per cent in other 
populations. Finally, the topotypic sample of alumina had a 
low frequency (5 per cent) of a slow allele of Malate dehydro- 
genase-1, a protein that was monomorphic for a faster allele in 
all other populations sur\eyed. 

The electrophoretic data provide support for the species status 
attributed to alumina on the basis of external morphological 
characters. Mayr (1970) and Selander and Johnson (1973) 
have noted the dangers in attempting to differentiate species on 
the basis of the number of protein differences between them: 
we have no independent method for assessing just how much 
genetic differentiation is involved in speciation even within a 
rather narrowly-defined taxon. It is, however, permissible to 
use Nei's index of genetic similarity to compare the distinctive- 
ness of populations of alumina, olssoni and semiUneatus. Paired 
comparisons of alumina and semiUneatus populations (Table 2) 
show that, on average, populations of these species are less simi- 
lar electrophoretically than are olssoni and semiUneatus; on the 
other hand, alumina and olssoni are much less similar to each 
other than either is to semiUneatus. 



14 BREVIORA No. 437 

The fact that only one of the nineteen proteins assayed can 
be called diagnostic of alumina is not an argument against the 
designation of alumina as a full species (see Lewontin, 1974: 
180-182, for a discussion of electrophoretic evidence of intro- 
gression in closely related species). Ayala and Powell (1972) 
and Prakash ( 1 969 ) have demonstrated that closely related 
species of Drosophila share many alleles, but that each species 
carries a particular allele at a different frequency. We find a 
similar situation with respect to three other proteins in aluyyiina 
and semilineatus: IDH-2, 6-PGD, and MDH-1. 

RELATIONSHIPS WITHIN THE SPECIES GROUP 

Williams (1961) has proposed a model for speciation within 
the seynilineatus species group which is based upon the Pleisto- 
cene di\'ision of Hispaniola into northern and southern islands. 
He suggested that seynilineatus is the autochthonous grass anole 
of the south island and that olssoni is the autochthon of the 
northern component. Williams then suggested, on the basis of 
somewhat limited distributional data, that semilineatus invaded 
the north, while olssoni, probably restricted by the absence of 
suitable habitat, did not successfully extend its range throughout 
the south. 

Our current knowledge about the distributions of the His- 
paniolan grass anoles, summarized in Figure 3, shows olssoni 
with a rather limited south island distribution. In fact, olssoni 
appears to be restricted on the south island to lowland areas 
within the currently known range of alumina on the Barahona 
Peninsula. The occurrence of olssoni on the Barahona Peninsula, 
however, does not invalidate Williams' scheme for differentiation 
within the group and subsequent invasion of the north by semi- 
lineatus. A. olssoni probably originated on the north island and 
then invaded the south, after the recession of a Pleistocene 
seaway, through the relatively dry lowland corridor at the east- 
ern edge of the Valle de Neiba near what is now the city of 
Barahona. Movement southward along the relatively mesic coast 
and westward across the deserts of the Barahona Peninsula and 
foothills of the Sierra de Baoruco probably followed its initial 
invasion of the southern island from the northeast. 

The differentiation of semilineatus from alumina can be at- 
tributed to a classical allopatric speciation phenomenon that 
requires only minor modification of Williams' (1961) scheme. 



1976 ANOLIS ALUMINA 15 

The range of alumina is separated from the western portion of 
the southern island by the Sierra de Baoruco-Massif de la Selle 
mountain range which comes close to the sea west of Belle-Anse, 
Haiti. The north slopes of these mountains appear to be suffi- 
ciently dry to limit the contact of the two species today; if the 
climate was similar during the Pleistocene, the arid areas may 
ha\e limited genetic exchange between the animals on either 
side of the tnountains and established a sufficient reproductive 
barrier to allow speciation. The presence of alumina at two 
localities near Cabral on the north side of the Sierra de Baoruco 
can be attributed to a very limited northward invasion through 
the same corridor through which ohsoni moved south into the 
Barahona Peninsula. 

The Barahona Peninsula and particularly the south slopes of 
the Sierra de Baoruco are becoming well known as a center of 
differentiation of the Hispaniolan saurofauna. Williams (1962, 
1963) recognized the biogeographic importance of the region 
with reference to Anolis barahonae and A. hendersoni. Schwartz 
(1964, 1967a, b, 1974), and Schwartz and Klinikowski (1966) 
have further documented differentiation there for the Anolis 
ricordi complex and the lizard genera Leiocephalus, Ameiva, 
and Diploglossus. 

On the electrophoretic evidence, alumina is genetically less 
similar to semilineatus than is ohsoni. If the degree of electro- 
phoretic dissimilarity of two species can be taken as an index of 
either their time of divergence or the magnitude of enx'iron- 
mental difference between their ranges, one can suppose that 
alumina and a "^proto'' -semilineatus diverged before Pleistocene 
seaways isolated the ancestors of semilineatus and olssoni or that 
the selecti\e regimes of the Barahona Peninsula at the time of 
differentiation of ahnnina and seynilineatus were perhaps more 
different from those on the rest of the southern island than were 
conditions on the northern island. Both suggestions are plausible 
and not incompatible. 

The less enlarged middorsal scales suggest that alujyiina is the 
most primitive of the three species in the semilineatus species 
group. A plausible and conservative model of evolution within 
the group would suggest that olssoni evolved from a "proto"- 
seyyiilineatus stock which had already di\erged from an ancestral 
alumina-Y\kt grass anole (lacking distinctly enlarged middorsal 
scales). The subsequent enlargement of dewlap scales in olssoni 



16 BREVIORA No. 437 

may have accompanied the dramatic change in dewlap color as 
a reproductive isolating mechanism to limit its hybridization 
with scyyiilineatus. The repeated changes in Pleistocene sea level 
would probably have allowed such sequential derivations to 
occur. 

Perhaps alumina and olssoni, species which may be evolution- 
arily one step remo\ed from each other, were different enough 
to allow ecological segregation and sympatry as olssoni invaded 
the eastern section of the south island. In contrast, an invasion 
by olssoni across the western end of the Cul-de-Sac Plain may 
have been inhibited by either unsuitable habitat or by com- 
petitive interactions with resident populations of semilineatus. 
Although semilineatus and olssoni are syntopic at a number of 
localities on the northern island (Hertz, in preparation), there 
is no reason to suggest that reciprocal invasions in either direc- 
tion should have equal probabilities of success. A. semilineatus 
may have undergone an ecological shift as it invaded the north- 
ern island, thereby allowing sympatry with olssoni. But indi- 
viduals of olssoni invading in the other direction may have been 
swamped by competition from the resident semilineatus. Clearly, 
we must know the ecology of both species in much greater detail 
before we can draw any firm conclusions about competitive pat- 
terns that may ha\'e shaped their current distributions. 

The description of a third species of Hispaniolan grass anole 
does not provide us with a necessarily complete analysis of the 
systematics of the semilineatus species group. There are several 
issues about which I must withhold final judgment pending the 
collection of additional data. Although I feel that I can with 
some confidence align the holotype of semilineatus with series 
of specimens from the Petionville area, I cannot confidently 
comment upon the systematic status of the grass anoles from the 
western portion of the southern island. The electrophoretic evi- 
dence tells us that populations of semilineatus near Petionville 
probably have had recent gene exchange with or are derived 
from populations on the north island; data from other studies 
(Williams, 1963; Schwartz, 1974) suggest that faunal leakage 
in this part of the Cul-de-Sac Plain may, in fact, be a common 
phenomenon. In fact, the animals found near Petionville may be 
directly derived from north island semilineatus and may not be 
conspecific with some other south island populations for which 
we ha\'e no electrophoretic data. 



1976 ANOLIS ALUMINA 17 

We cannot yet fully assess the possibility that north and south 
island populations of semilineatus differentiated after the semi- 
Uneatus invasion of the north through the western end of the 
Cul-de-Sac. Pleistocene sea le\el rose and fell a number of 
times, reseparating and rejoining the two component islands of 
Hispaniola with each creation and recession of the seaway. If 
semilineatus invaded the north during an early oceanic reces- 
sion, subsequent inundation of the low elevation plain may have 
split the range of semilineatus into two portions, allowing dif- 
ferentiation. North island animals may then have reinvaded the 
south near Petionville and Furcy, and may be distinct from 
animals at the distal end of the Tiburon Peninsula. Electro- 
phoretic data from semilineatus on the western end of the Tibu- 
ron Peninsula should clarify this point. 

ACKNO\\LEDGMENTS 

This study could never have been completed without the 
assistance and generosity of W. M. Smit and Alfredo Lebron 
of the Alcoa Exploration Company in the Dominican RepubKc. 

I thank Robert Holt, Ernest Williams, and especially Ray- 
mond B. Huey for assistance in the field. Ronald Crombie, 
x\lbert Schwartz, and Ernest Williams made suggestions during 
the course of the study. The late T. Preston Webster is re- 
sponsible for a substantial portion of the electrophoretic data 
and for teaching me electrophoretic techniques. Raymond B. 
Huey, Kenneth Miyata, Albert Schwartz, and Ernest Williams 
pro\'ided useful criticisms of drafts of the manuscript. 

Specimens examined in this study were borrowed from the 
American Museum of Natural Histor\' (AMNH), British Mu- 
seum of Natural History (BMNH), Museum of Comparative 
Zoology (MCZ), Albert Schwartz Field Series (ASFS), and the 
United States National Museum (USNM). I thank the cura- 
tors of these collections for allowing me to examine specimens 
in their care. 

Support for this study was provided by the Richmond Fund 
of the Department of Biology, Harvard University and National 
Science Foundation Grant GB-37731X to Ernest Williams. 
Specimens in the .\lbert Schwartz Field Series were collected 
under National Science Foundation Grants G-7977 and B- 
023603. 



18 BREVIORA No. 437 

LITERATURE CITED 

Ayala, F. J. AND J. R. Powell. 1972. Allozymes as diagnostic characters 
of sibling species of Drosopliila. Proc. Nat. Acad. Sci. 69: 1904-1906. 

Cope, E. D. 1864. Contributions to the herpetology of tropical America. 
Proc. Acad. Nat. Sci. Phila. 1864: 171. 

Lewontin, R. C. 1974. The Genetic Basis of Evolutionary Change. Co- 
lumbia University Press. New York. 

Mayr, E. 1970. Populations, Species, and Evolution. Belknap Press. Cam- 
bridge. 

Nei, M. 1972. Genetic distance between populations. .\mcr. Nat. 106: 
283-292. 

Prakash, S. 1969. Genie variation in a natural population of Drosophila 
persimilis. Proc. Nat. Acad. Sci. 62: 778-784. 

Schwartz. A. 1964. Diploglossus costatus Cope (Sauria, Anguidae) and its 
relatives in Hispaniola. Reading Publ. Museinn and Art Gallery Sci. 
Publ. 13:1-57. 

1967a. The Leiocephalus (Lacertilia, Iguanidae) of Hispan- 
iola. II. The Leiocephalus persoyiatus complex. Tulane Studies in 
Zoology 14: 1-53. 

1967b. The Anieiva (Lacertilia, Teiidae) of Hispaniola. III. 



Ameiva taeniura Cope. Bull. Mus. Comp. Zool. 135: 345-375. 

-1974. An analysis of variation in the Hispaniolan giant anole, 



Anolis ricordi Dumeril and Bibron. Bull. Mus. Comp. Zool. 146: 89-146. 
AND R. F. Klinikowski. 1966. The Ameiva (Lacertilia, Teii- 



dae) of Hispaniola. II. Geographic variation in Anieiva chrysolaema 
Cope. Bull. Mus. Comp. Zool. 133:425-487. 

Selander, R. K. and W. E. Johnson. 1973. Genetic variation among verte- 
brate species. Ann. Rev. Ecol. Syst. 4: 75-91. 

, M. H. Smith, S. Y. Yang, W. E. Johnson and J. B. Gentry. 

1971. Biochemical polymorphism and systematics in the genus Pero- 
myscus. 1. Variation in the old field mouse (Peromyscus polionotus) . 
Studies in Genetics VI: 49-90. Univ. Texas Publ. No. 7103. 

Webster, T. P. 1975. An electrophoretic comparison of the Hispaniolan 
lizards Anolis cy botes and A. tnarcanoi. Breviora No. 431: 1-8. 

Williams, E. E. 1961. Notes on Hispaniolan herpetology. 3. The evolu- 
tion and relationships of the Anolis semilineatus group. Breviora 
No. 136: 1-8. 

1962. Notes on Hispaniolan hcrpctolog\'. 6. The giant anoles. 



Breviora No. 155: 1-15. 



1963. Notes on Hispaniolan herpetology. 8. The forms re- 



lated to Anolis hendersoni Cochran. Breviora No. 186: 1-13. 



1976 ANOLIS ALUMINA 19 
1965. The species of Hispaniolan green anoles (Sauria, 



Iguanidae) . Breviora No. 227: 1-16. 

AND A. S. Rand. 1961. Notes on HisjDaniolan herpctology. 



2. A review of the Anolis seniilineattis group with the description of 
Anolis cochranae, a new species. Breviora No. 135: 1-11. 



>. ^GMP. 200L 
L/nr?ARY 



MAR 1 8 19,85 

B R E V I O R"A'' 

Miiseiim of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 30 July 1976 Number 438. 

CROCODILIANS FROM THE LATE TERTIARY 
OF NORTHWESTERN VENEZUELA: 
MELANOSUCHUS FISHERI SP. NOV. 

Carmen Julia Medina^ 

Abstract: Melanosucluts fishen n. sp., the first fossil record of this aUiga- 
torid genus, is based on two skulls from the Urumaco Formation of Huay- 
querian (Pliocene) age in the northern part of the State of Falcon, Venezuela. 
Relationships between M. fisheri and the living M. tiiger are uncertain. 
M. fisheri occurs together with five other crocodilian taxa of gigantic size. 
This assemblage and the numerous associated chelonians suggest that the 
Urumaco area in Huavqueiian times mav have borne some resemblance to 
the present \niazon basin. 

Rjesumen: Se describe Melanosurhus fnheri, una nueva especie de un 
genero aligat6rido hasta ahora sin un registro f6sil, basada sobre dos 
ejemplares tolcccionados en la Formaci6n Urumaco, de Edatl Huavquenense. 
en la parte norte del Estado Falcon, Venezuela. 

During the summer of 1972, a paleontological expedition 
from the Museum of Comparative Zoology, in collaboration 
with the Escuela de Geologia of the LIniversidad Central de 
Venezuela and the Ministerio de Minas e Hidrocarburos, worked 
in the vicinity of Urumaco, Distrito Democracia, Estado Fal- 
con, northwestern Venezuela. A rather large number of \'erte- 
brate fossils, most of them reptiles, were collected in the Llru- 
maco Formation, of Huayquerian age (Pascual and Diaz de 
Gamero, 1969; for a definition of Huayquerian, see Pascual and 
Odreman, 1973). Among the crocodilian materials are two 
skulls, representing two growth stages of a species of caiman, 
that have strikingly large orbits, extending as far forward as 
maxillary tooth 10, and a palatal exposure of the vomer. These 
features (and others) occur, among living crocodilians (Wer- 



iMuseo de Ciencias Naturales de Caracas 



2 BREVIORA No. 438 

muth, 1953), only in the black caiman, Melanosuchus niger 
(Spix). The two specimens are beyond doubt referable to the 
genus, of which they constitute the first fossil record, and are 
specifically distinct from the living form. 



Melanosuchus fisher? sp. nov. 

Type: Museo de Ciencias Naturales de Caracas No. 243, a 
skull lacking the pterygoids, parts of the ectopterygoids, pala- 
tines, jugals and premaxillaries, and parts of the left postfrontal 
and quadrate. 

Hypodigm: The type and MCZ No. 4336, a skull — lacking 
the pterygoids, the left quadrate and quadratojugal, and parts 
of the basioccipital and left squamosal — and the posterior two- 
thirds of both rami of the mandible. Both specimens have suf- 
fered some distortion, and, as in much of the Urumaco material, 
the bone surfaces are poorly preserved. 

Horizon: Upper part of the upper member of the Urumaco 
Formation in the "capa de huesos"' or "tortugas" of the field 
geologists. 

Localities: The type was found approximately /2 km north- 
west of Campo El Mamon, east of the El Jebe fault; MCZ 
No. 4336 about 4 km northeast of El Mamon, some 20 m west 
of the Chiguaje fault. Campo El Mamon is 2.5 km north of 
Urumaco. 

Diagnosis: Differing from M. niger as follows: skull more 
robust; snout deeper, hea\'ier; posterior portion of maxilla 
wider; interorbital bar more arched anteroposteriorly, not con- 
cave above, thicker dorsoxentrally; preorbital ridges on snout 
faint rather than strong; central portion of posterior border of 
cranial table transverse, not curved; palatine fenestrae shorter; 
12 maxillaPv' teeth, not 13-14, posterior ones larger; mandible 
heavier, thicker; external mandibular foramen not as deep. 
(For measurements see Table 1.) 

Discussion: I have examined six skulls of M. niger, fi\e in the 
American Museum of Natural History and one in the Museum 
of Comparative Zoologv. The skull of this species has been 
well described by Mook, (1921), Kahn (1933), and Medem 
( 1 963 ) . The last two authors figure and discuss growth stages, 
Kalin on the basis of 22 specimens ranging in occiput-snout 



iNamed for Dr. Daniel C. Fisher, finder of both specimens. By an odd 
coincidence, the type was the first and MCZ 4336 the last specimen to be 
collected from the Urumaco Formation bv the expedition. 



1976 MELANOSUCHUS FISHERI SP. NOV. 3 

length from 158.5 to 508 mm, Medem on 9 ranging from 99 
to 466 mm. Kalin's series came from Isla Marajo, north- 
eastern Brazil, Medem's from southern Colombia, thus nearly 
from opposite extremes of the species range. 

It can be stated with confidence that the diagnostic char- 
acters of AI. fisheri do not fall within the limits of individual 
or geographic \-ariation or of ontogenetic change in M. niger. 
The degree' of arching of the interorobital bar in the type 
(MCZ 4336 is crushed down in this region) is not approached 
even in the youngest specimen in Medem's series, and the depth 
of the bar evidently does not increase, relatively, with age in the 
living species. The conca\ity in the dorsal surface of this bar in 
M. niger tends to become shallow, although not to the point of 
disappearance, in large individuals, but in those comparable in 
size to the two specimens of M. fisheri it is \ery well defined. 
All specimens of the living species have very prominent ridges 
on the snout feven the youngest shows some traces of them), 
a conspicuous difference from the extinct form in which they 
are ver\' feebly expressed. The two last differences are asso- 
ciated; the ridges on the snout of M. niger that run postero- 
medially from the x'icinity of maxillary tooth 4 merge with the 
sides of the interorbital bar and contribute to its concavity. The 
posterior border of the cranial table is quite or nearly transverse 
in the youngest specimens of M. niger figured by Kalin and by 
Medem, but it rapidly becomes cur\-ed anteriorly with advanc- 
ing age, clearly differing from that of M. fisheri. In both speci- 
mens of the extinct species the number of maxillary teeth can 
be seen to be 12 in number, one or two less than in the living 
form, with the posterior ones being larger. The skull of M. 
fisheri is more robust than that of its relati\'e, as is shown by the 
more massi\e snout and posterior portion of the maxilla, the 
heavier mandible, and the more solidly constructed interorbital 
and ectopterygoid bars. The external mandibular foramen is as 
long as that of M. niger but is decidedly shallower, with the 
surangular in consequence being deeper. This feature and the 
smaller size of the palatine fenestra are probably also associated 
with the greater robustness of the skull of AI. fisheri. For the 
rest, the skulls of the two species are similar, the suture patterns, 
to the extent that these can be determined in the fossils, being 
essentially identical. The palatal portions of the \omers are 
clearly visible in MCZ No. 4336 as small, irregularly rhomboidal 
figures. The largest skull of AI. niger recorded by Kalin is 
508 mm in length. Whether or not Ai. fisheri attained to com- 
parable size, or exceeded it, is of course uncertain. 



4 BREVIORA No. 438 

Relationships between the two species must remain specula- 
tive for the present. As indicated above, nearly every character 
in which they differ is associated with the more robust skull of 
M. fisheri. If e\'olution within Melanosuchus proceeded in the 
direction of a lightening of the skull (in which case the more 
prominent ridges on the snout of M. niger could be interpreted 
as remnants of a prexiously more robust structure remaining 
along lines of stress) then an ancestor-descendant relationship 
would be likely. If not, the ancestor of the living species has 
yet to be found. 

Melanosuchus fisheri occurs together with fi\'e other croco- 
dilian species in the Urumaco Formation: Dinosuchus terror 
Gervais, Mourasuchus amazonensis Price, Gryposuchus jessei 
Giirich, Ikanogavialis gameroi Sill, and Balanerodus? sp. 
(Bryan Patterson, personal communication). Ikanogavialis is so 
far known only from the Urumaco. Of the others, none of which 
has hitherto been found in Venezuela, Balanerodus is known 
from the Miocene (Friasian) of Colombia (Langston, 1965) 
and the rest from the late Tertiary (Huayquerian?) of western 
Brazil. All are of very large to gigantic size. Compared to any 
of them Melanosuchus fisheri, even if it reached the dimensions 
of M. niger, was a pygmy. This crocodilian assemblage and 
the numerous chelonians associated with it suggest that the 
Urumaco area may at the time have had some resemblance to 
the Amazon basin, and bring some support to Wood and de 
Gamero's views (1971) concerning late Cenozoic geomorpho- 
logic changes in northwestern \^enezuela. 

Notes on the measurements (Table 1): Kalin's system of 
measuring (1931, fig. lA) has been followed, but additional 
measurements, those of Medem and those that illustrate certain 
diagnostic features, are also gixen. VV^here\er possible Kalin's 
indices (1931: 536-8, 663-679) have been calculated. With 
one exception, these fall within or very close to his ranges for 
M. niger based on specimens 200 mm and o\'er in skull length, 
the size at which adult proportions appear to be attained. The 
exception is snout length X 100/skull length (Kalin's No. 3), 
which is 51.21 in the' type and 52.34 in MCZ 4336, well below 
the range, 57.20' to 68.80 (n 13) for the Brazilian sample of 



iKalin lists an index of 47.70 for a skull 356..") mni in Iciigih. Either this 
specimen is decidedly anomalous (with a snout rclati\tiv shorter than 
Medem's 99.0 mm long skull) or a printer's error has occurred; I suspect 
the latter. 



1976 MELANOSUCHUS FISHERI SP. NOV. 5 

M. niger. This is too great a difference to be attributable to 
the distortion of the fossils; the snout is shorter relati\'e to skull 
length in M. fisheri. However, calculations of the index from 
Medem's measurements of specimens 200 mm and over show 
the Colombian sample fn 6) to range from 51.35 to 58.5 — 
thus on the whole shorter-snouted than the Brazilian — with 
the two fossils falling either just below or just within this. 

I also gi\'e measurements of MCZ 4043, the skull of AI. niger 
described by Mook, a specimen that has the advantage, from a 
comparati\e standpoint, of being close to MCZ 4336 in size. 
It was obtained by Louis Agassiz from "the Rio Madeira" 
(perhaps not far from its junction with the Amazon). 

Acknowledgments: I am most grateful to Dr. Jose F. Bona- 
parte, Institute Lillo, Tucuman, and Prof. Zulma de Gasparini, 
Museo de La Plata, for encouragement and advice; to Arnold 
D. Lewis for his help in preparing specimens; to Laszlo Meszoly 
for aid with the drawings; to Prof. Ernest E. VV^illiams and the 
American Museum of Natural History for access to specimens 
of Melanosuchus niger; and to Dr. R. C. Wood and Prof. Bryan 
Patterson for guidance and suggestions in preparing this paper. 
Prof. Maria Lourdes de Gamero, Universidad Central de Vene- 
zuela, Prof. Abdem Ramon Lancini, Director of the Museo de 
Ciencias Naturales de Caracas, and Prof. Eduardo Lira Espejo, 
Director of the International Department of INCIBA ^ Instituto 
nacional de Cultura y Bellas Artes de Venezuela) very kindly 
arranged the financial support that made my work at Har\-ard 
possible. Field work in \'enezuela by the Museum of Com- 
parati\e Zoology was financed by National Science Foundation 
Grant No. GB-32489 to Prof. Patterson. The photographs are 
by Alphonso Coleman. 




Figure 1. Melanosurhus fish 
showing the vomers. X i/^. 



si>. iiov. MCZ 4336. ventral view of skull 




Figure 2. Melanosuchus fisheri sp. nov. MCZ 4336, ventral view of skull. 
X Vz- The apparently small size of the premaxillarv foramen may be due 
in part to some overlap of the two sides of the specimen in this area. 



BREVIORA 



No. 438 




Figure 3. Melanosuchus fisheri sp. nov. Type, MCXC 243, dorsal view. X Vi- 



1976 



MELANOSUCHUS FISHERI SP. NOV. 




Figure 4. Melanosuchus fisheri sp. nov. Type, MCN'C 243. dorsal view. X Vz- 



10 



BREVIORA 



No. 438 




:5J 

X 



be 



1976 MELANOSUCHUS FISHERI SP. NOV. 11 




Figure 6. Melanosiichus fisheii sp. nov. Tvpe. MCNC 243. occipital view. 
XVi- 



12 



BREVIORA 



No. 438 



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References Cited 

RXlin, J. 1933. Beitrage zur vergleichenden Osteologie des Crocodiliden- 

schadels. Zool. Jahrb. (Anat. Ontog.) . 57: 535-714. 
Langston, W. 1965. Fossil crocodilians from Colombia and the Cenozoic 

history of the Crocodilia in South .America. I'niv. Calif. Pub. Geol. Sci. 

52: I-I57. 
Medem, F. 1963. Osteologfa craneal, distribuci6n geogrdfica y ecologfa de 

Melanosuchus niger (Spix) (Crocodilia, Alligatoridae) . Rev. Acad. 

Colomb. Sienc. Exact. Fis. Nat. 12 (45) : 5-19. 
MooK, C. 1921. Skull characters of Recent Crocodilia, with notes on the 

affinities of the Recent genera. Bull. Amer. Mus. Nat. Hist.. 44: 123-268. 
Pascual, R. and M. L. Diaz df Gamero. 1969. Sobre la presencia del 

genero Eiimegamys (Rodentia, Caviomorpha) en la FonTiaci6n Urumaco 

del Estado Falc6n (Venezuela) . Su significaci6n cronologica. Bol. In- 

formativo, Assoc. Venezolana Geol. Min. Pet., 12: 369-387. 
P.xscuAi , R. and O. Odrrman Ruas. 1973. Las unidades estratigrdficas del 

Terciario portadores de mamifcros, su distribuci6n y sus rclaciones con 

los acontecimientos diastroficos. .^ct, Quinto Congr. Geol. Argentine, 3: 

293-338. 
Wermiith. H. 1953. Svstematik der lezenten Krokodile. Mitt. Zool. Mus. 

Berlin, 29: 376-514. 
AVooD, R. C, AND M. L. Diaz de Gamero, 1971. Podocnemis iienezuelensis, 

a new pelomedusid (Testudines, Plcurodira) from the Pliocene of 

Venezuela and a review of the historv of Podocnemis in South .\merica. 

Breviora No. 376; 1-23. 



><^ ''^'''^'3. CCMP. ZOOL 

Linr?ARY 

MAR 1 8 1985 

"-^RVARD 

B R E V"1~0 R A 

Museum of Comparative Zoology 

us ISSN 0006-9698 



Cambridge, Mass. 30 July 1976 Number 439 

ON A NEW SPECIMEN OF THE 

LOWER CRETACEOUS THEROPOD DINOSAUR 

DEINONYCHUS ANTIRRHOPUS 

John H. Ostrom 

Abstract. A new spctimcii of Deinonychui antirrliopus, described here, 
provides morphological and dimensional data on skeletal elements missing 
in previously known specimens. Included are the femora, pubes. complete 
ilia and the sacrum. The pubis is normal in its morpholog)', but unusual in 
its length (more than twice that of the ischium) and orientation (appar- 
ently ventrad) . The femur is shorter than the tibia and features an unusual 
"posterior trochanter" that may have been the attachment site of muscles 
that poweret! the offensive leg kick and use of the pedal talon. 

INTRODUCTION 

My original report.s on the unusual theropod dinosaur Dei- 
nonychus antirrhopus (Ostrom, 1969a, 1969b) necessarily were 
incomplete because some skeletal elements were not then repre- 
sented in collections. Specifically, much of the po.storbital region 
of the skull was not known, nor were the femur, pubis or sacrum. 
These deficiencies now have been partially resolved thanks to 
the discovery in July, 1974 by a Harvard University expedition 
to Montana, of another specimen of Deinonychus antirrhopus. 
The purpose of this paper is to provide descriptions of the pre- 
viously unknown elements that are provided by the new speci- 
men, and to correct certain inaccuracies that were presented in 
my earlier studies of this remarkable species. 

Abbreviations of institutional names cited herein are as fol- 
lows: 

AMNH — American Museum of Natural History, New 
York City. 



BREVIORA 



No. 439 



MCZ — Museum of Comparative Zoology, Harvard Uni- 
versity. 

YPM — Pcabody Museum of Natural History, Yale Uni- 
versity. 

Materials: A partial skeleton (MCZ 4371, Field No. 74M 
#7) including fragments of the snout and mandibles with 
teeth, a nearly complete but poorly preserved vertebral series 
including the entire tail, the right forelimb and manus, frag- 
ments of the left manus, a complete pelvis and sacrum, both 
hind limbs and feet, and various rib and gastralia fragments; 
discovered by Steven Orzack and collected by F. A. Jenkins 
and party, July 1974. 

Geologic and Locality Data: Upper part of Unit V (see 
Ostrom, 1970), otherwise known (in part) as the Little Sheep 
Mudstone Member of the Cloverly Formation ( Moberly, 1 960 ) , 
Lower Cretaceous. The quarr\' site is situated 1 km south of 




Deinqnychus Localities 



Fig. 1. Locality map showing the Harvard Deinonychus 
site, together with the previously known American Museum 
sites in southern Mo"*ana. 



1976 



DEINONYCHUS ANTIRRHOPUS 







Si 






bo 



4 BREVIORA No. 439 

the Cashen Ranch house, NE 1/4 Sec. 32, T. 4 S., R. 29 E., Big 
Horn County, Montana. This site is within a few hundred 
meters of the two American Museum Deinonychus sites (see 
Fig. 1) reported previously (Ostrom, 1969b), and appears to 
be at approximately the same stratigraphic level as that of 
AMNH 3037. If my previous correlations (1969b, 1970) are 
correct, this level is well below that of the Yale Deinonychus 
Quarry near Bridger, Montana, which occurs in the lower part 
of Unit VII (equals part of the Himes Member of the Cloverly 
Formation as interpreted by Moberly, 1960). 

DESCRIPTION 

Excluding many isolated teeth of Deinonychus reported from 
numerous Cloverly sites (Ostrom, 1970), the present specimen 
brings the total minimum number of Deinonychus specimens to 
six individuals. The collections from the Yale Bridger Quarry, 
however, consist almost entirely of disarticulated remains that 
may well include more than the three individuals represented by 
the three articulated caudal series recovered there. As Table 1 
shows, the new specimen is the largest of all presently known 
specimens, and together with AMNH 3015 provides the first 
evidence of growth patterns and allometry in this taxon. The 
following descriptiv^e material deals primarily with those skeletal 
elements that were not known at the time of the earlier studies. 
These include the femur, pubis and sacrum, and certain aspects 
of the ilium, pes and metatarsus. 

Femur: Contrary to my earlier guess, the femur of Deinony- 
chus is shorter than the tibia by about 10 per cent. This is not 
an unusual difference, nor is it unusual for the tibia to exceed 
the femur length in theropods. But this new fact does require a 
reassessment of my earlier interpretation about the cursorial 
capacity of Deinonychus — a reassessment that will be discussed 
briefly later. 

The femur of Deinonychus is moderately robust, with a slight 
antero-posterior curvature and of hollow construction (Fig. 3). 
The head is sharply offset medially from the shaft by means of 
a stout but distinct neck. The axis of the neck projects trans- 
versely (parallel to the plane formed by the distal condyles) at 
about 100 degrees to the femoral shaft. The head is roughly 
ovoid and broadly convex, but with a slightly flattened area 



1976 



DEINONYCHUS ANTIRRHOPUS 

B C 



D 




Fig. 3. Left femur (restored) of Deinonychus antirrhopus 
(MCZ 4371) in posterior (A), medial (B), anterior (C), and 
lateral ( D ) views. Abbreviations : ec — external condyle ; gt — 
greater trochanter; he — head; ic — internal condyle; It — les- 
ser trochanter; pt — posterior trochanter. 



anterodorsally and with a distinct proximal posterorinferior lip 
or overhang that sets the head off from the inferior region of 
the neck. Externally, the neck expands dorsally into a large 
"greater trochanter," which presumably marks the attachment 
site of the M. iliotrochantericus. External and slightly anterior 
to this feature is a robust, but not large, "lesser trochanter" 
(anterior trochanter of some authors). Unlike some theropods 
(i.e., Allosaurus), this last structure is separated from the 
"greater trochanter" by only a narrow shallow groove rather 
than by a deep cleft. The "lesser trochanter" of theropods is 
generally considered as the insertion site of the M. ilio femoralis. 



6 BREVIORA No. 439 

Distal to these "trochanters" and posterior to the "lesser tro- 
chanter," is a distinct prominence or boss which projects latero- 
posteriorly. Similar, but much less prominent features occur on 
many other theropod femora, but it has received little or no 
attention and has not been given a formal designation. In this 
paper, for lack of a better term, it is referred to as the "posterior 
trochanter." Its significance is not known, but it is situated in 
the general region where we might expect the M. ischiotro- 
chantericus (= M. ischiofemoralis of birds?) to insert. More 
will be said of this later. 

The uncrushed portions (near midlength) of the shaft are 
nearly circular in section with a minimum transverse diameter 
of approximately 38 mm and an anteroposterior diameter of 
about 32 mm. Dimensions increase slightly toward both ends. 
Surprisingly, no recognizable fourth trochanter is preserved in 
either femur, nor is there any discernible scar indicating the 
insertion site of the M. caudifemoralis. Distally, the condyles 
are well developed and sharply separated in posterior (ventral) 
aspect by a deep intercondylar groove. The external condyle is 
nearly twice as large (both proximodistally and in anteropos- 
terior dimension) as the internal condyle. Femoral and other 
dimensions are given in Table I. 

Pubis: Perhaps the most important aspect of the new speci- 
men is the presence of both pubes. In my original study of 
Deinonychus (Ostrom, 1969b), I tentatively but incorrectly 
identified an isolated bone of curious shape as a right pubis, 
chiefly because it was found in the quarry immediately adjacent 
to a right ischium of about the same length. Subsequently I 
became convinced that it could not possibly be a pubis and 
after further study concluded that it was a right coracoid of 
unusually large size (Ostrom, 1974). 

The pubes of Deinonychus are of normal theropod design 
(Fig. 4), but of surprising length relative to the short length 
of the ischium. In fact, the pubis is more than double the 
length of the ischium, a condition that is unique among thero- 
pods. Although crushed in some regions, the major features are 
all discernible. The proximal end is moderately stout, but the 
actual articular surfaces for contact with the robust pubic pe- 
duncle of the ilium and the thinner peduncle of the ischium are 
not recognizable. There appears to have been a shallow obtu- 
rator notch immediately adjacent to the ischial suture, as shown 



1976 



DEINONYCHUS ANTIRRHOPUS 





Fig. 4. Pubes (restored) of Deinonychus antirrhopus (MCZ 
4371) in anterior (A) and left lateral (B) views. 



by remnants on the left side. The pubic shafts are oval in sec- 
tion, with the greatest diameter in the transverse direction. Just 
short of mid-length, the shafts expand medially and the two 
pubes join in a massive symphysis that measures 210 mm, or 
more than half the total pubic length. Over most of the sym- 
physial length, the pubes are quite broad, forming a transverse 
apron of 60 to 67 mm width. The distal extremity contrarily 
is narrow transversely, but expanded longitudinally into the typi- 
cal theropod "footUke" structure. This pubic "foot"' is approx- 
imately 135 mm long (anteroposteriorly), but only 35 mm in 
its maximum preserved width. 

There is no doubt any longer about the form of the pubis, but 
doubts still exist about the in vivo position of the pubes (and 



8 BREVIORA No. 439 

perhaps of the ischia as well) because of the manner in which 
the new specimen is preserved. The animal was buried lying on 
its ventrum, with the pubes pushed up and backward, tightly 
pressed against the ischia and the underside of the tail, and 
oriented parallel to the proximal caudals and the axis of the 
sacrum. The fact that both the pubes and the ischia are pre- 
served tightly pressed together and directed backward suggests 
that in life these two bones may not have been arranged as 
di\'ergently as in most other theropods. It further suggests that 
the pubes may have been oriented nearly perpendicular to the 
sacral axis, or perhaps even slightly posterior to that perpen- 
dicular. In the revised skeletal reconstruction (Fig. 2), I have 
so oriented the pubes at right angles to the long axis of the ilium 
and the sacrum. 

Ilium: Prior to the discovery of the Harvard specimen, only 
one incompletely preserved ilium (AMNH 3015) was known 
of Deinonychus. Fortunately, both ilia are preserved in MCZ 
4371, and a more accurate description is now possible. Un- 
expectedly, the ilium turns out to be much longer than I 
concluded from the American Museum specimen, with both 
extremities tapered or triangular, rather than rectangular in 
outline. In lateral aspect, the iliac blade is long, quite low, 
and with nearly uniform height throughout, except at the ex- 
tremities. The anterior process is longer than the posterioi^ 
process, as measured from the center of the acetabulum. Except 
for the tapered extremities, the long low profile and the off- 




Fig. 5. Left ilium (restored) of Deinonychus antirrhopus 
(MCZ 4371) in lateral view. Abbreviations: act — acetabu- 
lum; is — ischiac peduncle; pu — pubic peduncle. 



1976 



DEINONYCHUS ANTIRRHOPUS 



center position of the acetabulum resemble the conditions of 
coelurosaurs in general, and those of omithomimids in particular. 
As was noted in my earlier description of the American 
Museum ilium, the pubic peduncle is much more massive and 
longer than the ischiac peduncle, and, perhaps more important, 
it is directed downward and backward, rather than down and 
forward. The significance of this unusual orientation was not 
recognized 'then, but now in the light of the preserved position 
of the pubes in MCZ 4371, it seems reasonable to equate this 
with a possible ventrad or ventroposterior orientation of the 
pubes. Figure 6 shows my best estimate of the pelvic arrange- 
ment in Dcinonychus, compared with Strut hiomimus and Ty- 
rannosaurm. 

Ischium: Although both ischia are present in MCZ 4371, 
they add little to our knowledge of ischial morphology beyond 
that which was provided by previous specimens. However, the 






Fig. 6. Comparison of the pelves (lateral views) of Dcino- 
nychus, MCZ 4371, (A); Struthiomimus, AMNH 5339, (B); 
and Tyrannosaurus, AMNH 5027, (C); to show the unusually 
disproportionate lengths of the pubis and ischium in Dcinony- 
chus. The scale lines equal 10 cm. 



10 BREVIORA No. 439 

new specimen does establish that the ischia were united at their 
distal extremities by a well-developed symphysis. Whether the 
symphysis extended proximally to the level of the obturator 
flange, as I suggested previously (Ostrom, 1969b) on the basis 
of the surface texture of that process, cannot be established here. 
The anterior ischial surfaces are concealed by the broad pubic 
apron, which has been crushed against the ischia, and the pos- 
terior surfaces likewise are covered by the proximal caudal 
vertebrae that were pressed down against them. 

Sacrum: The sacral series, absent in previously known speci- 
mens, is difficult to decipher because the anterior portion is in- 
complete and the full dorsal vertebral count is not known in 
MCZ 4371. On the basis of the degree of co-ossification and 
the form of the sacral ribs, I conclude that the sacrum con- 
sisted of five segments in which the centra and neural spines 
were fused together. A sixth centrum, presumably representing 
the last dorsal, seems to be fused to the first sacral. This seg- 
ment, and the one behind it, has relatively weakly developed 
sacral ribs or transverse processes as compared with those of the 
following segments, but these anterior segments seem to have 
had firm articulation with the anterior processes of the ilia. On 
that basis, both might be considered as dorsal, rather than sacral 
vertebrae. 

Pes: Both feet and metatarsi are well preserved and appar- 
ently complete in the new specimen, offering verification of most 
of my original interpretations, but also providing a few new 
details. The new specimen suggests that the first metatarsal 
probably consisted of a distal portion only, rather than separated 
distal and proximal moieties. There is no evidence in either 
foot (both of which are otherwise complete) of a proximal 
portion of metatarsal I, as I suggested earlier (1969b, fig. 72). 
Since both feet and metatarsi in this specimen were preserved 
fully articulated, it seems unlikely that this particular element 
should be accidentally missing in both. 

Also, as preserved in MCZ 4371, the first digit may not have 
been reverted to the rear as far as I originally restored it ( 1 969b, 
Fig. 74 ) . However, since the first metatarsal was not fused to 
the second, and appears to have been ligamentously joined to 
the metatarsus, its preserved position (in the absence of any 
clear appositional scar) may or may not approximate the natural 
position in life. 



1976 DEINONYCHUS ANTIRRHOPUS 11 



MC Z 43 71 



Y PM 5205 



Fig. 7. Contrasting morphology of the second pedal ungual 
in the type specimen of Deinonychus (YPM 5205) and the 
new Harvard specimen. 



One final observation that pertains to the distinctive "Dei- 
nonychus" character preserved in the new specimen: that is the 
very different geometry of the foot talon. As Figure 7 shows, 
the claw in the type specimen (YPM 5205) is much more 
strongly curved than is that of MCZ 4371. Preservation of the 
type ungual is virtually perfect with no distortion or crushing 
and only a few small fragments missing. That is not true of the 
new specimen, but what distortion is apparent here does not 
seem to account for the conspicuous difference in shape. Barring 
preservational distortion, the difference could be due to individ- 
ual, ontogenetic, or sexual variation. At this point, I have no 
opinion on the correct explanation, but simply report the dif- 
ference. 



DISCUSSION 

In my original study (1969b) of Deinonychus, I assumed 
equal lengths for the femur and tibia, in the absence of any 
knowledge about the femur. The new specimen shows that the 
femur is shorter (approximately 90 per cent of tibia length) 
than I had thought and thus is similar to Strut hiomimus. This 
might indicate higher running speeds than I suggested in that 



12 BREVIORA No. 439 

study ( "moderately, but not unusually fast" ) . However, more 
important than fenun- ^tibia ratio as an index of cursorial ve- 
locity, is the lensjth of the metatarsus relative to the tibia. In 
the American Museum specimen (3015) of Deinonychus, this 
ratio is a surprisingly low .48. In MCZ 4371, it is .45. These 
values are well below those of anv other theropod now known, 
and when compared with those of such well-known fast-running 
birds as Strut hio (.95), Dromaius (.96) and Casiiarius (.85) 
(where the femur is onlv about half as long as the tibia) and 
the presumed fleet-footed Struthiomimus (.68), the only reason- 
able conclusion is that Deinonxchiis was not the most fleet-footed 
of theropods. 

This conclusion is reinforced bv other data revealed by the 
new specimen. MCZ 4371 is a larger individual than AMNH 
3015. but not imiformlv so. The forelimb is 20 per cent longer 
in MCZ 4371 and the tibia 18 per cent longer, but the meta- 
tarsus is only 12 per cent longer than in .-VMNH 3015. If the 
above .similar tibia and forelimb \alues represent an overall 
average difference in size between these two individuals, then 
metatarsal growth did not keep pace during ontogeny with the 
growth of other long bones. The metatarsal fraction of total 
hind limb length diminished with age, suggesting a correspond- 
ing decline in cursorial ability. At the time of mv earlier study, 
I was puzzled by the relative brevity of the metatarsus, but 
suggested that its unusuallv short length may have been cor- 
related with the specialized offensi\-e claw of the second toe. 
I am still of that opinion, especiallv in \icw of the apparent 
ontogenetic decline in relati\'e metatarsal length revealed bv the 
Har\-ard and American Museum specimens. 

The design and orientation of the sicklelike claw on digit II 
show that the offensi\'e or damaging stroke of this w'eapon was 
flexion — • a strong backward thrust of the claw and toe. This 
probablv was coordinated with a powerful backward kick of 
the entire hind leg. This means that the combined power of all 
the hind limb retractor muscles those that are usually invol\-ed 
in the locomotory power stroke) may have contributed to the 
action of this unusual and presumablv lethal device. Consider- 
ing the size and shape of the pedal talon, it is verv likelv that 
the toe and the metatarsus were subjected to considerable ex- 
tensional and shear stresses when this weapon was employed ■ — 
possibh- nuich higher stresses than usual during normal loco- 



1976 DEINONYCHUS ANTIRRHOPUS 13 

motory action of the foot and leg\ If so, what were the prin- 
cipal muscles available to counteract such stresses in these re- 
gions? Presumably, they were the equivalents of the M. gas- 
trocnemius and the digital flexors (particularly the M. flexor 
perferans et perforatus digiti II and M. flexor perforatus digit! 
II) of modern birds. As the primary flexor of the ankle, the 
gastrocnemius is the chief muscle available to oppose extensional 
stresses applied to the metatarsus and the foot. But as the chief 
ankle flexor, the mechanical requirements of the gastrocnemius 
dictate a proximal insertion on the posterior surfaces of the 
metatarsus. This arrangement, with its extremely short lever 
arm, provides little or no added strength to the metatarsal shafts 
against hyperextension or fracture. However, a shortened meta- 
tarsus would reduce the length of the resistant (hyperexten- 
sional) lever ami and thus can be considered as an adaptation 
to minimize the possibility of metatarsal hyperextension or frac- 
ture. This explanation is further supported by the fact that there 
is no positixe e\idence of an enlarged or particularly powerful 
"gastrocnemius." There are no tubercles evident on the femur 
distal extremity that might be the origin sites of such a muscle, 
and there is no hypotarsus-like structure, or any other evidence 
of muscle scars, on the proximal posterior surfaces of the meta- 
tarsals. 

Speculative though the above interpretations may be, they 
are further supported by a comparison of hind limb proportions 
in Deinonychus with those of both cursorial [Struthiomimus) 
and graviportal (Tyrannosaurus) theropods (see Fig. 8). The 
propodial — epipodial proportions of Deinonychus resemble 
those of Struthiomimus, but the metapodial — pes proportions 
resemble those of Tyrannosaurus (although not as massive). In 
the latter, the short metatarsus can be viewed as related to the 
great weight-bearing (and shear stress) problems in Tyranno- 
saurus (estimated live weight, 7,000 kg). In Deinonychus, how- 
ever, any excessi\'e shear stresses across the metatarsus must have 

lit is interesting that the Yale Deinonychus collection includes a damaged 
second phalanx of digit II which was fractured and healed during the ani- 
mal's lifetime. The broken extremities were displaced transversely and 
mended in this position, but perfectly aligned longitudinally. The longitu- 
dinal alignment apparently allowed the claw to function normally after 
healing, with only a sideways displacement of the arc of claw flexion. This 
specimen clearly indicates that the talon-bcai ing toe of Dt'inonvchus was 
subject to intense stress. 




Fig. 8. Comparison of hind limb proportions in Struthio- 
mimus, AMNH 5339, (A); Deinonychus, MCZ 4371, (B) ; 
and Tyrannosaurus, AMNH 5027, (C). Notice that the prox- 
imal elements of Deinonychus resemble those of cursorial 
Struthiomimus, but the distal proportions are more like those of 
graviportal Tyrannosaurus. The solid arrows indicate the loca- 
tion and approximate orientation of the chief locomotory mus- 
cle, the M. caudi-femoralis. The dashed arrow (B) indicates 
the possible position of the M. ischio-trochantericus, a second- 
ary femoral retractor which may have powered the offensive 
leg stroke. Scale lines equal 10 cm. 



1976 DEINONYCHUS ANTIRRHOPUS 15 

been derived from sources other than weight (which is estimated 
at less than 75 kg ) . Use of the foot talon against a larger prey 
animal might well have resulted in high stresses across the 
metatarsus, thereby favoring a shortened metapodial component. 
That could account for the peculiar distal proportions combined 
with the otherwise "cursorial" design of the tibia and femur. 

The above observations lead to one further speculation about 
the hind limb of Deinonychus and its probable actions. As 
noted in the descriptive section, there is a well-developed promi- 
nence (referred to here as the posterior trochanter) on the outer 
posterior proximal surface of the femur just distal to the "greater 
trochanter." This feature is present in many other theropods, 
where it is far less prominent, but as far as I am aware, it has 
not received any particular attention or interpretation. Even 
though any analogy here with modern birds may be question- 
able, it is interesting to note that a bony prominence also occurs 
in this same region in many modern birds. This avian feature 
is the site of insertion of the M. ischio-femoralis ( ^ the M. 
ischio-trochantericus of reptiles), one of the major retractors of 
the hind leg. Traditionally, the M. caudi-femoralis is considered 
the principal femoral retractor in modern reptiles (and espe- 
cially in the extinct bipedal theropods and ornithopods), pro- 
viding most of the stride power. The M. caudi-femoralis inserts 
on the fourth trochanter, or on a conspicuous scar near mid- 
shaft of the femur. This insertion site is well developed in 
virtually all reptiles, and particularly so in all theropods. The 
absence of a fourth trochanter in Deinonychus, or of any recog- 
nizable scar that can be equated with the caudi-femoralis, is a 
curious anomaly that I am unable to explain. 

I am not suggesting that the M. caudi-femoraHs was absent, 
or even reduced, or that the M. ischio-trochantericus (with its 
lesser leverage) had assumed part or all of the locomotory func- 
tion of that muscle. Rather, I wonder if the unusual prominence 
of the "posterior trochanter" here may not be related to the 
special predatory foot adaptations in Deinonychus which ob- 
viously were designed for application in a backward "killing" 
stroke — a motion quite separate and (perhaps necessarily) 
independent of limb movements concerned with locomotion. 
Turning our attention to another unusual feature of Deinony- 
chus, recall the extraordinary brevity of the ischium and the 
proximal placement of the obturator flange — a possible origin 
site of the M. ischio-trochantericus. I cannot help wondering 



16 BREVIORA No. 439 

about the possible connection between these three unusual fea- 
tures of Deinonychus: the offensive foot talon; the abbreviated 
ischium with a proximally situated obturator flange; and the 
prominent posterior trochanter of the femur. The last two 
features could well have been the origin and insertion sites of 
the ischio-trochantericus, which could have powered the offen- 
sive backward stroke of the hind leg, as opposed to the backward 
locomotory power stroke of the leg. TTie locomotory p)ower 
stroke almost certainly was produced by contraction of the 
caudi-femoralis, despite the apparent absence of an insertion 
scar or a fourth trochanter. 

Why should the offensive backward stroke of the femur be 
powered by a different set of muscles from the more usual loco- 
motory stroke? Perhaps the answer lies in the fact that the most 
important locomotory muscle of the reptilian hind leg is the 
M. caudi-femoralis, which originates on the proximal caudal 
vertebrae — an origin site which is not immovably fixed, and 
which also is the power base of the tail. The tail obviously is 
the critical balancing appendage, especially in bipedal reptiles 
(both obligate and facultative). Bipedal progression and dy- 
namic balance must be coordinated, so it is not surprising that 
it is the locomotory musculature (M. caudi-femoralis) that is 
involved with both the propulsive leg stroke and the balancing 
actions of the tail. On the other hand, the same kind of linkage 
between the mobile balancing appendage and an offensive stroke 
of the hind leg might be counterproductive. Precise equilibra- 
tion during any offensive leg stroke would be absolutely essential 
for an accurate and effective killing stroke, but if the same 
muscles contributed both to movements of the balancing tail 
and the offensive leg kick during an aggressive encounter, then 
the two actions could not be produced independently. Precision 
of both movements would be seriously impaired. If muscle 
contractions powering the offiensive leg stroke also produced 
deflection of the balancing tail, or vice versa, one could well 
be detrimental to the other. Thus it seems logical to conclude 
that the musculature that powered one action must have been 
freed as much as possible from producing or contributing to 
the other action. The M. caudi-femoralis, as it is organized in 
modern reptiles, cannot meet this requirement. A femoral 
retractor that does satisfy this constraint, though, is the M. 
ischio-trochantericus with its immobile origin on the ischium, 
rather than the tail. 



1976 DEINONYCHUS ANTIRRHOPUS 17 

SUMMARY 

There are no absolute explanations of fossil evidence. The 
preceding discussion has been offered only as possible reasons 
for the combination of unusual osteological characters that are 
peculiar to Deinonychus and other dromaeosaurid theropods 
(Dromaeosaurus, Velociraptor, Saurornithoides, Stenonychosau- 
rus). These' features are: 1) the sicklelike claw of the second 
toe; 2) the short metatarsus; 3) a prominent posterior tro- 
chanter; 4) a long pubis and an unusually short ischium with a 
proximally placed obturator flange; and 5) an unusual caudal 
series with extensive ossified tendons. The foot clearly functioned 
both for locomotion and as a predatory weapon. In that light, 
the hind limb proportions seem best explained as a compromise 
between high cursorial requirements (femur/tibia ratio) of a 
predator and the need for a strong metatarsal foundation for 
the foot weapon. The short ischium and prominent posterior 
trochanter are interpreted as the probable attachment sites of 
the offensive leg musculature (M. ischio-trochantericus), and 
the unusual caudal tendons (absent in non-dromaeosaurid 
theropods) reflect the extreme equilibration requirements of 
Deinonychus and its allies. 

ACKNOWLEDGEMENTS 

I am indebted to F. A. Jenkins, Jr., who recovered the speci- 
men herein described, and who very generously made it avail- 
able for me to study. 

LITERATURE CITED 

MoBERLv, R. I960. Morrison, Cloverly and Sykes Mountain formations, 
northern Bighorn Basin, Wyoming and Montana. Bull. Geol. Soc. 
Amer. 71: 1137-1176. 

OsTROM, J. H. 1969a. A new theropod dinosaur from the Lower Cretaceous 
of Montana. Postilla, Peabody Mus. Nat. Hist. 128: 1-17. 

1969b. Osteology of Deinonychus antirrhopus, an unusual 

theropod from the Lower Cretaceous of Montana. Bull. Peabody Mus. 
Nat. Hist. 30: 1-165. 

1970. Stratigraphy and paleontology of the Cloverly Forma- 
tion (Lower Cretaceous) of the Bighorn Basin area, Wyoming and 
Montana. Bull. Peabody Mus. Nat. Hist. 35: 1-234. 

1974. The pectoral girdle and forelimb function of Dei- 



nonychus (Reptilia: Saurischia) : A correction. Postilla, Peabody Mus 
Nat. Hist. 165: 1-11. 



18 



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^MivERSiTY 

Museum of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 30 July 1976 Number 440 

WEST INDIAN ANGLES: A TAXONOMIG 

AND EVOLUTIONARY SUMMARY 
1. INTRODUCTION AND A SPECIES LIST 

Ernest E. Williams^ 

Abstract. Accumulation of morphological, kaiyological and ecological 
data on West Indian anoline lizards permits and requires a taxonomic 
analysis more elaborate than usual and employing both formal and informal 
taxonomic categories. The categories are defined in this, the first paper of 
a series, and a species list of West Indian anolines displays the new arrange- 
ment. 

INTRODUCTION 

Since the pioneer study by Etheridge (1960) there has been 
a remarkable growth both in our factual knowledge and in the 
sophistication of our knowledge of one group of lizards — the 
anolines. This has been especially true in the islands of the 
West Indies. The mainland members have at the same time 
received less attention and have proved more refractory. (Cer- 
tainly the latter fact has influenced the first.) 

There is, however, nowhere any gathering together of the 
new knowledge. Partly this results from the continuing activity. 
Not even species lists have remained constant. A statement that 
Cuba or Hispaniola has x species is outdated before it is pub- 
lished. In the same way ecological information, ideas and theory 
have expanded far past the published record. It is not easy to 
keep on top of the field. 

Just because of this it is necessary that beginnings of a sum- 
man- be made. Both for old hands and for newcomers an 
exposition of how far we've come, where we are, and where 

iMuseum of Comparative Zoology, Harvard University, Cambridge, Massa- 
chusetts 02138 



2 BREVIORA No. 440 

we might go will be a useful thing. I propose as a first step a 
taxonomic-evolutionary summary. The summary is inevitably 
provisional and, more than that, intended to promote research, 
provoke criticism and encourage the search for further evidence 
and the endeavor for greater understanding. 

My objective has been to illuminate ecology and evolution. 
The taxonomic study I will here provide is one means to that 
end. The West Indian anoles are a group of enormous diversity, 
but the interest of this diversity is not in its bare existence but 
in its structure and origin - — ■ its balance, the interlocking of its 
parts and the historical paths by which this was achieved. 

This is no simple radiation — not just the checkerboard sub- 
division of some original widespread unit. Here the severed 
units have doubled back upon one another and are completely 
layered, juxtaposed and interdigitated. The fitting together 
of so many species is the problem. 

Such complexity involves several le\'els. One such level is 
generic. In the West Indies I recognize three very distinct 
genera, two genera autochthonous and old, species-poor and 
obviously relict, one genus species-rich beyond ordinary imagin- 
ing, a colonizing and expanding group, a newer invader from 
the mainlands that adjoin the West Indies to the west and 
south. The latter is, of course, the group, as full of problems 
as of interest, that must receive the maximal attention it deserves. 

In point of fact, there are few problems for the two old 
species-poor genera, but also little information. 

One of these two genera, ChamaeleoUs, is represented by two 
giant casque-headed arboreal species on Cuba which much re- 
semble the tree-crown giants of the Anolis equestris group with 
which they share the island (Garrido and Schwartz, 1968). 
The species of Chamaeleolis, however, are more primitive osteo- 
logically, more chameleon-like in moxement and appearance 
and apparently rarer than giant Anolis. 

The other of the genera, Chamaelinorops, initially erroneously 
reported from the tiny mile-square island of Navassa, west of 
southwestern Hispaniola, is, in fact, from the south island of 
Hispaniola, dwarf, ground-dwelling and extremely peculiar osteo- 
logically. Richard Thomas (1966) recognizes only a single 
species. 

Chamaeleolis and Chamaelinorops, except that they are true 
anoles sharing the characteristic adhesive pads of Anolis and 
the typical Anolis dewlap, are not close to each other nor to 



1976 WEST INDIAN ANGLES 1. 3 

Anolis. Their greatest interest lies in the possibility that they 
may represent an early (pre-Miocene ?) invasion of the Greater 
Antilles and may be relicts of an earUer island radiation of which 
we otherwise know nothing. 

The three species just mentioned apart, the remaining anoUnes 
of the West Indies are here regarded as belonging to the genus 
Anolis. Fortunately this overlarge taxon divides naturally, as 
Etheridge showed in 1960, into two sections called by Etheridge 
alpha and beta. Though this is a dichotomy based on an ap- 
parently trivial character, it makes excellent geographic sense. 
Savage (1973) has suggested that, instead of two sections, two 
genera be recognized — Anolis Daudin (type Anolis carolinensis 
Voigt) and Nor ops Wagler (type Anolis auratus Daudin). 
This would substitute formal designations for the currently in- 
formal ones but it would leave no formal (or informal) term 
for the two sections (or genera) taken together. Savage's action 
is well taken if alpha Anolis are closer to Charnaeleolis and 
Phenacosaiirm than to beta Anolis or the betas are closer to 
Chamaelinorops than to the alphas. This is a point I regard 
as at least doubtful, preferring to leave it in decent obscurity 
until there is more and better evidence. My own suggested 
phylogeny for anolines would have the alpha anoles the more 
primitive (as they certainly are in many respects), and the beta 
tvpe of caudal transverse process (which does not resemble those 
of other iguanids or of other lizard groups) arising secondarily, 
but only once, from the alpha condition in which the transverse 
process is absent. The transverse processes of Chamaelinorops 
have only a verbal similarity to those of beta anoles; I question 
the closeness of the relationship. Richard Etheridge would dis- 
agree with this scheme fundamentally. Very recent immuno- 
logical data fDessauer et al.) reported at the 1974 meetings of 
the -\mierican Society of Ichthyologists and Herpetologists ques- 
tion the fundamental distinction. I remain convinced of the 
reality of the two groups but, while so much remains contro- 
versial. I do not see the value of the formal designation; it is 
not even useful mnemonically. 

Means to Analysis of a Radiation: 
The Group Terms Utilized 

The two sections of Anolis have, according to my interpre- 
tation, provided three and only three invasions of the West 



4 BREVIORA No. 440 

Indies — one by betas into Jamaica, one by alphas into His- 
paniola, and a third by alphas into St. Lucia, (But see Yang, 
Soule and Gorman [1974] for the evidence for a landfall for 
the third invasion in Grenada instead.) All the extraordinary 
proliferation, diversity and complexity of Anolis in the West 
Indies has arisen out of these three stocks by intra-island radia- 
tion and inter-island interchange. There is therefore a formidable 
problem in analysis. 

Fortunately, part of the basic information is already available. 
A just published checklist of West Indian Amphibians and Rep- 
tiles by Schwartz and Thomas ( 1975) provides an informed and 
very careful list of Antillean taxa (including Anolis) with original 
citations and synonymies, as well as the distributions as known 
to the date of publication. The species are, however, listed 
alphabetically; no taxonomic arrangement or indication of rela- 
tionship is attempted. The taxonomic ordering presented below, 
in remedving this, endeavors to synthesize a great deal of bio- 
logical information. 

I have myself seen 72 species or members of superspecies of 
West Indian anoles in the field, six additional alive in captivity, 
and, 33 more as preserved specimens. Only three species, all 
very recently described from Cuba, are known to me only from 
descriptions {A. pygmaequestris, A. jiiangundlachi and A. fugi- 
t'wus). I have collected and studied anoles on all four of the 
Greater /Antilles, on several of the Lesser xA.ntilles and on one 
island in the Bahamas. This field knowledge I regard as basic 
to an undcstanding of the group. I have also participated in, 
encpuraged or aided studies at many other levels — osteology, 
karyotypes, electrophoresis, aut- and synecological studies. All 
of this information is utilized in the classification below. 

No classification can mirror perfectly the complexity of the 
evolutionary everits that have produced the more than 100 
species of West Indian Anolis. Nor, indeed, are the m'nuter 
details of relationship and evolutionarv sequence so well under- 
stood for likely to be) that we should attempt so perfect a 
system. Nonetheless the wealth of species to be allocated and the 
amount and variety of detail known about these same species 
seem to me to afford at once the possibility and the justification 
for an arrangement elaborate much beyond the usual. I there- 
fore utilize a number of informal terms, partly based on those 
emiployed by Etheridge in 1960, but descending into greater 
detail. I define these below. 



1976 west indian angles 1. 5 

Formal and Informal Categories Used 

Section. The primary dichotomy, a group osteologically de- 
fined at the highest level below the genus. Proposed by Etheridge 
(1960) for his alpha-beta division with Anolis osteologically 
defined on presence or absence of transverse processes on pos- 
terior caudal vertebrae. 

Subsection.^ A division setting off a major portion of a sec- 
tion, again osteologically defined. This term, not used by 
Etheridge, distinguishes punctetus and carolinensis subunits 
within the alpha section, basing them on the relationship of 
interclavicle and clavicle. In the shorthand terms used below, 
the relationship is described in terms of an arrow-shaped or T- 
shaped interclavicle. 

Series. A phyletic unit under the subsection definable on 
multiple characters. Osteological, chromosomal and even scale 
characters are utilized. Ordinarily this includes more than one 
species group and displays substantial morphological and even 
chromosomal diversity, i.e., products of a complex radiation 
that inferentially included several intermediate or annectent 
forms now extinct. 

I have found series to be the unit in terms of which evolution 
is most conveniently discussed. I shall, therefore, in succeeding 
papers mention more characters under this unit than under 
taxa at a higher or lower level. This permits higher compara- 
bility between series and puts on display also many of the 
features and conditions the evolution of which I will later trace. 

This is not to say that the greater number of characters makes 
the definition of series sharper or more rigorous. On the con- 
trary, precisely because series are the units within which evolu- 
tion is most readily seen, recognition of series is a matter of 
some subtlety — ^ as Tables 1-4 show. Morphological and 
karyological characters may broadly overlap (Table 1 ). Ecology 
and geography are major clues, but convergence in ecological 
adaptations is rampant (Table 2), and geography must be used 
with discretion. 

A balancing of all the evidence — not all of which is on record 
in Tables 1-4 — is the basis for the recognition of series. 

suhseries: a category utilized when a series has several recog- 
nizable subunits, Oidinarily including more than one species 
group. 



6 BREVIORA No. 440 

species group: the products of a simple radiation but often in- 
cluding species now widely sympatric (they may exhibit chro- 
mosomal diversity). I have sometimes used species group for 
a single species when that species is very distinct and may well 
be the last remnant of a radiation. 

species subgroup: employed when readily definable and sym- 
patric subgroups can be determined. 

superspecies: the products of a radiation, the representatives of 
which are still completely or mostly allopatric and usually 
chromosomally uniform. Species status often uncertain. (See 
below. ) 

species: the most recent evidence indicates that the recognition 
of valid species is much harder than was formally assumed. 
It cannot be routinely assumed that allopatry or parapatry 
imply subspecific status. As I indicate above, I have used 
the superspecies category for a number of equivocal or doubt- 
ful cases. 

subspecies: intraspecies differentiation differs so much in dif- 
ferent species that this unit is of very different value from 
one case to another. I do not report currently recognized 
subspecies in the species list below. I will do so in the de- 
tailed species group and species definitions to come, but I 
will do so without passing judgment on their validity. I imply 
only that the differentiation so indicated — in contrast with 
that implied by the use of superspecies — is clearly intra- 
specific. 



1976 



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BREVIORA 



No. 440 



c 



T3 
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c/D 






-a 










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u 


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c^ c^ c^ 

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c 

0; 


c 





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jj 




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c 


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> 



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jc 


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c 


TJ 


c 


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5i 


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s 


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be «H 

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Si >. 



" fa 



S s S « j2 



1976 WEST INDIAN ANGLES 1. 



o 


O 


CJ 


u 






S 


5 


o 


O 


t: 


t 


<u 


CJ 


3 


3 


Ph 


Ph 





a 


(t3 


73. 


r2 


'E 




o 


u 




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c 




"^ 


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i: _c 



ex 



c^ (i hU c^ 



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to 

00 

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^ in 

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^ 




c 




c3 


J^ 


m 


c 




c^ 


j3 


m 








.2 



O "^ tH HH CO 

o^ -Q 3 3^ ^ r~- 

^ ^ g Q 2 t- ^ 



3r^^ cr- S^'^^^-M'-^, , i-]Op,co (Up 



«a^^~« Sc3-h-Q 



10 BREVIORA No. 440 



^ 






JJt 




c 






c 










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u 


rS 




lU c^ 




O 

c 


C 
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Montserrat 
Giiadeloup 
Marie Gal 


03 
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o 

p 



c 

nPQ 
5 ^ « JS 
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^ 


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'S 


C 


'S 


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03 


r3 


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03 


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CI- 


G, 


O- 


a 


D- 


& 


.22 


IT) 


.22 


(70 


.22 


_l/3 



CO 
CO 



CO 
CO 



o 

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•^.Sdr;-,^ ^co<^J cococng^^ 

_ a; C - § 00 Q. 5-.S^ t;^ = ^> 3o2prtC-g;:3 

c b £ '-^ Q. b,r^ '^^ ^^ "t: L_] 13 =^ , fc S-O o o ^ "^ 



-P3 



s 






1976 WEST INDIAN ANGLES 1. 11 



C 



^ 








c^ 




M 




M 




o 




CJ 




O 'rt 


O 


5^ 


■c5 


O § 


J5 5 


i- be 


c h 


'^ 03 


o «-> 


P-i )— 1 


S<g 



o 



r^ 



lO 



CO Q^ 

o 

m 
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in to J ^ 

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i "' --^ -^ ^ -2 P- — 



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2 =s 1 -^ s"-^ s 

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12 BREViORA No. 440 



c 



T3.2 'o'o 'oc'oo'o 



a 


^ 






o 


"o 






c 


'S 


cC 


ci 


a 


D- 






rt rt 


rt c3 cS 03 


^ -Q 


_C ^ ^ JD 


3 P 


3 3 S 3 



K ffi ffi Xffi ffiKE uuuuuu 









CO CM CO 



rh 






^ '^^S -G ^ J §3^ 2 ^K d < - ^ sS S "^ -g -5 






o 



*v ^^ ^j — i .^r^-' . ^ '^^ '^-'.^ "nj ?-- ^ t^%.^ V- (^ ^4J :^^ "o p^ ^*j 






1976 



WEST INDIAN ANGLES 1. 



13 



^ a 


C^ 


f3 








2 '° 


O 


O 








c 'c 


c 


c 


a a 


rS 


{^ 


D, a 


O- 


a 


1/3 O! 


1/5 


c/: 



XX XX 



d d ci 
^ [c [o 
'c 'c 'c 

rt rt rt 
CI. O- D, 

K E E 



c 








03 








m 








a 




C 




g 




c3 




c^ 




£ 




J5 




>. 




rt 




rt 




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13 

C/3 


c 




c3 ti 




OJ 


rt 


> 


■^ 


> 


3 t: 


rt 


.ti 


r3 



u o u hj ^ i 



QJ ^ 



CO 
00 



PQ 

qj '"^ 

•n — . CM 

O., flJ CO 

52 £^ 

s s ^ 

s a s 

?^. ^ ~ 

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--: -c o . 



CO 









CO 
CO 

C75 



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c 



ex. 

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CO 

to 

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2 a 

O 
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a. «« 









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cr. CO 

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o 



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14 BREVIORA No. 440 







>^ 






C3 






U 






c 






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s 






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13 






K 






in 


^ 




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ca 


pq 






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pq 


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p 


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5: .52-^ s cj-jR 5« 5>, g ^ V c^-- ^ 2^.;^ "S ^ ;s ^ "S i:2 -H 



1976 



WEST INDIAN ANGLES 1. 



15 



s 



Pi 03 rt rt ct 

JO ^ ^ ^ -O 

3 3 3 3 3 

U U O U U 






C/3 

o 

lO HH 

I — , HH 
Cn [-H 

r-. m <-N Q 
n; c^ f^ ^ ^ 



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^ d d 
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a ri n 

Oh O- Q, 



U d X'XX 



CM 

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C/2 





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03 






O 


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CI, 


:r.:B 



^ t •-> i C/^ 

■Ci, 52 S S ^ 



3 Ph O 



o "~^ 

Ph 

ex ^ 






CO 



CO 



i:; c 






.S 




C rt 






Si 




t- f- 










■H' 3 






u 




C >, 






J= 




•" c3 






H 




cfU 


Pi 


Pi «3 






^ -,- 


_CJ 


CJ CJ 


^^ 




•— T3 






c 




Ki c 


'rt 


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c 


J* 


£ 2 


£ 


£ £ 




c 


-i2,0 


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Pi r3 


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re 

a. 


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CT; 


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S 






m 


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cr. 


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c/: p^ <H 



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c: c 
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16 BREViORA No. 440 









c 














£ .a 














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.^f^ 














Bahamas 
areas of 
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c 




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isl 


£ 


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c 


03 cJ r3 rt 


rt c^ rt 03 


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^ 


3 (^ -S 

U S.£ 


rt 


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u u 


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u 



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c/^ S^-ii J! ^ 3 b ^ O^^ N -< -C ^' ^ S ^ txq 

^■S S ^^ ^« w -^ :^ ^ f < §-1 ^ S 



s 



S f^ '5 -? c cj -Q* 3 i: =^ ^ --, 



•s-ighl I ill Hill" II 5.2 






a 



H 



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1976 



WEST INDIAN ANGLES 1, 



17 



a re 
t/D re 



re <u 



Qj re 
^ r 



^ 




QJ 




Q 




^^ 




re 


lA 


3 
re 
U 




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S-H 


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^ 


iy> 


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a.— 
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♦"I e X 
t« o -^ 



c <u 

CD '— 



a; C 

re Dh 
7^ o 



be m .E w ^ 



00 00 ^D t^ 


""--:::; 00 — H 00 






'O 


<o'^ " 


•a 


^^^=^ 


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CO l> 


!> lO 


W 


eridge, 19 
cd or floa 
gives fust 


c 
3 

a. 


^,_^ 








re 


js y. 




TJ-< CM Tt^ -^ 


«« ^3^ 




■^ Tfl 


JZ 






""^ '**'^ ^"\ "^'^^ 


so «*^ lf5 so \ 


■^ lO 


^•■^^^^^ 


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w 


CO CO CO CO 


OO 




OO QO 


Pi 


o s: 




1^ 










.2 ElT 


S" 


T^ "^ ^ "^ 


T^ ^ tJH Tt^ "^ '^ "^ 


^ tH 




— s3 
•0=2 


'1- 


(M CM CM CM 


CM CM CM CM CM CM CM 


CM CM 


> 


'c ^ ^ 










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T3 


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c 
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a. T 
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C 


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C 


t' 5 " 
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I + +1 +1 I I I + 1 I I 



+ 11 ++. I + 



o °? 



CM 



•J-; to CO CO 10 
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00 

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CO \,cO CO CO CO CO 
00 rv. 00 CO 00 CO eo 



^ ^ 



X3 — , — ( 



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II <^ 



rL. ("L. rv. 



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+ + ++:5 +1 1 + + + + + 



s.is 



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18 



BREVIORA 



No. 440 







o c -S 






ii rt o 






^ 






c 






3 






hi 






'4-1 






3 
fo-Q 






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fac 






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t! u -a 




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04 


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w 

pq 
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gg 


h 





fciD^ 




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b/D 



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c/2 



+ 



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lology and with 
syntopic, but m 
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c °-= §. 














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r- <— Cfl 














-p C O 














c . >- 














Ra 
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are 
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o^^^-S, 














aj P Q< re 














K fc- i-c 














r- "" P bO 

C t; 3 P 


s « 












t« 2 S IJ 

p-S -55 


o -2 


<o s 




<^ 






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s 


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52 






^ 




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G 


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a 


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-2 






.SQ=i 



1976 



WEST INDIAN ANGLES 1. 



19 



C 
< 



+ + 






3 



+ + + 






'■i-> e 
J2 



+ + + 



+ + + + 



A 



X5 

3 



3 



+ + 



3 

u 



w 

I— I 

w 



S h '^ 

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+ + + + 



+ 



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rt 












Tj 












i-c 












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s 


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ll 




u 
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=^ji 




^ h o 


S V- 


C 


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« b/3 


1 


<o « J: 


C2- =3 


3 


f--^ 


b « 


1 


^ ^^ 




« 


ia/3? 



20 



BREVIORA 



No. 440 



+ 






c 



J3 



a. 



^1 







^ 


C 


-*! 


u-i 


c 

3 


o 


u 


O 


u 


b 


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!/5 






h 


OJ 




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bo 








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3 








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u 


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+ 



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5 « S 



+ ;§ 



bO 



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S 3 




<3 -2 
S 3 



^. X 



o 

c 


is 


•thei 
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0- 




i^< 


1/3 



1976 WEST INDIAN ANGLES 1. 21 

ACKNOWLEDGMENTS 

The new data on West Indian anoHnes is the work of many 
hands. The singularities of the present taxonomic arrangement 
are the author's own. Financial support for the work result- 
ing in the present paper has been provided by the National 
Science Foundation under grant GB-37731X and prior grants 
to E. E. Williams. 

> REFERENCES CITED 

Etheridge, R. 1960. The relationships of the ancles (Reptilia: Sauria, 
Iguanidae) : an interpretation based on skeletal morphology. Ann 
Arbor, Michigan: University Microfilms, xiv -|- 236 pp. 

. 1965. The abdominal skeleton of lizards in the family Igua- 
nidae. Herpetologica 21: 161-168. 

G.^RRiDO. O. H. AND A. Schwartz. 1968. Cuban lizards of the genus Cha- 
maeleoli% Quart. Jour. Florida Acad. Sci. 30: 197-220. 

Gorman, G. C. 1965. Interspecific karyotypic variation as a svstematic 
character in the genus Anolis (Sauria, Iguanidae) . Nature 208: 95-97. 

— . 1974. The chromosomes of the Reptilia, a cvtotaxonomic 

interpretation. In A. B. Chiarelli and E. Capanna eds., Cvtotaxonomy 
and Vertebrate Evolution, pp. 349-424. Academic Press, New York. 

Gorman, G. C. and Brad Stamm. 1975. The Anolis lizards of Mona, 
Redonda, and La Blanquilla: chromosomes, relationships, and natural 
history notes. Jour. Herp. 9: 197-205. 

Rand. .\. S. and E. E. Williams. 1969. The anoles of La Palma: aspects 
of their ecological relationships. Breviora Mus. Comp. Zool. No. 327: 
1-17. 

Savage, J. 1973. A preliminary handlist of the herpetofauna of Costa Rica. 
Los Angles, 17 pp. 

Schwartz, A. and R. Thomas. 1975. A checklist of "W'est Indian am- 
phibians and reptiles. Special Publication Carnegie Museum No. 1: 
1-216. 

Thomas, R. 1966. A reassessment of the fauna of Navassa Island. Jour. 
Ohio Herp. Soc. 5: 73-89. 

Williams, E. E. 1972. The origin of faiuias. Evolution of lizard congeners 
in a complex island fauna: a trial analvsis. Evolutionarv Biology 6: 
47-89. 

Yang, S. Y., M. Sovxt and G. C. Gorman. 1974. Anolis lizards of the 
eastern Caribbean: a case study in evolution. 1. Genetic relationships, 
phvlogeny and colonization sequence of the roquet group. Syst. Zool. 
23: 387-399. 



IV1U3. COM?. ZOOL 
LinrjARY 



MAR 18 1985 

iiseum of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 27 December 1976 Number 441 



LIZARD KARYOTYPES FROM THE 

GALAPAGOS ISLANDS: 

CHROMOSOMES IN PHYLOGENY 

AND EVOLUTION 

D. Paull,^ E. E. Williams^ and W. P. Hall- 
Abstract. The iguanid lizards, Conolophus subscristatus, Tropidxirus albe- 
marlensis, T. delanonis, and T. duncanensis have similar 2n =:z 36 karyotypes. 
C. subcristatus has a 12 metacentric macrochromosome and 24 microchromo- 
some karyotype that is here shown to be primitive for the Iguanidae and 
probably for all lizards, while the three Galapagos Tropidurus have iden- 
tical patterns to Tropidurus species from eastern South America and differ 
from the primitive karyotype of C. subcristatus by non-Robertsonian modi- 
fications of three pairs of macrochromosomes. 

All available karyotypic data for the Iguanidae are summarized and used 
to discuss how one may determine which karyotypes are "primitive" within 
radiations and what possible roles Robertsonian karyotypic variation may 
play in the process of evolution. Analysis of karyotypic and systematic in- 
formation suggests a causal relationship between karyotypic differentiation 
and the rapid proliferation of new species, such that the need for geographic 
isolation seems to be minimized by the chromosomal differentiation. 

INTRODUCTION 

Among the karyotypic ally well-studied families of squamate 
reptiles, the iguanid lizards are known to show great chromo- 
somal diversity, with most of the variation apparently resulting 
from Robertsonian mutations (centric fusions and/or fissions) 
(Gorman, 1973; HaU, 1973). To more fully understand the 
biological significance and evolution of this diversity, workers 

^Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 
02138. 

^Department of EPO Biology, University of Colorado, Boulder, Colo. 
80309. 



2 BREVIORA No. 441 

at the Museum of Comparative Zoology have been accumu- 
lating karyotypes for as many and diverse iguanid species as 
possible. As part of this program, during the winter of 1969-70 
Paull was able to karyotype four iguanid species from the 
Galapagos Islands. Two of the four principal phyletic branches 
of the family, the "iguanines" and the "tropidurines" (Savage, 
1958; Etheridge, 1964) are represented in these islands, and both 
were sampled in the study. All four species showed a 2n == 36, 
1 2 metacentric macrochromosome, 24 microchromosome pat- 
tern which is belie\ed by some to be primitixe for the Iguanidae 
or, indeed, for all hzards (Gorman et al., 1967, 1969; Webster 
et al., 1972; Gorman, 1973). 

Addition of karyotypes for these four species to our data base 
provides the occasion to discuss the e\'idencc of "primiti\'eness" 
for karyotypes within a radiation and the role Robertsonian 
karyotypic variation may play in the process of evolution. How- 
ever, we must first describe the karyotypes of the Galapagos 
lizards sampled. 

KARYOTYPES OF GALAPAGOS IGUANIDS 
(D. Paull and W. P. Hall) 

Specimens examined: Representatives of two distantly re- 
lated branches of the Iguanidae have reached the Galapagos, 
presumably by overwater colonization. The iguanine species in 
the Galapagos belong to the endemic genera Amblyrhynchus 
(one species, the marine iguana) and Conolophus (two species 
of land iguanas). Of these, Conolophus siibcristatus was karyo- 
typed. The tropidurine radiation is represented in the archi- 
pelago by eight endemic species of Tropidurus (lava lizards). 
Additionally, Tropidurus has a South American continental 
radiation of 12 species (Etheridge in Peters and Donoso-Barros, 
1970). Island species karyotyped were Tropidurus albemarlen- 
sis, T. delanonis and T. duncanensis. Table 1 lists the species 
karyotyped and their collection localities. 

Methods: All chromosome preparations were made in the 
Galapagos Islands using laboratory facilities kindly supplied by 
the Charles Darwin Research Station on Santa Cruz Island. 
Cells were spread for karyotyping by air drying smears of metha- 
nol : acetic acid (3:1) fixed suspensions of testis, bone marrow 
or spleen tissues prepared directly from colchicine pretreated 
animals. The techniques used were similar to those of Evans et 



1976 



CHROMOSOME EVOLUTION 



species 



Table 1 
Galapagos specimens karyotyped 

locality 



Conolophus subcristatus 
Tropidurus albemarlcnsis 
Tropidurus albeynarlensis 
Tropidurus albemarlensis 

Tropidurus delanonis 
Tropidurus duncanensis 



individuals 
karyotyped 



South Plazas Id., 

Las Plazas 1 

South Plazas Id., 

Las Plazas 6 

Bartolome Id., 

Sullivan Bay 4 

Santa Cruz Id., between 

Galapagos Hotel and 

Darwin Station 6 

Hood Id., behind beach, 

Gardiner's Bay 5 

Duncan Id., small 

cove on NE end 2 



al. (1964), Bianchi and Contreras (1967) and Patton and 
Hsu (1969). 

Results: All species had 36 chromosomes, with 12 biarmed 
macrochromosomes and 24 microchromosomes (Fig. 1). No 
cytologically distinct sex chromosomes or intrageneric variation 
of any kind was seen. However, conspicuous differences in arm 
ratios and relative sizes of the macrochromosomes were noted 
between the genera ( Fig. 1 ) . 

In Conolophus subcristatus (Fig. 1, lower), taking the macro- 
chromosomes in order of size, beginning with the largest, pair 
one is very slightly submetacentric; pair two is distinctly sub- 
metacentric, with the long arm slightly less than twice as long 
as the short; pairs three and four are almost exactly metacentric 
and, in many spreads, indistinguishable in size; pair five is 
nearly metacentric; and pair six is submetacentric, with the long 
arm about 1.5 to 2.0 times the length of the short arm. Pairs 
one and two are similar in length, three and four are slightly 
but distinguishably shorter than two, five is distinguishably 
shorter than four, and six is conspicuously shorter than five. 
Some of the microchromosomes seem to be metacentric or sub- 
metacentric, but our preparations do not resolve their structures 
well enough to allow them to be unequivocally paired. 



BREVIORA 



No. 441 



S 




'*i:o 



«* 



9 
i 



• 
1 



5 u 



2 o 



1 i 

C IC 



ai 



1976 



CHROMOSOME EVOLUTION 




Figure 2. Comparison of male Tropidurus karyotypes. The species are 
T. delanonis (left) , T. duncanensis (middle) , and T. albemarlensis (right) . 
The karyotypes are printed to the same magnification. 



6 BREVIORA No. 441 

In Tropidurus (Fig. 1, upper; Fig. 2) pairs one through five 
show a fairly even gradation in length, with pair six being 
conspicuously smaller than five. Comparing the arm ratios to 
those of Conolophus, Tropidurus pair two is more submetacen- 
tric, with the long arm being slightly more than twice the length 
of the short; and Tropidurus pair five, rather than being meta- 
centric, is almost subacrocentric, with the long arm about 2.5 
times the length of the short. Again, some of the microchromo- 
somes appear to be metacentric or submetacentric, but they are 
not adequately resolved to allow accurate pairing. 

PRIMITIVE KARYOTYPES 

(E. E. Williams and W. P. Hall) 

Comparisons with other iguanid genera and other families of 
lizards suggest that the Tropidurus pattern is derived with re- 
spect to the Conolophus pattern, which may be primitive for 
several families, including the Iguanidae. 

As Gorman (1973) in the latest review of reptilian chromo- 
somes has emphasized, our knowledge of hzard karyotypes and 
especially of iguanid karyotypes has increased immensely since 
the early work of Matthey (1931, 1933). In the iguanids even 
such very speciose genera as Sceloporus and Anolis have now 
been sampled very extensively and at least one or two species 
have been examined in all major subgroups of the family 
(Table 2). The more species studied, the more widely one 
chromosomal arrangement is demonstrated: that with a 2n of 
36, with 12 metacentric macrochromosomes and 24 micro- 
chromosomes. 

The kind of 2n = 36 karyotype characteristic of the Galapa- 
gos Tropidurus (i.e. 2n, arm ratios, etc.) has also been reported 
for representatives of this genus in eastern South America (Gor- 
man et al., 1967; Peccinini, 1969; and Bccak et al., 1972); 
however, we are not aware of its occurrence in any other genera. 
On the other hand, the 2n = 36 Conolophus karv'otype, or at 
least the details of its macrochromosomal pattern, is found in 
many different lizard groups. In the Iguanidae (Table 2) pre- 
cisely this macrochromosomal pattern is found in such diverse 
groups as the anolines (Gorman, 1973), sceloporines (Cole, 
1970; Pennock et al, 1969; Gorman, 1973; Hall, 1973), 
Crotaphytus (Montanucci, 1970), iguanines (Cohen et al., 
1967; Gorman et al., 1967; Robinson, 1974), oplurines (Gor- 



1976 



CHROMOSOME EVOLUTION 



Table 2 

Genera are those in Peters and Donoso-Barros, 1970 unless 
otherwise noted. Reference for number of species karyotyped 
and diploid number is Gorman 1973 or this paper unless other- 
wise noted. 



Genus 

Arnblyrhyncfiiis 

Anisolepis 

Anolis 

Aperopristis^ 

Aptycholaeynus 

Basilisfus 

Brachylophiis 

CaUisaurus 

Chalnrodon 

ChamaeleoUs 

Cham aelinorops 

Conolophus 

Corytophanes 

Crotahhytus 

Ctenoblepharis 

Ctenosaura^' 

Cyclura 

Dil)lolae?nus 

Dipsosaurus 

Enyalioides 

Enyaliiis 

Holbrookia 

Hoplocercus 

Ifliiana 

Laemanctus 

Leiocephnlus^'^ 

Leiosaurus 

Liolaemus 

Morunasaurus 

Ophryoessoides 

Opiums 

Petrosaurus 

Phenacosaurus 



Number of 

species 

in genus 



6^ 

2 
320 



Number of 

species 
karyotyped 



1 

2 


1 


ca 200'^ 


80 + = 


1 






1 






4 


1 


95 




c 




2G 


2 


V 






2s 


1 


r 


1 


2 


1 


3 






510 


3 


8^^ 






7 


1 


3:3 


1 


3 






314 


1 


7 


1 


815 






316 


2 


1 






2 


1 


2 








20 


5 


4 






50 + ^^ 


8 


2 






Q19 





Range of 
known 2n 



2n= 



=36 



2n=25-48 



2n=36 
2n=34 



2n= 



36 

2n=36=^ 
2n=36 



2n=36 

2n=36 
2n=36 



2n=36-^ 
2n=36^ 

2n=34-^ 

2n=34 

2n=32-36=^ 



2n=30-40 



2n=36 

2n=34^ 

2n=36 



BREVIORA 



No. 441 



Table 2 (Continued" 





Number of 


Number of 






Genus 


species 
in genus 


species 
karyotyped 


Range of 
known 2n 


Phrynosaura 

Phrynosoma 

Phymatunis 

Plica 

Platynotus'-^ 

Polychrus 

Pristidactylus"^ 

Proctotretus 

Sator 

Sauro?naIus 

Sceloporus 

Stenocercus^^ 

Strobilurus^° 

Tropidurus 

Uma 

Urocentron 

Uranoscodon 

Urosaurus 

Urostrophus 

Uta 


3 

1421 

222 

2 
1 

624 

4 

3 
2« 

7« 

64 + ^^ 
29 
1 

20 

531 

430 

4 

10« 

3 

632 








9 


2n= 


=34 


1 


2n= 


=40 


4 
1 

1 

428 

45 + 


2n- 
2n= 

2n= 
2n= 
2n= 


= 20-30=' 
=36 

=3427 

=36 

=22-46^ 








4 
3 


2n= 
2n= 


=36 
=34 








5 


2n- 


=34^ 


6 


2n= 


=34 



Because they do not appear on the Etheridge dendrogram, 
two recently described genera have been omitted from the table: 
Vilcunia Donoso-Barros and Cei, J. Herp. 5: 90. 1971. 
Pelusaurus Donoso-Barros, Neotropica 19: 132. 1973. 
Both are said to be allied to Liolacmus and Proctotretus. 
Etheridge (personal communication) infers from their descrip- 
tions that they are allied to the Liolaeynus-Phrynosaura-Cteno- 
blepharis complex and not to Proctotretus. 



lEibl-Eibesfeldt, I. 1962. Neue Unterarten der Meerechse, Amblyrhynchus 
cristatus, nebst weiteien Angaben zur Biologic der Art. Senckenbergiana, 
Biol. 43: 177-199. 

2This is only a rough approximation. Continental members of the genus 
are in general poorly understood, and species are still being discovered in 
Hispaniola and Cuba. 

-We include unpublished data from material in the Museum of Com- 
parative Zoology. 



1976 CHROMOSOME EVOLUTION 9 

4Etheridge (personal communication) after examination of specimens 
prefers to recojjnize the genus as distinct from Leiosaiints. Cf Miiller, L., 
1922: i'ber Aperopristis paronae Peracca und die Genera Aperopristis 
Peiacca und Leiosatirus Dumeril and Bibron. Senckenbcrgiana 4: 153-159. 
sAvery, D. and W. \V. Tanner, 1970. Speciation in the Fijian and Tongan 
iguana Brnrh\Inphns CSauria. Iguanidae) with description of a new 
species. Great Basin \at. 30: 166-172. 

•■Smith, H. and E. H. Taylor, 1950. An annotated checkHst and kev to 
the reptiles of Mexico, exclusive of the snakes. Bull. l\ S. Nat. Mas. 199: 
1-253. Cophosffiirw; (Holbrookia) texana is placed with CaHisnnrus here. 
See Norris, K. S., 1958. The evolution and systcmatics of the iguanid genus 
Uma and its relation to the evolution of other North American desert 
reptiles. Btdl. Amer. Mus. Nat. Hist. 114: 247-326, and Axtell, quoted by 
Norris. The only important character separating the two genera is the 
condition of the car opening, which is evaluated as less important than 
character.^ separating other sceloporine genera. In fact, it would not be 
unreasonable to lump the tliree sand-swimming genera, Callisiuinis. Hol- 
brookia and Uina. This larger genus would still have only 10 species. 

'Angel, F. 1942. I.es Lczards dc Madagascar. Mem. L'Acad. Malgache 
M: 1-193. 

"^Garrido, O. H. and A. Schwartz. 1968. Cuban lizards of the genus 
Ckamaeleolis. Quart. J. Fla. .\cad. Sci. 30: 197-220. 

^'Thomas, R. 1966. \ reassessment of the herpetofauna of Na\assa Island. 
J. Ohio Herp. Soc. 5: 73-89. 

loMontanncci, R. 1969. Remarks on the Crotaphytiis-Gambelia contro- 
versy (^Sauria: Iguanidae) . Herpetologica 25: 308-314. Montanucci, R. 1970. 
Analysis of hybridization between Crotnphytus un'slizenii and Crotaphxtus 
silus (Sauria. Iguanidae) in California. Copeia 1970: 104-123. 

iiCei, J. 1974. Two new species of Ctenoblepharis (Rcptilia. Iguanidae) 
from the arid cnviionments of Central Argentina (Mcndoza Province). 
J. Hcrp. 8: 71. In contrast to Cci. ^vc continue to recognize Phtynosaura 
as distinct from Ctenoblepharis. On the key character of juxtaposed versus 
imbricate dorsals, Cei's two new sjjccies arc Ctrnoblepharis. 

i^Etheridge follows an unpublished MS bv Clavton E. Rav in uniting 
Enyaliosaiiriis and Ctenosaiira. 

i^'fide Albert Schwartz and Michael Carev, who arc preparing a revision 
of the genus. 

i^Smith and Taylor 1950, as in footnote 6 above. 

I'^Etheridge, R. 1969. A review of the iguanid genus Enxaliiis. Bull. 
Brit. Mus. (N.H.) , Zool. 18: 233-260. 

ifiWilliams, K. L. and H. M. Smith, 1958. Herpetologica 13: 265-267. 
i^Etheridge. R. 1966 (The systematic relationships of ^V'est Indian and 
South American lizards referred to the iguanid genus Eeiocephalus. Copeia 
1966: 79-91) listed 16 species in his restricted genits Leiocephalus. Subse- 
quent to 1966, Schwartz, by reinterprctation of already described forms, has 
raised the number to 20. O. H. Garrido, 1973 (Nueva especies de Leio- 



10 BREVIORA No. 441 

cephalus (Lacertilia, Iguanidae) para Cuba. Poeyana No. 116: 1-19) has 
added a 21st species. 

I'^Richard Sage (personal communication) believes the taxonomy of Lio- 
laemiis to be still in a very primitive state. He suggests that the genus may 
contain as many as 100 biological species. 

loFritts, T. H. 1974. A multivariate evolutionary analysis of the Andean 
iguanid lizards of the genus Stenocercus. Mem. San Diego Soc. Nat. Hist. 
7: 1-89. Fritts reallocates to Stenocercus a number of the species formerly 
placed in Ophryoessoides, leaving only six described and two undescribed 
species in the genus. 

-oLazell, J. D. 1969. The genus Phenamsaurus (Sauria: Iguanidae) . 
Breviora No. 325: 1-24. 

2iPresch, W. 1969. Evolutionary osteology and relationships of the horned 
lizard genus Phrynosoma (family Iguanidae) . Copeia 1969: 250-275. 

22Cei, J. M. and L. P. Castro, 1973. Taxonomic and serological researches 
on the Phymaturus patagonicus complex. J. Herp. 7: 237-247. 

-sEtheridge (in a letter) : "I know of no reference that presents evidence 
for or against recognition of this genus. Most, but not all, recent authors 
include the species semitaeniatns in Tropidiirus. 

24Gorman, G. C, R. B. Hucy and E. E. Williams, 1969. Cytotaxonomic 
studies on some iniusual iguanid lizards assigned to the genera Chamae- 
leolis, Polychrus, Polyrhroides and PheJiacosnurns, with behavioral notes. 
Breviora No. 316: 1-17. Polychroides svnonymizcd with Polychrus. 

lispeccinini, D. 1969. Variarao nos cromossomos do lagarto Polychrus 
marmoratus (Sauria, Iguanidae) de diferentes localidodes (Nota preliminar) . 
Rev. Brasil. Biol. 30: 1-4. (The Polychrus from Sao Paulo with 2n^0 is 
actually P. acutirostris [P. Vanzolini, pers. comm.]. See also Peccinini, D., 
Be^ak, O. Frota-Pessoa and Iris Ferrari, 1971. Sex determination of the 
"pseudo-XO/XX" type in the Brazilian lizard Polychrus sp. (Sauria, 
Iguanidae). Caryologia 24: 129-139, and Becak, M. L., "\V. Becak and L. 
Denaro, 1972. Chromosome polymorphism, geographical variation and karyo- 
types in Sauria. Caryologia 25: 313-326. 

2'5"Barrio (1969) has demonstrated Cupriguatuis araucaniis Gallardo 1964 
is conspecific with Leiosaurus fascintus Dorbigny in Dumeril and Bibron 
1837. Barrio used the combination Cupriguanus fasciatus. However, 
Leiosaurus fasciatus was in 1843 designated the type of Pristidactylus by 
Fitzinger. If fosciatus is congeneric with achnlevsis, the type of Cupriguanus, 
the latter becomes a jimior synonvm of Pristidactylus which then would 
be regarded as having foin^ species, fasciatus Dorbigny 1837, scapulatus 
Burmeister 1861, achalensis Gallardo 1964 and casuliatiensis Gallardo 1968." 
Etheridge (personal commimication) . 

27Lowe, C. H. and M. D. Robinson. 1971. The chromosome pattern in 
Sator grandnexnts (Reptilia: Iguanidae) , Baja California, Mexico. J. Arizona 
Acad. Sci. 6: 282. 

28Robinson, M. D. 1974. Chromosomes of the insular species of the 



1976 CHROMOSOME EVOLUTION 11 

chuckwalla lizards Cgeniis Samomalus) in the (iulf of California. Mexico. 
Herpetologica 30: 162-167. 

20Sixty-foiir Sceloporus species are currently recognized. When taxonomic 
revisions suggested bv Hall, 1973 are incorporated, this number will be 
raised to about 72. with considerable uncertainty remaining concerning the 
number of biological species to be recognized within the forquatus and 
formosus species groups. 

soEtheridge, R. 1968. A review of the iguanid lizard genera Vracentron 
and Strohilunis. Bull. Brit. Mus. (N.H.) . Zool. 17: 48-64. 

^nVilliams, K, L.. P. S. Chrapliwy and H. M. Smith. 1959. A new fringe- 
footed lizard from Mexico. Trans. Kans. Acad. Sci. 62: 166-162; Mavhew, 
VV. W. 1964. Taxonomic status of California populations of the lizard 
genus Uma. Herpetologica 20: 170-183. 

s-'Ballinger. R. E. and D. \V. Tinkle, 1972. Svstcmatics and evolution of 
the genus IHa fSauria: Iguanidae) . ^fisc. Publ. ^fus. Zool. Univ. Michigan 
No. 141: 1-83. 



man et al., 1967) and the tronidurines ^Gorman et al., 1967). 
In other familie.s f Table 3) thi.s pattern ha.s been demonstrated 
in the Aeamidae fArronet, 1965; Gorman and Schochat, 1972; 
Hall. 1970; Sokolov.skv, 1972 V in the Teiidae fGorman, 1970) , 
in the Gerrho.sauridae fMat<;hey, 193.3; Hall, unpub.) and in 
the Amtohisbaenidae fHuana; et al., 1967). 

This most widely distributed Conolohhus-Yikt karyotype is a 
source of controversy. On the one hand, it has been called 
"primitive" f Gorman and others). On the other hand, it has 
been in*:erpreted, as by Co'e f1970. 1971b), as derived in at 
least some iguanids, or, as by M. J. T). White, as a possible 
example of an exceptionally .stable confis^uration that has been 
repeatedly evolved within a group, f White at one time called 
this "the principle of homologous change," but he now prefers 
to call it "karyotypic orthoselection" [see White, 1973 for dis- 
cussion].) 

Those who deny the primiti\-eness of the 2n =^ 36 pattern 
hold very firmly to the concept that primitive karyotypes in 
lizards consist entirely of acrocentrics with karyotypic evolution 
then occurring by centric fusions of them. 

This view, that acrocentrics are prima facie primiti\'e, ha.s 
rested on the belief that fusion is cytologically much easier than 
fis,sion and hence much more common (Matthey, 1949; White, 
1954, 1959; Reiger et al., 1968). In particular, the generation 
of a new centromere, which supposedly occurs in fission, has 
seemed to lack any mechanism that would readily permit the 



12 



BREVIORA 



No. 441 



event, while fusion has been interpreted as the result of the 
(conceptually) less difficult process of reciprocal translocation 
followed by loss of a centromere carrying small segments of one 
or both chromosomes. 

However, there are now many cases for which fission is an 
obligatory explanation of the origin of the karyotypes of highly 
derived groups and species — too many cases to allow any doubt 
of the reality of fission as one possible path of karyotypic evolu- 
tion — whatever its mechanism. Even White (1973) now ad- 
mits its existence under the name "centric dissociation" in cer- 
tain cases. Morescalchi (1973) finds fission the hypothesis of 
choice for the origin of the karyotypes of certain species of Hyla 
and Eleutherodactylus. In reptiles, Webster et al. (1972) 



Table 3 
Karyotypes in Non-Iguanid Lizard Families 



12V + 24m or 
direct derivatives 
occur in some member 



Only karyotypes not readily 
related to 12V + 24m known 



Agamidae 

Chamaeleontidae 

Xantusiidae 

(6V +121 + 24m) 

Gerrhosauridae 

Teiidae 

(see Gorman, 1970) 

Anguidae 

Helodermatidae 
(lOV + 41 + 24m) 

Varanidae 

Scincidae 

(see Greer et al., in prep.) 



Gekkonidae 
Pygopodidae 
Lacertidae 
Annielhdae 



Based on karyotypic data in Gorman (1973), unless other- 
wise noted. 



1976 CHROMOSOME EVOLUTION 13 

presented evidence for the highly deri\ed phyletic position of 
Afiolis jnonticola, the one species within that very large genus 
that has a 2n as high as 48. It is similarly inescapable that 
fissioning has occurred several times in Sceloporus (Hall and 
Selander, 1973; Hall, 1973). A recent discussion in the journal 
Evolution has summarized some of the mammalian evidence 
for fission (Lawlor, 1974; Baker et al, 1975). It is no longer 
reasonable to peremptorily reject fission as a plausible mode of 
Robertsonian karyotypic e\'olution. 

The argument for the primiti\eness of the 12 macro- and 
24 microchromosome pattern for iguanids and for lizards, how- 
ever, does not depend on the supposed plausibiHty or implausi- 
bility of fission. The argument becomes easier to accept if fission 
is admitted in certain cases, but primiti\'eness for a karyotype, 
as for any other character state, can be determined on its own 
merits, independent of any theoretical mechanism for the evoHi- 
tion of that character state. 

A large literature now exists dealing with objective recogni- 
tion of primitiveness. Kluge and Farris fl969) may stand as 
an example. They would use the following criteria (1969:5), 
listed in order of reliability: 

( 1 ) The primitive state for any particular group 
is likely to be present in many representatives of closely 
related groups. 

(2) A primitive state is more likely to be wide- 
spread within a group than is any one more ad\'anced 
state. 

(3) The primitive state is likely to be associated 
with states of other characters known from other 
evidence to be primitive. 

They add that "closely related groups can be selected through 
estimates of oxerall similarity that make no assumptions about 
primitive conditions." 

"Widespread" they define not by counting taxa but as 
occurring in several taxa that otherwise would ha\e little in 
common. They would also use "available fossil material." 

At least in intention these criteria have merit, but, in general, 
such criteria are especially difficult to use in our present stage 
of knowledge of karyotypes. Fossils are clearly unavailable. It 
is still rare for karyotypes to be known for even a substantial 
number of any group and, on the contrary, those at hand may 
be a very biased sample. The problem of real similarity may 



14 BREVIORA No. 441 

be serious; diploid number by itself is meaningless; there must 
be near identity in chromosome morphology paralleling taxo- 
nomic relationships inferred on other grounds. 

When a group has been as well sampled as the Iguanidae 
now are, however, the Kluge and Farris criteria begin to be 
applicable and the comparative method can lead to sound re- 
sults when appropriately applied. Cole misrepresents, indeed 
caricatures, the comparative approach when, in opposing the 
concept of the 12 + 24 karyotype as ancestral in iguanids, he 
says (1970:31): 

"These conclusions are based on the assumption that the general 
karyotypic condition found in the majority of species that were 
available for sampling, at whatever level of the taxonomic hierarchy 
one happened to be working with, was. therefore, the most primi- 
tive," 

and, again (1970:31): 

"If I were to simply employ the principal [sic] on which these 
authors' argmnents are based, I would reach a rather diiferent con- 
clusion for the spiuosus group, for in this group the karyotype of 
the liindelli subgroup (12 biarmed macrochromosomes plus 10 
smaller chromosomes, most of which are clearly biarmed; a 12 + 
10 karyotype) would then be considered ancestral because it occurs 
in five of the nine species in the species group and none of the 
three remaining general kaiyotypes of the group is represented in 
more than two species." 

Our own reasoning has a quite difTerent base than that por- 
trayed by Cole. We start from a base that is broad and quite 
independent of the ideas of Cole or ourselves. 

Figure 3 provides a dendrogram of the phylogenetic relation- 
ships of the genera of the Iguanidae which represents the present 

Figure 3. Phylogeny and karyotype distribution in the family Iguanidae. 
The base dendrogram summarizes phylogenetic conclusions of Richard 
Etheridge from his osteological studies and is reproduced here with his 
permission from a privately circulated figure. The available karyotypic 
information for the family (references cited in Tabic 3) is superimposed 
on this dendrogram and indicates (1) the basic diploid karyotype for each 
genus chromosomed or (2) in karyotypically variable genera, the most 
primitive karyotype in the genus plus the range of 2n known for it. "V" 
chromosomes are metacentric macrochromosomes, "I" arc acrocentric macro- 
chromosomes, and "m" are microchromosomes. Thus, an 8V,81-(-24m 
karyotype has a 2n of 40 with eight metacentric macrochromosomes, eight 
acrocentric macrochromosomes, and 24 microchromosomes. 



1976 



CHROMOSOME EVOLUTION 



15 



Uracentron 

Strobilurus 

Platynotus 

Plica 

8V,8I'24m 



Ophryoessoides 
Stenocercus 



Phrynosaura 
Ctenoblephans 



Uranoscodon 



a Anolis 

2n. 25-48 
(inc.l2V*24m) 
SOUTH AMERICA 
CENTRAL AMERICA, 
WEST INDIES 



Apfycho/aemus 
Anisolepis 



Urosfrophus 




Sceloporus 

2n. 22-4' 
(inc I2V*2 



eastern Tropidvrus 

l2V'24nn 



Phenacosaurus 

l2V»24m 
NORTHERN 
SOUTH AMERICA 



Enyalius 




fi Anolis 

2n 29-40 

MEXICO TO 

SOUTH AMERICA, 

CUBA, JAMAICA 

Chamaelinorops 

l2V»24m 
HISPANIOLA 

Chamaeleolis 

l2V*24m 
CUBA 



SOUTH AMERICA 
CENTRAL AMERICA 



Diplolaemus 

Leiosaurus 

Aperopristis 

Pristidacfylus 

l2Vt24m 





Hoplocercus 

Morunasaurus 

Enyalioides 

12V •24m 
SOUTH AMERICA 



oplunnes 



NORTH AMERICA, 

CENTRAL AMERICA, 

NORTHERN SOUTH 

AMERICA, WEST INCHES 

Corythophanes 

Laemanctus 

Basiliscus 

l2V24m 
CENTRAL AMERICA, 
NORTHERN SOUTH 
h AMERICA 



Opiurus 

12 V ♦24m 

Chalarodon 

MADAGASCAR 



16 • BREVIORA No. 441 

views of Richard Etheridsje and Richard Estes, extending those 
of Savage (1958), Etheridge (1964) and Presch (1969). This 
has been based wholly on osteology and thus it gives us a picture 
of relationships constructed without knowledge of or reference 
to karyotypes. We have then superimposed on this dendrogram 
the range of karyotypes for each iguanid genus for which these 
are known. We use both the published reports summarized by 
Gorman, 1973 (most references cited by Gorman are not re- 
peated) and our still unpublished data. We use the latter 
despite the lack of formal (and especially pictorial) documenta- 
tion because it fills out much of the picture of karyotypic varia- 
tion in the Iguanidae without significantly altering it. 

The Etheridge scheme recognizes a basal stock (called 
"monmasaurines" by Estes and Price, 1973) from which seven 
major lineages arise (Polychrus, the AnoUs-Enyalius-Diplolaemus 
lineage, the "tropidurines," the "sceloporines" plus Crotaphytus, 
the "iguanines," the "basiliscines," and the "oplurines" ) . 

Karyotypes are known for one genus of morunasaurines (one 
species of EnyaUoides — our unpublished data ) ; in this and in 
six of the seven derived lineages {all except Polychrus) the 
"primiti\'e" 1 2 + 24 karyotype is known to occur or is the only 
karyotype known. Furthermore, in every one of the derived 
lineages (except Polychrus) those genera closest to the base on 
Etheridge's diagram — i.e. those believed for osteological reasons 
to be the more primitive members of each lineage (and, in fact, 
each sublineage ) — ha\'e either the 1 2 + 24 karyotype or a 
12 + 22 karyotype that differs from the primitive condition by 
the absence (by loss or fusion) of a pair of microchromosomes. 
In the iguanine line, for example, all genera in the sublineage 
containing Iguana, Ctenosaura, Dipsosaurus and Sauromalus 
have a 12 + 24 or (Iguana) 12 + 22 karyotype. 

The sceloporine line is especially instructi\'e in this and other 
respects. Crotaphytus, an early offshoot of this line, has 12 + 
24 (Cohen et al., 1967; Montanucci, 1970). Every primitive 
sceloporine and some Sceloporus have 12 + 22 (several authors, 
summarized in Hall, 1973). Within Sceloporus (a morphologi- 
cally more derived genus) numbers range from 12 + 10 to 
24 + 22; if the 12 + 22 pattern is accepted as primitive for 
the genus, it is clear that both fusions and fissions must have 
been involved in the evolution of its karyotypic diversity. 

The tropidurine line is not well sampled, but as we have 
shown above, Tropidurus from eastern South America and the 



1976 CHROMOSOME EVOLUTION 17 

Galapagoan Tropidurus (derived from western South American 
stock) have the basic 12 + 24 pattern, although their karyo- 
types are slightly derived in non-Robertsonian ways. The only 
West Indian Leiocephalus published (Gorman et al., 1967) also 
has the basic pattern. Besides confirming the 12 + 24 pattern 
in Hispaniolan species. Hall (unpub.) has found representatives 
of the Cuban branch of the genus to have 12 + 20 patterns. 
In the speciose genus Liolaemus, the single species so far re- 
ported (Gorman et al., 1967) has 12 + 22. although shdes 
made by Richard Sage and examined by Hall indicate the 
presence of considerable karvotvpic variation in this genus 
(2n — 30-40)\ 

In the line leading to AnoUs. the primiti\-e genera Chamaeleo- 
Us, Phcnacosaurus and Chamaelinorops (Hall and Williams, in 
preparation) have 12 + 24 karyotypes, as do many of the alpha 
section of the genus Anolis itself. The 12 + 24 pattern is also 
found in the related lineage including the genera Pristidactylus 
and A7ii sole pis. 

Given this evidence, it is difficult to contest the hypothesis 
that the .12 + 24 pattern is primitive in the Iguanidae. It is 
possible to go to greater detail : arguments similar to those above 
support the idea that even the detailed chromosome size and 
arm ratios found in Conolophus must be primitive for iguanids. 

Additionally, when we notice ( 1 ) that quite similar 1 2 + 24 
karyotypes are found in other lizard families (Table 3), both in 
families that e\'eryone agrees are closely related to the Iguani- 
dae, i.e. the Agamidae and the Chamaeleontidae, and in fam- 
ilies that are just as universally regarded as not closely related 
(Gerrhosauridae, Anguidae, Amphisbaenidae) ; (2) that still 
other families (Scincidae, Helodermatidae, Varanidae) have 
karyotypes easily derivable from 12V + 24m (Gorman, 1973), 
it then becomes clear that, using the most neutral descriptive 
terms, the 12 + 24 karyotype is an extraordinarily stable and 
conservative pattern. To us it is evident that the most careful 
and skeptical use of the Kluge and Farris criteria of primiti\e- 

lAlthough variation in chiomosonie n.nmber was clearly demonstrated in 
this material, the preparations were not of good quality and the data were 
complicated by the inclusion of unnamed taxa (clearly Liolaemus, however) . 
Further work will be required before publication is warranted. However, 
we think that the existence of substantial karyotypic variation in this genus 
should be noted. 



18 BREVIORA No. 441 

ness points unequivocally to the ancestral position of the 12 + 
24 karyotype not only for iguanids, but for all lizards. 

On the other hand, there is no doubt that some lizard families 
and at least one group within the Iguanidae have karyotypes 
that are very difficult to reconcile with derivation from a 12 + 
24 pattern (Gorman, 1973). The Gekkonidae will sers^e as an 
example of the first case; Polychrus is, of course, the second. 
We are impressed that these deviant families and groups are in 
general isolated cases, neither closely related to one another, nor 
arguable as ancestral to forms with the 12 + 24 karyotype. In 
all of these cases, morphological and other evidence suggests 
long separation from plausible basal stocks, and hence leads all 
the more strongly to the conclusion that the Conolophus-like 
12 + 24 pattern is the one primitive for lizards. 

EVOLUTIONARY ROLE OF 
ROBERTSONIAN CHANGE 

(W. P. Hall and E. E. Williams) 

Let us point out immediately that there is no antithesis be- 
tween primitiveness and stability — the two explanations that 
have been proposed for the iteration of one chromosome pat- 
tern throughout a large number of species. On the contrary, 
the genera which show only the widespread chromosome pat- 
terns which we believe to be primitixe seem not to be the ones 
that have radiated widely. This is a point which we want to 
stress, especially for the Iguanidae. Karyotypically conservative 
groups, so far as we can see, have produced only few species. 
Of the more than 50 genera of iguanids, only three — Anolis, 
Sceloporus and Liolaemus — are very large, each including more 
than 50 species (several times 50 in Anolis), or have produced 
high levels of sympatry (5+ syntopic species in several areas of 
the ranges of Anolis or Sceloporus) . Of the others, only Steno- 
cerus (as revised by Fritts, 1974), Tropidurus (if the Galapagos 
.species are included ) , and Leiocephalus ( a purely insular radi- 
ation in the West Indies) have as many as 20 species, and no 
others have as many as 15. Excepting Polychrus, whose several 
karyotypes bear little obvious relationship to one another and 
none to any other iguanid, the other small, conservatively spe- 
ciating genera show on current evidence little or no intrageneric 
variation in karyotypes, and indeed very little variation among 
genera. Of the 14 non-sceloporine small iguanid genera (in- 



1976 CHROMOSOME EVOLUTION 19 

eluding Crotaphytus) sampled, 11 have the 12 + 24 pattern. All 
of the primitive sceloporines (eight genera) and Iguana among 
the iguanines have 12 + 22. Only Plica, aside from Polychrus, 
stands out in showing a notably different karyotype (16 + 24), 
and its modifications seem relatively simple (presumably fissions 
of four of the primitive metacentric macrochromosomes) . Of 
the genera of middle size (20-29 species), the four sampled 
species of Tropidurus have again the 12 + 24 pattern but differ 
somewhat in arm ratios from the usual condition, and while 
some Leiocephalus have the 12 + 24 pattern, others have 12 + 
20 (reduction in two pairs of microchromosomes — Hall, un- 
published). Stenocercus has not yet been sampled. 

Contrasting strongly with this picture of conservative specia- 
tion and karyotypic evolution in the small iguanid genera is a 
picture showing extensive, usually Robertsonian karyotypic vari- 
ation in each of the three prolifically speciose genera. In Anolis 
2n's range from 25 to 48 (Gorman, 1973; Hall, unpub.), in 
Sceloporus they range from 22 to 46 (Gorman, 1973; Hall, 
1973), and in a few Liolaemus they range from 30 to 40 (Sage 
and Hall, unpublished). 

The apparent association of conservative speciation with con- 
servative karyotypic evolution, and prolific speciation with re- 
markable karyotypic diversity suggests the possibility of an 
evolutionarily important causal relationship between karyotypic 
differentiation and speciation. Though there are undoubtedly 
other possibihties and explanations that might be raised, it is 
this possibility of causal relationship that we here want to evalu- 
ate. We offer the following arguments to demonstrate that 
the relationship between speciation and karyotypic diversity is 
genuine. 

( 1 ) Intrageneric variation in karyotypes. Since few small 
genera from six of the seven major iguanid lineages are repre- 
sented by karyotypes from more than one species, we must agree 
that we cannot safely compare the amounts of m^rageneric vari- 
ation between small and large genera in these lineages. This 
defect, however, most certainly does not apply to the sceloporine 
lineage: all nine sceloporine genera and the related Crotaphytus 
are cytologically well known. Half or more of the species from 
each of these 10 genera have been karyotyped: 3/5 from Crota- 
phytus, 2/2 from Petrosaurus, 9/14 from Phrynosoyna, 2/2 
from Callisaurus, 3/3 from Uma, 2/3 from Holbrookia, 1/2 
from Sator, 6/6 from Uta, 5/10 from Urosaurus, and 45 + / 



20 



BREVIORA 



No. 441 



Table 4 

Distribution of karyotypic x'ariation in Iguanidae: interspecies 
comparisons (large genera except Sceloporus omitted). 



9 



in 



Other Other small 
scelo- iguanid 

Sceloporus^ porines" genera 



All small 
iguanid 
eenera 



48 
46 
44 
42 
40 
38 
36 
34 
32 
30 

28 

26 

24 
22 
20 

species 
karyotyped 

species known 

% karyotyped 
species with 
2n=36or34 




2 
2 
1 
3 

1 
18 
15 
2 



3 

13 


45 + 
64 + 

42% 










33 











33 
49 



>Vb 








1 Plica plica 

13 
1 

1 



Polychrus 
marmoratus 



1 Polychrus 
peruvianus 

1 Polychrus 
femoralis 





1 Polychrus 
acutirostris 



19 

122 

74% 








1 



13 

34 



1 





1 

52 
171 

90^ 



^includes polymorj)hisnis within ciniently recognized "species." 
-here inchiding Crotaphytus. 



1976 CHROMOSOME EVOLUTION 21 

64+ from Sceloporus (data summarized from Table 2). None 
of these genera (except Sceloporus) shows any intrageneric vari- 
ation, and the only intergeneric difTerence is between the 12 + 
24 Crotaphytus and the 12 + 22 sceloporines. Within Scelo- 
porus only 1 3 species ( 1 5 after taxonomic revisions by Hall ) 
are known to have the primitive sceloporine condition ( 2n = 
34), while the remaining 32 (40 or 41 after revisions) karyo- 
typed species have derived patterns — and most of these belong 
to the phylogenetically more advanced large-scaled branch as 
defined by Smith (1939). In the sceloporine lineage (Table 4), 
the correlation between chromosomal diversity and prolific spe- 
ciation is clear cut and does indeed appear to be fundamental. 
And even with our poor sampUng of the small non-sceloporine 
genera, the association between chromosomal conservation and 
few species per genus is, at the least, suggestive. 

(2) Intergeneric diver sity in karyotypes. Although compara- 
tively few species of the small, non-sceloporine genera have been 
karyotyped, still there is less intergeneric diversity observed than 
we would expect if variation were randomly distributed in the 
family. Phylogenetic relationships inferred from morphology 
( Fig. 3 ) show that many of these genera must have been evolv- 
ing as independent lineages for comparatively long times, possi- 
bly since the Cretaceous (Estes and Price, 1973). Given so 
long a period of evolution, they show remarkably little evidence 
of the acquisition or accumulation of chromosomal differentia- 
tion. As we have said, there are \tv\ few known differences 
among genera, and, in fact, few departures from the 12 + 24 
pattern. In the 25 small genera sampled (Table 5), the few 
observed cases of m/^rgeneric variation are slight indeed com- 
pared to the known m^rageneric variation of the phylogenetically 
more recent large genera. Unless the sampling of the small 
genera has been biased in some unknown way, this should be 
quite significant. 

The deviations from the 12 + 24 pattern among the small 
genera are again : Plica ( 1 6 + 24 in no more than four muta- 
tional events, and possibly in only one, fide Todd, 1970), 
Iguana (12 + 22 in one event), all of the "primitive" scelo- 
porines (12 + 22 in one event in the common ancestry- for all 
species), and Polychrus (2n's= 20-30 resulting from an un- 
determined number of events producing karyotypes derived in 
relation both to one another and the 12 + 24 pattern). Con- 
trasted to the limited intergeneric variation in the family as a 



22 BREVIORA No. 441 

whole is the remarkable interspecific diversity involving many 
mutational events found within each of the three especially 
speciose genera (cf. Fig. 3). Again, this relationship is clearest 
in the well-investigated sceloporine Hneage (Hall, 1973, in 
prep.). 



Table 5 
Distribution of intergeneric comparisons in the Iguanidae 

small genera large genera 

CM CM 

CM CM 

T3 + T3 + 

r-\ C ^ ^ -i-'r^C'-i 

.^ -5 -^ c t' .*^ -B^ 

linpso-p O o5'5c^J d O^-SCM 

nneage c c<o-- c g<o-h 



u 


O 


u 


>-^ 


C 


u 


<u 


c^ 


bTj 


-^iS 


o 


O 


C 


C 



basal stock 


3 


1 








- 


— 


oplurines 


2 


1 








- 


- 


basiliscines 


3 


1 








- 


- 


iguanines 


8 


6 








- 


- 


sceloporines 
& Crotaphytus 


9 


9 





1 


1 


1 


tropidurines^ 


11 


1 


1 


4 


3 


2 


anolines 


3 


2 





1 


1 


1 


anoline relatives 


8 


3 








- 


- 


Polychrus 


1 


1 


1 





- 


- 


Totals 


49 


25 


2 


6 


5 


4 



iThe number of genera in the lineage for which karyotypic data exist. 
-The number of genera in the lineage containing one or more species with 

a karyotype deviating from the 12 + 24 or 12 + 22 pattern. 
'■'■Stenocercus, Tropidurus, Leiocephalus, and Liolacmtts, each with 20 or 

more species, are treated as large genera. 



1976 CHROMOSOME EVOLUTION 23 

(3) Relative recency of species and of karyotypic diversity. 
If rates of fixation of Robertsonian mutations were independent 
of the process of speciation, one expectation might be that many 
of the older genera would accumulate karyotypic variants while 
phylogenetically recent groups might show little variation, even 
though they include many species. 

Hall (1973) would adduce Sceloporus as a counter-example, 
since he believes it to be a phylogenetically quite recent genus. 
We summarize Hall's views and arguments here. 

(A) While the Iguanidae may have originated in the Cre- 
taceous (Estes and Price, 1973), the differentiation of the 
present sceloporine genera probably did not antedate the de- 
velopment of the xeric habitats in North America during the 
Miocene (Axelrod, 1950, 1958). This conclusion is supported 
by the facts: (1) that Crotaphytus and all sceloporine genera 
are largely or entirely restricted to the North American deserts 
(except Sceloporus itself, which has extensively radiated in 
mesic habitats as well as in deserts); (2) that all sceloporine 
genera except for the osteologically primitive Crotaphytus and 
Petrosaurus (Etheridge, 1964; Presch, 1969) and Sator (which 
simoly has not been tested) will "shimmy bury" (Axtell, 1956) 
in loose soil or sand for escape or sleeping cover (Stebbins, 
1943, 1948; Axtell, 1956; Norris, 1958), a behavior not seen 
in any of the other North American or West Indian iguanids 
(we know nothing about South American iguanids in this re- 
spect) ; and (3) that all sceloporines but no other North Ameri- 
can iguanid genera (including Crotaphytus) show a "sink-trap" 
type of nasal passage and almost always an associated valvular 
nostriP (Stebbins, 1943, 1948; Savage, 1958; Hall, unpub.), 
which clearly seems to have been evolved in relationship to the 
use of shimmy burial for cover in xeric environments where 
loose soil is likely to be more readily available for cover than 
that provided by plants or permanent burrows in firm soil. 

(B) Within the sceloporines, Sceloporus seems to be one of 
the most recently differentiated genera. Osteological data clearly 
show four groups within the sceloporines (Savage, 1958; Ethe- 
ridge, 1964; Presch, 1969) : the primitive Petrosaurus; the 
speciaHzed Phrynosoma; the group of "sand-swimming" (Norris, 
1958) genera, Callisaurus, Holbrookia and Utna; and the group 

^Petrosaurus, which Hves in xeric habitats but which does not shimmy 
bury under experimental conditions, has the nasal sink tuap but lacks the 
nasal valve (Hall, personal observation) . 



24 BREVIORA No. 441 

of four genera, Sceloporus, Urosaurus, Sator and Uta. Accord- 
ing to Etheridge (1964), these last four genera cannot be dis- 
tinguished osteologically. Howex-er, comparisons suggest that 
Sceloporus, with its mucronate imbricate scales developed to a 
degree not found in any of the other North American iguanids 
(where granular scales seem to be the primitive condition) and 
with its loss of the gular fold found in all other sceloporines, is 
the phylogenetically most recent of these four genera. 

(C) Within Sceloporus, again based on characteristics of the 
squamation, it seems clear that Smith's (1939) small-sized, 
small-scaled species are more closely related to the other 
sceloporines than is the large-sized, large-scaled branch. To 
summarize the cytosystematics of these two divisions, the phylo- 
genetically more primitive small-scaled branch contains 20 spe- 
cies by present taxonomy: 12 of these are karyotypically con- 
servative,^ five have not been karyotyped, and the three that 
are karyotypically derived are also highly derived ecologically 
{merriami, 2n ^ 46, is a specialized cliff -face dweller and 
scalaris and aeneus, 2n = 24, are specialized montane bunch 
grass dwellers ) . Within the morphologically more advanced 
large-scale branch, according to present taxonomy, only the 
single species, .S". orcutti, has the primitive 12 + 22 pattern, 
while all other karyotyped species are derived. At present (very 
conservative taxonomy) this branch contains a minimum of 43 
species, of which all but 13 (all 13 are in chromosomally highly 
derived species groups) have been karyotyped. 

This phylogenetically most recent radiation of the sceloporines 
(the large-scaled Sceloporus) has co\ered the entire ecological 
and geographical range of lizards in North America (Smith, 
1939) and shows simultaneously a truly remarkable karyotypic 
diversity (2n's from 22 to 46). It is also notable that the most 
ecologically differentiated small-scaled species in the genus (the 
scalaris group species [2n = 24] and the merriami [2n = 46] ) 
are also among the karyotypically most highly derixed forms. 

The one egregious example of chromosomal diversity in a small 
genus, Polychrus, seems in the very fact of its uniqueness equally 
a counter-example to the generality of the proposition that 
deviant karyotypes tend to accumulate in all genera with time. 

iKaryotypically at least to a first approximation: Cole (1971) notes that 
maculosus has a 2n r= 31, X^X.,Y|3 (based on three specimens, only one a 
male) and tliat in pxrocfphnlus chromosome 1 shows a jjericentric inversion. 



1976 CHROMOSOME EVOLUTION 25 

Polyrhrus is seen on Etheridge's diagram as an isolated basal 
twig, truly \ery old and very distinct, entirely suitable as a group 
in which de\iant karyotypes might accumulate. But each of 
the other six major groups is as old in Etheridgean terms. If 
karyotypic di\ersity is a product only of time, even a random 
and superficial sampling of the other small genera should, so it 
seems to us, ha\e resulted in more cases of highly deri\ed karyo- 
types than are in fact in front of us. 

Our own surmise regarding Polyrhrus is that the six forms 
currently recognized, all highly arboreal, may represent only the 
few sur\'i\ors of an old and formerly more prolific lineage of 
tree dwellers that, perhaps, has been largely replaced by the 
radiation of Anolis in the arboreal habitat. 

Chromosomal evolution in the Iguanidae: 
two major patterns 

We do not deny that the history of karyotypic change is not 
now and ne\"er will be known from direct e\'idence, that the 
real and unique historical process must be inferred from its 
products, nor that the survey of iguanid karyotypes, though it 
is already impressive, is incomplete. We insist, however, that 
the present sample is large enough to justify conjecture and to 
point to the kinds of evidence that will verify or negate postu- 
lated sequences. 

Our picture of chromosomal evolution in lizards, and per- 
haps also in many other groups, is that there are both periods 
of chromosome conservatism with usually slow geographic modes 
of speciation ( Mayr, 1 963 ) and episodes of karyotypic instability 
associated with rapid proliferations of new species (Hall, 1973). 
We believe that both Anolis and Sceloporus exhibit these phe- 
nomena (cf. especially Webster et al, 1972; Williams and Web- 
ster, 1973; and Hall and Selander, 1973), and presumably 
Liolaemus also does. Todd (1970) suggests a similar relation- 
ship between karyotypic diversification and prolific speciation 
in the Canidae. 

To us the comparative data strongly suggest that karyotypic 
diversification and speciation are in many cases functionally 
related, such that the temporal and/or geographic requirement 
for the separation of populations is somehow minimized (not 
eliminated but very greatly reduced) when chromosomal differ- 
ences become fixed between them. White's model of "stasi- 



26 BREVioRA No. 441 

patric" speciation (White et al., 1967; White, 1968; Key, 1968) 
offers one mechanism, and others are possible (Hall, 1973; in 
prep.). Here we wish to emphasize only that Robertsonian 
mutations frequently are found fixed between species of rapidly 
proHferatinff groups but only rarely are found as intrapopula- 
tion polymorphisms. (Wallace, 1959, provided an early note of 
this phenomenon in Drosophila.) Given this distribution of 
Robertsonian mutations, we think it especially significant that, 
among the varieties of chromosomal rearrangements, the Robert- 
sonian ones probably ha\'e the least impact on the meiotic as- 
sortment or recombination of balanced genomes; but, on the 
other hand, at least in mammals where breeding and cytological 
studies have been made, these mutations are increasingly impli- 
cated as a significant source of chromosomal malassortment in 
meiosis serving to reduce the effective fertility of chromosomally 
heterozygous individuals (Polani et al., 1965; Gustavsson, 1971a, 
b; Cattanach and Mosely, 1973). Once a chromosomal differ- 
ence is established, reduced heterozygote fertility could then 
serve in appropriate circumstances as an intrinsic partial barrier 
to gene flow between karyotypically differentiated homozygous 
populations, thereby reducing the requirement for extrinsic bar- 
riers to gene flow before speciation could ensue (Hall, 1973; in 
prep.). 

Then, assuming some model of chromosomal speciation based 
on cytogenetically reduced fertility in heterozygotes, the proba- 
bility or frequency of such speciation in given lineages should 
be highly dependent on parameters of their genetic systems such 
as: mutation rates, malassortment rates, population structures, 
mating systems, ets. Chromosomal speciation might then be 
precluded in some lineages because of unfavorable genetic sys- 
tems that would allow speciation only by conservative geographic 
modes; on the other hand, genetic systems of other lineages may 
especially fa\'or chromosomal speciation, and thus allow great 
proliferations of species, even in the absence of strong extrinsic 
barriers to gene flow. Such a chromosomal speciation theory can 
easily account for the associations of karyotypic diversity and 
prolific speciation found in Sceloporus, Anolis, and apparently 
in Liolaemiis. 

The test of the chromosomal speciation model of karyotypic 
evolution as it pertains to the iguanids will be found in the still 
unsampled or inadequately sampled iguanid radiations, par- 
ticularly those of South America. Stenocercus, now with 29 



1976 CHROMOSOME EVOLUTION 27 

recognized species and with notable sympatry, is certainly cru- 
cial. The karyotypic \ariation in Liolaemiis and Leiocephahis 
must be confinued, and the karvotypic patterns in these two 
genera adequately documented. We suggest that the species of 
mainland Tropidurus, which seem to have rather complicated 
distributions (Peters and Donoso-Barros, 1970), may also repay 
careful attention. Only such a wider survey of the kar\'otypes 
of the Iguanidae can provide either a x'erification of the evolu- 
tionary patterns we ha\'e suggested here, or, by demonstrating 
new patterns, require alternative models. 

RESUMEN 

Los iguanidos: Conolophus subcristatiis, Tropidurus albemar- 
lensis, T. delanonis, y T. duncanensis tienen cariotipos similares 
de 36 cromosomas. C. subcristatus, con 12 macrocromosomas 
metacentricos y 24 microcromosomas, tiene un cariotipo que se 
demuenstra ser "primitivo" dentro la familia Iguanidae, y que 
probablemente es tambien primitivo entre todos de los lagartijos. 
Los cariotipos 2n = 36 de los tres Tropidurus son iguales y 
tambien al Tropidurus del este de Suramerica, pero ellos son 
diferentes del cariotipo primiti\o porque hay modificaciones 
"no-Robertsonianas" de tres pares de los macrocromosomas. 

La filogenia y todos de los datos cromosomicos de la Iguani- 
dae estan resumidos para una discusion sobre la determinacion 
de que cariotipos son "primitivos" dentro radiaciones de espe- 
cics, y tambien sobre los funciones que sir\'en las mutaciones de 
Robertson en el proceso de evolucion. Analisis de la informacion 
sobre los cariotipos y sistematica demuenstra una conexion 
causal y cerca entre la diferenciacion cariotipica y la prolifera- 
cion rapida de especies nuevas, donde el requisito para aislami- 
ento geografico se minimiza a causa de la diferenciacion cromo- 
somica. 

ACKNOWLEDGMENTS 

We are grateful to Captain Sir Thomas Barlow and Mr. 
Roger Perry for permission to collect on the Galapagos Islands 
and for the opportunity to use the facilities of the Charles Dar- 
win Research Station, and to Professor James J. Hoff and 
Richard C. Paull for assistance in the field. We thank R. B. 
Stamm for his assistance in the lab and T. P. Webster, Richard 
Etheridge, Richard Estes and James J. Jackson for comments 



28 BREVIORA No. 441 

and criticisms. Richard Etheridge generously provided the 
dendrogram of iguanid relationships and has amplified or an- 
notated our statements on generic size in Table 2. The research 
was partially supported by NSF grants GB 1980 IX and GB 
37731 to E.E. Williams and GB 27911 (1969-70) to R. Rol- 
lins of the Committee on Evolutionary Biology at Harvard and 
NIH grant RR 8102 administered by the Division of Research 
Resources. 

REFERENCES 

Ari'ONEt, v. N. 1965. Description of tlic karyotypes of Againa caucasica 

and Ph)j)iocephalus helioscapus (Aganridae, Rcptilia) [in Russian, 

English summary]. Tsitologiya 1: 237-239. 
AxELROD, D. I. 1950. Evolution of desert vegetation in western North 

America. Carnegie Inst. Wash., Publ. 590: 215-360. 
. 1958. Evolution of the Madro-Tertiary gcoflora. Bot. Rev. 

24: 433-509. 
AxTELL, R. W. 1956. A solution to the long neglected Holbrookia laceuita 

problenr and the description of two new subspecies of Holbrookia. 

Bull. Chicago Acad. Sci. 10: 163-179. 
Baker, R. J., J. H. BowrRS, .\nd M. H. SxirrH. 1975. Reply to cotument 

on "Chromosomal evolution in Peroniyscus." Evolution 29: 189. 
Barrio, .\. 1969. Sobre la real ubicacion generica de Leiosaurus fasriatus 

D Orbigny (Lacertilia, Iguanidae) . Physis 29: 268-270. 
Becak, M. L., W. Becak, and L. Denaro. 1972. Chromosome ])olymorphism, 

geographical variation and karyotypes in Sauria. Caryologia 25: 313-326. 
BiANCHt, N. O. AND J. R. CoNTRERAs. 1967. The chromosomes of the field 

mouse Akodon azarac (Cricctidae, Rodentia) witli special reference to 

sex chromosome anomalies. Cytogenetics 6: 306-313. 
Bury, R. B., G. C. Gorman, and J. F. Lvnoh. 1969. Karyotypic data for 

five species of anguid lizards. Expcrentia 25: 314 316. 
Cattanach, B. M. and H. Mosr.i.v. 1973. Nondisjunction and reduced 

fertility caused by the toliacco mouse metacentric chromosomes. Cyto- 

genet. Cell Genet. 12: 264 287. 
Cohen, M. \f., C. C. Huang, and H. F. Ci.ark. 1967. The somatic chromo- 
somes of three lizard species, Gekko gecko. Iguana iguana, and Cro- 

taphytiis collaris. Experentia 23: 769-771. 
Coi.E, C. J. 1970. Karyotypes and evolution of the spinosus group of 

lizards in the genus Sccloporus. Amer. Afns. Novitates No. 2431: 1-47. 
. 1971a. Karvotypcs of the five monotypic species gioups of 

lizards in the genus Sceloporus. .Amer. Mus. Novitates No. 2450: 1-17. 
. 1971b. Karyotypes and relationships of the pxrorephalus 



group of lizards in the genus Sceloporns. Herpetologica 27: 1-8. 
EsTES, R. and L. I. Price. 1973. Iguanid lizard from the Upper Cretaceous 
of Brazil. Science 180: 748 751. 



1976 CHROMOSOME EVOLUTION 29 

Etheridge, R. 1964. The skeletal inoiphologv and systematic relationships 

of sceloporinc lizards. Copeia 1964: 610-631. 
Gorman, G. C. 1970. Chromosomes and the systematics of the family 

Teiidae (Sauria, Reptilia) . Copeia 1970: 230-245. 
. 1973. The chromosomes of the Reptilia, a cytotaxonomic 

interpretation. In A. B. Chiarelli and E. Capanna eds., Cytotaxonomy 

and Vertebrate Evolution, pp. 349 424. Academic Press, New York. 
, L. Atkins, and T. Hoi.zincer. 1967. New karyotypic data 



on 15 genera of lizards in the family Iguanidae with a discussion of 
cytologic^ and taxonomic information. Cytogenetics 6: 286-299. 
. AND D. Shochat. 1972. A taxonomic interpretation of chromo- 



somal and electrophoretic data on the agamid lizards of Israel with 
notes on some East African species. Herpctologica 28: 10&-112. 

GusTAvssoN, I. 1971a. Cliromosomes of repeat-breeder heifers. Hereditas 
68: 331-332. 

. 1971b. Distribution of the 1/29 translocation in the A. I. 

bull population of Swedish Red and AVhite cattle. Hereditas 69: 101- 
106. 

Hall. \V. P. 1970. Three probable cases of parthenogenesis in lizards 
(.\gamidae, Chamaeleontidac, Gekkonidae) . Experentia 26: 1271-1273. 

. Comparative population cytogenetics, speciation, and evolu- 
tion of the iguanid genus Sceloponis. Unpublished Ph.D. thesis. Har- 
vard University. 

AND R. K. Selandfr. 1973. Hybridization of karyotypically 



differentiated populations in the Sceloponis grammicus complex (Iguani- 
dae) . Evolution 26: 226-242. 

Hsu, T. C. AND J. L. Patton. 1969. Bone marrow preparations for chromo- 
some studies. In K. Benirschke ed.. Comparative Mammalian Cyto- 
genetics, pp. 454-460. Springer-Verlag, New York. 

Huang, C. C, H. F. Clark, and C. Gans. 1967. Karyological studies on 
fifteen forms of amphisbaenians (.\mphisbaenia-Reptilia) . Chromo- 
soma 22: 1-15. 

Key. K. H. L. 1968. The concept of stasipatric speciation. Syst. Zool. 17: 
14-22. 

Kluge. a. G. and J. S. Farris. 1969. Quantitative phyletics and the evolu- 
tion of anurans. Syst. Zool. 18: 1-32. 

Law lor, T. E. 1974. Chromosomal evolution in Peromyscus. Evolution 
28: 688-691. 

Matthev, R. 1931. Chromosomes des Reptiles, Sauriens, Ophidiens, Che- 
loniens. La evolution de la formule chromosomiale chez les Sauriens. 
Rev. Suisse Zool. 38: 117-186. 

. 1933. Nouvelle contribution a I'etude des chromosomes chez 

les Sauriens Rev. Suisse Zool. 40: 281-316. 

. 1949. Les Chromosomes des Vertebres. 353 pp. F. Rouge, 



Lausanne. 



30 BREVIORA No. 441 

Mayr, t. 11-63. Animal Species and Evolution. 797 pp. Harvard Belknap 
Press, Cambridge, Mass. 

MoNTANUcci, R. R. 1970. Analysis of hybridization between Crotapliytus 
wislizenii and Crotapliytus silus (Sauria, Iguanidae) in California. 
Copeia 1970: 104-123. 

MoRESCALCHi, A. 1973. Amphibia. In A. R. Chiarelli and E. Capanna, 
eds., Cytotaxonomy and Vertebrate Evokition, pp. 233-248. Academic 
Press, New York. 

NoRRis, K. S. 1958. The evolution and systematics of the iguanid genus 
Unia and its relation to the evolution of other North American desert 
reptiles. Bull. Amer. Mus. Nat. Hist. 114: 247-326. 

Peccinini, D. M. V. M. 1969. Cariotipo e mecanismo de determinacao de 
sexo em algumas especies de lacertilios Brasileros (Iguanidae e Teiidae) . 
Master's Thesis, Universidad de Sao Paulo, 46 pp. 

Pennock, L. a., D. W. Tinkle, and M. W. Shaw. 1969. Minute Y chromo- 
some in the lizard genus Uta (Family Iguanidae) . Cytogenetics 8: 9-19. 

Peters, J. A. and R. Donoso-Barrgs. 1970. Catalogue of the neotropical 
Squamata. Part II. Lizards and amphisbaenians. 293 pp. Smithsonian 
Institution, Washington, D. C. 

POLANi, P. E., J. L. Hamlrton, F. Giannei.li, and C. O. Carter. 1965. 
Cytogenetics of Down's Syndrome (Mongolism) . II. Frequency of 
interchange trisomies and mutation rate of chromosome interchanges. 
Cytogenetics 4: 193-206. 

Presch, W. 1969. Evolutionary osteology anrl relationships of the horned 
lizard genus PIu\)iosoi)ia (Family Iguanidae) . Copeia 1969: 250-275. 

Rieger, R., a. Michaelis and M. M. Green. 1969. A Glossary of Genetics 
and Cytogenetics: Classical and Molecular. 3rd ed.. Revised. Springer- 
Verlag, New York. 507 pp. 

Robinson, M. D. 1974. Chromosomes of the insular species of the chuck- 
walla li7ai(ls (genus Sninoinniiis) in the Gulf of California, Mexico. 
Hcrpetologica 30: 162-167. 

Sa\ace, J. M. 1958. The iguanid lizard genera Urosainiis and Uta with 
remarks on related gioups. Zoologica 43: 41-54. 

Smith, H. M. 1939. The Mexican and Central American li/anls of the 
genus Scrlojioitis. Field Mus. Nat. Hist., Zool. Ser. 26: 1-397. 

SoKOLOvsK^■, V. V. 1972. Comparative karyology of the reptiles [in Rus- 
sian]. Viniti — .411 Union Institute of Scientific and Technical Informa- 
tion, Moscow. 51 pp. 

Stebbins, R. C. 19)3. .Adaptations in the nasal passages for sand burrow- 
ing in rlie sainian genus Ihna. Amer. Nat. 77: 38-52. 

. 1948. Nasal structure in lizards \\ith reference to olfaction 

and conditioning of ihe inspired air. Anui. J. \nat. 82: 183-222. 

Todd, N. B. 1970. Karyotypic fissioning and canid pliylogenv. J. Tlicor. 
Biol. 2fi: 445-480. 

\V\i,iACE, B. 1959. Influence of genetic svstems on geographical distribu- 
tion Cold Spring Harbor Svmp. Quant. Biol. 24: 193-204. 



1976 CHROMOSOME EVOLUTION 31 

Webster, T. P., ^V. P. Hall, and E. E. Williams. 1972. Fission in the 
evolution of a lizard karyotype. Science 177: 611-613. 

White, M. ]. D. 1954. Animal Cytology and Evolution. 2nd Ed. Cam- 
bridge Univ. Press. 

. 1959. Speciation in Animals. Australian J. Sci. 22: 32~39. 

. 1968. Models of speciation. Science 159: 1065-1070. 

. 1973. Animal Cytology and Evolution. 3rd Ed. Cambridge 

Univ. Press. 961 pp. 

-, R. E. Blackith, R. M. Blackith, and ]. Cheney. 1967. 



Cytogenetics of the viatica group of morabine grasshoppers. L The 
"coastal" species. .Vustralian J. Zool. 15: 263-302. 
Williams, E. E. and T. P. Webster. 1974. Auolis rupinae. new species, 
a syntopic sibling of A. monticola Shreve. Breviora Mus. Comp. Zool. 
429: 1-22. 

NOTE ADDED IN PRESS 

Since we wrote the above, Vegni Talluri et al. (1975) pub- 
lished kanotypes identical to those Tropidiirus karyotypes given 
here for the additional two species, T. jacobi (James Bay, James 
Id.) and T. indejatigabilis (Academy Bay, Indefatigable Id.); 
and for two additional populations of T. alhemarlensis (Vil- 
lamil, Albemarle Id. and Punta Espinoza, Narborough Id.). 
These data further support our conclusion that chromosomal 
differentiation plays no functional role in classical geographic 
speciation and add contrast to the situations of frequent as.so- 
ciation between the fixation of chromosomal differences and 
speciation which does not involve obvious geographic separa- 
tion. 

ADDITIONAL REFERENCE 

Vegni Talluri, M., R. Dallai, and B. Lanza. 1975. The karyotype of 
some Tropidiirus (Reptilia, Iguanidae) from the Galapagos Islands. 
[published reprint from: Galapagos, Studi e Ricerche — Spedizione 
"L. Mares — . G.R.S.T.S."]. 9pp. Gruppo Ricerche Scientiliche e 
Tecniche Subacquee, Firen.:e (Italy) . 



^'^^S. CCMP. ZOOL 



B R E V I 0„,]R A 

Museiiiii of Comparative Zoology 



us ISSN 0006-9698 



Cambridge. Mass. 27 May 1977 Number 442 

A LABORATORY STUDY OF THE TURKISH HAMSTER 
MESOCRICETUS BRANDTI 

Charles P. Lyman' and Regina C. O'Brien' 

Abstract. Ten years of laboratory observations of a breeding colony of Meso- 
cricetus from central Turkey are summarized. From morphological and karyological 
evidence, as well as cross-breeding experiments, it is concluded that Mesocricetus 
brandti is distributed in specific areas throughout Asiatic Turkey, south into Pales- 
tine, Syria, the northern part of Iraq, northwest Iran, and over the Caucasus into 
Daghestan. The chromosomal polymorphism noted in this species may be due to the 
isolation of groups to these specific types of terrain. 

The reproductive cycle, growth, and care and behavior in the laboratory are de- 
scribed. Records of hibernation are detailed and compared with those of Mesocri- 
cetus auratus. It is emphasized that M. brandti offers a unique opportunity to study 
the factors which influence hibernation using matched animals of known age and 
lineage. 

INTRODUCTION 

The study of hibernation in mammals has long been hampered by 
the lack of an animal which could be bred and raised readily under 
laboratory conditions. Ground squirrels, marmots, dormice, Euro- 
pean hamsters, hedgehogs and microchiropteran bats have been used 
extensively as experimental animals, but all of these species are 
usually obtained by collecting them in the wild. 

The Syrian hamster is easily raised in the laboratory and has been 
used as an experimental animal for more than forty years. How- 
ever, hibernation in this species is extremely unpredictable, and 
many individuals fail to hibernate even when exposed to cold for 
many months (Lyman, 1948, 1954). 



'Department of Anatomy, Harvard Medical School 

Boston, Massachusetts, 02115 and 

Museum of Comparative Zoology 

Harvard University, Cambridge 

Massachusetts 02138 



2 BREVIORA No. 442 

In August, 1965, we obtained 13 hamsters trapped in central 
Turkey.' These animals hibernated readily when exposed to cold 
(5 ± 2°C) the following winter and bred in the laboratory during 
the next spring. Since that time, the colony has been continued, 
detailed notes have been maintained, and various experiments have 
been undertaken. In 1971, we received an additional 29 animals. 
We maintained and bred these animals separately from the original 
group for comparative data. 

The two groups resembled each other closely, and clearly differed 
from laboratory specimens of the Syrian or golden hamster, Meso- 
cricetus auratus. As information concerning the Turkish hamsters 
accumulated, it became apparent that this species is the best experi- 
mental animal for the study of certain aspects of hibernation, par- 
ticularly those which require animals of known history and Uneage. 
Offspring of the two original colonies have been distributed to sev- 
eral investigators and we hope that the Turkish hamster is now 
established as an experimental animal. For this reason, our knowl- 
edge concerning this hamster is detailed below. 

Identification is, of course, of critical importance in animals used 
for experimental studies. However, because of the Hmited amount 
of comparative material available, comparisons of skins and skulls 
have led to conflicting opinions on the taxonomy of Mesocricetus 
(Ognev and Heptner, 1927; Aharoni, 1932; Ellerman, 1941, 1948; 
Vinogradov and Argiropulo, 1941; Ellerman and Morrison-Scott, 
1951; Vereshchagin, 1959; Hamar and Schutowa, 1966; Ivanov, 
1969; Raicu et al., 1969; Vorontsov and Krjukova, 1969; Todd et al., 
1972). The genus exhibits chromosomal polymorphism while show- 
ing relatively few morphological differences over a wide geographic 
area. Emphasis by some investigators on the analysis of karyotypes 
has led to divergent taxonomic views which can not be fully recon- 
ciled until more definitive data such as chromosomal banding studies 
and cross-breeding experiments in the laboratory have accumulated. 
For this reason we have limited ourselves to a consideration of the 
identity of the Turkish animals and have attempted to delineate 
their distribution in their natural habitat. 



'We are grateful to Dr. Bahtiye Mursaloglu of the University of Ankara, Ankara, 
Turkey for making arrangements to obtain the hamsters, and to Mr. Haluk Anat of 
the same university for trapping the hamsters and supplying us with information con- 
cerning their natural history. 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 3 

TAXONOMIC MATERIALS 

The animals which formed our original colony in 1965 were cap- 
tured by Mr. Haluk Anat in Malya, near Kirsehir (39°N, 34°E)^ 
about 125 kilometers (km) southeast of Ankara, Turkey (Fig. 1). 
Mr. Anat trapped the second group six years later near the city of 
Ankara. We refer to these two groups collectively as "Turkish" 
hamsters in this text. 

Skins and skulls of hamsters from Malya which had been trapped 
alive and kept in captivity for varying periods of time were used for 
comparison with the specimens of Mesocricetus in the British Mu- 
seum.^ As far as we have been able to determine, the British Mu- 
seum collection is the only representative collection of this genus in 
the United Kingdom or the United States. It contains specimens 
from a now defunct colony in the Wistar Institute which were ob- 
tained from Pirbadan [m] (35° N, 48° E), Iranian Kurdistan, a 
small village 125 km north-northwest of Hamadan, Iran (Fig. 1). 
Hamsters from this colony have been the subject of karyological 
and breeding studies and have been referred to as "Kurdistan" or 
"K" hamsters^ (Lehman and Macpherson, 1967; Palm, Silvers 
and Billingham, 1967; Todd et al.. 1972; Raicu et al., 1972). 

We observed four hamsters originating from a colony of Ru- 
manian hamsters maintained by Dr. Petre Raicu, University of 
Budapest', alive in our laboratory. We used two skins and skulls 
from the Rumanian colony to compare with the specimens in the 
British Museum. In addition, hamsters trapped for Dr. Michael 
Murphy in Aleppo, Syria were maintained in our laboratory for 
comparative studies and breeding experiments. 

Breeding crosses were usually attempted by placing the female in 
the cage of the male, though occasionally the reverse process was 
used. In many instances we used vaginal smears to determine the 
estrous cycle, and exposed the female to the male in the afternoon 



2The report that these animals came from Aksehir, Turkey (Todd et al., 1972) is an 
error derived from faulty original information. 

3We thank Drs. J. E. Hill and G. B. Corbet of the British Museum for making these 
specimens available 

"All place names in this paper are those given in the original articles. 

5We are grateful to Dr. William Nixon of Randolph, Mass. for supplying these 
animals. 

*Dr. Murphy made a trip to Aleppo to obtain this colony and observe their be- 
havior. We acknowledge his gift with thanks. 



BREVIORA 



No. 442 




SC*L£ I "~"~ I 



Figure 1. Map showing distribution of A/cvorr/cf/Rv. 

prior to estrus. If the animals fought, they were separated tem- 
porarily, and pairing was attempted at later dates. Otherwise, the 
animals were left together for three or four days and then separated. 
The animals were observed for copulatory behavior and vaginal 
smears were often made if copulation was suspected but not ob- 
served. 

We prepared karyotypes from metaphase spreads obtained from 
normally dividing primary skin cultures. The cells were harvested 
on the seventh to ninth day of culture. After exposure to colchicine 
(final dilution 36 micrograms per milliliter of medium) for five 
hours, and hypotonic solution for 10—15 minutes, cells were fixed 
in 3:1 methanol:glacial acetic acid and stained with Wright-Giemsa 
stain. 

Chromosomes were counted in at least 100 cells per animal. Se- 
lected metaphase spreads were photographed and homologous 
chromosomes were paired in the conventional manner according 
to their length and position of the centromere. 

DISCUSSION OF TAXONOMY 

It is generally agreed that the distribution of the genus Meso- 
cricetus includes the eastern parts of Rumania and Bulgaria, 
across Anatolian Turkey into the Caucasus, and south into Syria, 
the northern tip of Iraq and the northwest part of Iran. 

Hamar and Schutowa (1966) have presented the most complete 
taxonomic investigation of the genus in recent time. They had 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 5 

available for comparison 102 specimens from the Moscow and 
Leningrad Zoology Museums as well as the Humbolt Museum in 
Berlin. Based on morphological data, they recognized three species 
of one group. They proposed that Mesocricetus newtoni was found 
in eastern Rumania and Bulgaria and that Mesocricetus auratus 
was confined to the type locality of Aleppo, Syria. Mesocricetus 
brandti appeared to occupy the rest of the range of Mesocricetus 
except for an area north of the Caucasus which was occupied by 
Mesocricetus raddei and M. r. nigriculus. 

Based on cranial measurements, they found little difference be- 
tween newtoni, auratus and brandti, but the two forms of raddei were 
distinctly different, with relatively massive skulls and larger molar 
teeth. The bodies of animals of the raddei group were also larger 
in size, and the black color of their bellies contrasted with the grey 
or white of the smaller newtoni-auratus-brandti group. 

Our examination of specimens of raddei from the Caucasus in the 
British Museum and the Museum of Comparative Zoology con- 
firmed the observations of Hamar and Schutowa (1966). These 
authors sumjnarized their findings by suggesting that raddei should 
be separated from the other group at the subgeneric level, and the 
evidence presented for this view is convincing. 

When compared to specimens from Rumania or Turkey, the dor- 
sal pelage of the Syrian or golden hamster is notably different. The 
color of the latter is a rich Sanford's brown (Ridgway, 1912) while 
our animals from Turkey are tawny-olive to Saccardo's umber dor- 
sally, with an overlay of black-tipped guard and pile hair which 
darkens the pelage. Although Hamar and Schutowa do not empha- 
size this, our animals from Rumania were darker dorsally than our 
animals from central Turkey, as were several of the seven Rumanian 
specimens in the British Museum. The subauricular stripe in the 
Syrian animals is poorly defined, being a mixture of brown and black 
hairs, while in the Turkish and Rumanian animals it is pure black. 
There is a difference in the coloring of the chest, for in the Syrian 
hamster it is brownish with a narrow white mid-stripe, while in the 
Turkish and Rumanian animals there is a pronounced bar of black, 
as illustrated by Hamar and Schutowa (1966). The Kurdistan or "K" 
specimens were paler on the dorsal surface but otherwise closely re- 
sembled our animals, which were trapped approximately 1 350 km to 
the west-northwest. 

It is of incidental interest that the pelage of the type specimen 
of M. auratus in the British Museum is indistinguishable from speci- 



6 BREVIORA No. 442 

mens which Dr. Murphy collected in Syria in 1971 and these in turn 
match the common laboratory Syrian hamster. Thus, laboratory 
breeding since 1930, when the first animals were captured (Adler, 
1948), has not altered the natural coat color of this animal, though 
many mutations in coat color have occurred. 

More than one thousand Turkish hamsters have been raised in our 
laboratory, and slight differences in the pelage have been observed. 
Some individuals have darker dorsal pelage than others, and the 
ventral surface may vary from grey-white to grey, particularly on 
the abdomen. There are no consistent differences in color between 
the colony which originated from Malya and the colony from An- 
kara. 

The four living animals from Rumania were about the same size 
as laboratory strains of Syrian hamsters, but our Turkish hamsters 
were bigger as full-grown adults. Twenty-one two-year-old Turkish 
hamsters chosen at random averaged 163 grams (g) with a range of 
137 to 258 g. In comparison. 111 two-year-old Syrian hamsters 
averaged 105 g with a range of 97 to 113 g (Ahman and Dittmer, 
1964). The heads of the living animals from Rumania differed from 
the others, for the face was more pointed and ratlike, though the 
nasal portion of the skulls of Rumanian animals is not narrower. 
Laboratory interbreeding experiments have been detailed by Todd 
et al. (1972) and can be briefly summarized. No offspring have re- 
sulted frorr ratings between our animals from Malya or Ankara 
in Turkey laboratory animals originating in Syria (Todd et al., 
1972; presen study), though copulation has been observed (Mur- 
phy, persi dl communication; present study). Attempts to cross 
Kurdistan a limals with laboratory Syrian hamsters have been simi- 
larly unsuccessful (Palm et al., 1967). 

The cross between Rumanian female and Malya male hamsters 
has produced viable offspring, but the reciprocal cross has not been 
attempted. Todd et al. (1972) report the survival of five females, 
with a male living only two weeks. In our laboratory, a female killed 
her first litter and two females survived in the second litter, both 
of which had developed diabetes mellitus by the age of nine weeks. 
Diabetes was not the invariable result of this cross, however, for we 
tested a female raised by Todd et al. and found the urine negative 
for glucose. The hybrids were larger than either parent, with one 
weighing 293 g as an . .ilt. T j et al. report copulation between 
a hybrid Rumanian-TurkisV female and ? Malya male, but presented 
no evidence that the hybrid was fertile. 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 7 

Viable hybrids have been produced in the cross between female 
hamsters from Kurdistan and males from Rumania, but the recip- 
rocal cross was unsuccessful (Raicu et al., 1972). (Dr. M. Bah- 
manyar of the Pasteur Institute of Iran collected both Dr. Raicu's 
and the Wistar Institute hamsters. He writes that the former ani- 
mals were trapped 25 km west of Pirbadan.) The number of young 
per litter was small and averaged about 50 per cent of the number 
expected in non-hybrid litters. Histological examination revealed 
that both fhe male and female reproductive organs of the adult 
hybrids were atrophic, and on this basis both sexes were consid- 
ered to be totally sterile. 

Raicu and Bratosin (1968) and Raicu et al. (1969) have obtained 
reciprocal crosses between Syrian and Rumanian hamsters. Though 
the litters were small, the individual animals were large and the 
gonads of the hybrids appeared histologically atrophic, leading to 
the conclusion that the animals were sterile. Todd et al. (1972) found 
that the hybrid females resulting from crosses between Rumanian 
males and Syrian females would mate with Syrian males. No viable 
young were produced, though reabsorbing embryos were found at 
autopsy in three cases. 

In contrast, there was no indication of lack of fertility between 
the two groups of animals from Turkey. Reciprocal crosses in our 
laboratory between hamsters originating in the Ankara and Malya 
areas result in normal healthy litters and the Fi generations are fer- 
tile, producing normal young 

The karyotypes of the various groups of hamsters, including 
hamsters in our laboratory which originated in Malya, have been 
reported by Todd et al. (1972). Since that time we have established 
the karyotype of hamsters from the area of Ankara. The latter 
animals have a diploid number (2N) of 44 chromosomes with a 
fundamental number of autosomal arms (FN) of 80 (Fig. 2), while 
in the hamsters from Malya 2N = 42 and FN = 78 (Fig. 3). Hybrids 
of both sexes were examined and in these 2N = 43 and FN = 79. 

In the animals from Ankara, the autosomal complement is com- 
posed of two pairs of metacentric chromosomes (#3 and #13), two 
pairs of acrocentric chromosomes (#19 and #21) and the remaining 
16 pairs, which are a graduated series of submetacentric and subte- 
locentric chromosomes. Because of the similarity and gradation in 
size of the majority of the chromosomes, it was not possible to 
identify with complete certainty many individual chromosomes, 
including X and Y. 



BREVIORA No. 442 



a II n u n n 



X Y 



115 H -5X U M 



lii u X2I u fin 



IX fii kK AA. 



* « no 



Figure 2. Representative karyotype of Mesocricetus brandti from the area of Ankara, Turkey. 
Autosomes are arranged sequentially from upper left to lower right. 



In comparing the karyotype of the Ankara hamsters with that ot 
the Malya group the same difficuhy obtained, yet it is obvious that 
the difference in diploid number between the two groups is the result 
of the absence of the acrocentric pair #19 in the Malya hamsters 
leaving only one pair of acrocentric chromosomes. The hybrid ani- 
mals have three acrocentric chromosomes, two small ones and one 
slightly larger. The X chromosome of the Malya hamster is unmis- 
takably larger than that of the hamster from Ankara. 

Comparison of the karyotype of the Ankara hamster with that of 
the Kurdistan hamster (Lehman and Macpherson, 1967) reveals 
certain differences. In the Kurdistan hamster 2N = 42 and FN = 80. 
The smallest autosomal pair appears similar to #20 in the Ankara 
karyotype and there are no acrocentric elements. The Y chromo- 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 9 

i U u u u u 

XY 

:)i u n n u 



H n ih u u 



hh SX H IB 



AO 



Figure 3. Representative karyotype of Mesocricetus brandtiiiom Malya, Turkey. 
Autosomes are arranged sequentially from upper left to lower right. 



some of the Kurdistan hamster is obviously much larger than that 
of the Ankara animals. 

From the evidence which has accumulated to date, a few conclu- 
sions can be drawn. It is apparent that the hamsters of the genus 
Mesocricetus from Anatolia to at least as far east as Iran are not 
subspecies of the Syrian hamster, Mesocricetus auratus, as some 
authors have suggested (Ellerman and Morrison-Scott, 1951; 
Vereshchagin, 1959). While wild-caught Syrian, Turkish, or Ru- 
manian hamsters breed readily in captivity with animals from the 
same area, all attempts to cross the Syrian with the Turkish or 
Kurdistan hamster have failed. If the animals will not breed in the 
laboratory under a wide series of conditions, it is unlikely that they 
would breed in the field. The bright golden brown color of the back. 



10 BREVIORA No. 442 

the absence of black subauricular and chest patches and the creamy 
white belly plus the smaller size of the Syrian hamster reinforce the 
concept that M. auratus is a separate species. 

When Syrian and Rumanian hamsters are compared, the differ- 
ence in pelage is equally striking. These animals are similar in size, 
but the Rumanian animals are much darker dorsally and on the 
belly, have darker chest patches, and the ratlike face is obvious in 
the live animals. Although viable crosses have been produced in 
the laboratory, both histological data and attempts at breeding 
indicate that the Fi generation is sterile. On the basis of karyologi- 
cal studies (Raicu and Bratosin, 1968) as well as electrophoretic 
and chromatographic analysis of the blood serum, Raicu et al. (1968, 
1969) stated that the Rumanian hamster, Mesocricetus newtoni, 
was a different species from the Syrian hamster, Mesocricetus 
auratus, and the data given above reinforce this conclusion. 

The Rumanian hamster is smaller and darker than the Turkish and 
Kurdistan animals, and the facial appearance in the live Rumanian 
animals is distinctive, but these differences are not sufficient for 
specific separation. Crosses between Rumanian hamsters and ani- 
mals from Malya in central Turkey or from Kurdistan produce 
viable young, but the Fi generation is sterile. Furthermore, the 
occurrence of diabetes in the Rumanian-Turkish cross, and the 
small litters and failure of reciprocal crossing in the Rumanian- 
Kurdistan matings, are evidence that the crosses encounter biologi- 
cal barriers sufficient to separate them as species. 

When Lehman and Macpherson (1967) described the karyotype 
of their hamsters from Kurdistan, the animals were tentatively iden- 
tified by the British Museum as Mesocricetus brandti from skins 
and skulls. Our subsequent comparison with skins and skulls of 
hamsters from Malya reveal that the two groups resemble each other 
closely, though the Kurdistan animals are lighter on the dorsal sur- 
face. Color photographs of these animals alive, lent through the 
kindness of Dr. R. E. Billingham of the Wistar Institute, indicate 
that the animals do not possess the ratlike face which seems to be 
typical of hamsters from Rumania. After karyotypic examination 
of hamsters from Kurdistan, Malya, Rumania and Syria, Todd et 
al. (1972) concluded that the karyotypes of animals from Kurdistan 
and Malya most closely resembled one another. 

The evidence is persuasive that the animals from Malya and An- 
kara and the Kurdistan hamsters are all Mesocricetus brandti. 
The lighter dorsal pelage of the Kurdistan animals, when compared 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER H 

to the two colonies from central Turkey and the specimens in the 
British Museum, suggest that the Kurdistan animals may be a sub- 
species of M. brandti. In this regard, it is of interest that Mr. Anat, 
who is a trained field naturalist, has written us that the animals in 
the northeast area of Turkey near Kars are larger and darker than 
hamsters from central Turkey. Several subspecies of M. brandti 
may exist in Turkey, Armenia and northwest Iran, separated by 
various geographic barriers. 

We conclude that there are three distinct species of small hamsters 
of the genus Mesocricetus in areas south and west of the Caucasus. 
Mesocricetus newtoni inhabits the Dobruja area in the eastern part 
of Rumania and Bulgaria and, according to Hamar and Schutowa 
(1966), probably is not found west of the Danube in Rumania. 
Mesocricetus brandti ranges across Anatolian Turkey into the 
northern part of Iraq and the northwestern portion of Iran. Hamar 
and Schutowa (1966) and Aharoni (1932) report that M. brandti 
is found south of Aleppo and Aharoni indicates the southern limit 
is at the latitude of the Sea of Galilee. In contrast to this wide dis- 
tribution, M. auratus appears to be restricted to the area of the type 
locality of Aleppo, Syria. 

For definitive information on the distribution of Mesocricetus 
in and north of the Caucasus, one must turn to investigators who had 
access to live specimens or collections in Russia. Although there 
are differences of opinion on the number of subspecies involved, 
Ognev and Heptner (1927), Vinogradov and Argiropulo (1941), 
Hamar and Schutowa (1966) and Gromov et al. (1963) all agree that 
the genus in this area is clearly separable into two species, referred 
to here as M. brandti and M. raddei. M. brandti are smaller than 
M. raddei and there is no black on their bellies. In their karyological 
studies, Vorontsov and Krjukova (1969) and Ivanov (1969) also 
concluded that M. brandti and M. raddei were specifically different. 
The M. brandti studied by Ivanov came from the Daghestan area 
and Gromov et al. (1963) state that this species is found in the plains 
and foothills of Daghestan. Gromov et al. (1963) and several other 
Russian investigators remark on the discontinuity of the distribution 
of M. brandti in the Caucasus, and we find nothing in the Hterature 
to indicate that it is actually sympatric with M. raddei in any area. 

Todd et al. (1972) reviewed the reported karyotypes of M. brandti 
from Daghestan, Armenia, Iranian Azerbajzan and Erevan, and of 
a specimen which was probably from Iran. They presented a haploid 
idiogram comparing the karyotypes of the Kurdistan animals with 



12 BREVIORA No. 442 

our hamsters from Malya, M. newtoni from Rumania and labora- 
tory M. auratus. The diploid number (2N) of M. newtoni was 38, 
that of M. auratus was 44, and that of the various specimens of 
M. brandti was 42. They concluded that the differences in the 
karyotypes of the samples of M. brandti, including the fundamental 
numbers, was sufficient to suggest that the designation M. brandti 
was being applied to a group of "cryptic" species. At the time their 
data was collected, it was not known that the karyotypes of hamsters 
from Malya and Ankara differed both in diploid and in fundamental 
numbers, though the animals appeared phenotypically similar and 
produced fertile crosses in the laboratory. Zimmerman (1970) 
emphasizes the frailty of karyological data when used by itself to 
determine taxonomic relationships and the breeding success be- 
tween the Malya and Ankara group reinforces this concept. 

Recent evidence indicates that some rodents may be genetically 
isolated though phenotypically similar. Spala.x (Nevo and Shkol- 
nik, 1974), Perognathus {Patton, 1972), Thomomys (Panon, 1973) 
and Peromyscus (Schmidly and Schroeter, 1974) are some of the 
rodents which have marked chromosomal differences in groups from 
restricted, contiguous areas. Patton and Dingman (1970) point 
out that the diversity in karyology often occurs in burrowing rodents 
with limited dispersibility and small breeding populations. Field 
reports, particularly those of Argiropulo (1939), andlour observa- 
tions of "field" conditions in the laboratory indicate that hamsters 
are fossorial rodents, though their way of life is not comparable to 
that of an animal such as Thomomys, which spends virtually its 
whole existence underground. Aharoni (1932) and Argiropulo 
(1939) both emphasize that Mesocricetus is found in very specific 
types of habitat and Argiropulo states that the distribution oi brandti 
in Armenia is limited to specific altitudes. Mr. Anat has written us 
that there was no available open water in the areas where he trapped 
our animals, yet we find that hamsters in the laboratory which are 
fed laboratory rat chow cannot survive without water to drink. 
Thus, brandti in some areas must depend on metabolic water and 
water in their food, and this must limit their distribution. Further- 
more, Argiropulo states that they never live in wooded areas. These 
factors, coupled with their slow locomotion, may limit brandti to 
very specific types of habitat and isolate one group from another, 
which may explain why the karyotype of each population of this 
species examined to date differs from the other. 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 13 

Aharoni (1932) points out that the Syrian hamster is confined to 
the immediate vicinity of Aleppo and suggests that the very dry 
climate has resulted in the recognizably different M. auratus. This 
population appears to have been isolated long enough to become 
specifically distinct, a conclusion that is supported by the cross- 
breeding experiments. Evolutionary factors of less influence may 
be causing the differences in karyotypes of M. brandti without 
producing reproductive isolation or obvious differences in pheno- 
type. 

NATURAL HISTORY 

Published observations of M. brandti under natural conditions 
are scanty, the most complete being that of Argiropulo (1939) who 
studied this species in the Caucasus. Because his paper is written 
in Russian, a brief summary is included here, supplemented with 
field notes taken by Mr. Anat when he was trapping hamsters for 
us in Turkey. Both sources agree that the habitat of M. brandti 
consists of dry, rocky steppe country sometimes bordering culti- 
vated fields. These hamsters do not inhabit wooded or bushy areas, 
and appear to avoid areas of high humidity and dampness. In the 
Caucasus they are found at altitudes as high as 2800 meters. Al- 
though they may be seen at dawn and dusk, they are mainly noc- 
turnal and feed principally at night. In the wild, they are relatively 
fearless, for Argiropulo observed that they were apt to stand on 
their hind legs facing an intruder in the agonistic posture so often 
seen in the Syrian hamster. 

In order to observe these animals under as natural conditions as 
possible, a compound measuring 160 X 280 centimeters (cm) and 
filled with clay soil to an average of 30 cm was set up in our labo- 
ratory. One part of the area was made higher than the remainder 
by the addition of rocks up to three kilograms in weight, and part 
of the flatter surface was planted with ground pine. A sunken dish 
served as a water supply and the bare soil was moistened periodically. 
During the month of July, two male and three female five-week- 
old M. brandti were released simultaneously into the compound. 
Within a few hours there were numerous holes in the soil and all 
of the animals were underground. For the next four months, no 
animal was ever seen in the broad daylight, though they were seen 
above ground when light was failing in the evening or under dim 
artificial illumination at night. 



14 BREVIORA No. 442 

During the period of observation, the animals were given Purina 
laboratory chow (Ralston Purina Co., St. Louis, Mo.) as food, which 
was thrown at random on the ground. Most of the food which was 
not eaten was stored underground during the night. Argiropulo 
noted that M. brandti stores both grain and grasses in its burrows. 
M. R. Murphy (personal communication) reports that Mesocricetus 
auratus obtains grain by standing on its hind legs, grasping the stalk 
with its front feet or pushing it down, and cutting the stalk with its in- 
cisors. The animal then eats the fallen grain or puts it in its cheek 
pouches. Numerous stalks of tall grasses with ripened ears were 
pushed into the soil of the compound to simulate growing grain and 
the animals were observed in the evening under dim illumination. In 
spite of the fact that these hamsters were the sixth generation of ani- 
mals raised in the laboratory, with no exposure to any growing grain, 
they cut down the stalk and consumed the kernels as described above. 

After the five hamsters had been in the compound for 130 days, 
they were removed using box traps, and a diagram of their tunnel 
system was constructed by squirting silicone foam (Froth Pak, 
Insta-foam Products Inc., Addison, 111.) down the various holes 
and permitting it to harden. Once hardened, the earth was scraped 
away and a positive cast of the underground galleries was obtained. 
Although these casts show several points of interest, it must be 
borne in mind that the hamsters were limited in the depth of their 
digging by the floor of the room. 

At the time that the casts of the burrows were made, we were 
unaware o^ the work of Argiropulo (1939) with detailed descrip- 
tions of burrows which he excavated in the Caucasus. However, 
within the limitations imposed by the artificiality of the com-' 
pound, the two series of burrows were remarkably similar. Both 
Argiropulo and Anat note that the main entrance of the burrow, , 
when on a flat surface, is an almost vertical shaft extending a 
meter or more in depth. The vertical shaft usually curves quite 
abruptly to a horizontal plane, and the galleries and rooms branch 
from this shaft. Mr. Anat indicates that the hamster burrows 
can always be distinguished from those of Microtus or Citellus, 
which may inhabit the same area, because the burrows of the 
latter two enter the ground at a slant. When a hamster burrow 
was made in a bank, Argiropulo found that it was nearly hori- 
zontal. This type of tunnel was usually abandoned in a few 
days after other tunnels had been dug and the space filled in 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 15 

with dirt and feces. Both in the artificial situation and in the natural 
state there were several underground chambers. One or more of 
these chambers were used for storage of food. Another chamber 
was a nesting area which in the natural state was filled with dried 
grass. In the artificial compound, the nesting material was ground 
pine and shreds of a cardboard box that had been placed in the 
area as a surface shelter. Argiropulo found that one chamber was 
used exclusively as a latrine, but no such chamber was found in the 
artificial compound. 

To observe the method of digging employed by the hamster, a 
narrow, rectangular, clear Plexiglas box measuring 62 X 31 X 3.5 cm 
(inside dimensions) was constructed. The box was placed on edge 
with its long side down and filled with earth to a depth of 17 cm. A 
hamster placed on top of the earth was squeezed so that both of its 
sides touched the sides of the box. The animal was capable of mov-^ 
ing forward or backward, or of turning around, but it could only 
rest comfortably with its body parallel to the long axis of the box. 

After several minutes of exploration, the hamster always com- 
menced digging in the earth. The rapidly moving front feet were 
used to displace the earth, with motions like a dog burying a bone. 
The loose dirt accumulated under the chest and abdomen. To move 
this dirt, the hind feet were thrust backward together, while the 
front feet were braced. During this maneuver, the hamster arched 
its back and lifted its head. If the animal was in a tunnel, the head- 
lifting served to tamp the earth on the roof of the tunnel. As the 
tunnel or the surface excavation became longer, the earth at the 
head of the excavation was moved to the tail by a series of parallel 
kicks of the hind feet accompanied by back arching and head lifting, 
as described above. After each set of kicks, the hamster moved 
backward about one-half the length of its body and resumed moving 
the earth. Use of the mouth and cheek pouches for moving dirt 
was not observed, though the animals often took small quantities 
of earth in their mouth as if to taste it. 

The rapidity with which hamsters can move earth and dig tunnels 
marks these rodents as truly fossorial, and indicates that hamsters 
kept in laboratory cages are denied a significant part of their nor- 
mal behavioral pattern. The lifting of the head to tamp the roof 
of the tunnel apparently has functional significance, for Argiropulo 
observed that the inside of the tunnels which he excavated were 
very smooth. 



16 BREVIORA No. 442 

LABORATORY CARE AND REPRODUCTION 

Turkish hamsters in this colony are now kept in individual 17 X 
17 X 24 cm floored cages and given ample shavings for bedding. 
During breeding 23 X 23 X 38 cm cages are used. The hamsters do 
not thrive in wire bottomed cages. The animals are given water 
and Purina laboratory rat chow ad lib. with a sUce of raw apple 
and 30 g rolled oats weekly. 

In 1968-1969 the colony was greatly reduced due to "wet tail" diar- 
rhea and osteomalacia. The diarrhea was partially controlled by 
adding 0.05 per cent tetracycline hydrochloride (Polyotic, Cyana- 
mid) to the drinking water for a period of not longer than two weeks. 
Various diets, one consisting wholly of Purina mouse breeder 
chow, have been experimentally tested, but the diet outlined above 
has been most satisfactory. The cause of osteomalacia has not 
been determined, but since it no longer occurs in the colony, it proba- 
bly was due to an insufficiency in the diet exacerbated by an intes- 
tinal infection which, in its most virulent form, resulted in "wet 
tail" and death. 

Although it rarely occurs in M. auratus, several M. brandti suf- 
fered from malocclusion, so that the upper and lower incisors grew 
unchecked by wear. The unrestrained growth was remarkably rapid. 
In a three-month-old hamster the upper incisors had grown slightly 
more than a full circle with one incisor penetrating and exiting 
from the roof of the mouth. The lower incisors of a one-year-old 
hamster with malocclusion were trimmed periodically and the ex- 
cess tooth material was measured. Growth averaged 1 cm per month 
compared to growth of approximately 0.4 cm per month reported 
by Sarnat and Hook (1941) in the much larger thirteen-lined ground 
squirrel {Citellus tridecemlineatus). 

Pairs of hamsters from five weeks to two years of age were exposed 
to each other for breeding in every month of the year. The young- 
est female to produce a litter was 50 days old when she gave birth, 
and animals seven to eight weeks of age reproduce successfully 
though they have not yet attained full growth. Hamsters reach 
their peak of fecundity at about one year of age, and most of our 
breeding is carried out with animals oi this age. The tests of one- 
year-old males are large during the breeding season and histological 
examination shows active spermatogenesis. Females of this age 
almost invariably have normal estrous cycles. In the second year, 
the female estrous cycles are less regular during the breeding season 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 17 

though active spermatogenesis usually occurs in the males. Breed- 
ing is notably less successful in these animals. We have not at- 
tempted to breed three-year-old hamsters. 

In our laboratory (Boston, Mass.), animals exposed to natural 
day lenpth usually were not in breeding condition between the 
months of November and March. During this period the females 
which were examined were apt to have anestrous vaginal smears 
with a preponderance of non-nucleated squamous cells. The testes 
of some iftales were atrophic with no histological evidence of sper- 
matogenesis, and no stored sperm in the epididymides. Other 
animals of both sexes had all the anatomical indications of full 
breeding condition. The testes were the same size as those of sum- 
mer animals and histological examination revealed spermatogenesis, 
while biopsy of the epididymides showed motile sperm. Vaginal 
smears indicated normal estrous cycles. 

In 1971, pairs of hamsters were exposed to each other from Janu- 
ary to September. As was expected, successful breeding was low 
for the first three months of the year, improved in the next two 
months and then declined. Thus, from January 20 to April 3 there 
were 1 1 litters out of 33 exposures. Successful matings peaked in 
May, with seven litters born out of 13 exposures. From June 1 to 
September 3, there were only 1 1 litters born in 34 exposures. 

From 1966 to 1971 various changes in daily illumination and diet 
were carried out in order to discover the optimum conditions for 
breeding this species in the laboratory. During the first two years 
after their arrival, the animals were maintained on 10 hours of il- 
lumination in each 24 hour period. Breeding success was high dur- 
ing the first summer (1966) with a total of 109 young in 16 litters 
being produced by 23 pairings from the wild-caught group of six 
males and seven females. During the next summer there was a 
notable decline in successful matings. Of a total of 79 pairings with 
42 males and 45 females, only 1 1 1 young in 15 litters were produced. 
In the case of unsuccessful pairings, the female always appeared 
to be the unwilling one of the pair. 

In the difficult years of 1968 and 1969 the colony barely main- 
tained a steady population, but by 1970 a successful diet had been 
developed, the daily lighting had been standardized to the illumina- 
tion of this latitude, and attempted breedings were usually limited 
to the spring and summer months. Under this regime, breeding 
success improved, with 17 litters in 34 matings in 1974. 



18 BREVIORA No. 442 

Hoffman and Reiter (1965) and Elliott et a!. (1972) have reported 
that the reproductive cycle of the male Syrian hamster is responsive 
to photoperiod. A controlled experiment to test this responsiveness 
in the Turkish hamster has not been carried out, but the birth of 
litters in January and February, with the day length near the mini- 
mum, shows that the photoperiodic effect is not absolute. Data 
presented below indicate that hibernation is involved in a much 
larger segment of the yearly life of the Turkish hamster than is the 
case in the Syrian species. Turkish hamsters which are exposed to 
a constant temperature of 22°Clin the laboratory during the winter 
months are living in an environment which they would never en- 
counter for long periods in the natural state. It is not surprising 
that photoperiod alone does not control the seasonal sexual cycle 
under these circumstances. 

The breeding behavior of the Syrian hamster has been described 
by Murphy and Schneider (1970) and the behavior of the Turkish 
hamster is similar. When the animals were paired in the afternoon, 
there was usually some preliminary exploration, but not much ac- 
tivity occurred until nightfall. In some cases, however, an immedi- 
ate antipathy developed and violent fighting occurred at once. 

After the first half hour of exposure, fighting was not as frequent, 
but it could occur at any time. If the animals were kept together for 
more than four days the chances of fighting increased, perhaps be- 
cause the female was pregnant. Fighting was always violent, with 
the female biting the male near the head region and then clawing 
him with both front and rear feet. The male, in spite of the punish- 
ment he received, often followed the female and attempted to copu- 
late, only to be attacked again. If the male retreated, the female 
often pursued him and bit him on the flanks and hind quarters. 
Attempts to reduce the damage to the male by providing various 
shelters were unsuccessful because of the male's persistence in pur- 
suing the female. Often the punishment received by the male was 
fatal. At autopsy the cuts and bruises did not seem sufficiently 
severe to cause death and it is possible that exhaustion was a con- 
tributing cause. 

The estrous cycle of the Turkish hamster is similar to that de- 
scribed by Kent (1968) for the golden hamster. The cycle is of four 
days duration and estrus is characterized by the appearance of 
large numbers of nucleated epithelial cells in the vaginal smear. 

Gestation is 15 days in length and the litter size has varied from 
one to 13 with the average being six (N young— 1021,Nlitters= 177). 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 19 

No female has produced more than three litters during one breeding 
season and Argiropulo (1939) reports that two litters a year is maxi- 
mum for wild animals in the Caucasus. The female becomes ex- 
tremely apprehensive before the birth of her litter, and partial 
darkness and ample bedding are a necessity. Even if left undisturbed 
she may kill and eat the young, and if she is handled during parturi- 
tion the destruction of the litter is inevitable. Perhaps a littering 
cage which approximated the fossorial existence in the wild would 
reduce thfs nervousness, but this has not been tested. 

Hamsters are born blind and naked, but they quickly grow a 
protective coat. By the twelfth to thirteenth day the eyes open and 
the young eat rolled oats, ground laboratory chow and drink from 
the water bottle. The young are weaned by the mother before the 
twentieth day, and we routinely separate the young from the 
mother at four weeks of age. 

The young live together amicably until about the seventh week, 
when fighting starts to occur. Usually one animal is the aggressor, 
but if it is removed from the litter, another soon takes up the role. 
The attacked animal is bitten violently, resulting in large wounds 
from tears in the tender skin, and these are often fatal. Presumably, 
under natural conditions this aggressive behavior causes dispersion 
of the litter with httle harm to its members, but in the laboratory 
it is a serious problem. Although we have used several types of 
shelters, we have not been able to keep Turkish hamsters in collec- 
tive cages and must separate and maintain them in single cages at 
about six weeks of age. Obviously this is a serious disadvantage 
for maintaining a large colony. 

The young hamsters grow rapidly, and healthy litters average 
91 g in weight with a range of 50 to 141 g (N = 41 1) at six to seven 
weeks of age. Animals born after June do not attain sexual maturity 
until the following spring. 

AGING 

Turkish hamsters in captivity tend to become obese after the 
first year of Hfe and sometimes weigh over 200 g, with our heaviest 
recorded animal attaining 285 g. It is unlikely that hamsters in the 
wild, with limited food supply and an active life, would ever reach 
this weight. 

The median hfe-span of 43 animals which were bred and kept in 
the laboratory at a room temperature of 22 ± 3°C was 670 days 
with a range of 231 to 1399 days. After about two years of age, the 



20 BREVIORA No. 442 

pelage usually loses its glossy smoothness and the animals present 
a scruffy appearance. The effect of hibernation on aging in this or 
any other species of mammal is unknown and an experiment is in 
progress to determine if these hamsters age less rapidly during 
hibernation. 

HIBERNATION 



During the past ten years, quite complete records of the colony 
of M. brandti have been maintained, so that some information can be 
gleaned concerning the putative factors which control hibernation. 
Previous observations on M. auratus in this laboratory (Lyman, 
1954; Lyman, unpublished observations) provide comparison be- 
tween the two species. 

In those studies, all hamsters were moved to a cold room kept at 
5 ± 2°C and lighted 8 to 10 hours a day. The animals were housed 
in individual cages with ample shavings and food and water ad lib. 
Hibernation was monitored by dusting the backs of hibernating 
Syrian hamsters with shavings. If the shavings were still in place 
at the next observation, hibernation was assumed to be continuous. 
This technique is common among students of hibernation (Mrosov- 
sky, 1971) but is not always accurate. Because hamsters often bur- 
row in their bedding before hibernation, shavings occasionally be- 
come lodged on the back and the hamster may be erroneously scored 
as being in continuous hibernation. Following the technique of 
Johnson (1931), extraneous material (rolled oats) was placed on 
the backs of the hibernating Turkish hamsters. Arousal from hiber- 
nation invariably displaced this material. The result of this differ- 
ence in technique is to make the Syrian hamsters appear to be "bet- 
ter" hibernators than they actually are. 

The timing and period of cold exposure was not the same for 
M. auratus and M. brandti. The former were moved to the cold 
at various times of year and remained there until they hibernated 
or died. In contrast, because it was necessary to enlarge the M. 
brandti colony, most of these animals were exposed to cold in the 
autumn and removed to the warm room for breeding in the spring. 
The fact that the M. auratus were moved to the cold in all seasons 
of the year should have no effect on the comparisons given below. 
Smit-vis and Smit (1963) have theorized that there is a seasonal 
preparation in this species which results in a greater tendency for 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 21 

hibernation in February. The evidence is not persuasive, however, 
for it is based on 26 animals, two of which are excluded from the 
basic computations. The paper relates the month in which the ani- 
mals were exposed to cold to the lapse of time before hibernation 
occurred, but there are no data for April through July or for Sep- 
tember and October, and there is data for only one animal in August 
and two in March. From our records of over 2800 Syrian hamsters, 
placed in the cold in every month of the year, we adduce no con- 
vincing evidence that the tendency to hibernate is seasonal in this 
species. 

In studies such as this, an as yet unresolved problem concerns 
the criteria which should be used to distinguish "good" from "poor" 
hibernators. There are at least five interdependent factors which 
can be considered in such a comparison. These are: 1) the number 
of animals in a group which hibernate during the period of exposure 
to cold, 2) the span of time between cold exposure and hibernation, 
3) the length of the hibernating season, 4) the amount of time spent 
in hibernation compared to the amount of time in the cold, and 
5) the length of the period during which an animal remains in con- 
tinuous hibernation. Concerning the fifth factor, Pengelley and 
Fisher (1961) have shown that the lengths of the periods or bouts 
of hibernation are quite predictable with individual Citellus lateralis, 
but these lengths may vary greatly in a group. We find that the same 
situation obtains in Mesocricetus so that the maximum time of un- 
interrupted hibernation is more definitive than an average of the 
lengths of the bouts of hibernation. Until more is known about the 
subtle physiological nuances which determine whether an animal 
does or does not hibernate, it is not practical to attempt to assign 
meaningful values to the five factors listed above, nor to make 
statistical comparisons. Therefore, in the following studies as many 
factors as possible are considered but no attempt is made to weight 
their importance. 

In a group of 373 Syrian hamsters exposed to the cold, 68 percent 
or 252 hibernated and 121 animals died before ever entering the 
hibernating state. The shortest period before hibernation occurred 
was three days of cold exposure and the longest was 218 days (Ly- 
man, 1954). Once hibernation occurred, the period during which 
M. auratus hibernated lasted about three months. An accurate 
figure on the percentage of time spent in hibernation cannot be 
obtained because many of the animals were used in acute experi- 
ments and the best hibernators were usually chosen. The longest 



22 BREVIORA No. 442 

recorded bout of hibernation was 21 days and the usual period of 
hibernation was a week or less (Lyman, 1954). 

The hamsters from which these data were collected were orig- 
inally obtained from commercial dealers and hence must be de- 
scendants of the trio which were the foundation stock of the labo- 
ratory hamsters used today (Adler, 1948). Twelve M. auratus, 
which were the first generation of animals obtained in the field near 
Aleppo by Dr. M. Murphy, were exposed to the cold in the autumn 
of 1971 under the conditions described above. Although the sample 
is small, there was no detectable difference in the tendency of this 
group of animals to hibernate when compared to laboratory ani- 
mals. Thus, forty years of domestic breeding with no selection pres- 
sure for hibernation has not changed the pattern of hibernation in 
M. auratus. 

These "native" Syrian hamsters provide some information con- 
cerning the percentage of time spent in hibernation. The group 
was moved to the cold on November 15, 1971 and taken from the 
cold on May 26, 1972. No animal hibernated before January and 
hibernation occurred only 11.3 per cent of the total time. The best 
months for hibernation were January through April, during which 
time the animals were in hibernation 13.7 per cent of the time. 

Between the years 1971 and 1975, a group of 318 M. brandti v>iqvq 
exposed to cold for periods of 145 to 172 days. Two hundred ninety- 
three or 92 per cent hibernated at some time during this period. 
Eight animals died without hibernating and the remaining 17 did 
not hibernate at all. Thirty-one animals entered hibernation within 
24 hours after exposure to cold. Once hibernation started, the 
period during which this species hibernated lasted about five months, 
but for some animals the hibernating season was as long as 10 
months. In the entire group, 36 per cent of the time in the cold be- 
tween November 15 and April 15 was spent in hibernation. The 
longest period of unmterrupted hibernation was 28 days, with the 
average bout of approximately the same duration as M. auratus. 
Thus M. brandti is a "better" hibernator than M. auratus in all cate- 
gories except the average length of the bout of hibernation. 

The records of over 400 M. brandti which have been exposed to 
cold reveal no obvious factors which control or influence hiberna- 
tion in this species. However, they do give a clearer picture of the 
pattern of hibernation and its variability, and obviate the need for 
further research in some aspects of the problem. 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 23 

As reported for M. auratus (Lyman, 1948), M. brandti does not 
undergo the period of autumnal fattening and lethargy which is so 
typical of the Marmotini that hibernate. Like M. auratus, M. 
brandti almost invariably lose weight during the first two or three 
weeks of exposure to cold, with the heavier animals losing the 
greater amount. Some animals, especially the young of the year, are 
apt to regain and exceed the original weight if kept in the cold for 
several months. The amount of hibernation which occurs during 
this time is not correlated with the change in body weight. 

Examples from the spring and summer months demonstrate that 
this species will hibernate at any time of the year. Certainly, if com- 
pared to Marmotini such as Citellus lateralis or C. tridecemlineatus, 
any seasonal change in the tendency to hibernate is very poorly de- 
fined. These citellids, if experimentally naive, rarely hibernate 
when exposed to cold during the spring and summer months and 
almost invariably hibernate in the autumn. In contrast, of 66 M. 
brandti which were in the cold in June, 75 per cent hibernated at 
some time during the period and, in July, 77 per cent of 53 cold- 
exposed animals hibernated. 

The majority of our animals were placed in the hibernaculum 
between October 1 and January 2. In a sample of 257 animals, no 
particular one of the four months was favored for onset of hiberna- 
tion or for the number of days spent in the hibernating state. Wheth- 
er exposure to cold was begun in early October or in January, hi- 
bernation became much less frequent by the beginning of April 
and, when removed from the cold room in May, most of the animals 
had ceased hibernating. 

No organized experiment to test the effect of photoperiod has 
been carried out, but it is clear that the timing of the onse* of hiberna- 
tion does not depend exclusively on this factor, since hamsters hi- 
bernated in June when the daily illumination was close to its peak. 

Although 9.7 per cent of a sample of 3 1 8 animals hibernated with- 
in 24 hours after being exposed to cold, these animals did not neces- 
sarily hibernate for a greater number of days than animals which 
started hibernation at a later date. Age does not affect the ability 
to hibernate, for a comparison between animals ranging from 3.5 to 
36 months of age revealed no differences in the onset or pattern of 
hibernation. 

In spite of efforts to standardize conditions, the frequency and 
pattern of hibernation changed from year to year in the colony as 



24 BREVIORA No. 442 

a whole and with individual animals. There is no simple explana- 
tion for the fact that individuals failed to hibernate with six months 
of cold exposure one year, and hibernated under the same condi- 
tions the following year. Nor can the observation that the colony 
exposed to cold hibernated only 16 per cent of the time in 1970 and 
39 per cent in 1971 be readily explained. Conditions in the cold 
room are maintained as identically as possible from year to year, 
but changes such as shifts in caretaking personnel are inevitable. 
Even differences in handling the animals during cleaning of the 
cages might influence the onset of hibernation. 

It was observed that the first shipment of M. brandti hibernated 
67 per cent of the first sojourn in the cold and only 42 per cent in the 
second. This suggested that environmental factors might play an 
important role m the control of hibernation. The ability of ani- 
mals to store food had already been shown to have an effect on hi- 
bernation in M. auratus (Lyman, 1954) and this factor was included 
in the experimental design. 

M. brandti between five weeks and six months of age were placed 
in the following experimental conditions. As often as possible, 
sexes and litter mates were matched. On July 8 five animals were 
placed in the "natural" compound described above. For all other 
groups, each individual was put in a separate cage (23 X 23 X 38 cm) 
and given ample shavings for bedding, and water ad lib. A control 
group of 13 animals was given Purina rat chow pellets ad lib., which 
they were able to store in a corner of the cage, while another group 
of six was given ground chow in unspillable food cups which pre- 
vented these animals from storing food. Twelve other cages were 
fitted with exercise wheels, and the number of revolutions for each 
wheel was monitored by an event recorder (model A620X, Esterline 
Angus Instrument Co., Indianapolis, Ind.). The animals in six of 
these cages were given rat chow pellets ad lib. while the others were 
fed from unspillable food cups. All animals received 14 hours of 
light daily at the start of the experiment, and this was reduced by 
one hour approximately every 1 6 days. On November 1 2 all animals 
were moved to smaller individual cages (17 X 17 X 24 cm) with 
shavings, water and rat chow pellets ad lib., and on November 15 
all were moved in these cages to the cold room, where the daily il- 
lumination was 9 hours. Thus, after the preparation period, all ani- 
mals were exposed to the same conditions. Records of hibernation 
were kept for the following 137 days. 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 25 

As has been reported for many other species, the use of the ac- 
tivity wheel varied greatly from animal to animal. The nocturnal 
habits of M. brand ti were verified, for virtually no activity was 
recorded during the daylight hours. There was no correlation be- 
tween the tendency of an animal to run in the wheel and its subse- 
quent hibernation. All of the hamsters hibernated at some time 
during their exposure to cold, but no one group showed a tendency 
to enter hibernation earlier or to have longer bouts of hibernation. 
The percentage of time spent in hibernation was as follows: "natu- 
ral" enclosure, 56 per cent; activity wheel, food hoarding, 43 per cent; 
no wheel, food hoarding, 37 per cent; wheel, no hoardmg, 34 per 
cent; no wheel, no hoarding, 25 per cent. These results suggest that 
the conditions prior to cold exposure may have an influence on the 
subsequent amount of hibernation and that "natural" conditions 
increase the tendency to hibernate. However, none of the factors 
tested are crucial to the onset of hibernation. 

Experiments such as these are easily carried out with M. brandti. 
Because this animal hibernates readily and can be bred in the labo- 
ratory, it offers a unique opportunity to study the effects of various 
factors using matched pairs of known age and lineage. 

ACKNOWLEDGMENTS 

This research was supported by grants from the National Institutes 
of Health and contracts with U.S. Air Force. 

In this ten year study many people contributed their help. We are 
particularly grateful to Jane L. Bihldorff, Michael F. Carr, Helen 
Cheever, and Delphi M. Toth. We owe great thanks to Dr. N. Kala- 
bukhov of the Far East Scientific Center of the Academy of Sciences, 
Vladivostok, USSR, who has been most helpful in giving advice and 
references concerning the species of Mesocricetus in the USSR. We 
are indebted to Barbara Lawrence for her critical review of the manu- 
script, but any errors or ambiguities are uniquely our own. 

LITERATURE CITED 

Adler, S. 1948. Origin of the golden hamster Cr/c^/uiawrami as a laboratory ani- 
mal. Nature, 162: 256-257. 

Aharoni, B. 1932. Die Muriden von Palastina und Syrien. Z. Saugetierk., 7: 
166-240. 

Altman, p. L., and D. S. Dittmer. 1964. Biology data book. Fed. Amer. Soc. 
Exp. Biol., Washington, D. C. p. 95. 



26 BREVIORA No. 442 

Argiropulo, a. I. 1939. The distribution and ecology of certain mammals of 
Armenia, (translation) (Russ.) Acad. Sci. U.S.S.R., Armenian Branch, Trans. 
Biol. Inst., 3: 47-57. 

Ellerman, J. R. 1941. The families and genera of living rodents, Vol.2. London: 
British Museum (Nat. Hist.) pp. 1-690. 

1948. Key to the rodents of South-West Asia in the British Museum col- 
lection. Proc. Zool. Soc. London, 118: 765-816. 

Ellerman, J. R., and T. C. S. Morrison-Scott. 1951. Checklist of Palaearctic 
and Indian mammals 1758 to 1946. London: British Museum (Nat. Hist.) 
pp. 1 810. 

Elliott, J. A., M. H. Stetson, and M. Menaker. 1972. Regulation of testis func- 
tion in golden hamsters: a circadian clock measures photoperiodic time. 
Science, 178: 11\-117,. 

Gromov, L M., a. a. Gureev, G. A. Novikov, L L Sokolov, P. P. Strelkov and K. K. 
Chapskii. 1963. Mlekopitaiushchie fauny, SSSR (Mammal Fauna of the 
USSR) Part 1. Izdatelstvo akademii nauk SSSR. Moscow, Leningrad. (Aka- 
\demiia Nauk SSSR. Zoologicheskii Institut. Opredeliieli po faune SSSRff^l). 

Hamar, M., and M. Schutowa. 1966. Neue Daten iiber die geographische 
Veranderlichkeit und die Entwicklung der Gattung Mesocricetus Nehring, 1898 
(Glires, Mammalia). Z. Saugetierk., 31: 237-251. 

Hoffman, R. A., and R. J. REtrER. 1965. Pineal gland: influence on gonads of 
male hamsters. Science, 148: 1609-1611. 

Ivanov, V. G. 1969. The chromosome complements of A/f.somre/w.s of the Cauca- 
sus and their ta.xonomic position. In N. N. Vorontsov (ed.), Tlie mammals, 
evolution, karyology, ta.xonomy, fauna (translation). Acad. Sci. USSR, Si- 
berian Branch, pp. 105-106. 

Johnson, G. E. 1931. Hibernation in mammals. Quart. Rev. Biol., 6: 439-461. 

Kent, G. C, Jr. 1968. Physiology of reproduction. In R.A. Hoffman, P. F. 
Robinson, and H. Magalhaes (eds.). The golden hamster, its biology and use in 
medical research. Ames, Iowa: Iowa State University Press, pp. 119-138. 

Lehman, J. M.,and J. Macpherson. 1967. Thekaryotypeof Kurdistan hamster. J. 
Hered., 5«: 29-31. 

Lyman, C. P. 1948. The oxygen consumption and temperature regulation of 
hibernating hamsters. J. Exp. Zool., 109: 55 78. 

1954. Activity, food consumption and hoarding in hibernators. J. 

Mamm., 35: 545-552. 

Mrosovsky, N. 1971. Hibernation and the hypothalamus. New York: Appleton- 
Century-Crofts. vi + 287 pp. 

Murphy, M. R.,and G. E. Schneider. 1970. Olfactory bulb removal eliminates 
mating behavior in the male golden hamster. Science, 167: 302-304. 

Nevo, E.,and a. Shkolnik. 1974. Adaptive metabolic variation of chromosome 
forms in mole rats, Spala.x. Experientia, 30: 724-726. 

Ognev, S. I.,and V. G. Heptner. 1927. A new subspecies of hamster from Daghe- 
stan {Mesocricetus raddei, Nehr.) and some remarks on the Russian species of 
Mesocricetus (with a key to them). Ann. Mag. Nat. Hist. ser. 9, 19: 142-150. 

Palm, J., W. K. Silvers, and R. E. Biliingham. 1967. The problem of histo- 
compatibility in wild hamsters. J. Hered., 58: 40-44. 

Patton, J. L. 1972. A complex system of chromosomal variation in the pocket 
mouse, Perognathus bailey i Merriam. Chromosoma (Berl.), 36: 241-255. 

1973. Patterns of geographic variation in karyotype in the pocket gopher. 



1977 LABORATORY STUDY OF THE TURKISH HAMSTER 27 

Thomomys bottae (Eydoux and Gervais). Evol., 26: 574-586. 

Patton, J. L., AND R. E. DiNGMAN. 1970. Chromosome studies of pocket gophers, 
genus Thomomys. W. Variation in T. bottae in the American southwest. Cyto- 
genetics, 9: 139-151. 

Pengelley, E. T., and K. C. Fisher. 1961. Rhythmical arousal from hibernation 
in the golden-mantled ground squirrel, Citellus lateralis tescorum. Can. J. Zool., 
39: 105-120. 

Raicu, p., and S. Bratosin. 1968. Interspecific reciprocal hybrids between 
Mesocricetus auratus and M. newtoni. Genet. Res. Camb., II: 113 114. 

Raici', p., M. Hamar, S. Bratosin, and I. Borsan. 1968. Cytogenetical and bio- 
chemical researches in the Rumanian hamster (Mesocricetus newtoni). Z. 
Saugetierk., ii: 186-192. 

Raicu, P., M. Ionfscu-Varo, and D. Duma. 1969. Interspecific crosses between 
Rumanian and Syrian hamsters. J. Hered., 60: 149-152. 

Raicu, P., M. Ionescu-Varo, M. Nicolaescu, and M. Kirillova. 1972. Inter- 
specific hybrids between Romanian and Kurdistan hamsters. Genetica, 43: 
223-230. 

Ridgway, R. 1912. Color standards and color nomenclature. Washington, D. C: 
Ridgway, 53 plates & 44 pp. 

Sarnat, B. G., and W. E. Hook. 1941. Effect of hibernation on rate of eruption 
and dentin apposition in the ground squirrel. Proc. Soc. Exp. Biol. Med., 46: 
685-688. 

S'cHMiDLY, D. J., AND G. L. ScHROETER. 1974. Karyotypic variation in Peromyscus 
boylii (Rodentia: Cricetidae) from Mexico and corresponding taxonomic im- 
plications. Systematic Zool., 23: 333-342. 

Smit-vis, J. H., AND G. J. Smit. 1963. Occurrence of hibernation in the golden 
hamster, Mesocricetus auratus Waterhouse. Experientia, 19: 363-364. 

Todd, N. B., C. W. Nixon, D. A. Mulvaney, and M. E. Connelly. 1972. Karyo- 
types oi Mesocricetus brandti and hybridization within the genus. J. Hered., <53: 
73-77. 

Vereshchagin, N. K. 1959. The mammals of the Caucasus. Academy of Sciences 
of the USSR, Zoological Institute. Academy of Sciences of the Azerbaidzhan 
SSR. Jerusalem: Israel Program for Scientific Translation, pp. 1-816. 

Vinogradov, B. S., AND A. I. Argiropulo. 1941. Fauna of the USSR. Mammals. 
Key to rodents. Zoological Inst. Acad. Sci. USSR. Jerusalem: Israel Program 
for Scientific Translation. 230 pp. 

VoRONTsov, N. N., AND E. P. Krjukova. 1969. The karyotypical differentiation of 
Mesocricetus raddei and taxonomical relationships of members of the genus 
Mesocricetus. In N. N. Vorontsov (ed.). The mammals, evolution, karyology, 
taxonomy, fauna (translation). Acad. Sci. USSR, Siberian Branch, pp. 107- 
109. 

Zimmerman, E. G. 1970. Karyology, systematics and chromosomal evolution in 
the rodent genus, Sigmodon. Publ. Mus. Mich. State Univ., Biol. Ser. 4{9): 
389-454. 



"-»r\ J 



MAR 3 8 )985 

B E E V I (9rfi° A 



TV 

Museum of Comparative Zoology 

us ISSN 0006 9698 

Cambridge, Mass. 27 May 1977 Number 443 

EPIGONUS TREWAVASAE POLL, A JUNIOR SYNONYM 

OF EPIGOMS CONSTASCIAE (GIGLIOLI) 

.(PERCIFORMES, APOGONIDAE) 

Garry F. Mayer' and Enrico Tortonese^ 

Abstract The holotvpe of Pomatuinuhtlus consiaiuiae Giglioli was reexam- 
ined and compared with specimens of Epigonus irewavasae Poll from equatorial 
west Africa, northwest Africa. Madeira, and the western Mediterranean. The two 
nominal forms were not distinguishable at the species level; thus, the name Epi^onus 
consianciae should be used for the species under consideration. A description and 
discussion of the geographic variability of E. consianciae is provided. 

INTRODUCTION 

Questions about the status of Pomatoniichthys constanciae 
have recently arisen as the result of increased interest in deep-sea 
members of the family Apogonidae. P. constanciae was originally 
described from a single specimen from the Straits of Messina 
(Giglioli, 1880). The description was incomplete, unillustrated 
and included several inaccuracies. Although Giglioli designated 
the fish a new species, he indicated that the specimen resembled 
juveniles of the more common Mediterranean apogonid, Epi- 
gonus lelescopus (Risso). 

Perhaps because of Giglioli's comment and because of the 
dearth of information about Mediterranean continental slope 
ichthyofaunas, P. constanciae was generally accepted as a junior 



'MESA New York Bight Project. National Oceanic and Atmospheric Admin- 
istration. Old Biology Building. State University of New York. Stony Brook, NY. 
11794. U.S.A. 

-Museo Civico di Storia Naturale. "Giacomo Doria." Via Brigata Liguria 9. 
Genoa. ItaK. 



2 BREVIORA No. 443 

synonym of E. telescopus (e.g., Jordan and Evermann, 1896; Jor- 
dan, Evermann and Clark, 1930; Schultz, 1940; Tortonese and 
Queirolo, 1970; Tortonese, 1973). Goode and Bean (1896) were 
the last authors of a comprehensive treatise to give P. constanciae 
consideration as a separate species. Cavaliere (1956) claimed to 
have compared the juveniles of Epigonus telescopus dndPoma- 
tomichthys constanciae, but Tortonese and Queirolo (1970) and 
Tortonese (1973) deemed the study of doubtful validity. 

In 1974, Mayer published a revision of the genus Epigonus. 
Although the paper included P. constanciae as a synonym of 
E. telescopus, it pointed out that several features of P. constanciae 
were reminiscent of traits characteristic of Epigonus trewavasae 
Poll. The same work reported E. trewavasae from the Mediter- 
ranean for the first time and urged that further studies be under- 
taken on the relationship between E. trewavasae and P. con- 
stanciae. 

The current paper stems from a reexamination of the P. con- 
stanciae holotype and a comparison of this specimen with E. tre- 
wavasae from equatorial west Africa, Mauritania, Madeira and 
the western Mediterranean. Our findings indicate that P. constan- 
ciae and E. trewavasae are conspecific and that the correct name 
for the species should be Epigonus constanciae (Giglioli). 

METHODS 

Specimens were examined according to techniques outlined by 
Mayer (1974, 1975) and by Hubbs and Lagler (1958). Statistical 
analyses were performed on a Wang Model 720 Programmable 
Calculator at the Department of Marine Science, University of 
South Florida. A package program was used to calculate routine 
meristic parameters, such as means and standard deviations. 
Linear regression analyses were undertaken on mensural data 
using Ricker's method for Functional (GM) Regressions (1973: 
412^14). To insure linearity, scatter diagrams were plotted for 
all morphometric data. Characters exhibiting non-linear scatters, 
as determined by inspection, were excluded from consideration. 

Meristic characters exhibiting slight variability are reported 
in the text as the value of the character followed by the number 
of individuals exhibiting that character. Data exhibiting greater 
variability are presented in tabular form. Mensural data are 



1977 



E. CONSTANCIAE DESCRIPTION 



presented as linear regressions and/or as ratios on standard 
length (SL) or head length (HL). 

Specimens for this study were obtained from the following 
institutions: 

IRSN: Institut Royal des Sciences Naturelles de Belgique, 
Brussels 

ISH: Institut fiir Seefischerei, Hamburg 

ISTPM: Institut Scientifique et Technique des Peches Maritimes, 
L9 Rochelle 

MCZ: Museum of Comparative Zoology, Cambridge, Massa- 
chusetts 

MF: Museo Municipal do Funchal, Madeira 

MSNG: Museo Civico di Storia Naturale, Genoa 

MZF: Zoological Museum, Florence 

USNM: National Museum of Natural History, Washington. D.C. 

ZMB: Zoologisches Museum der Humboldt Universitat, Berlin 

SYSTEMATICS 

Mayer (1974: 183-186) provided a comprehensive description 
of Epigonus trewavasae. The following discussion of E. comtan- 
ciae represents a recapitulation of the former account, modified 
to reflect data obtained from a larger, more representative sam- 
ple of individuals. Readers are referred to Mayer (1974) for 
synonymies, accounts of ontogenetic change and in-depth com- 
parisons with congeners. 




Figure I. Epigonus consianciae holotype. MZF 3089, 115 mm SL. 



4 RRi-vioRA No. 443 

h'pii^onus consianiiae {Cj'\gVio\\. ISSO). Figure I. 

PoDiatomiihihys consfanciae Giglioli, 1880: p. 20; original descrip- 
tion; Straits of Messina. Holotype e.xamined: MZF 3089. 

Ei-ni^oniis iri'wavasae Poll, 1954: p. 91; figure 27; original descrip- 
tion; NOORENDE III, Sta. 52: 06 08'S, 1 1^30'E, 280 290 m. 
Holotype e.xamined: IRSN 209. 

Matfriai Examinhd 

IRSN 209 (I: 153.9 mm, holotype Epigonus trewavasae. NO- 
ORDENDE III Sta. 52. 06°08'S, 1P30'E. 280 290 m); IRSN 
16549 (I: 97.1 mm, Golfe de Cagliari, 540 m); IRSN 16550 (4: 
75.5-98.6 mm, Golfe d'Oristano, no depth recorded); ISH 386 64 
(3: 144.6-153.2 mm. WALTHER HERWIG Sta. 137 64, 15°5rN, 
I6°51'W, 200 m); ISTPM uncatalogued (10: 129.9-158.9 mm, 
THALASSA Sta. U30, I7°56'N, 16°32'W, 224 322 m); ISTPM 
uncatalogued (1: 149.0 mm, THALASSA Sta. X007, 20°59'N, 
I7°37'W, 250-330 m); ISTPM uncatalogued (1: 148.0 mm. THA- 
LASSA Sta. X020. 19°08'N. 16°48'W. 280-468 m); MCZ 48849 
(1: 76.9 mm. LA RAFALE Cr. I. Sta. 27. 11°25'N. 17°21'W, 
200 m); MF 3694 (1: 117.6 mm. from stomach of Polymixia 
nohilis): MF 7231 (1: no SL. from stomach of Aphanopus carbo); 
MF 8213 (1: no SL. from stomach of P. nohilis): MF 12518 (I: 
124.5 mm. from stomach of P. nohilis); MF 14695 (1: 131.3 mm, 
from stomach of A. carho): MF 15650 (1: no SL, from stomach 
of A. carho); MF 16067 (2: 120-127 mm. from stomach of A. 
carho); MF 17088 (1: 125.5 mm. from stomach of A. carho): 
MF 18204 (1: 97.0 mm. from stomach of A. carho); MF 20599 
(1: no SL. from stomach of A. carho); MSNG 38155 (I: 89 mm. 
Messina. Sicily); MZF 3089 (1: 117 mm. holotype of Pomaio- 
michthvs consianciae, Straits of Messina); USNM 207723 (2: 
98.6 100.9 mm, LA RAFALE Cr. I. Sta. 27, ir25'N, 17°2I'W, 
200 m); USNM 207724 (1: 70.9 mm, LA RAFALE Cr. I, Sta. 27. 
11°25'N, 17°2I'W, 200 m); ZMB 23101 (1: 148.9 mm, MORS 
Sta. 136 72, I8°36'S, 1 L'29'E, 275 m); ZMB 23102 ( 1: 116.5 mm, 
MORS Sta. 175 72, 18^05'S, 1 r-^27'E. 360 m). 

Diagnosis 

In the past, E. consianciae has been mistaken for E. iclescopus 
(Risso) and E. pandionis (Goode and Bean). The distribution 



1977 E. CONSTANCIAE DESCRIPTION t> 

of E. constanciae also makes confusion likely with E. denticulatus 
Dieuzeide. E. constanciae may be distinguished from the above 
species by the presence of VII-I, 9 dorsal fin elements, a pungent 
opercular spine. lingual teeth and six to eight pyloric caeca. 

E. constanciae most closely resembles E. pectinifer Mayer, a 
species found in the Caribbean Sea. the Gulf of Mexico and the 
western Pacific Ocean. The former is distinguished by the pres- 
ence of numerous awl-like fill rakers (29-36) and elongate D:I 
and All fin spines (12.7 16.5 per cent and 13.5-16.8 percent SL, 
respectively). 

Description 

See Tables 1 and 2 for morphometric and meristic data. 

Body elongate, fusiform in larger individuals; body depth 
21.1-30.0 per cent SL. Head moderate, length 32.2-38.9 per cent 
SL; height 16.2-18.7 per cent SL. Snout tapering to a rounded 
point. 7.0 9.1 per cent SL. Lower jaw 13.9-17.6 per cent SL; 
equal to or projecting slightly beyond upper jaw; bearing two 
nubs of variable prominence on anterior surface of mandible 
near symphysis. Maxilla reaching 13-12 length of eye; ventro- 
lateral surface of maxillary head bearing sharp, posteriorly re- 



Table I. Epi^onus constanciae regression parameters, u = functional (GM) 
regression coefficient; \- Y intercept of GM regression line. Y = uX + v; 
b = predictive regression coefficient ±95 per cent confidence interval; 
a = Y intercept of predictive regression line, Y = bX + a; n = number 
of specimens. 



HL 


0.391 


4.033 


0.348 ±0,026 


Head height 


0.191 


-1.810 


O.I89±0.01i 


Body depth 


0.332 


8.043 


0.324 ±0.027 


Eye diameter 


0.168 


0.976 


0.1 64 ±0.0 15 


Snout length 


0.078 


0.333 


0.074 ±0.010 


Maxillary length 


0.I6I 


-0.384 


O.I57±O.OI3 


Lower jaw length 


0.174 


1.280 


0.167±0.017 


Interorbital width 


0.090 


0.910 


0.087 ± 0.008 


Caudal peduncle length 


0.269 


0.333 


0.264 ±0.018 


Caudal peduncle depth 


0.145 


3.418 


0.1 39 ±0.0 15 


D:I 


0.156 


1.219 


150 ±0.0 15 


All 


0.156 


0.547 


0.149 ±0.019 


P:I 


0.145 


0.298 


0.1 42 ±0.009 



3.240 


33 


1.566 


30 


7.062 


33 


0..367 


32 


0.922 


32 


0.076 


34 


0.428 


36 


1.300 


36 


0.242 


35 


2.673 


34 


0.546 


31 


0.282 


27 


0.233 


35 



6 HRi \ loR \ No. 443 

Tabic 2. /:'/>/!,'("»/> iiiii^uinciae meristic data. \ = mean; SI) = standard de\ia- 
tmn: n - number o\ specimens. 





;::^ 


Ranae 


SD 


n 


Sectoral fin rays 


17.65 


16 


19 


0.62 


40 


Gill rakers 












IcUal 


.^2.43 


29 


36 


1.75 


40 


I'pper arch 


9. IS 


S 


10 


64 


39 


1 ower arch 


22..^! 


20 


25 


l.-^O 


39 


Lateral line scales 


47,74 


46 


50 


0,9.^ 


34 


P\k)ric caeca 


7.00 


6 


X 


0.64 


35 



cur\ed process. ma\ be obscured b\' o\erl\ing skin. Eye longer 
than high, reaching upper profile ot head. 4 1. 1 49.1 per cent HL. 
Interorbital width K.7 10.9 per cent SL. Caudal peduncle mod- 
erate, length 24.3 28.8 per cent SL; depth 9.7-13.8 per cent SL. 

Dentition subject to ontogenetic and geographic variation 
(see Mayer. 1974: 185 and below). Teeth small, conical: present 
on premaxillae. mandibles and vomer; palatines edentulous or 
dcntigerous; tongue dentigerous. bearing triangular patch of 
glossohsal teeth along midline and broad. closeK -spaced patches 
of teeth parallel to margins; endo- and ectopter\goids edentulous. 

Opercle dominated b\ pungent. bon\ spine; surmounted by 
one or more poorly ossified spinelets. Preopercle well-ossified; 
angle narrowly produced; ornamentation subject to geographic 
variation (see below). Interopercle and subopercle weakly ser- 
rated or unserratcd. 

Gill rakers awl-like. 29 36 on first gill arch. 8-10 on upper 
portion. 20 25 on lower portion (see Table 2). 

First dorsal fm VTI (38); second dorsal fin 1. S (2). 1. 9 (36). 
1. 10 ( I ). Anal fin I. 8 ( I). 11. 9 (36); \entral fms 1. 5 (40). soft rays 
extending to le\el of anus. F'ectoral tin 16 19 (see lable 2). 
D I \ariable. 1.7 3.2 per cent SL; DT. .ML and PI long and 
pungent: 12.7 16.5 per cent. 13.5 16.8 per cent and 13.0 15.5 
per cent SL. respecti\el\ . 

Prescr\ed specimens \ell(n\-bro\\ n; mouth light; branchial 
and peiitoneal membranes black: dorsal tin membranes dark. 
\'iseer;i olten sheathetl b\ heaw. \ellow-brown fat deposits. 

Deseription based on tort\ specimens 70.9 158.9 mm SL. 



1977 e. constanciae dfscrif^non 7 

Notes on the Hoeot^ pe of E. constanciae 

As mentioned earlier. Giglioli's description of Pomaromichthvs 
constanciae is incomplete and includes a number of inaccuracies. 
Tortonese and Queirolo (1970) figured and redescribed the holo- 
t\pe but did not full\' characterize the fish. One aspect of the 
present investigation involved reexamining Giglioli's type speci- 
men, now in rather poor condition. Measurements and counts 
obtained from the holotype appear in the Appendix. Qualitative 
features not mentioned in previous descriptions include: 

1. Lower jaw slightly prominent with two nubs at symphysis. 

2. Small teeth on premaxillae. mandibles, vomer, palatines 
and tongue; lingual dentition composed of a central tooth 
patch and a series of elongate tooth patches parallel to the 
borders of the tongue. 

3. Opercular spine well-developed; two poorly developed spine- 
lets above, but well-separated from the spine. 

4. Dil and AI spines short; D:I, All and P:I spines long. 
Longest ventral fin rays reach anus. 

If previous E. irewavasae descriptions (e.g.. Mayer, 1974; Poll. 
1954) are compared with P. constanciae data presented in the 
Appendix and abo\e, it becomes apparent that the two forms 
are synonymous. Giglioli's holotype bears all characters diag- 
nostic of E. trewavasae including VII-L 9 dorsal fin elements, 
long fin spines, 18 pectoral rays, numerous awl-like gill rakers, 
a pungent, bony opercular spine, mandibular chin nubs and 
lingual teeth. The species is correctly called E. constanciae be- 
cause Pomatomichthvs is a junior synonym of Epigonns. and 
constanciae has precedence over trewavasae. 

The fish was named after Giglioli's wife, Costanza. 

DISTRIBUTION 

E. constanciae is found from the coast of equatorial west 
Arfica to the Straits of Messina. The species has also been taken 
off Madeira and probabl\' occurs off the Canaries and Azores. 
E. constanciae appears most abundant on upper portions of the 
continental slope between 200 and 400 m. 



8 BREVIORA No. 443 

GEOGRAPHIC VARIATION 

Mayer (1974: 185-186) briefly commented on geographic vari- 
ation in Epigonus trewavasae and suggested that Mediterranean 
and African populations may represent subspecies. At the time, 
sufficient material was not available to draw firm conclusions 
about the validity of infraspecific categories. Nevertheless, 
variations were noted in qualitative characters such as the promi- 
nence of chin nubs, the development of vomerine and palatine 
teeth and the occurrence of preopercular armor. 

The consolidation of Pomatomichthys constanciae and Epi- 
gonus trewavasae brings the question of geographic variation 
into sharper focus, because holotypes of the nominal species 
were taken from opposite ends of the E. constanciae range. An 
analysis of geographic variation in E. constanciae was accom- 
plished by dividing the specimens examined for this study into 
three geographic groups: a Mediterranean series, a Madeiran 
series and an African series. The African series was the largest 
group and included fishes taken along the west African coast 
from 18°S to 21° N. The lumping of west-central African and 
northwest African populations follows Maurin's conclusion (1968: 
78) that the Cape Blanc-Arguin Bank region represents a major 
northern faunal limit for demersal forms with tropical affinities. 

Table 3. Comparisons of meristic data from African, Mediterranean and 
Madeiran populations of Epij^onus constanciae. Afr. = African popu- 
lation: Med. = Mediterranean population: Mad. = Madeiran popula- 
tion. See text for explanation. 

COEFFICIENTS OF DIFFERENCE 

Afr. vs. Med. Afr. vs. Mad. Med. vs. Mad. 

Pectoral fin rays 0.33 0.48 0.20 

Gill rakers 

Total 0.63 1.24 0.26 

Upper arch 0.28 0.59 0.14 

Lower arch 0.57 1.27 0.39 

Lateral line scales 0.50 0.61 0.16 

Pvioric caeca 0.12 0.38 0.09 



1977 e. constanciae description 9 

Meristic Characters 

Meristic data comparisons were accomplished by calculating 
Coefficients of Difference (C.D.'s) (Mayr. 1969: 189-193) for the 
three geographic groups. The index indicates the degree of non- 
overlap between two populations for a given character. Popula- 
tions exhibiting a non-overlap of 90 per cent or more (C.D.^ 1 .28) 
are considered distinct at the subspecies level. Table 3 presents 
data from the current study. .As indicated, none of the differences 
observed are significant at the subspecies level. 

MORPHOMETRIC CHARACTERS 

Analogous examinations of mensural data were undertaken 
using regression techniques. Individual values obtained from 
Mediterranean and Madeiran specimens were compared to re- 
gression data from African populations by means of Student's 
t-tests. Results of the tests are summarized below. 

Mediterranean and African populations are very similar and 
do not appear distinct at the subspecies level. Only one Mediter- 
ranean data point fell outside the 99 per cent confidence limits 
of the African group and most data points fell within the 95 per 
cent or 98 per cent limits. 

The Madeiran population is more distinct. Madeiran E. con- 
stanciae appear characterized by shallower bodies, shorter heads 
and shorter jaws than African forms. However, part of the di- 
vergence may be an artifact of the Madeiran specimens' poor 
condition. All were obtained from the stomachs of large, pred- 
atory fishes and many were considerably damaged. Additional 
data are required to assess the taxonomic significance of differences 
exhibited by the Madeiran specimens. 

Qualitative Characters 

Geographic variability reported by Mayer (1974) for qualita- 
tive characters was confirmed by the present study. Character 
states such as prominent chin nubs, weakly armored preopercular 
margins, edentulous palatines and vomerine tooth patches lack- 
ing posterior median extensions are prevalent, but not uniform 
among fishes from equatorial west Africa. In contrast, poorly 



10 F^R[ \ lOR \ No. 443 

dc\ duped chin nubs, spintnis picopcrcular margins, dentigerous 
palatines ami \iinicrmc tcnnli patclics witti posterior median 
extensions are eommon among Mediterranean specimens. The 
trequenex ot "northern" and "southern" character states \aries 
with latitude. Matieiran and Mauritanian specimens tend to be 
intermediate, with the former more closeK resembling Mediter- 
ranean tisli and the latter more similar to specimens from equa- 
torial Africa. Because characters exhibit continuous variation 
along the range o\ the species, and because populations at spe- 
cies' distribution limits exhibit a mixture of character states and 
are not umformh' "northern" or "scvathern". qualitative characters 
do not support the division of /:'. conMoiuiae into formal sub- 
species. 

CONCLUSIONS 

The picture that emerges for Epi^onus constanciae is one of a 
polvniorphic species whose range extends from the eastern equa- 
torial Atlantic to the western Mediterranean. This pattern has 
been reported for other organisms bv manv authors (e.g.. Briggs, 
1974: Maurin. I96S: Tortonese. 1964. I960: Ekman. 195.^). The 
prevalence o{ polymorphism among Atlantic and Mediterranean 
populations has been discussed by Tortonese (1964: 103-104: 1959 
387). Although genetic differences between populations cannot 
be ruled out. Tortonese suggests that Atlantic-Mediterranean phe- 
notvpic dissimilarities mav be induced bv regional differences in 
temperature regime and food availabilitv. 

It has been suggested elsewhere (Maver. 1972) that E. consian- 
ciac may be among the least vagile species ot the genus Hpiiionus. 
Adults appear stronglv associated with the substrate. Neither 
pelagic larvae nor pelagic juveniles are known tor this species, 
although such torms have been discovered tor several congeners. 
The smallest known specimen o{ I'., consiam uw is a 29.8 mm SL 
juvenile (IRSN 16548) taken bv a bottom trawl off Sardinia. 
1 he latter data suggest that /:'. c nnsia/hic/c mav have either a 
short-lived pelagic phase or mav lack a pelagic phase entirelv. 

I he hvpothesis ot limited vagilitv tits the pattern of phenotvpic 
variabilitv observed iov /:'. i oiisunii inc. Individual populations 
adapt to local conditions and evolve characteristic traits. Breeding 
mav beas>umed to occur primarilv between members ot the same or 



1977 E. CONSTANCIAF DESCRIf'TION 11 

neighboring populations, thus preser\ing local phenotvpes. Rela- 
ti\eiy isolated populations, such as that from Madeira, are more 
unique in appearance but still conform to phenotypic patterns 
resulting from latitudinal gradients in environmental or genetic 
factors. 

ACKNOWLEDGEMENTS 

This work was initiated while the senior author was an Asso- 
ciate in IchthyoIog\' at the Museum of Comparative Zoology and 
was completed during his tenure at the Department of Marine 
Science. University of South Florida. We are indebted to these 
institutions as well as to the Museo Civico di Storia Naturale for 
logistical support and encouragement. The authors also wish 
to thank the following institutions and individuals for providing 
study materials: X. Missionne, Institut Roval des Sciences Natu- 
relles de Belgique; G. Krefft. Institut fiir Seefischerei: C. Maurin 
and J. C. Quero. Institut des Peches Maritimes; K. F. Liem and 
R. Schoknecht. Museum of Comparative Zoology; G. E. Maul. 
Museo Municipal do Funchal: B. Lanza and M. Bucciarelli. Flor- 
ence Zoological Museum: E. A. Lachner. Smithsonian Institu- 
tion: and C. Karrer, Zoologisches Museum. Berlin. Special thanks 
go to J. C. Briggs. R. C. Baird. G. Krefft and G. Smith for criti- 
callv reviewing the manuscript. 



BIBLIOGRAPHY 

Bri(,(,s J. C. 1974. Marine Zoogeography. New York: McGraw-Hill Book 

Co. 475 pp. 
CwMHKi. A. 1956. Su rari stadi lar\ali e gio\anili di PonuUonuchihvs con- 
Mane uic Ciigl. c relaii\a difteren/ia/ione da stadi giovanili della specie affine 
Pcnuuonnis !c/e\(i>/vuni Risso. Boll. Pesca Piscicult. Idrobiol.. (n.s.) //(I): 

1:1 i:". 

Ek\i\\ S. 1953. Zoogeographv of the Sea. London: Sidgwick and Jackson. 

I td. 4P pp. 
Gi( 1 1 F. H. ISNO. Flenco dei Manimiteri. degli Iccelli e dei Rettili ittiofagi 

.Appartenenti alia Fauna Italica e Catalogo degli Antihi e dei F\-sci Italiani. 

Firen/c: Stanipcria Keale. S5 pp. 
Gcxini (1, B.. \\i) \ IF Bi w 1S9^. (Oceanic lchth\ology, Washington. 

n.C.: Smithsonian Institution. 55.^ pp. 
Hi nn^ G 1 vM. K 1 1 \(,l FR, 19Sn. Fishes oI the C.reat 1 akes Region, 

Ann Arbor: i ni\. \1ich. Press. 213 pp. 



12 BRr\ lORA No. 443 

.loKDW 1), S. \\i) H W . Fmkmwn IS96. The lishcs ot North and Middle 
America. Bull. l.S. Nat. \1us.. 41 \): 1 1240. 

\M) H. W. Ci \Kk \^M). Check List ot the Fishes of North 

and Middle America. .Appendix .\. Report to the United States Commis- 
sioner of Fisheries for I92S. Washington. DC: U.S. Covernment Printing 
Office. 670 pp. 

M.Ai RiN. C. 196S. Ecologie ichthsologiques des fonds Chalutables atlantiques 
(de la baie ibero-marocaine a la Mauritanie) et de la Mediterranee occidentale. 
Theses presentees a la Faciilte des Sciences de l'Uni\ersite de Nancy. No. 
d'Ordre: A. O. 2 1S2. 14,"^ pp. 

M'\>FR. G. F. 1972. Systematics. functional anatomy, and ecology of the 
cardinalfish genus Epii(onus ( Apogonidae). Ph.L'i. Thesis. Har\ard Uni- 
versity. 190 pp. 

1974. \ revision of the cardinalfish genus Epii^ofius ( Perciformes. 

Apogonidae). with descriptions of two new species. Bull. Mus. Comp. Zool.. 
1460Y 147-203. 

1975. Results of the research cruises of FRV "Walther Herwig" to 



South America. XXXI.X. The epigonine fauna of the South Atlantic, with 

a key to the genera and a redescription of Roscnhlaiiia rchuMa Mead and 

De Falla. Arch. Fi.sch. Wiss.. 26{\): 13 28. 
MwR. E. 1969 Principles of Systematic Zoology. New York: McGraw-Hill. 

Inc. 42S pp. 
Pom M. 1954. Poissons. IV. Teleosteens acanthopterygiens (premiere partie). 

Res. Sci. E.xped. Oceanogr. Beige Eaux Cot. Afr. Atl. Sud. ■^t3A): 1-390. 
Ru Ki R \V. E. 1973. Linear regressions in fishcr\ research. J. Fish. Res. Bd. 

Canada. Jd: 409 434. 
Sciii 1 I/. I.. P. 1940. Two new genera and three new species of cheilodipterid 

fishes, with notes on other genera of the famiU. F'roc. U.S. Nat. Mus.. 

SS{}0X5). 403 423. 
ToRiDMsi E. 1959. General remarks on the Mediterranean deep-sea fishes. 

In: International Oceanographic Contercnce. 1959. Preprints. M. Sears (ed.). 

Washington. D.C.: Amer. Assoc. ,Ad\. Sci. 386 387. 
1964. Ihe main biogeographical features and problems of the Med- 
iterranean livh fauna. Copcia. l'^64{\): 98 107. 
. 1473. .Apogonidae. In: Check-list of the Fishes of the Northeastern 

Atlantic and of the Mediterranean. Vol I. ,1, C. Hureaii and Ih. Monod 

(eds,). Pans: UNESCO 365 367. 
WD L. C. Qi fiRoio. 19'70. Contributo alio studio dell' ittiofauna del 

Mar 1 liiure orientale. Ann. Mus, St. Nat. Genova. 7,V; 21 46, 



1977 



E. CONSTANCIAH DESC RIPTION 



13 



APPENDIX 

Meristic and morphometric data obtained from the holot\pe ot Epiiianus 
insicmcicif. MZF 3089. Measurements are in millmieters. 



Meristic data 
Dorsal fm 
Anal fin 
Pectoral fin 



\ II I. 9 
II. 9 

18 



Pehic fin 
Lateral line 
Gill rakers 



1.5 

46 
30 



Morphometric data 

SL ' 115 

HL 37 

Head height 19.5 

Body depth 30 

Eye 17 

Snout 8 



Interorbital width 


12.5 


Maxilla 


16 


Caudal peduncle length 


29 


Caudal peduncle depth 


13 


D:I 


16 


All 


17 


P2I 


15 



B R E V I ■■0m" A 

MAR 18 1985 

Museium of ConipaFa^il^^QZoology 



us ISSN 0006-9698 ^^"Ty 



Cambrihge. Mass. 27 May 1977 Number 444 

STATIONS OF THE THAYER EXPEDITION TO BRAZIL 

1865 - 1866 

Myvanwy M. Dick' 

Abstract. In April of 1865, a group of scientists from the Museum of Compara- 
tive Zoology led by Louis Agassiz embarked for Brazil. The purpose of this expedi- 
tion was to study, in situ, the fauna and flora of that country. Members of the ex- 
pedition, divided into three groups, explored the major part of Brazil from the Rio 
Tocantins eastward to the coast. Although the major part of the work was done on 
the distribution and relationships of the fresh water fishes (Agassiz's prime interest) 
many superb collections were made of plants, invertebrates and vertebrates. Thor- 
ough geological surveys provided much information on the structure of Brazil. This 
paper traces and maps the numerous stations where specimens were collected. Based 
on modem and nineteenth century maps, and on notes and papers recording the 
routes followed and collections made by the expedition, it represents a compilation 
of all this information into a single, indexed directory. This directory should facili- 
tate the future labors of individuals working with the Thayer Expedition material. 

INTRODUCTION 

Louis Agassiz had for many years wanted to observe in situ the 
fauna and flora of Brazil, a study that he knew would be substan- 
tially encouraged by the Brazilian government. The opportunity 
to do so. however, had never presented itself. This cherished dream 
became a reality in 1865 through the understanding and generosity 
of Nathaniel Thayer, a Boston businessman keenly interested in 
science. Thayer assumed the expenses of Professor and Mrs. Agas- 
siz and six assistants for an expedition to Brazil lasting several 
months. 

The scientific staff from the Museum of Comparative Zoology 



'Museum of Comparative Zoology 



BREVIORA No. 444 



included John G. Anthony, Assistant Curator of Molluscs (who 
was with Agassiz in Rio de Janeiro and made extensive collections 
in this area before illness forced his return to Cambridge); Joel 
Asaph Allen, Assistant Curator of Birds (later the noted Curator 
of Birds at the American Museum of Natural History); Frederick 
C. Hartt and Orestes St. John, geologists trained by Agassiz; George 
Sceva, Preparator and collector primarily of fossil material; and 
James Burkhardt, an artist who had worked for many years with 
Aggassiz. In spite of poor health, Burkhardt was an indefatigable 
worker and made, according to Agassiz, over eight hundred illus- 
trations during the expedition. Most of these unfortunately have 
disappeared, but the remaining examples, which are in the Museum 
of Comparative Zoology, indicate the high quality of his draft- 
manship. 

Agassiz also had extremely able and enthusiastic young volun- 
teers in his party: Edward Copeland, who accompanied Hartt 
from Rio de Janeiro to Bahia; Newton Dexter; Walter Hunnewell; 
William James, the future great professor of philosophy at Harvard 
University; Stephen Thayer, the son of Nathaniel Thayer; and 
Thomas Ward. 

On 2 April 1865 the group left Boston aboard the S.S. Colorado, 
guests of the Pacific Mail Steamship Company. While at sea, 
Agassiz gave a series of lectures giving background for the work 
planned for the months ahead. On 23 April 1865 the S.S. Colorado 
docked in Rio de Janeiro. Agassiz had arrived at the threshold of 
his longed-for goal. 

The area covered by the members of the expedition was extensive. 
They divided into three groups and explored the major part of Brazil 
from the Rio Tocantins eastward to the coast; they also explored 
some of its tributaries, to the borders of Colombia and Peru. Al- 
though the major part of the work was done on the distribution 
and relationships of the fresh water fishes ( Agassiz's prime interest), 
many superb collections were made of plants, invertebrates and 
vertebrates. Thorough geological surveys provided much infor- 
mation on the structure of Brazil. 

The numerous stations where specimens were collected have 
been difficult to locate because several were small villages or clusters 
of dwellings on the edge of a river or lake. Many no longer exist 
and some have names other than those used in 1865. Rivers and 



1977 THAYER EXPEDITION 



lakes that were recognized a hundred years ago are frequently 
overlooked in recent atlases. Researchers involved with the Thayer 
Expedition have devoted many hours to tracing these stations and 
collections. Using both modern and nineteenth century maps, it 
has been possible to locate them fairly accurately. Information 
concerning the routes followed and the collections made by mem- 
bers and associates of the expedition was obtained from publica- 
tions listed in the bibliography. The locality names used in the text 
and on the maps are almost always those employed during the 
period of the Thayer Expedition. Subsequent changes in locaHty 
names or spelling are italicized in the index. Having this informa- 
tion incorporated in a single directory should facilitate the future 
labors of individuals working with the Thayer Expedition material. 
Much work needs to be done with the material collected by 
Agassiz and his corps, especially with the collections of fishes. 
Many other groups, both plant and animal, also need comprehen- 
sive study and evaluation to complete the survey of Brazil envis- 
aged by Louis Agassiz so long ago. 

MEMBERS OF THE THAYER EXPEDITION 

1865 — 1866 

Museum Staff: 

Louis Agassiz 
Elizabeth Gary Agassiz 

Joel A. Allen Ornithologist 

John G. Anthony Malacologist 

James Burkhardt Artist 

Frederick G. Hartt Geologist 

George Sceva Preparator 

Orestes St. John Geologist 

Volunteers: 
Edward Copeland 
Newton Dexter 
Waher Hunnewell 
William James 
Stephen Van R. Thayer 
Thomas Ward 



BREVIORA 



No. 444 



Members added in Brazil 
Major M. Coutinho 

Monsieur D. Bourget 

Mr. Talisman 

Collections also made by: 

Captain Anacleto 

Senhor Augustinho 

Senhor Barrosa 

Colonel Bentos 

Senhor S. E. Pimento Bueno 

Dr. Pacheco de Silva 
Major Estolano 
Senhor Felice 

Senhor JoaoBaptistadaFonseca 

Senhor Glaziou 

Senhor Honorio 

Dr. Justa 

Mr. Kaulfuss 

Senhor Antonio de Lacerda 

Senhor Mariano P. F. Lage 

Senhor Cicero de Lima 
Dr. Couto de Magalhaes 

Dr. Malcher 

Dr. Mendes 

Senhor Penna 

Senhor Joachim Rodriguez 

Senhor Sepeda 

Mr. Charles Taylor 

Father Torquata 
Senhor Vinhas 



Brazilian Government Corps of 

Engineers 
French naturalist residing in Rio 

de Janeiro 
Officer, Amazonian Steamship 

Company 

Fishes, Amazon River 
Fishes, Amazon River 
Miscellaneous, Amazon River 
Fishes, Rio Trombetas 
Forest and river fishes, Amazon 

River 
Fishes, Rio de Janeiro 
Fishes, Amazon River 
Geological observations, miscel- 
laneous, Ceara 
Fishes, mammals, environs of Rio 

de Janeiro 
Plants, palms. Organ Mountains 
Miscellaneous, Amazon River 
Fishes, Rio Parahyba do Norte 
Andean fossils 
Miscellaneous, Bahia 
Plants, animals, fishes, Rio Novo, 

basin Rio Parahyba 
Fishes, insects, interior Brazil 
Miscellaneous, at his direction, 

upper Amazon River 
Birds, Amazon River 
Miscellaneous, Ceara 
Fishes, Amazon River 
Fishes, Santarem 
Miscellaneous. Amazon River 
Drawings and collections of fishes 

and insects. Posse 
Fishes, Amazon River 
Fishes, Rio Xingu 



1977 THAYER EXPEDITION 5 

The Emperor, Dom Pedro II, had extensive collections of fishes 
made for Professor Agassiz from several rivers in southern Brazil. 

Thomas G. Cary, Mrs. Agassiz's brother, though not connected 
with the expedition, contributed material from Montevideo, Buenos 
Aires and other localities. 

ACKNOWLEDGMENTS 

The suggestions made by Dr. Ernst Mayr and Dr. Raymond A. 
Paynter, Jr., Museum of Comparative Zoology, concerning the 
presentation ef material have been greatly appreciated. The gen- 
erous assistance of Dr. P. E. Vanzolini, Museu de Zoologia da 
Universidade de Sao Paulo, has been invaluable and I am indeed 
grateful. Dr. Naercio Menezes, also of the Museu de Zoologia, 
kindly located six stations that had completely eluded me. To 
Dr. Bruce B. Collette, United States National Museum, I am in- 
debted for his critical reading of the manuscript. These kind people 
all have my sincere thanks. 

THE THAYER EXPEDITION 

Louis Agassiz established a laboratory in Rio de Janeiro which 
the members of the expedition used as a base for their work during 
the weeks prior to their departure for other areas of Brazil. Nu- 
merous brief trips were made at this time, including one over the 
route of Dom Pedro II Railroad, which Agassiz believed would 
be of great future value for the transportation of large geological 
and zoological specimens. Zoologists and geologists thoroughly 
investigated the railroad from its origin in Rio de Janeiro to its 
terminus in Parahyba, a distance of one hundred miles. On 27 
April Professor and Mrs. Agassiz enjoyed their first journey on 
the railroad as guests of Major Ellison, chief engineer. Two speci- 
mens from the fresh waters of Brazil were collected at this time, 
inaugurating what was to be an outstanding assemblage of Bra- 
zilian fishes. 

An excursion of several days duration was made by the Agassizes 
in May, 1865; it was the first extending any distance from Rio de 
Janeiro. They went by ferry to Maua then by railroad and post 
coach to Petropolis. From Petropolis they continued along the 
valley of the Rio Piabanha to Posse, crossed the Rio Parahyba at 



6 BRI-VIORA No. 444 

Entre Rios, soon reached the Rio Parahybuna. then went on to Juiz 
de Fora. This trip provided an excellent opportunity to study the 
plant life and geological formations and to collect numerous speci- 
mens of fishes and other vertebrates. 

Examples of the peculiarities of Brazilian geological formations 
were observed in Tijuca, about eight miles from Rio de Janeiro, 
26-27 May. On 21 June the Agassizes again went to Juiz de Fora, 
arrived there 22 June, then continued thirty miles farther where 
they were guests of Senhor Lage at his fazenda. Senhor Lage ac- 
companied the Agassizes to the Serra da Babylonia. On 27 June 
they returned to Rio de Janeiro. 

In June the party divided into units to facilitate the exploration 
of Brazil. St. John, Allen, Ward and Sceva left for the interior 
the first week of June and Hartt and Copeland left to explore the 
coast from the Rio Parahyba to Bahia the following wee. On 
25 July, the Agassizes, Major Coutinho, Burkhardt, Monsieur 
Bourget (a French naturalist residing in Rio de Janeiro engaged 
by Agassiz as a collector and preparator), Hunnewell and James 
left Rio de Janeiro on the "Cruzeiro do Sul" for Bahia, where 
they arrived 28 July. In Bahia they were joined by Dexter and 
Thayer, who had preceded them. Antonio de Lacerda, their host, 
had greatly assisted Dexter and Thayer in their collecting. The 
next collecting station was Maceio on 30 July, then Pernambuco 
31 July and on 2 August the Parahyba do Norte as far as the town 
of Parahyba. They arrived at Ceara 5 August, the guests of Dr. 
Mendes, and were at Maranhao 6 August. They arrived at Para 
10 August, where Mr. Pimenta Bueno arranged living and working 
quarters from 10 August to 19 August, and the use of a steamer, 
the "Icambiaba", for a month between Para and Manaos. In Para, 
Mr. Talisman, a young Brazilian officer of the Amazonian Steam- 
ship Company, joined the group. Breves, 20 August, was the first 
station in Amazonas. Leaving Aturia, they passed into the Rio 
Tajapuru and stopped at the village of Tajapuru. 

On 2 1 August, after two days spent skirting the island of Marajo, 
they left the Para River and turned into the mainstream of the 
Amazon. They went to Gurupa, 22 August, where a collection of 
forest fishes was made, and on to Porto de Moz on the Rio Xingu, 

23 August. A brief non-collecting halt was made at Prainha on 

24 August, and they were at Monte Alegre. at the mouth of the Rio 
Gurupatuba on 25 August. 



1977 THAYER EXPEDiriON 7 

From Santarem, at the mouth of the Rio Tapajoz, Dexter. James 
and TaHsman left. 26 August, to explore the Rio Tapajoz. Bour- 
get and Hunnewell remained in Santarem. Bourget to make col- 
lections, Hunnewell to repair his photographic equipment. Agas- 
siz, Burkhardt, Thayer and Coutinho continued on to Obydos. 
Before leaving the Rio Tapajoz to join the Amazon, the "Icambiaba" 
detoured through a narrow channel. Igarape-Assu. A brief stop 
was made at Obydos, 26 August. On 27 August they landed at Villa 
Bella, at the mouth of the Rio Tupinambaranas. By canoe, the 
group left Villa Bella to explore Lago Jose Assu until 30 August, 
proceeding from there to a branch of the Rio Ramos, which con- 
nects the Amazon to the Rio Madeira. They returned to Villa Bella 
on 2 September. 

A rest period was taken from 4 to 1 1 September at Manaos. on 
the Rio Negro. There they were joined by Dexter, James and Talis- 
man. A brief halt was made at Barreira das Cudajas on 12 Septem- 
ber and on 13 September they made another short halt at Coarl. 
Teffe stands beside a small lake formed by the Rio Teffe before it 
joins the Amazon. This site was visited on 14 September, then 
Fonte Boa, 15 September and Sao Paulo de Oliven^a, 17 Septem- 
ber. Here James and Talisman left the group to continue from 
there to the Rios I^a and Hyutahy by canoe. The farthest point 
reached by Agassiz was Tabatinga, 19 September. Bourget re- 
mained in Tabatinga for a month, collecting there and in the sur- 
rounding country. Agassiz and the rest of the party returned to 
Teffe, remaining there until 22 October. Agassiz and Coutinho 
were at a station on a branch of the Rio Solimoens from 4 October 
to 6 October. James and Talisman rejoined them at Teffe, 16 Oc- 
tober and Bourget boarded the "Icambiaba" 22 October. 

While in Teffe, Dexter made an extensive collection of birds 
and, with the assistance of Hunnewell, Thayer and local sportsmen, 
added to the reptile and animal specimens. Agassiz also purchased 
a large collection of insects at this station. 

Manaos was reached on 23 October. While at this station, Agas- 
siz, Coutinho, Burkhardt, Dexter and James, preceded by their 
host, Senhor Honorio, had a two day, 27 October to 29 October, 
collecting excursion to Lago Hyanuary on the western side of the 
Rio Negro. Talisman and Dexter explored the Branco and Negro 
rivers for six weeks, returning to Manaos 10 December. Thayer 
and Bourget were at Lago Cudajas for ten days, 27 November to 



8 BREVIORA No. 444 

6 December (Thayer also collected at Lago Alexo), and James 
went to Manacapuru for ten days. 

The Agassizes, Coutinho and Burkhardt left Manaos 10 De- 
cember to spen ten days at Mauhes. They proceeded down the 
Rio Ramos to the Rio Mauhes, which runs almost parallel with 
the Amazon, to the Rio Madeira (which joins the Amazon oppo- 
site Serpa). The land encircled by these rivers is the island of Tu- 
pinambaranas. Mucaja-Tuba and another small settlement on 
the Rio Mauhes were visited 13 15 December, and on 21 Decem- 
ber the group returned to Manaos. 

On 26 December the party left Manaos and went up the Rio 
Negro as far as Pedreira on the "Ibicuhy," arriving there 28 De- 
cember. A collection of palms had been made at Taua Peassu and 
they were picked up on the return to Manaos 31 December. 

The stay at Manaos ended about 12 January 1866 and the party 
returned to Villa Bella 16 January. Collections were made 18 Janu- 
ary at Lago Maximo, a short walk overland from Villa Bella. A 
narrow outlet connects this lake to the Rio Ramos. On 21 January 
the party was again at Obydos, and 22 January at Santarem, which 
is situated on a point of land separating the Rio Tapajos and the 
Amazon. They left Santarem 24 January for Monte Alegre, where 
they stayed until 29 January. Agassiz, Coutinho and a few friends 
went to the Serra d'Erere, northwest of Monte Alegre, to see the 
geological formations. Coutinho went by horseback and Agassiz 
by canoe, along the Rio Gurupatuba, up a narrow stream known 
as the Rio Erere to a point on a line with the Serra and from there 
proceeded on foot. 

Senhor Vinhas, a resident of Porto de Moz, had a collection of 
fishes from the Rio Xingu ready for Agassiz upon his arrival 29 
January, so the group quickly continued on to Gurupa, arriving 

30 January. They reached Tajapuru 31 January and spent two 
days there. 

From 4-27 February they were at Para. On 28 February they 
left on the steamer "Tabatinga" to explore the great island of Marajo. 
They visited Soures, then, 29 February to 4 March, Vigia, Baia do 
Sul and the small island of Tatuatuba. They returned to Para 5 
March. 

On 26 March the party left Para for Ceara where they arrived 

31 March. A stop was made en route at Maranhao so that Agassiz 
and Coutinho might examine the coastal geology with more care 



1977 THAYER EXPEDITION 9 

than on their previous visit. An excursion to Pacatuba and the 
Serra da Aratanha, 6-12 April, included the Agassizes. Coutinho 
and Senhor Pompeo, Government Engineer of the Province. Col- 
lections were made in the vicinity of Ceara by various people in- 
cluding Senhor Felice and Senhor Cicero de Lima. They left Ceara 
16 April and continued, with a brief stop at Pernambuco, to Rio 
de Janeiro, where they arrived 25 April. 

During the time in Rio de Janeiro, 25 April to 2 July, collections 
were packed and shipped to Cambridge. A few short trips were 
made, to Petropolis and again along the Dom Pedro 11 Railroad. 
The last excursion made by Agassiz was 9-12 June, to the Organ 
Mountains, accompanied by Mrs. Agassiz, Mr. Glaziou and Dr. 
Nageli. The group went by boat to Piedade, then walked from 
there to Theresopolis, collecting along the way. 

On 2 July Burkhardt, Hartt, Copeland, Sceva, Dexter, James, 
Hunnewell and Thayer embarked for the United States. 

Allen, Ward and Sceva left Rio de Janeiro 9 June 1865 under 
the leadership of Orestes St. John. Their route led to Juiz de Fora, 
then across the Serra da Mantiqueira to Barbacena (where Ward 
left the group) through Lagoa Dourada, across the Rio Caranda- 
hy and the Rio Paraopeba just above the water gap of the Serras da 
Piedade and Itatiaiassu, to the village of Morro Velho. They tra- 
versed the basins of the Rio Parahyba, Rio Grande (Rio de La 
Plata) and Rio Sao Francisco. By way of Sabura, Santa Luzia, 
Lagoa Sancta and Sette Lagoas they reached Gequitiba. 

Sceva remained in Lagoa Santa to search for fossils in the caves 
of the region, but a previous collector had taken most of the ma- 
terial. He did, nonetheless, prepare an excellent series of mam- 
mals before returning to Rio de Janeiro. After a few days in Rio 
de Janeiro, devoted to preparing and packing specimens collected 
by other members of the expedition, Sceva went to Cantagallo to 
collect until he rejoined Agassiz in Rio de Janeiro to return to the 
United States. 

St. John and Allen left Gequitiba 31 July 1865 and travelled by 
canoe down the Rio das Velhas, arriving at the junction of the Rio 
Sao Francisco 28 August. After three days here they continued by 
boat to Januaria, reaching there 9 September and remaining until 
19 September. 

Allen, in ill health, decided at this point to continue on alone 
to Bahia, taking with him the material so far collected. Still trav- 



10 BREVIORA No. 444 

elling by boat he followed the Rio Sao Francisco past Urubii to 
Chique-Chique. where he arrived 8 October and stayed for several 
weeks. From Chique-Chique his route led overland to the coast 
by way of Engenho Velho, Jacare and Olhos d'Agua, across an 
arid plain, to Taboleiro de Jacobina, down the "Tombador" (a 
precipitous defile) to Jacobina valley. From Jacobina he went to 
the Serra da Terra Dura, Arraial do Riacho do Jacuhype, Feira 
de Sta. Anna, Espelto (Espelho?) and Cachoeira, then to Bahia, a 
journey of many weeks. He did not reach Bahia until the end of 
November. On 1 5 December Allen left Bahia for Cambridge. 

St. John followed the Rio Sao Francisco to Barra, then went 
along the valley of the Rio Grande to the Rio Preto, Santa Rita 
and Paranagua. Several days were spent at Paranagua before con- 
tinuing on along the valley of the Rio Gurgueia to Manga. The 
party followed the Rio Paranahyba to Sao Gonqallo where many 
specimens (primarily birds, reptiles and insects) were taken. There- 
zina, capitol of the state of Piaui, was the next station and excellent 
collections of fishes were made from the Rio Poty, a tributary of 
the Rio Parnahyba. From Therezina St. John continued on to 
Caxias, proceeded to Maranhao by way of the Rio Itapicuru and 
arrived at Maranhao 8 January 1866. Illness kept St. John in 
Maranhao until February, when he left to join Agassiz in Para. 
His geological observations were as careful and as valuable as his 
zoological collections. 

After Ward separated from his associates at Barbacena he con- 
tinued on to Ouro-Preto, Mariana and Santa Barbara, then down 
the Rio Piracicaba to the Rio Doce. He followed the river almost 
to the point where it meets the Rio Antonio. Crossing the Serra 
das Esmeraldas he arrived at the basin of the Rio Jequitinhonha 
and explored several branches of the river after passing Diaman- 
tina. He proceeded cross-country to the town of Rio Pardo and the 
Rio Pardo. He crossed the Rio Sao Francisco at Januaria, then 
continued northwestward until he reached the Rio Tocantins, which 
he followed to Para. From Para Ward returned to the United 
States. 

Hartt visited Brazil twice, first as a member of the Thayer Expe- 
dition, and, later, on a private expedition. Although he and Cope- 
land made extensive collections, particularly of marine inverte- 
brates and fishes, their major interest was the geology of Brazil. 
This has been comprehensively treated by Hartt, both from per- 



1977 THAYER EXPEDITION 11 

sonal observation and from the work of other geologists, in his 
book. The Scientific Results of a Journey in Brazil. 1870. Con- 
siderable information is also included on the common species of 
plants and animals and areas of fossil beds. 

Hartt and Copeland did not go directly from Rio de Janeiro to 
Bahia as planned. Their first attempt took them as far as Nova 
Almeida north of Victoria, but they were forced to return to Rio 
de Janeiro for lack of mules and money. After again working near 
Rio de Janeiro they embarked on a sailing vessel for Sao Matheos, 
stopping wherever possible along the coast. From Sao Matheos 
they went to the Rio Doce, returned to its mouth and then back to 
Sao Matheos. They proceeded north to Belmonte via the Rio 
Jequitinhonha and then returned to Porto Alegre, then north to 
Bahia. 

Rather than complicate the survey of the area covered by Hartt 
and Copeland the following route description is given (with the 
exception of their return to Porto Alegre) in direct sequence. 

The environs of Rio de Janeiro were thoroughly explored by 
Hartt and Copeland. They examined the area of the Serra do 
Mar. which borders the coast from the state of Sao Paola to the 
state of Rio de Janeiro, and also the Serra da Mantiqueira. partly 
separated from the Serra do Mar by the valley of the Rio Parahyba 
do Sul. Within the bay of Rio de Janeiro the Ilha de Paqueta, 
Ilha das Cobras, Ilha das Enxadas and Ilha do Governador were 
explored. They visited several places near Rio de Janeiro, includ- 
ing the hills that run westward to Lagoa de Freitas, separated 
from Corcovado by the valley of Botafogo. They also explored 
Tres Irmaos, southwest of Corcovado, the Gaira to the west 
and the Tijuca range to the north, which is separated from 
Corcovado by the pass of Boa Vista. They covered the entire length 
of the Dom Pedro II railroad. The Rio Macacu was followed to 
Porto Villa Nova, then along the Cantagallo Railway to Porto 
das Caixas and the terminus at Cachoeria. 

Between Rio Janeiro and Cabo Frio observations were made at 
Marica, Lagoa de Marica, Lagoa de Saquarema (east of Porto 
Negra) and Lagoa de Araruama. After Cabo Frio they stopped 
at Os Buzios. Cabo de Sao Thome, Macahe, the Ilhas de Santa Ana 
(which are a few miles offshore from Macahe) and Campos on the 
Rio Parahyba do Sul. Just south of Campos is the Lagoa Feia into 
which the Rio Macacu empties. Several lagoons have formed 



12 BREVIORA No. 444 

along this coast, including large ones such as Rio Iguassu and 
Lagoa do Campello. 

They followed the Rio Muriahe. a tributary of the Rio Parahyba, 
for several miles then followed the Rio Parahyba from Campos 
to Sao Fidelis. From here they travelled northward through the 
valley of Vallao Grande to Bom Jesus on the Rio Itabapuana, 
which divides the states of Rio de Janeiro and Espiritu Santo. 
They went down the Rio Itabapuana, past Porto da Limeira, to 
the mouth of the river. 

Hartt and Copeland continued northward to Lago Maroba, 
then along the coast to the town and river of Itapemirim and the 
surrounding area, then on to Piuma, the Rio Benevente and Guara- 
pary. Stops were made at Ponta da Fructa and Ponta de Jecii en 
route to the bay of Espiritu Santo, Villa Telha (on the bay of Espi- 
ritu Santo) and Victoria. Fifty miles northwest of Victoria they ex- 
plored the Rio Santa Maria and Sao Leopoldina then went on to 
Carapina, Santa Cruz and the Rio Doce, which they traversed from 
its mouth to Porto do Souza. They went down the river to Linhares 
and Lagoa Juparanaa (connected to the Rio Doce by a deep chan- 
nel called the Rio Juparanaa). 

Returning to the coast they continued northward, stayed briefly 
during November 1865 in the vicinity of Barra Secca, then passed 
the Lagoa Marircu and the Rio Azeites at As Azeites on the way 
to Sao Matheos. Hartt journeyed to the Fazenda do Capitao 
Grande on the Rio Bra^o do Norte, which joins with the Brago do Sul 
to form the Ric Sao Matheos. On the return trip he mapped the river 
as far as Sao Matheos. Further north along the coast they noted 
the Rio Itahu.ias. 

From Porto Alegre they went up the Rio Mucury to Santa Clara 
and from that point travelled westward, using the Minas road, to 
the headwaters of the Rio Mucury near Pote. Farther west they 
passed the watershed dividing the Rio Mucury and the Rio Jequi- 
tinhonha. They arrived at the valley of Jequitinhonha in April 
1866 via the Rio Setubal. A detour was made from the Rio Setubal 
to Alto do Bois before they proceeded cross-country to Calhao. 
After arriving at Calhao, Hartt made a visit to Minas Novas, re- 
turned to Calhao, and with Copeland, went down the Rio Aras- 
suahy and the Rio Jequitinhonha to Belmonte. From Belmonte, 
Hartt and Copeland returned to Porto Alegre, before again follow- 
ing the coast north. 



1977 THAYER EXPEDITION 13 

Observations were made of the terrain between the Mucury and 
Peruhype rivers. They visited Sao Leopoldina, about ten miles 
upstream, and Villa Vi9osa, four miles below. The route continued 
northward along the coast to the Rio Caravellas, Prado, the Rio 
Jucurucu, Porto Seguro and Santa Cruz. After again reaching 
Belmonte they went on to Cannavieiras and the Rio Pardo. They 
ascended the river to the head of navigation, passing Cachoeirinha 
do Rio Pardo and the fazenda of Sisterio. 

They stopped briefly in Ilheos on the Rio Cachoeira and in 
Camamu. en route to Bahia. From Bahia they ascended the Rio 
Paraguassu as far as Cachoeira. passing the valley of Iguape. The 
area surrounding Bahia received close attention before Hartt and 
Copeland returned to Rio de Janeiro, where they joined Agassiz 
and other members of the expedition for departure to the United 
States. 



14 BREVIORA No. 444 

INDEX 

Name Map 



Abrolhos. Parcel dos 


3 A 


Alto dos Bois 


3A 


Aniarcmtc (San Gonqallo) 


5 


Antonio (So/z/o Antonio), Rio 


7 


Arancho 


11 


Arary (Arari) 


11 


Arassuahy (Arac^uai), Rio 


3A 


Armaqao 


3B 


Arraial do Riacho do Jacuhype 


6 


Arupuana, Rio 


9 


As Azeites 


3B 


Assii Grande 


9 


Aturia, Rio (Furo do Aturid) 


9 


Azeites, Rio 


3B 


Bagro do Rio das Velhas 


4, 6 


Bahia, (Salvador) 


3A, 4, 6, 11 


Barbacena 


4, 7 


Barra (Itabapoana) 


3B 


Barra (Villa do Barra) 


4, 6 


Barra Secca {Seca) 


3B 


Barreira das Cudajas, (Codajds) 


10 


Belmonte 


3A, 6 


Belmonte, (Jequitinhonha), Rio 


3 A 


Benevente, Rio 


3B 


Boa Vista, Pass 


1 


Bom Jardin (Jardim) 


6 


Bom Jesus (Bom Jesus do Itabapoana) 


3B 


Botafogo 


1 


Bra^o do Norte, Rio 


3A 


Braqo do Sul (Mantena), Rio 


3A 


Branco, Rio 


10 


Breves 


9 


Cabo de Sao Thome (Tome) 


3B 


Cabo Frio 


3B, 6 


Cabrdlia (Santa Cruz) 


3B 


Cachoeira (Caxoeira) 


3A, 6 


Calhao, (Itira) 


3A 



1977 THAYER EXPEDITION 15 

Name Map 

Camamu 3A 

Cameta 9 

Campos 3B 

Caninde, Rio 5 

Cannavieiras 3A 

Cantagallo (Cantagalo) 2 

Cantagallo (Cantagalo), Railroad 2 

Capita© Grande 3B 

Carandahy (Carandai), Rio 4 

Caravellas (Caravelas), Rio 3A 

Carolina 8 

Caxias 5 

Caxoeira (Cachoeira) 3A 

Caxoeira, Rio 3A 

Ceara (Forialeza) 11 

Chique Chique (Xique Xique) 4, 5, 6 

Coari 10 

Codajds (Barriera das Cudajas) 10 

Coeira, Rio 3A 

Co Ionia Leopoldina (Sao Leopoldina) 3A 

Contas, Rio de 3A 

Corcovado I 

Cudajas 10 

CuruQa 9 

Curupira, Lago 9 

Diamantina 4, 6, 7 

Doce, Rio 3B, 7 

Dom Pedro II Railroad 2 

Ega (Teffe) ( Tefe) 10 

Engenho Velho 6 

Entre Rios 2 

Erere, Rio 9 

Espelto (Espelho?) 6 

Espiritu Santo, Bay 3B 

Esta^So d'Entre Rios {Tres Rios) 2 

Fatuarana 9 

Fazenda Sen. Lage 3B 

Feira de Sant'Anna {Santana) 6 



16 BREVIORA No. 444 

Name Map 



Fonte Boa 


10 


Formosa 


7 


Fortaleza (Ceara) 


11 


Furo do Atiirid (Rio Aturia) 


9 


Furo do Tajapuru (Rio Tajapuru) 


9 


Gaira 


1 


Gavia (Gdvea) 


1 


Gequitiba {Jequitihd) 


4,6 


Gongogi, Rio 


3A 


Grande, Rio 


6 


Guarapary (Guarapari) 


3B 


Guaxindiba or Itahunas (ftaunas), Rio 


3B 


Gurugueia (Gurgueia). Rio 


5 


Gurupatuba {Parieutuba), Rio 


9 


Gurupu (Gurupd) 


9 


Hyanuary, Lago 


10 


Hyapura {Japurd), Rio 


10 


Hyavary, (Javary) (Javari), Rio 


10 


Iqa, Rio 


10 


Igarape-Assu (A9U) 


9 


Igarape Grande 


9 


Iguape, Rio 


3A 


Iguassu (Igua<^u), Rio 


3B 


Ilha das Cobras 


1 


Ilha das Enxadas 


1 


Ilha de Marajo 


9 


Ilha do Governador 


1 


Ilha Grande do Jutai 


8 


Ilha Paqueta 


1 


Ilhas de Santa Ana (Santaua) (Anna) 


3B 


Ilheos (Ilheus) 


3A, 6 


Itahapoana (Barra) 


3B 


Itabapuana (Itahapoana), Rio 


3B 


Itacoatiare (Serpa) 


9 


Itapemerim (Itapemirim). Rio 


3B 


Itapicuru, Rio 


5, 11 


Itatiaiassu (Itatiaiu^u), Serra do 


4, 6 


Itai'mas (Guaxindiba or Itahunas), Rio 


3B 



1977 THAYER EXPEDITION 17 

Name Map 

Itim. Rio 5 

Jacare. Rio 6 

Jacobina 6 

Jacuipe (Jacuhype), Rio 3A 

Januaria 4, 6, 7 

ya/7wra' (Hyapura). Rio 10 

Jatuarana 9 

Javary (Javari) (Hyavary), Rio 10 

Jequitibd (Gequitiba) 4, 6 

Jequitinhonha (Belmonte), Rio 3 A, 7 

Jose Assu (/4(w), Lago 9 

Jose Fernandez, Lago 10 

Jucurucu, Rio 3A 

Juiz de Fora 3B, 4, 6, 7 

Juparanaa, Rio (Lagoa Juparana) 3B 

Jutahy {Jutai), Rio 10 

Lagoa de Araruama 2 

Lagoa de Freitas (Lagoa Rodrigo de Freitas) 1 

Lagoa de Marica 2 

Lagoa de Saquarema 2 

Lagoa do Campello (Campelo) 3B 

Lagoa do Maximo (Lago Maximo) 9 

Lagoa Dourada 4, 6 

Lagoa Feia 3B 

Lagoa Juparana (Rio Juparanaa) 3B 

Lagoa Maricu 3B 

Lagoa Sancta (Santa) 4, 6 

Lago Cudajas (CtxVa/a'i'j 10 

Lago Curupira 9 

Lago do Bota 10 

Lago Hyanuary (January) 10 

Lago Irandubo 10 

Lago January (Hyanuary) 10 

Lago Jose Assii (A(^u) 9 

Lago Jose Fernandes 10 

Lago Manacapuru 10 

Lago Maroba 3B 

Lago Saraca 9 



18 BREVIORA No. 444 

Name Map 

La Plata (Grandel), Rio 6 

Linhares 3B 

Macacos, Rio dos 9 

Macacu. Rio 2 

Macahe (Macae) 3B 

Maceio 11 

Madeira, Rio 9 

Mamuru (Tupinambaranas), Rio 9 

Manacapuru 10 

Manaos (Manaus) 9, 10 

Manga 5 

Manhuassu {Manhuaqu), Rio 3B 

Mautena{^v2i<;o do Sul), Rio 3 A 

Marajo, Ilha de 9 

Maranhao (Sao Luis) 11 

Maranon, Rio 10 

Mariana 7 

Marica, Lagoa do, Ilhas 3B 

Mateus, Rio 3A 

Maua (Barao de Maud) 2 

Mauhes (Maues), Rio 9 

Minas Novas 3A 

Monte Alegre 9 

Morro Velho 4, 6 

Moura (Pedreira) 10 

Mucaja-Tuba (Mucajd) 9 

Mucury (Mucuri). Rio 3A 

Mucury Pequeno, Rio 3A 

Mugiquisaba (Mogiquic^aha) 3A 

Muriache (Muriahe) (Muriae). Rio 3B 

Mutum, Rio 3B 

Nazareth 9 

Negro, Rio 10 

Novo, Rio 3B 

Obydos (Obidos) 9 

Olhos d'Agua 6 

Organ Mountains (Serra dos Orgaos) 2 

Os Buzios (Caho dos Buzios) 3B 



1977 THAYER EXPEDITION 19 

Name Map 

Ouro-Preto (Ouro Preto) 7 

Pacatuba 1 1 

Paqueta, Ilha de 1^3 

Para (Beleni) 8, 9 

Parahyba (Paraiha) do Norte. Rio 1 1 

Parahyba {Paraiha) do Sul, Rio 3B, 4 

Parahybuna (Paraihuna), Rio 3B 

Paramirim, Rio 6 

Paranagua 5 

Paraopeba, Rio 6 

Parcel dos Abrolhos 3A 

Pardo 3A, 7 

Pardo, Rio 3A, 7 

Parguassu (Parguac^u), Rio 3 A 

Pahntim (Villa Bella) 9 

Parnaiba (Parnahyba), Rio 5 

Pedreira (Moura) 10 

Pernamhuco (Recife) 11 

Peruhype (Peruipe), Rio 3A 

Petropolis 2, 6 

Philadelphia ( Teqfilo Otoni) 3A 

Piabanha. Rio 2, 33 

Piauhy (Piaui), Rio 3 A 

Piedade 1 

Piracicaba, Rio 7 

Pirahy (Pirai), Rio 2 

Po-Assu {Canal de Poac^u) 3 A 

Ponta da Fructa {Fruta) 3B 

Ponta de Jecu 3B 

Ponta Negra 2 

Porto Alegre 3A 

Porto da Limeira 3B 

Porto das Caixas 2 

Porto de Moz 9 

Porto do Souza 3B 

Porto Nacional 7, 8 

Porto Seguro 3A 

Posse 3B 



20 BREVIORA No. 444 

Name Map 

Pote 3A 

Poty (Puty) (Poll), Rio 5 

Prado 3A 

Prados 4 

Prainha 9 

Preto, Rio 2, 4 

Puty (Poti), (Poty), Rio 5 

Ramos, Rio (Parana do Ramos) 9 

/^iT//^ (Pernambuco) 11 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Ri 

Rio de Contas 3A 

Rio de Janeiro 1, 3, 4, 6, 7 

Rio Doce 3B, 7 

Rio dos Macacos 9 

Rio Erere 9 

Rio Gongogi 3A 

Rio Grande 6 

Rio Grande (La Plata?) 4 

Rio Guaxindiba or Itahunas (Itaunas) 3B 

Rio Giirgueia (Gurugueia) 5 

Rio Gurupatuba (Parieuiuha) 9 

Rio Hyapura (Japwd) 10 

Rio Hyavary (Javary) (Javari) 10 



o Antonio (Santo Antonio) 1 

o Arupuana 9 

o Arassuahy (Ara(^uai) 3 A 

o Azeites 3B 

o Aturia (Furo do At una) 9 

o Belmonte (Jequitinhonha) 3A, 7 

o Benevente 3B 

o Bra^o do Norte 3A 

o Bra(jo do Sul (Rio Mantena) 3 A 

o Branco 10 

o Caninde 5 

o Carandahy (Carandai) 4 

o Caravellas (Caravelas) 3 A 

o Caxoeira 3A 

o Coeira 3A 

o das Velhas 3A, 4, 6, 7 



1977 THAYER EXPEDITION 21 

Name Map 



Rio Hyutahy 


10 


Rio I^a 


10 


Rio Iguape 


3A 


Rio Iguassii 


3B 


Rio Itabapuana {Itabapoana) 


3B 


Rio Itanheim 


3A 


Rio Itapemerim 


3B 


Rio Itapicurii 


5, 11 


Rio Itaimas'iliSihnnsiS or Guaxindiba) 


3B 


Rio Itim 


5 


Rio Jacare 


6 


Rio Jacuipe (Jacuhype) 


3A 


Rio Japurd (Hyapura) 


10 


Rio Javary (Hyavary) (Javari) 


10 


Rio Jequitinhonha (Belmonte) 


3A, 7 


Rio Jucururii 


3A 


Rio Juparanaa {Lagoa Juparana) 


3B 


Rio Jutahy (Jutai) 


10 


Rio Macacu 


2 


Rio Madeira 


9 


Rio Manhua^u (Manhuassu) 


3B 


Rio Mantena (Bra^o do Sul) 


3A 


Rio Maranon 


10 


Rio Mateus 


3B 


Rio Mauhes (Maues) 


9 


Rio Mucury (Mucuri) 


3A 


Rio Mucury Pequeno 


3A 


Rio Muriahe (Muriae) (Muriache) 


3B 


Rio Mutum 


3B 


Rio Negro 


10 


Rio Novo 


3B 


Rio Paraguassu 


3A 


Rio Parahyba (P^raiha) do Norte 


11 


Rio Parahyba (Paraiba) do Sul 


2, 3B, 4 


Rio Parahybuna (Paraibuna) 


3B 


Rio Paramirim 


6 


Rio Parnahyba (Parnaiba) 


11 


Rio Paraopeba 


6 



22 BREVIORA No. 444 

Name Map 



o Pardo 3 A 

o Parnaiha (Parnahyba) 5 

o Peruhype ( Av7///u') 3A 

o Piabanha 2, 3A 

o Piauhy 3A, 5 

o Piracicaba 7, 

o Pirahy 2 

o Piuma 3B 

o Poty (Poti) (Puty) 5 

o Preto 2. 3A, 4 

o Ramos ( Parana de Ramus) 9 

o Sanianihaia (Sambaia) 2 

o Santo Antonio (Antonio) 7 

o Sao Francisco 3A, 4, 5, 6, 7 



o Sao Matheos (Sao Mateus) 3B 

o Setubal 3A 

o Solimoens (Solimoes) 10 

o Tajipuru (Furo do Tajipnru) 9 

o Tapa]07{TapaJds) 9 

o Tocantins 7, 8 

o Trombetas 9 

o Tupinambaranas (Mamurn) 9 

o Una 3B 

o Vacaria 7 

o XingLi 9 

Sabura (Sahara) 4 

Salvaterra 9 

Saniamhaia (Sambaia). Rio 2 

San Gon(^allo (Amarante) 5 

San Luiz (Maranhao) 11 

San Mateo (5'^7r^ A/^;/f'z/.v) ( Matheos) 3B 

San Paolo (Scio Paulo de Olivem^a) 10 

Santa Anna Islands (//has de Santana) 3B 

Santa Barbara 3A, 38, 7 

Santa Caterina (Catherina) 3B 

Santa Clara 3A 

Santa Cru? (Cahra/ia) 3B 

Santa Luzia 4, 6 



1977 THAYER EXPEDITION 23 

Name Map 



Santarem 


9 


Santa Rita de Cassia (Villa da Santa Rita) 


4 


Santo Antonio (Antonio), Rio 


7 


Sao Fidelis 


3B 


Sao Francisco. Rio 


3A,4,5, 


Sao Leopoldina (ColSnia Leopoldina) 


3B 


Sao Matheos (Sao Mateus) 


3B 


Sao Paulo de Oliven^a (San Paola) 


10 


Sao Tome, Cabo de 


3B 


Serpa (Itacoatiare) 


9 


Serra da Aratanha 


11 


Serra da Babylonia (Babilonia) 


3B 


Serra da Mantiqueira 


3B, 4, 7 


Serra da Piedade 


4, 6 


Serra das Esmaraldas 


7 


Serra da Terra Dura 


6 


Serra de Cupati 


10 


Serra de Monguba 


11 


Serra d'Erere (Erere) 


9 


Serra do Itatiaiassu (Itatiaiuqu) 


4, 6 


Serra do Mar 


3B 


Serra dos Orgaos (Organ Mountains) 


3B 


Sette Lassas 


4, 6 


Setubal. Rio 


3A 


Silva 


9 


Sisterio 


3A 


Solimoens (Solimoes), Rio 


10 


Soures (Soure) 


9 


Tabatinga 


10 


Taboleiro de Jacobina 


6 


Tajipuru, Rio (Furo do Tajipuru) 


9 


Tapajoz (Tapajds), Rio 


9 


Tatuatuba 


9 


Taua Peassu ( Tauape(;a(^u) 


10 


Teffe (Ega) ( Tefe^ 


10 


Teofilo O/OA?/ (Philadelphia) 


3A 


Teresopolir. (Theresopolis) 


3B 


Therezina ( Teresina) 


5 



6,7 



24 BREVIORA No. 444 

Name Map 

Tijuca 1 

Tocantins, Rio 7, 8 

Tombador 6 

Tonantins 10 

Tres Irmaos 1 

Tres Rios (Esta^ao d'Entre Rios) 2 

Trombetas, Rio 9 

Tupinambaranas ( Tupinambarama) 9 

Tupinambaranas (Mamuru), Rio 9 

Una, Rio 3B 

Urubu 6 

Vacaria, Rio 7 

Vallao Grande ( Valao Grande) 3B 

Velhas, Rio das 3A, 4, 6, 7 

Vereda do Mocambo (Mocambo) 4 

Victoria ( Viioria) 6 

Vigia 9 

Villa Bella (/'an/?///?.?) 9 

Villa da Santa Rita {Santa Rita de Cassia) 4 

Villa do Barra {Barra) 4, 6 

Villa ( Vila) Velha 3B 

Villa (F/7o) Viqosa 3 A 

Vitoria (Victoria) 6 

Xingu, Rio 9 

Xique Xique (Chique Chique) 4, 5, 6 

Ypiranga {Ipiranga) 2 



1977 THAYER EXPEDITION 25 

REFERENCES 

Agassiz, LoL'is, ANO El IZABETH Carv. 1868. Journey in Brazil. Boston, Ticknor 

and Fields. 540 pp. 
Allen, Joel A. 1916. Autobiographical Notes and Bibliography of Scientific 

Publications. New York, American Museum of Natural History, pp. 11, 215. 
Cambridge. Harvard University Archives. William James papers. 
Museum of Comparative Zoology. Scientific catalogs and specimens from 

various departments. 

Museum of Comparative Zoology Archives. Louis Agassiz papers. 

Museum of Comparative Zoology Archives. "List of Fishes" (by Orestes 



St. John). 

Eigenmann, C'. H. 1917. The American Characidae. Memoirs of the Museum 
of Comparative Zoology, 43. 

Hartt, C. F. 1870. Scientific Results of a Journey in Brazil. Boston, Fields, Os- 
good and Company, xxiii + 620 pp. 

Institlto Brasileiro de Geografia. 1972. Carta do Brasil Ao Milionesimo. Rio 
de Janeiro. 

LuRiE, Edward. 1960. A Life in Science. Chicago, University of Chicago Press, 
pp. 345-350. 

Tharp, LoriSE Hall. 1959. Adventurous Alliance. Boston, Little, Brown and 
Company, pp. 16 190, 204-210. 

Times Atlas. 1957. Volume 5, Cambridge, Houghton, Mifflin Co. Maps 115, 
116, 117, pp. xii, 23, 57. 

United States Board on Geographic Names. 1963. Gazetteer, Brazil. Wash- 
ington, D.C., U.S. Government Printing Office. 915 pp. 

Vanzolini, P. E. AND N. Papavero. 1968. Indice Dos Toponimos Contidos Na 
Carta Do Brazil. Sao Paulo, Funda^ao de Ampara a Pequisa do Estado de Sao 
Paulo. 197 pp. 

WiLGUS, A. Curtis. 1972. Nineteenth Century Travelers: Louis and Elizabeth 
Agassiz. Americas 24 (2) : 25-32. 



26 



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No. 444 




1977 



IHAVl R FXPEIMIION 



27 




28 



BREVIORA 



No. 444 




1977 



THAYER EXPEDITION 



29 




u 



30 



BREVIORA 



No. 444 




Figure 4. Rio de Janeiro to Paranagua - St. John 



THAYER EXPEDITION 



31 




Figure 5. Paranagua to Maranhao — St. John 



32 



BREVIORA 



No. 444 



CHIQUE CMIQUE 




Figure 6. Rio dc Janeiro. Chique Chique. to Bahia — Allen 



1977 



THAYER EXPEDITION 



33 





Figure 7. Rio de Janeiro to Rio Tocantins — Ward 



34 



BREVIORA 



No. 444 




PORTO NACIONAL 



Figure 8. Rio Tocantins to Para — Ward 



1977 



THAYER EXPEDITION 



35 




36 



BREVIORA 



No. 444 




1977 



THAYER EXPEDITION 



37 



ARARY 




N 



PERNAMBUCO 



MACEIO 



BAHIA 



Figure 1 1. Coast. Maranhao to Bahia — Agassiz et al 



-f^^* ' 



MU3. COM?. ZOOL 
LIOrJARY 



MAR 1 8 1985 

B R E V^'I-S©PvR A 

Museum of Comparative Zoology 



LIS ISSN 0006-9698 



Cambridge. Mass. 30 November 1977 Number 445 

NATURAL HISTORY OF CERION. VII. 

GEOGRAPHIC VARIATION OF CERION (MOLLUSCA: 

PULMONATA) FROM THE EASTERN END OF ITS RANGE 

(HISPANIOLA TO THE VIRGIN ISLANDS): COHERENT 

PATTERNS AND TAXONOMIC SIMPLIFICATION. 

Stephen Jay Gould' and Charles Paull- 

Abstract. The eastern cerions (Hispaniola to the Virgin Islands) have, following 
the conventional practice for this diverse genus, been split into seven allopatric taxa. 
We reject this traditional scheme on the basis of a multivariate morphometric analy- 
sis ( 19 characters, 20 shells per sample) of 23 samples from all major areas of Cerion's 
eastern range. We first show that eastern cerions are distinct from other members 
of the genus by patterns of covariation among their morphometric measures. We 
then demonstrate by canonical analysis that populations of each island have a dis- 
tinct morphology. Were there no coherent patterns among islands, current nomen- 
clature might be supported. However, the first canonical axis (59 per cent of all 
information) for an analysis by islands arrays the populations in perfect geographic 
order — from egg-shaped, finely and copiously ribbed shells in the east, to more 
cylindrical, apically pointed shells with fewer, stronger ribs in the west. The mor- 
phological direction of this cline approaches common cerion forms further west 
(particularly on Cuba); unique morphologies are isolated at the eastern extreme 
of the range. In addition, a plot of Mahalanbbis vs. geographical distance shows 
a remarkably tight relationship (r = 0.96) between geographic and morphological 
distance. Therefore, we synonymize all living eastern cerions into the single taxon 
C. sirlatellum ("Ferrusac" Guerin-Meneville), reserving C. rude (Pfeiffer) for the 
rather different St. Croix fossil. 

I. INTRODUCTION 

Cerion has attracted the attention of eminent naturaHsts ever 
since Linnaeus designated its type species in 1758. W. H. Dall 
(1905). H. A. Pilsbry (1902). P. Bartsch (1920). Ludwig Plate 



'Museum of Comparative Zoology. Harvard University. 
^Rosenstiel School of Marine Science, Universitv of Miami. 



2 BREVIORA No. 445 

(1906, 1907). W. J. Clench (1957), and Ernst Mayr (1956, 1963) 
invested substantial segments of their careers trying to understand 
the protean diversity of this unusual genus containing more than 
600 described species and subspecies. So far Cerion has stood firm 
against all efforts to find coherence in the geographic distribution 
of its exuberant morphological variation. Elegant, oversimplified 
stories have collapsed (Plate, 1906), while more circumspect evo- 
lutionists speak of "crazy-quilt" distributions and random trans- 
port, by hurricanes (Mayr and Rosen, 1956). The burden of avail- 
able names has generally precluded a search for order and simplicity 
in the areas of Cerion's greatest diversity — Cuba, the Bahamas, 
and the Cayman Islands. Yet order and simplicity demonstrably 
exist when we trade museum trays and lists of names for the map- 
ping of recognizable elements and their interactions in the field 
(Gould and Woodruff, in press, and in preparation on Little Ba- 
hama Bank, New Providence, Great Exuma, and Long Island, 
Bahamas). 

As another strategy, we might wage some preliminary skirmishes 
before attacking the central areas of Cuba and the Bahamas, since 
Cerion displays markedly less variation in three peripheral areas 
of its distribution. The isolated populations on Aruba, Bonaire, 
and Curasao have received much attention (Baker, 1924; Hum- 
melinck, 1940; de Vries, 1974; Gould, 1969). Geographic variation 
in the single species, Cerion uva (L.), reflects both isolation (each 
island has a characteristic, if subtly differentiated, morphology) 
and local habitat; patterns have been stable for at least 50 years 
(Baker, 1924; Gould, 1969). This coherence allows hope for sensi- 
ble interpretations elsewhere. The second peripheral area is con- 
tiguous with central regions: the Florida Keys with their single 
species, Cerion incanum (Binney). The third peripheral area is 
larger, more diverse, also contiguous to major centers, and equally 
unstudied: the essentially linear array of islands running from 
Hispaniola through Mona Island and Puerto Rico, to Necker and 
Anegada in the Virgin Islands (Fig. 1). Here 11 names are avail- 
able for a basic morphology that all students of Cerion have recog- 
nized as unique to this Eastern area (Pilsbry, 1902). These eastern 
cerions provide a superb opportunity for modern study; they repre- 
sent a situation intermediate between the relative simplicity of 
Cerion uva in the Leeward Islands (where we cannot practice the 



1977 



EASTERN CERIONS 



art of taxonomic rectification, since no one ever successfully split 
the single species) to the complexity of New Providence Island with 
its 82 named species. 




Fig. 1 . The distribution of eastern cerions. Crosses indicate localities used in 
this study for the large islands only. 



Moreover, the linear arrangement of islands leads to simple hy- 
potheses of clinal distribution in our search for coherence, while 
their geographic range (980 km from Western Haiti to Anegada) 
permits a truly regional not merely local, study. 

II. SYSTEMATICS OF EASTERN CERIONS 

Collectors of eastern cerions report that these snails share the 
habitat conventionally assigned to the genus (though our own ob- 
servations of Bahamian forms demonstrate a much wider range); 
they live near the coast in limestone regions. Traditional wisdom 
proclaims that the animals are active on moist evenings, feeding 
predominantly on fungal mycelia in decaying vegetation; in drier 
conditions, they secrete an epiphragm and attach to plants, remain- 
ing immobile for long periods (several months in dry laboratory 
conditions). Beyond these basic facts of distribution, we know vir- 
tually nothing about the ecology and life history of any Cerion, 
including the eastern forms. (D. S. Woodruff has been studying 
two populations on Abaco Island, Bahamas during the past three 
and one-half years; our first adequate data will soon be available.) 

Heretofore, the eastern cerions have been defined by concho- 
logical characters alone. These snails have a unique and consistent 
morphology (Fig. 2), rendered in most taxonomic descriptions by 
two characters: 



BRFVIORA 



No. 445 



1) An unusually obtuse apex, giving the entire shell a barrel or 
egg-shaped appearance (contrasting with the more common cylin- 
drical form of species with pointed apices and rapid attainment in 
ontogeny of a definitive width; or the triangular form of species 
with pointed tops and continually increasing width). 

2) Fine, abundant and regular ribs, evenly covering the entire 
surface of the shell. 




Fig. 2. Representative eastern cerions. Top row Hispaniola: second row, Mona 
Island (left) to Anegada (right). The large specimen at the extreme right is fossil 
C. rude from St. Croix. Note the major clinal trends from coarsely ribbed, apically 
pointed species in the west (Hispaniola) to copiously and finely ribbed, barrel- 
shaped forms in the east. Note also, the peculiarities of individual islands as de- 
scribed in the text — i.e., the nearly circular aperture of Puerto Rican specimens. 
Top row, starting from left: 1) holotype of "C. tortuga" (Acad. Nat. Sci. Phil. No. 
146701 — locality 22 of this study); 2) holotype of "C. yumaense ferruginum" 
(M.C.Z. No. 76230): 3) paratype of "f. vunuwnsc" (M.C.Z. No. 185945 locality 
18 of this study): 4) holotype of "C. vunuicn^f sa<ina" (Acad. Nat. Sci. Phil. No. 
47200). Bottom row: 1) holotvpc ol "(". nunuicnsc" from Mona Island (M.C.Z. 
No. 171019 —locality 12 of this studv ): I'ucrto Rican specimen of ('. \iii(iicllum 
from locality 7: 3) from Nccker Island. Virgin Islands, localitv I: 4) from Anegada. 
Virgin Islands, locality 2. C\ rude is from lot No. 60564 Acad. Nat. Sci. Phil. C. 
ruilc specimen is 34.7 mm in height 



1977 EASFERN CFRIONS 5 

Pilsbry (1902), in the last comprehensive attempt to estabhsh 
subgroups within Cerion, defined the eastern forms as one section 
among 1 1 in the subgenus Strophiops. He included two extrane- 
ous forms within the group: 

1. Cerion antonii reported by Kiister (1847) from the South 
American coast in Guyana. This species has been dutifully listed 
by all students of Cerion: comments about its anomalous geogra- 
phy abound. But we do not believe that it exists. The shells were 
described from a collector's cabinet, not from field observations. 
Mistaken labels in such cabinets once gave other cerions a proud 
range from India to China. We have no field records of living 
Cerion in South America. Moreover, Kiister's shells have not been 
seen, so far as we can tell, by any subsequent author. All records 
of C antonii depend upon the single poor figure published by 
Kiister and reproduced by Pilsbry (1902) in making his assignment. 

2. C. caymanense Pilsbry from Grand Cayman. According to 
Pilsbry (1902), C. caymanense differs from other members of the 
eastern group by its less obtuse apex and stouter ribs. But these 
are the very characters that should place it out of the group; for 
obtuse apices and fine ribs are the defining traits of eastern cerions. 
We do not understand why Pilsbry didn't include C. caymanense 
with the more similar Cuban group of C. cyclostomum (Kiister). 

The remaining taxa of Pilsbry's eastern group all reside m the 
geographic area of this study. Current nomenclature divides the 
eastern cerions according to accepted criteria within the genus: 
specific names have been awarded to allopatric populations of 
recognizable morphology. Moving from east to west among living 
species, cerions of the Virgin Islands (Anegada and Necker) and 
Puerto Rico belong to the same taxon. Cerion siriatellum — al- 
though the phenetic distance between Puerto Rican and Virgin 
Island samples is greater than that between most other species in 
the group (see section VI). Some confusion has surrounded the 
appropriate name for this taxon. Pupa striatella. attributed to 
Ferussac but never described by him. was defined by Guerin- 
Meneville in the 1829 Rci^ne Animal of Cuvier (MoUusca. p. 16, 
plate 6. Hg. 12). But Guerin-Mene\ille listed only "the Antilles" for 
a locality, and the task of deciding which cerions merited the desig- 
nation fell to later authors. Kiister ( 1 847) and many others attributed 
Guerin-Mene\ille"s name to the Puerto Rican specimens, but Poey 



6 BRFVIORA No. 445 

and others applied C\ stnaiclluin to a species from Cabo Cruz. Cuba 
and used So\\erb\'s (l(S75) ('. crassilahrc tor Puerto Rico. Piisbry 
(1902) followed Poey, but changed his mind later (1943), awarded 
a new name to the Cuban species (C. cahocnizense Piisbry and de 
la Torre), and preserved the earlier C. striatelluni for the Puerto 
Rican form. We follow Pilsbry's later opinion and consider C 
crassilabre as a synonym of C striatellum. This is a happy reso- 
lution since we will synonymize all living eastern cerions under this 
name, and since thick lips are decidedly not characteristic of many 
populations, whereas all are abundantly ribbed. 

Moving westward from Puerto Rico, we next encounter Cerion 
on Mona Island. Clench (1951, p. 274) awarded the Mona Island 
population their own name (C monaense) for their coarser costae 
and smaller size compared with Puerto Rican relatives. 

Taxonomy of the Hispaniolan forms is more exuberant, but the 
defining criteria — size, ribbing, relative width, and shape of apex 
— have not been enlarged. Kiister's name (1847). Pupa striatella 
minor from Haiti, is the earliest available, but all subsequent au- 
thors have ignored Kiister's inadequate description, and we will not 
challenge this tradition. Piisbry and Vanatta described the first 
full species in 1895 (p. 210); Cerion yumaeme from the Dominican 
Republic — not so wide, more pointed apically, and more coarsely 
ribbed than the Puerto Rican populations. Maynard (1896) fol- 
lowed soon afterwards with Strophia ferruginea from Haiti, gen- 
erally ranked as a subspecies of C. ywnaense by later authors (e.g., 
by Piisbry, 1902). Two additional subspecies accrued in later years: 
C ywnaense sallei (Piisbry and Vanatta, 1896, p. 325; first attrib- 
uted to C. striatellum, but later, and more appropriately, switched 
to C ywnaense by Piisbry, 1902), and C. yumacnse saona (Va- 
natta, 1923). Definitions involve little more than size: sallei is quite 
small, while saona is larger, with coarser costae than either sallei or 
the nominate subspecies. Finally, Piisbry and Vanatta ( 1 928) erected 
Cerion tortuga for more strongly colored, thicker lipped specimens 
from the nearby island of Tortuga. 

In summary, living eastern cerions are generally ranked in seven 
taxa: C. striatellum ("Ferussac" Guerin-Meneville) for Puerto Rico 
and the Virgin Islands, C. monaense Clench for Mona Island (be- 
tween Puerto Rico and Hispaniola), and C. yumaense Piisbry and 
Vanatta [with subspecies C. v. vumaense. C. y. ferruginum (May- 



1977 EASTERN CERIONS 7 

nard). C r. sallei Pilsbry and Vanatta. and C. y. saona Vanatta] 
and C lonuga Pilsbry and Vanatta for Hispaniola and associated 
islands. They are distinguished by little more than variation in the 
key characters used to define the entire eastern group: coarseness 
of ribbing, obtusity of the apex, and width of the shell. 

A somewhat different fossil form has long been known from St. 
Croix Cerion rude (Pfeiffer). [Pfeiffer's (1855) second taxon, C 
latilahre. has been properly synonymized by all later authors as 
nothing more than a short specimen of C. rude.'] C. rude is consid- 
erably larger than other eastern forms. It also departs from its near- 
est neighbors in the very characters distinguishing them from other 
Cerion — C. rude has a more pointed apex and fewer, coarser ribs 
than most living eastern Cerion. Yet its general appearance is still 
closer to the living eastern cerions than to any other group within 
the genus. Jacobson (1968) states that the fossil shells are buried 
in soils and often appear at the surface after preparation of the 
land for cultivation of cane. He attributes their extinction to cul- 
tivation and burning (itself, primarily, for cultivation), but we have 
no firmly documented record of St. Croix Cerion living in historic 
times. 

III. MATERIALS, METHODS, AND INTENTIONS 

Following the protocol of Gould. Woodruff, and Martin (1974), 
we made 19 measurements on 20 specimens (when available) for 23 
samples of eastern cerions. All taxa but C. yumaense sallei are 
represented in our samples. Our measures attempt to capture the 
standard taxonomic characters of aduh size, whorl sizes and num- 
bers, size and shape of the aperture, shape of the spire, and pat- 
terns of ribbing. With the exception of one sample personally col- 
lected on Anegada (by C.P.), all shells come from museum collec- 
tions (see Table 1). In all but two cases (Maynard's paratypes of 
C yumaense ferruginum, and fossil C. rude — where we amalga- 
mated whatever we could find from the collections of four museums 
to form an adequate sample), we restricted ourselves to large, recent 
samples collected en masse without obvious preference for large, 
unusual or attractive specimens. We gathered these data for a mul- 
tivariate study of geographic variation in eastern cerions. We 
wished to address the following issues of distinction (at several 
levels) and coherence. 



8 RRFMORA No. 443 

Table 1. Samples used in this stiid\.* 

1. Necker Island, 220391 

2. Anegada. 229017 

3. Anegada, collected C. Paull 

4. Anegada, 203778 

5. Near Tower Guanica Insular Forest, Puerto Rico, 212303 

6. Tamarindo Beach. Puerto Rico, 212302 

7. Guanica Insular Forest, Tower Road, Puerto Rico, 212293 

8. Cabo Rojo Colony No. 2, Puerto Rico, 212298 

9. Pt. Criollo, Guanica, Puerto Rico, 216702 

10. Cayo Maguey, La Paruera, 212292 

11. Cabo Rojo Colony No. 1, 212297 

12. Mona Island paratypes, 171020 

13. Mona Island, 190144 

14. Mona Island, Isabella Anchorage paratypes, 184052 

15. N.W. Saona Is., Hispaniola, 98829 

16. Lighthouse Cabo Engano, Altagracia, Hispaniola, 251314 

17. North Shore Bahia de Yuma, Hispaniola, 251313 

18. Yuma River, paratypes, Hispaniola, 181967 

19. Romana Prov. 0.7 mi. east of Macao, Hispaniola, 250638 

20. N.E. Grande Cayemites Is., Hispaniola, 251 161 

21. Juanilo, Altagracia, Hispaniola, 25132 

22. [ortuga Island ANSP 146701 and 146702 

23. St. Croi.x [composite sample: MCZ. Harvard; ANSP, AMNH (NY.) and Field 
Museum (Chicago)]. 

♦Samples 1-21 (with the exception of sample 3 collected by C. Paull) are from the 
Dept. of Molluscs, Museum of Comparative Zoology (lot number indicated). Sam- 
ple 22 is from the Academy of Natural Sciences, Philadelphia. Conventional iden- 
tifications follow: samples 1-1 1, C striatellum; 12-14, C. monaense; 15. C. vumaense 
saona: 16-19, and 21, C vumaense: 20, C. feniif^inuni: 22, C. lortu^a: 23, C rude. 



1. Distinction of eastern cerions from other sections of the 
genus: Are evident differences in morphology matched by pat- 
terns of covariation not encountered in other cerions? 

2. Distinction among islands: Do the morphologies of eastern 
cerions record unambiguously the island of their occurrence? If 
so, we may identify isolation by geography as the primary corre- 
late of morphological variation in these animals. 

3. Distinction among samples: A long tradition m Cerion studies 
(amply affirmed by all our qualitative observations) holds that 
nearly every local population has developed its own recognizable 
morphology. (The older conchologists chose to recognize this pri- 



1977 EASTERN CERIONS 9 

mary observation by granting each form its own specific name.) 
Does such local differentiation exist among the eastern cerions? 

4. Coherence: If cerions are distinct by island (issue 2 above), 
can we detect any interisland pattern (clinal or otherwise) that 
might suggest a common control correlated with geography (selec- 
tion along climatic gradients, gene flow, or isolation by distance, 
for example)? The discovery of such a pattern will not permit us 
to distinguish among these potential explanations, for a pattern of 
coherence does not specify its cause, and museum specimens will 
not settle the issue. If we find no su'ch pattern, current taxonomy 
granting autonomy to each major island must be maintained. If 
overall patterns exist, some consolidation may be in order, since 
hypotheses of direct interaction or similar responses of a single 
system to common factors would gain strong support from such 
patterns. 

The subsequent definition of our measures follows Gould et al. 
(1974. pp. 522-524) except for 6 and 1 1. Cerion is a biometrician's 
delight for three reasons: 

1) It preserves its entire ontogeny on fully exposed whorls. 

2) It ceases growth with a definitive adult aperture. Thus we 
measure patterns of covariation among truly standardized adults. 

3) We can specify comparable stages of ontogeny among speci- 
mens because the boundary between protoconch and accretionary 
shell provides an unambiguous, biological criterion for numbering 
whorls. Our characters can record both terminal sizes (for reason 
2 above) and standardized traits of intermediate growth stages. 

Our measures include: 

1. Width of protoconch. 

2. Width at the end of the fourth whorl. 

3. Total number of whorls of the adult shell (with the termina- 
tion of the protoconch taken as the 0th whorl). 

4. Number of ribs on the fourth whorl. 

5. Number of ribs on the sixth whorl. 

6. Number of ribs in 40 micrometer units (2.22 mm) at the end 
of the first whorl. 

7. Length of the adult shell, apex to lower apertural tip. 

8. Maximum width of the adult shell. 

9. Height of the protoconch. 

10. Total height of the shell at the end of the fourth whorl. 



10 BKf NIOKA No. 445 

I I. Height trom the end (^t uhorl 4 to the end o\ whorl 6. 

12. Width of the umbilicus. 

13. Width of the apertural lip at its widest point (measured 
parallel to the plane of the aperture). 

14. Thickness of the apertural lip at its thickest point (measured 
perpendicular to the plane of the aperture). 

15. Height of the aperture. 

16. Width of the aperture. 

17. Protrusion of the aperture beyond the junction of aperture 
and terminal whorl (see fig. 5 of Gould et al., 1974) — a 
measure of both adult size and intensity in change of adult 
coiling. 

18. Tilt of the aperture — a "pure" (ratio) measure of change 
in coiling to produce the adult aperture. 

19. Weight of the shell. 

IV. DISTINCTION OF EASTERN FORMS FROM OTHER 
CERIONS: PATTERNS OF COVARIATION 

The distinction of eastern cerions on traditional criteria of static 
adult morphology has long been recognized (Pilsbry, 1902). We 
have been experimenting with the promising, but rarely-used cri- 
teria of "dynamic morphology," or differences in the structure of 
covariation among variables (Gould, 1969; Gould et al., 1974). We 
have, so far, studied in detail only the C. bendaJIi Pilsbry and Va- 
natta group of Abaco cerions (Gould et al., 1974). We wish to re- 
port some interesting differences between these and the eastern 
cerions. 

For seven samples of C. hendalli from Abaco we studied factor 
loadings on oblique axes (following varimax rotation) in separate 
R-mode analyses of each sample. We found that five axis solutions 
(recovering 70 to 75 per cent of the total information) yielded con- 
sistent and sensible interpretations with sufficient reduction of di- 
mensions to form interesting clusters. We grouped each variable 
with the axis of its highest loading. In all seven samples, we ob- 
tained the same five groups, each defined by the unvarying asso- 
ciation of two measures (see previous list of measures), and the 
more variable inclusion of others: 

1. Adult size with whorl number (3) and shell height (7) as its 
focus. 



1977 FASIFRN CFRIONS II 

2. Ribbing, with ribs on the fourth (4) and ribs on the sixth (5) 
whorl. This group is by far the most distinct among Abaco shells. 
It always occupies the second or third axis and never includes other 
measures (except, in two cases, the anomalous ratio measure of 
apertural tilt, where there can be no consistency, for measure 18 
loads once positively, once negatively, and both times weakly). 

3. Early whorl heights, with protoconch height (9) and height 
at the fourth whorl (10). 

4. Later whorl sizes, with height of the middle whorls (11) and 
width at the fourth whorl (2). 

5. Apertural lip, with lip width (13) and thickness (14). 

We performed similar R-mode oblique factor analyses for 21 
eastern samples with sufficient specimens, using the DUVAP pro- 
gram (with provision for missing data) as in Gould et al., 1974. 
Five axes encompass 71 to 90 per cent of all information in these 
samples. No new groupings appear with any consistency or fre- 
quency. Associations three through five occur with less regularity 
but similar composition (in 14, 10, and 11 samples respectively). 
We note two outstanding differences between eastern and Abaco 
cerions: 

1. In Abaco samples, whorl number and shell height are invari- 
ably associated as the focus of a group representing adult size. We 
find this association in only 9 samples of eastern cerions. In 16 
samples (including some of these 9), whorl number appears in neg- 
ative association with one or more measures of shell size at stand- 
ardized early whorls (measures 1, 2, 9, 10, and 1 1). Such a negative 
association appears in no sample of Abaco C. bendalli. We assume 
that this disparity reflects a difference in the control exerted via 
growth over final adult size. A shell may reach large size either by 
growing more whorls or by growing larger whorls. If final size is 
strictly controlled, we anticipate a negative interaction of whorl 
number and whorl size — as in the eastern cerions (and in our one 
sample of a second Abaconian species, Gould et al., 1974, p. 528). 
In a jigsaw puzzle with rigid borders, increase in the size of one 
piece must be matched by decreasing size in others. But if size is 
not so strictly controlled, and if all factors that increase it work in 
concert (or at least independently), then we expect no negative 
interaction — as in the seven samples of Abaconian C. bendalli. 
If the borders of the jigsaw puzzle can expand, then all pieces 
might increase together. 



12 «RF\ lORA No. 445 

We do not know which of our characters best measures the 
elusive property of overall "size," but let us consider shell weight 
as a possible surrogate: For all nine samples of Abaconian C 
hendalli (only seven were factor analyzed), the mean coefficient of 
variation for shell weight is 21.00 (range from 15.51 to 25.85). 
Twenty-one eastern samples yield a mean C.V. of 16.82 (range 
from 9.9 to 30.2, with only three samples above 20.0). This differ- 
ence is significant at the 5 per cent level (t = 2.33 at 28 d.f.). Adult 
weight within samples is less variable for eastern than for Abaco 
cerions. Final size may be more strictly controlled in the eastern 
cerions. 

C.V.'s for weight exhibit an interesting heterogeneity among 
eastern cerions. Hispaniolan samples have a mean C.V. of only 
12.41 (range from 9.9 to 14.3), while all other samples average 
19.02 (range from 15.0 to 30.2, no overlap with Hispaniola). The 
lower value for Hispaniola is correlated with stronger negative in- 
teraction between whorl number and whorl size. In factor analyses 
of all seven Hispaniolan samples, for example, whorl number (3) 
always sorts in negative association with shell height from the 
fourth through sixth whorls (11). We find this negative interac- 
tion in only 5 of the remaining 14 groups. 

2. In Abaco Cerion. the ribbing measures always form a tight 
cluster, completely distinct from all other variables. Their asso- 
ciation in eastern cerions is equally strong, but their group often 
includes other measures, almost invariably of adult size. In 18 of 
21 samples, other measures group with rib numbers. Shell width 
(8) and umbilical width (12) join in four samples, width at the 
fourth whorl (2) and lip width (13) in three (but never lip thick- 
ness), apertural width (16) and apertural height (15) in two. This 
discovery supports our intuition that the "ribs" of eastern cerions 
— with their even spacing, complete coverage of the shell, and 
regularity of form — are not controlled in the same manner as the 
stronger, more irregular, and more widely spaced ribs of most other 
cerions. The more common pattern of other species may record 
environmental fluctuations and the pauses in growth that accom- 
modate to them. Perhaps the ribbing in eastern cerions reflects a 
more regular, internal metabolic pattern correlated with general 
growth (and final size) of the animal. 



1977 EASTERN CERIONS 13 

We gain some support for this speculation from coefficients of 
variation. In Abaco C. hendalli, mean C.V. for ribs on the fourth 
whorl is 14.75 (range from 12.27 to 20.94), and for ribs on the sixth 
whorl, 14.83 (range from 11.93 to 17.00). Comparable figures for 
the same measures in eastern cerions are 8.57 (range from 3.67 to 
13.59) and 8.68 (range from 3.42 to 1 1.27). This distinction cannot 
be explained by differences in absolute numbers of ribs, for the 
ranges are comparable in both groups (mean rib number varies 
from 33 to 80 in Abaco samples and from 25 to 78 in eastern sam- 
ples). 

V. DISTINCTIONS AMONG ISLANDS FOR 
EASTERN CERIONS 

The traditional taxonomy of eastern cerions records primarily a 
distinction among islands (though we note with some surprise that 
no one ever split the Virgin Island from the Puerto Rican cerions). 
We have already reported an interesting difference in variability 
of shell weight between Hispaniola and other areas. Differences 
in variation and covariation should be pursued more vigorously 
(if only because they are so widely neglected); but for now we 
proceed along traditional lines and examine differences in form. 

We performed a canonical analysis on all 23 samples (373 speci- 
mens) using D DA, a program written by John Rhoads, Depart- 
ment of Anthropology, Yale University (see Gould et al., 1974). 
We treated each sample separately, without any prior identification 
of its island. Table 2 records the order of discriminating power for 
variables expressed in univariate ANOVA. We find a perfect cor- 
respondence between this order and the traditional criteria of tax- 
onomic distinction: strength of ribbing (variables 4, 5), width of 
the shell (8, 13, and 16), and obtusity of the apex (10). (Shells 
bearing obtuse apices are high at the end of the fourth whorl; 
whorls enlarge quickly and begin their rapid translation down the 
axis of coiling early in ontogeny. Triangular apices are lower at 
the end of the fourth whorl: the "triangle" records a slower increase 
in v\ horl size and a later incidence of the strong allometry that gives 
Cerion its characteristic shape by increasing height relative to width 
during post-juvenile to pre-adult growth.) The matrix of means for 
islands also records the rough order of discrimination. (Table 3 — 



14 HRFVIORA No. 445 

Table 2. Univariate ANOVA: discriminatory power ot individual variables for 
all 23 samples F-ratios with 22 and 350 degrees of freedom. All are 
significant. 

Name Number F-ratio 

4th ribs 4 208.5 

6th ribs 5 157.0 

aperture width 16 102.9 

width 8 102.9 

4th height 10 101.8 

lip width 13 67.2 

aperture height 15 54.1 

height 7 53.0 

1st ribs 6 52.4 

4-6th height 11 52.1 

umbilicus 12 45.2 

4th width 2 40.4 

whorls 3 33.6 

lip thickness 14 25.2 

protrusion 17 22.7 

protoconch width 1 19.5 

protoconch height 9 18.8 

tilt 18 3.3 



not the mean of sample means, but the mean of ungrouped shells 
for each island.) 

The first three canonical axes account for 87 per cent of the vari- 
ance among sample means (45.1, 29.6 and 12.3 per cent respec- 
tively; the fourth axis falls below 5 per cent). Figure 3 displays 
the projection of sample centroids upon the first two axes. Sam- 
ples clearly group by islands; all islands are widely separated with 
the exception of Hispaniola and Mona. The introduction of axis 
three completes the pattern; Mona and Hispaniola are now distinct 
(Fig. 4). We need only three components of variation to sort 23 
samples unambiguously into five larger groups. The strength of 
determination by island is well recorded in the classification matrix: 
309 of 373 specimens are correctly classified, and every misclassi- 
fied specimen sorts with another sample of its island. If we redo 
the analysis with islands rather than samples, as groups, not a 
single specimen is misclassified. Each lies closest to the centroid 
of its own island. Geographic isolation seems to be the primary 
correlate of morphological variation in eastern cerions. 



1977 



EASTFRN CF RIONS 



15 



Table 3. Matrix of mean values tor islands. Weight in grams. Linear measures 
in mm. 











Puerto 




His- 


St. 




Number of 


Necker 


Anegada 


Rico 


Mona 


paniola 


Croi.x 


















specimens 


6 


59 


140 


40 


110 


19 


1. 


protoconch 
















width 


3.35 


3.01 


3.12 


3.41 


2.96 


3.52 


2. 


4th whorl 
















width 


9.48 


8.70 


8.96 


9.48 


8.18 


10.44 


3. 


total whorls 


7.58 


6.95 


7.26 


6.42 


6.80 


8.41 


4. 


4th ribs 


74.33 


68.78 


56.67 


30.47 


33.05 


36.47 


5. 


6th ribs 


60.50 


53.78 


51.09 


26.02 


29.70 


34.68 


6. 


1st ribs 


13.92 


16.97 


15.85 


11.94 


9.23 


12.03 


7. 


height 


24.70 


20.03 


22.29 


22.32 


20.17 


28.60 


8. 


width 


10.82 


9.34 


10.25 


9.93 


8.85 


12.40 


9. 


protoconch 
















height 


1.96 


1.96 


2.08 


2.64 


2.53 


2.71 


10. 


4th height 


6.63 


6.48 


4.10 


5.32 


4.51 


4.61 


II. 


4th-6th height 


7.56 


7.01 


5.19 


7.34 


5.72 


4.69 


12. 


umbilical 
















width 


4.19 


4.05 


4.16 


4.33 


3.83 


6.23 


13. 


lip width 


.66 


.48 


1.02 


1.02 


.63 


1.22 


14. 


lip thickness 


.99 


.55 


.92 


1.08 


.59 


.87 


15. 


aperture 
















height 


9.58 


7.89 


8.99 


8.59 


7.82 


10,59 


16. 


aperture 
















width 


7.63 


6.59 


8.10 


7.23 


6.40 


9.87 


17. 


protrusion 


2.45 


2.01 


2.78 


2.21 


2.04 


3.35 


18. 


tilt 


2.10 


1.95 


2.07 


1.85 


2.12 


2.08 


19. 


weight 


1.20 


.55 


.77 


.80 


.56 





The matrix of factor pattern for the analysis by samples (Ta- 
ble 4) defines the uniqueness of each island and exhibits the mor- 
phological bases of separations in Figures 3 and 4. (The factor 
pattern matrix of standardized partial regression coefficients per- 
mits us to discern how the original measures vary together in dis- 
tinguishing samples along the canonical axes.) We did not include 
shell weight (variable 19) in this analysis because we could not 
weigh the St. Croix fossils with their permineralized shells and 
filled interiors. 



16 



BRFVIORA 



No. 445 







-6 




2 4 

AXIS I 




Fig. 3. Plots of centroids for all samples on first two axes of a canonical analy- 
sis (74.7 per cent of all information) to show distinction of islands and coherence 
within an island. Islands are circumscribed by minimum convex polygons. A- 
Anegada, N-Necker, P-Puerto Rico. M-Mona. S-St. Croix, and H-Hispaniola. 
U(-Cen'on tortuga (locality 23), Hf-C ywnaense ferruginum (locality 20). H.-C. 
yumaense saona (locality 15). 




J L. 



2 4 

AXIS I 




12 



•n 



.S -6^ 

Fig. 4. Axis 1 vs. axis 3 for the same canonical anahsis depicted in Figure 3. 
Code letters same as in Figure 3. Note that Mona and Hispaniola (overlapping in 
Fig. 3) are distinguished by the third axis. 



1977 



EASTERN CERIONS 



17 



Table 4. Factor pattern (structure of covariance) for original variables on first 
fi\e canonical axes for all 23 samples. 



No. 


Name 


1 


2 


3 


4 


5 


1 


protoconch width 


-0.0430 


0.1082 


-0.3254 


0.2116 


-0.0632 


2 


4th width 


-0.0239 


0.2443 


-0.4745 


0.1357 


-0.1085 


3 


whorls 


-0.0047 


0.2851 


-0.1552 


0.4931 


-0.1116 


4 


4th ribs 


0.6196 


0.4860 


0.2486 


-0.0291 


-0.1228 


5 


6th ribs ' 


0.4526 


0.5397 


0.2748 


-0.0553 


-0.2221 


6 


1st ribs 


0.2427 


0.3124 


0.0459 


0.2449 


0.4016 


7 


height 


-0.0968 


0.3027 


-0.4599 


-0.1611 


-0.4270 


8 


width 


-0.0607 


0,5167 


-0.5713 


-0.1082 


-0.2853 


9 


protoconch height 


-0.1590 


-0.1151 


-0.1863 


-0.0339 


-0.0540 


10 


4th height 


0.3893 


-0.2690 


-0.4891 


0.1371 


0.1004 


11 


4-6th height 


0.1832 


-0.2792 


-0.2773 


0.4856 


-0.2967 


12 


umbilicus 


-0.0624 


0.2278 


-0.4813 


-0.3020 


0.2144 


13 


lip width 


-0.2000 


0.4035 


-0.2115 


0.3895 


0.1607 


14 


lip thickness 


-0.0726 


0.2054 


-0.1453 


0.4387 


-0.1463 


15 


aperture height 


-0.0788 


0.3720 


-0.3568 


-0.0482 


-0.4348 


16 


aperture width 


0.1126 


0.5692 


-0.3769 


-0.0336 


-0.1512 


17 


protrusion 


-0.0602 


0.2759 


-0.0882 


-0.0274 


-0.0037 


18 


tilt 


-0.0180 


0.0311 


0.0417 


-0.1369 


-0.1053 



The first axis separates Virgin Island samples from all others (see 
Figs. 3, 4). Its factor pattern records high values for measures of 
ribbing and whorl heights (10 and 11). Indeed, the Virgin Island 
samples exceed all others in their number of ribs and the obtuseness 
of their apices (Fig. 2). The negative loading of protoconch height 
(9) also reflects the obtuse apex of Virgin Island cerions (see mean 
values in Table 3). High protoconchs are followed by slowly ex- 
panding whorls that join the embryonic shell to produce a pointed 
apex with modest whorl heights; low protoconchs precede rapid 
expansion (high values of 10-11) and build more obtuse apices. 

The second axis separates Puerto Rican samples (positive pro- 
jections) from Mona Island and Hispaniola (negative projections). 
The factor pattern contains high positive loadings for two groups 
of variables: measures of ribbing (4 and 5) and shell widths, par- 
ticularly aperture width (8 and 16). Whorl heights (10 and 11) 
load negatively. The cerions of Puerto Rico can be distinguished 
visually by their more circular apertures; this feature translates to 
high values for aperture width in our quantitative analysis. The 
St. Croix specimens (which also project strongly and positively on 



18 HKJ MORA No. 445 

axis two) are wide as a simple function of their large size. The 
ribbing measures set the major contrast between Puerto Rican 
samples and Mona and Hispaniolan samples. Puerto Rican cerions 
are abundantly ribbed (though not so abundantly as Virgin Island 
samples), while Mona and Hispaniolan cerions have the fewest 
(though strongest) ribs among eastern cerions. Puerto Rican spe- 
cimens also have fairly triangular apices. This feature translates 
as low whorl height (10 and 1 1) in our measures; the negative load- 
ing of measures 10 and 11 assures the high positive projection of 
Puerto Rican samples on the second axis. 

The third axis separates Mona and Hispaniola to complete the 
distinction among islands. Mona and Hispaniolan samples fell 
together with negative projections on the first two axes for essen- 
tially the same reason — their small number of ribs. Several meas- 
ures of size — particularly of widths (2, 8, 12, and 16 — load 
strongly and negatively upon the third axis. Hispaniolan shells 
are small and especially narrow (note low mean values of 2 and 8 
in Table 3); they project positively upon an axis with negative 
loadings for these measures. Mona samples are large and wide; 
since measures for size and width have negative loadings, these 
samples project negatively upon the axis. 

Subsequent axes continue the patterns of distinction by island. 
The fourth axis, for example, separates Mona, having high pro- 
jections, from St. Croix, having low projections. Highest loadings 
are for height from whorl 4 through 6 (positive) and number of 
whorls (negative). St. Croix specimens with their maximal num- 
ber of whorls and minimal fourth to sixth whorl heights project 
negatively. Mona samples with their distinctively low number of 
whorls project positively. The fifth axis separates Virgin Island 
samples into their two areas of Necker and Anegada. The Necker 
Island sample projects strongly and negatively upon an axis with 
positive loadings for its low number of ribs on the first whorl (rela- 
tive to Anegada), and high negative loadings for various measures 
of shell size. 

VI. DISTINCTION AMONG SAMPLES 

A persistent tradition of Cerion studies holds that virtually every 
local population bears its own distinctive morphology. This re- 
markable phenomenon has inspired the burden of an exuberant 



1977 FASrERN CFRIONS 19 

specific nomenclature for these animals. Nothing we have seen in 
several years of study contradicts this conventional observation. 
We therefore predicted that samples within islands should be dis- 
tinct in separate canonical analyses for each island. Our results 
confirm this prediction with great accuracy. The classification ma- 
trix for four Virgin Island samples correctly assigns 63 of 65 spe- 
cimens to their correct sample (96.9 per cent). Only 3 of 40 speci- 
mens from Mona Island lie closest to the centroid of a sample not 
their own (92.5 per cent are correctly classified). Of specimens 
from Hispaniola, 101 of 110 are correctly allocated among eight 
samples (91.8 per cent). Only Puerto Rico falls below 90 per cent 
success — 1 16 of 139 specimens in seven samples (83.5 per cent). 

We have, in summary, established a hierarchy of distinction 
among the eastern cerions. Their basic design is strikingly different 
— both in form and in pattern of covariance — from other ceri- 
ons. Islands are distinct. Samples within islands can also be sep- 
arated with little overlap. 

VII. A UNIFYING THEME AND A TAXONOMIC PROPOSAL 

The distinction of islands does not preclude the existence of pat- 
tern among them. We concluded that geographic separation serves 
as the primary correlate of morphological distinction among the 
eastern cerions. But does the degree of morphological separation 
reflect geographic distance? If so, we might expect a clinal distri- 
bution of morphology, since the islands form an essentially linear 
array. If we fmd no coherent pattern, taxonomic distinction by 
island should be maintained (with, God forbid, a new taxon for 
the Virgin Islands). 

The output of our canonical analysis for islands includes a ma- 
trix of Mahalanobis distances (D^) — a measure of separation be- 
tween centroids with adjustments for variance and covariance (Ta- 
ble 5). Each entry is a measure of morphological distinctness 
(based on all variables) between shells of two islands. If we plot 
D- (see Sneath and Sokal, 1973, pp. 127, 128) against geographic 
separation (defined, for any island pair, as the distance between 
our two closest sample localities), we obtain a remarkably tight 
and linear relationship (r = .96). This calculation (Fig. 5) excludes 
the St. Croix fossils as representatives of another time with no 
expected relationship to current trends. When we plot the St. Croix 



20 



BREVIORA 



No. 445 



Table 5. Matrix of Mahalanobis distances (D-) for analysis using all specimens 
grouped b\' islands. 



Necker 











Anegada 


18.87 









Puerto Rico 


125.33 


102.90 







Mona 


155.33 


134.73 


54.66 





Hispaniola 


164.32 


131.32 


38.00 


31.23 


St. Croix 


214.23 


187.83 


65.86 


65.29 



57. 









X 


NS 












200 










xas 








NHo^^ 


r>, 150 


- 














NM 


o ^/^ 


Q 
















^ 


-^M 


CO 
O 














oNP ^^ 




° °AH 


<ioo 


- 












^°AP 






_J 
< 

X 

< 

50 

n 


X 


'^NA 




^ 


°PH 


^PS 


^MS 




Xhs 

1 1 



100 200 300 400 500 

DISTANCE IN KM 



Fig. 5. Mahalanobis distance (morphological) vs. geographic separation for 
eastern cerions. Note the tight correlation and the departure of St. Croix fossils 
from the trend. A-Anegada, N-Necker, P-Puerto Rico, M-Mona. H-Hispaniola, 
and S-St. Croix. 



1977 EASTERN CERIONS 21 

S H M P AN 

jV V Vi V I I I I I \|/ \1/ I 

-4 -2 2 4 6 8 10 

Fig. 6. Projections on the first canonical axis for a discriminant analysis by 
islands. This axis explains more than half (58.8 per cent) of all information. Note 
the perfect correspondence of morphological gradation and geographic position for 
the modern samples. Only the St. Croix fossils depart from the trend. Code letters 
as in Figure 5. 

distances on Figure 5. they depart markedly from the regression 
Une of modern samples. Geographic and morphological distance 
are closely correlated in living eastern cerions. But this relation- 
ship does not guarantee any graded trend of morphology along the 
linear array of islands. In fact, we already have a test for clines in 
the results of our canonical analysis. 

In our canonical analysis for islands, the most significant dis- 
criminator (the first axis) explains 58.8 per cent of all information. 
Figure 6 shows the array of projections. Living cerions are arranged 
in perfect geographic order; only the fossils of St. Croix depart 
from the pattern. The morphological gradation runs from egg- 
shaped, finely and copiously ribbed shells with very obtuse apices 
(Virgin Islands) to more cylindrical, apically pointed shells with 
fewer, stronger ribs (Hispaniola) — see the discussion of factor 
patterns in Section V. The direction of the cline also meets our 
expectations. Virgin Island shells represent the extreme expression 
of morphological features that characterize eastern cerions; they 
are also geographically furthest from any contact with non-eastern 
congeners. The Hispaniolan specimens, situated in closest prox- 
imity to non-eastern cerions, depart most strongly from modal east- 
ern morphology towards the more "normal" form of the genus. 

We are not addicted to speculative biogeography, and we offer 
no preferred hypothesis for this pattern; we do not see how mu- 
seum specimens can distinguish among such alternatives as selective 
gradients and gene flow, and we know virtually nothing about the 
history of colonization. We only note that the strikingly discordant 
position of the St. Croix fossils may indicate a fairly recent origin 
for the graded series of modern forms. (The St. Croix fossils are 
morphologically furthest from their nearest modern relatives on 
Necker and Anegada.) 



22 HRI \ loRA No. 445 

In the light of this pattern, we see no reason to maintain the 
taxonomic distinction among Hving eastern forms. Some common 
factor accounts for most of the morphological variabiity among 
these populations. We therefore consider all of the following names 
as synonyms of the first-named form, Cerlon striatellum ("Ferru- 
sac" Guerin-Meneville), 1829: C. nionaen.se Clench, 1951; C. vu- 
niaen.se Pilsbry and Vanatta, 1895 (with its four subspecies); C. 
ferruginiini (Maynard, 1896 — ranked as a subspecies of C. yu- 
niaen.se by most later authors); and C. tortuga Pilsbry and Vanatta, 
1928. We maintain, for now, Cerlon rude (Pfeiffer). 1855 for the 
St. Croi.x fossils. The St. Croix shells depart completely from 
modern trends in geographic variation. Indeed, this departure 
serves as a strong support for taxonomic identity of recent popu- 
lations — for it argues against any appreciable antiquity for the 
modern pattern. Moreover, the St. Croix shells, while closer to 
the eastern mode than to any other morphotype of Cerion, vary 
from this mode in the direction of more conventional Bahamian 
and Cuban forms — shells are larger, more cylindrical, and more 
sparsely and strongly ribbed. As another argument for the tax- 
onomic unity of modern forms, we note that the morphological 
distances among islands are trifling compared with the distinction 
of several forms that interbreed freely throughout the Bahamas. 
In fact, and incredible as it may seem in the light of Cerion s fan- 
tastic morphological variety, we have located no unambiguous case 
of sympatry without interbreeding for the entire genus (two of nine 
reported cases have a fair chance of validation in our opinion). 

VIII. CONCLUSION 

Studies of Cerion have been dominated by the conviction that 
no regular patterns of geographic variation exist. All leading stu- 
dents have invoked haphazard transport by hurricanes to support 
a notion of "crazy-quilt" distributions (Mayr and Rosen, 1956). 
We now present the first evidence for simple, graded, coherent 
variation over a large part of Cerion's range. Moreover, S. J. 
Gould and D. S. Woodruff, working in the northern Bahamas 
during the past four years, have found strikingly consistent pat- 
terns of within-island variation on all islands visited (Abaco, 
Grand Bahama, New Providence, Great Exuma, and Long). The 
"ribby" morphotype, for example (called C. glans (KUster) and 



1977 EASIFRN C FKIONS 23 

about 50 synonyms on New Providence, C. ahacoense Pilsbry and 
Vanatta and 3 synonyms on Abaco and Grand Bahama, etc.), al- 
ways inhabits coastal areas at the edges of the Pleistocene banks; 
while the mottled morphotype inhabits interior coasts (with respect 
to the banks) and interior areas. The two morphotypes intergrade 
in a variety of interesting ways at their contacts. The discovery of 
simple and coherent patterns, both within and between islands, 
leads to a hope even an expectation — that some sense will yet 
be made of Cerion. 

IX. ACKNOWLEDGMENT 

This work was supported by NSF Grant BMS 72-02213. 



References 

Baker, H. B., 1924. Land and freshwater molluscs of the Dutch Leeward Islands. 

Occ. Pap. Mus. Zool. Univ. Mich. No. 152, 158 pp. 
Bartsch, p., 1920. Experiments in the breeding of Cerions. Papers Dept. Marine 

Biol. Carnegie Inst. Wash. No. 282, 55 pp. 
Clench, W. J., 1951. Land shells of Mona Island, Puerto Rico. J. Conchyliologie 

90: 269-276. 
1957. A catalogue of the Cerionidae(Mollusca — Pulmonata). Bull. Mus. 

Comp. Zool. 116: 121 169. 
Dall, W. H., 1905. Fossils of the Bahama Islands, with a list of the nonmarine 

moUusks. //; G. B. Shuttuck (ed.). The Bahama Islands. N. Y., MacMillan, 

pp. 23-47. 
DE Vries. W.. 1974. Caribbean land molluscs; notes on Cerionidae. Stud. Fauna 

Cura(jao and other Carib. Islands 45: 81-117. 
Goci [), S. J., 1969. Character variation in two land snails from the Dutch Lee- 
ward Islands: geography, environment, and evolution. Syst. Zool. 18: 185-200. 
GoDi D, S. J., D. S. Woodruff, and J. P. Martin, 1974. Genetics and morpho- 
metries of Cerion at Pongo Carpet: a new systematic approach to this enigmatic 

land snail. Syst. Zool. 23: 518-535. 
Gifrin-Menevii 1 E, F. E., 1829. Iconographie du regne animal de G. Cuvier, 

Mollusques. Paris. Fortin. 
Hi \1\1E1 INCK, P. W.. 1940. Mollusks of the genera Cericni and Tiulora. Stud. 

Fauna Curasao, Aruba. Bonaire, and Venezuelan Islands 2: 43-82. 
Jacobson, M. K.. 1968. The land Mollusca of St. Croix, Virgin Islands. Sterkiana 

32: 18-28. 
Ki'STFR, H. D.. 1847. Die Gattungen Pupa. .\fe,i;a\pira. Balca und Tonuiit'llma. 

Conchylien-Cabinet (2). 1. pt. 15. 194 pp. 
Maynari). C. .!.. 1896. Monograph of the genus Strophia. Contributions to 

Science, vol. 3. NewtonviUe. Mass. 



24 BREVIORA No. 445 

Mayr, E., 1963. Animal species and evolution. Cambridge, Mass., Harvard Univ. 

Press, 797 pp. 
Mayr, E., and C. B. Rosen, 1956. Geographic variation and hybridization in 

populations of Bahama snails (Cerion). Novitates Am. Mus. Nat. Hist. No. 

1806,48 pp. 
Pfeiffer, L., 1855. Beitrage zur Molluskenfauna Westindiens. Malak. Blat. 2: 

98-106. 
PiLSBRY, H. A., 1902. Family Cerionidae. Manual of Conchology, Second Series, 

Pulmonata, Vol. 14. Philadelphia, Acad. Nat. Sci., pp. 174-281. 

1943. Note on Cerion striatelluni. Nautilus 57: 34-35. 

PiLSBRY, H. A. AND E. G. Vanatta, 1895. New species of the genus Cerion. Proc. 

Acad. Nat. Sci. Phila. 47: 206-210. 
1896. Catalogue of the species of Cerion, with descriptions of new forms. 

Proc. Acad. Nat. Sci. Phila. 48: 315-338. 
1928. Land shells of Tortuga Island, Haiti, and a new Haitian Oleacina. 



Proc. Acad, Nat. Sci. Phila. 80: 475-478. 
PtAfE, L., 1906. Die Artbildung bei den Cerion-Landschnecken der Bahamas. 

Verhandl. deutsch. zool. Ges. 16: 127-136. 
1907. Die Variabilitat und die Artbildung nach dem Prinzip geographi- 

scher Formenketten bei den Cerion-Landschnecken der Bahama-Inseln. Arch. 

Rassen Gesellsch. Biol. 4: 433-470. 581-614. 
Sneath, p. H. a., and R. R. Sokai, 1973. Numerical Taxonomy. San Francisco, 

W. H. Freeman, 573 pp. 
SowERBY, G. B., 1875. Monograph of the genus Pupa. Conchologia Iconica, 

vol. 20. 
Vanatta, E. G., 1923. Two new Santo Domingan land shells. Proc. Acad. Nat. 

Sci. Phila. 75: 359-360. 



''^^L.'S. CCMP. ZOOL 

B R E V I "ax A 

Museum of Comparative Zoology 

us ISSN 0006-9698 

Cambridge, Mas s. 20 December 1978 Number 446~ 

7 ^ - . ~ 

A NEW INTERPRETATION OF THE MAMMALIAN TEETH 

OF TRIBOSPHENIC PATTERN FROM THE 

ALBIAN OF TEXAS 

Percy M. Butler' 

Abstract. A reappraisal has been made of tribosphenic teeth from the Albian of 
Texas, previously described by Patterson, Slaughter and Turnbull. At least six 
genera are represented. Pappotherium and Holoclemensia, the most common gen- 
era; are sufficiently alike to be placed in the same family, Pappotheriidae. They 
have four molars as in marsupials and complex premolars as in placentals. They 
cannot be classified either as marsupials or placentals but are considered to belong 
to a separate evolutionary line (Pappotherida, new order). Two forms of smaller 
size, Kermackia texana and Trinititherium slaughter! gen. et sp. nov., whose lower 
molars are at a more primitive stage of evolution than the Pappotheriidae, are 
placed in a new family Kermackiidae (Aegialodontia, new order). Slaughteria erup- 
lens gen. et sp. nov., the only species for which there is evidence of tooth-replace- 
ment, is placed provisionally in the same family. A new infraclass, Tribotheria, is 
proposed to include mammals with tribosphenic molars that are not classifiable as 
marsupials or placentals. 

INTRODUCTION 

Patterson (1956) reported on a collection of teeth found in the 
Greenwood Canyon, near Forestburg in eastern Texas. Many of 
the molar teeth are of tribosphenic pattern. They are mid-Creta- 
ceous (Albian), the oldest tribosphenic molars known at that time. 
The Greenwood Canyon fauna is intermediate in age between the 
late Jurassic Purbeck and Morrison faunas and the late Cretaceous 
Djadokhta fauna of Mongolia. During this long interval of about 
50 million years, the Jurassic mammal orders were largely replaced 



'Zoology Department, Royal Holloway College, Englefield Green, Surrey TW20 
9TY England. 



2 BREVIORA No. 446 

by marsupials and placentals. Only the multituberculates survived 
into the late Cretaceous in more than smaU numbers. 

The differentiation between marsupials and placentals is believed 
to have taken place during the earlier part of the Cretaceous. Both 
marsupials and placentals share the tribosphenic molar pattern. 
With only the evidence of isolated teeth, Patterson was unable to 
decide in which infraclass his specimens belonged or whether they 
represented a group from which both marsupials and placentals 
would arise. Accordingly, he classified them as "Therian mammals 
of uncertain infraclass affinities but of Metatherian-Eutherian 
grade." 

Between 1965 and 1971, Slaughter published descriptions of im- 
portant additional material. He obtained it from Butler Farm, a 
locality of similar age to the Greenwood Canyon locality. He 
named two species based on upper molars: Pappotherium patter- 
soni and Holoclemensia texana. The holotype of P. pattersoni is 
a fragment containing the last two molars. The holotype of H. 
texana is a penultimate molar of which the protocone shelf is lack- 
ing; Slaughter used a complete last molar as a paratype and re- 
ferred a lower molar to this species. He named a third species, 
Kermackia texana from another isolated lower molar. 

Holoclemensia differs from Pappotherium in the arrangement of 
the buccal stylar cusps of its upper molars (Fig. 1). In Pappother- 
ium there is a large stylocone (Style B), about as high as the meta- 
cone. In Holoclemensia, the stylocone is much smaller, but there 
is a large conical cusp (Style C) opposite the notch between the 
paracone and the metacone. In the incomplete material at his dis- 
posal Patterson (1956) mistook Style C for the stylocone. Style C 
occurs in many late Cretaceous marsupials; it is higher than the 
stylocone in Albertatherium and Aquiladelphis (Fox, 1971). 

Slaughter (1968b) regarded Holoclemensia as a marsupial, not 
only because of the presence of Style C, but also because the meta- 
cone is less reduced than in Pappotherium. In 1968a he described 
some partly molariform premolars, unlike those of any marsupial. 
In 1971, he referred to Pappotherium a mandible that contained a 
newly erupted and a partially erupted premolar together with worn 
molars. This specimen indicates diphyodonty of the anterior den- 
tition, characteristic of placentals. He therefore regarded Pappo- 
therium as a placental. Subsequently Fox (1975) deduced from the 



1978 



MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 




Figure 1. P, last two upper molars of Pappotherium pattersoni, SMF-SMU 
61725, and buccal view of penultimate molar. H, the same, Holoclemensia texana, 
SMP-SMU 62099 and 62147. A, buccal view of upper molar of Alphadon marshi. 
Stylar cusp c is indicated by an arrow. 



shapes of the last two teeth that Pappotherium had only three 
molars. 

Lillegraven (1969, 1974) and Hoffstetter (1970, 1972) believe that 
the marsupial-placental differentiation was due to isolation on dif- 
ferent continents. Placentals in Asia and marsupials in North 
America suggest that the placentals that appear in North America 
towards the end of the Cretaceous are immigrants from Asia. This 
opinion does not accord with Slaughter's view that both infraclasses 
were present on the continent as early as the Albian. It seemed 
desirable to reexamine the evidence provided by the Texas Albian 
fauna that affects a crucial stage of mammalian evolution. 



4 BREVIORA No. 446 

During visits to the United States in 1966 and 1974 I studied 
specimens from Greenwood Canyon that Patterson described (1956) 
as well as additional material that Turnbull (1971) described. The 
latter was subsequently collected from the same locality. The 
Greenwood Canyon material is housed in the Field Museum of 
Natural History, Chicago (registration numbers PM — ). Of the 
Butler Farm collection, I saw the specimens described by Slaughter 
(1965, 1968a) and the jaw containing premolars with molars 
(Slaughter, 1971). I studied other Butler Farm specimens from 
casts. The material from Butler Farm is housed in the Shuler 
Museum of Paleontology, Southern Methodist University, Dallas, 
Texas (registration numbers SMP-SMU — ). I used a Wild M5 
microscope to make camera lucida drawings, at known magnifica- 
tions, of all the specimens studied. 

Nearly all the specimens consist of isolated teeth. A primary aim 
of the investigation was to find the most probable way in which 
they could be combined into dentitions. The relationships of the 
species believed to be present in the fauna could then be considered. 

UPPER MOLARS 

All the upper molars that I have seen are drawn in crown view 
in Fig. 2. Except in the holotype of Pappotherium, all are isolated, 
and their position in the series must be judged from their shape 
alone. Last molars, whether M^ or M'', are easily recognizable. 
Reduction of the metacone and loss of the metastyle accompany a 
reduction of the posterobuccal lobe of the outline. These features 
are widespread and almost certainly primitive among mammals 
with tribosphenic molars. On passing forward along the molar 
series, both in marsupials and placentals, the posterobuccal lobe 
becomes relatively larger, so that eventually it projects buccally 
beyond the level of the parastyle; the metacone also enlarges and 
the metastylar crest becomes longer. The degree of difference be- 
tween adjacent molars varies considerably between genera and 
families and to some extent individually. The penultimate molar 
is preserved in the holotypes of Pappotherium and Holoclemensia 
(Fig. 2, d, 1). In both, the posterobuccal lobe is less prominent 
than the parastylar region, though both possess the metastylar crest. 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 5 

When measured by the relative distances of the metacone and par- 
acone from the buccal edge, Holoclemensia has a rather less prom- 
inent posterobuccal lobe, but the difference between the two genera 
is not great. Holoclemensia also has a higher metacone than Pap- 
potherium, a difference that is repeated on the last molar. 

Of the isolated last molars, PM 1075 (Fig. 2, c), though smaller 
than the holotype of Pappothehum, agrees with that specimen in 
structure; PM 1015 (Fig. 2, b) has a less reduced posterobuccal 
lobe and a larger metacone; PM 1287 (Fig. 2, p) has a smaller pro- 
tocone shelf than Pappotherium. The posterior surface of the stylo- 
cone of PM 1015 has a small cusp, indistinctly indicated on PM 
1075. Whether this cusp is the equivalent of Style C is uncertain. 
Style C is comparatively well developed on the last molar of Holo- 
clemensia. but Turnbull (1971, Fig. 1) misidentified it as the meta- 
style. PM 1015 and PM 1075 both differ from Holoclemensia and 
resemble Pappotherium in having a concavity of the anterior out- 
hne lingually to the parastyle. PM 1287, with its small protocone, 
is clearlv distinct from both Holoclemensia and Pappotherium. 

Several upper molars have the posterobuccal region less reduced 
than on the penultimate molars of Holoclemensia and Pappothe- 
rium, and they must be considered as examples of more anterior 
molars. PM 1000 (Fig. 2, o) has a well preserved buccal stylar shelf 
on which Style C is the highest cusp. This identifies the tooth as 
Holoclemensia, but the metacone is farther from the buccal edge 
than the paracone is. PM 886 (Fig. 2, m) has lost the parastylar 
region, but Style C, which is preserved, is as high as the metacone. 
This tooth is larger than PM 1000 and differs from it in that the 
paracone and metacone are nearly equidistant from the buccal 
edge. PM 1004 (Fig. 2, n), poorly preserved, agrees with PM 886 
in comparable parts. There is evidence that Holoclemensia had two 
molars anterior to the penultimate tooth and four molars with all 
included (Fig. 5, H). 

PM 999 (Fig. 2, h) resembles the penultimate molar of Pap- 
potherium in the enlarged stylocone and in the metacone, which 
is less high than the paracone; however, the posterobuccal lobe of 
PM 999 is prominent and bears a strong metastylar crest. PM 1238 
(Fig. 2, i), of which the anterior part of the stylar shelf is missing, 
agrees with PM 999 in the metacone and metastyle and is almost 



BREVIORA 



No. 446 








1 mm 
> < 



Figure 2. Crown views of upper molars, a, SMP-SMU 61725 (holotype of 
Pappotherium pattersoni); b, PM 1015; c, PM 1075; d, SMP-SMU 61725 (holotype 
of P. pattersoni); e, PM 884; f, PM 1749; g, PM 1325? fi, PM 999; i, PM 1238; 
j, SMP-SMU 62402; k, SMP-SMU 62099 (paratype of Holoclemensia texana); 1, 
SMP-SMU 61947 (holotype of H. lexana), m, PM 886; n, PM 1004; o, PM 1000; 
p, PM 1287. All drawn as right teeth: b, c, e, f, i, o, p have been reversed. 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 7 

certainly another example of the same tooth. PM 884 and PM 
1749 (Fig. 2, e, f) represent a different type of molar. The meta- 
stylar region has broken off in both specimens. As in Pappothe- 
rium, the stylocone is high and the metacone is lower than the para- 
cone. These teeth are larger than PM 999 and PM 1238 (Fig. 2, h, 
i), and based on the incomplete crowns, the metacone and para- 
cone were nearly equidistant from the buccal margin. On PM 999, 
PM 884 ajid PM 1749 there is a slight elevation of the marginal 
ridge in the region corresponding to Style C, but this elevation is 
very different from the enlarged Style C of Holoclemensia. 

There are therefore two types of upper molar, with a pattern 
similar to Pappotherium, that correspond to the supposed first two 
molars of Holoclemensia — a larger, more symmetrical M^ and a 
smaller M' with a more prominent metastylar corner. The evidence 
suggests that Pappotherium has four molars like Holoclemensia 
(Fig. 5, P). 

PM 1325 (Fig. 2, g) is the lingual part of an upper molar which 
seems from its size to be more probably referable to Pappotherium 
than to Holoclemensia. 

SMP-SMU 62402 (Fig. 2, j) resembles PM 999 and PM 1238 in 
pattern but differs from them in its smaller size. This tooth may 
be M' of a smaller species, of which PM 1075 might be the last 
molar. 

LOWER MOLARS 

The allocation of lower to upper molars is particularly difficult 
in the case of the therians from the Albian of Texas. This difficulty 
arises because Pappotherium and Holoclemensia have been dis- 
tinguished mainly by the buccal stylar shelf, which does not occlude 
with the lower teeth. The protocone, which does occlude, is missing 
from most upper molars. 

Patterson and Slaughter have distinguished six types of lower 
molars. Types 1, 2, and 3 by Patterson (unworn examples illustrated 
in Fig. 3), Types 4, 5 and 6 by Slaughter. Combinations of some of 
these types seem probable. 

Teeth cf Type 1 are characterized by the anteroposteriorly short 
trigonid, in which the paraconid is lower than the metaconid, and a 
comparatively wide talonid on which the hypoconid is elevated and 
placed nearly in line with the protoconid. In Type 1 Patterson 



8 



BREVIORA 



No. 446 



placed PM 1005 (Fig. 3, H) and the trigonid PM 887; to these may 
be added the trigonid PM 3877. Slaughter (1965) classified three 
teeth from Butler Farm as Type 5 (SMP-SMU 62131, 61727, and 
61735). These differ from Type 1 in the less elevated trigonid, 
which has a greater anterior tilt, and in the narrower talonid. Teeth 






K 





1 mm 



Figure 3. P, H, K, and T, left lower molars drawn in crown, lingual and pos- 
terior views. P, Type 2 (PM 965, reversed), referred to Pappoiheriiim: H, Type 1 
(PM 1005, reversed), referred to Holoclemensia; K, Type 3 (PM 1245, reversed), 
Kermackia: T, Type 6 (SMP-SMU 61728), holotype of Triniiitherium slaughteri. 
S, SMP-SMU 61992, holotype of Slaughieria erupiens. crown and lingual views. 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 9 

of Type 5 were referred by Slaughter (1968b) to Holoclemensia. 
SMP-SMU 61726, belonging to Slaughter's Type 4, has a relatively 
long and narrow talonid with a less elevated hypoconid; it has lost 
the tops of the trigonid cusps. 

The differences between Types 1, 4, and 5 could be due to serial 
position in a single species — Type 4 being the last molar. Type 5 
the penultimate, and Type 1 the most anterior molar of the three. 
A trigonid< PM 966, agrees with Type 1 in the relative height of the 
paraconid and metaconid, but the trigonid is narrower transversely 
and less compressed anteroposteriorly. Probably because of its 
more open trigonid, Patterson included this specimen with Type 2, 
but it could be a first molar standing in the same dentition as Type 

1 and bringing the number of molars to four (Fig. 5, H). 

Teeth of Patterson's Type 2 have a more open trigonid than those 
of Type 1. The metaconid is smaller; it is similar in height to the 
paraconid. The talonid is narrower than in Type 1. The hypoconid 
is comparatively low, and it is not placed so far buccally in relation 
to the protoconid. The talonid is preserved completely only in PM 
965 (Fig. 3, P); in PM 948 the hypoconulid has been broken off. 
PM 930, PM 660, PM 1249, and PM 1119 are isolated trigonids. 
A feature of Type 2 teeth, not noted by Patterson, is the oblique 
crest that continues along up the posterior wall of the trigonid al- 
most to the top; it ends in an intermediate position between the tip 
of the metaconid and the notch dividing that cusp from the proto- 
conid. In teeth of Types 1 and 5, and also in PM 966, the oblique 
crest is confined to the lower part of the trigonid wall. Type 2 teeth 
are smaller than those of Types I, 4 and 5: their trigonid widths 
range from 0.91 to 1.16 mm compared with 1.26 to 1.46 mm in the 
other types (measurements from Turnbull, 1971). No teeth of Type 

2 have been reported from Butler Farm. 

It is probable that Types 1, 4, and 5 are referable to Holocle- 
mensia and Type 2 to Pappotherium. The upper molars of Holo- 
clemensia are larger, and they have larger metacones which might 
correlate with large and buccally situated hypoconids on the lower 
molars. The upper molars referred to Pappotherium are too small 
to fit lower molars of Types 1, 4, and 5, but their occlusion with 
Type 2 lower molars is feasible. PM 930, the Type 2 tooth with 
the smallest and most open trigonid, would be a first molar. The 
largest examples of Type 2, PM 965 and PM 660, would be second 



10 BREVIORA No. 446 

molars, corresponding to the largest upper molars. The smaller 
PM 948 and PM 1249 would be third or perhaps fourth molars 
(Fig. 5, P). It is to be noted that Crompton (1971) used a Type 4 
tooth to illustrate the occlusion of Pappotherium. 

Patterson's Type 3 was represented by only one damaged tooth 
(PM 922), but an unworn specimen (PM 1245; Fig. 3, K) has been 
recovered subsequently. It is very similar to the holotype of Ker- 
mackia texana, from which it differs by its higher and more pointed 
protoconid and metaconid. This difference is due to loss of enamel 
from the trigonid of the holotype. The trigonid is widely open, and 
the paraconid points anteriorly. The protoconid is the highest tri- 
gonid cusp, the paraconid the lowest. From the metaconid arises 
a strong posterior crest (named Distal Metacristid by Fox, 1975), 
continuous with the crista obliqua. The talonid is narrower than 
the trigonid. The hypoconid stands only slightly more buccally 
than the hypoconulid. The entoconid is small, and the talonid basin 
is open on the lingual side between the entoconid and the meta- 
conid. 

Slaughter's Type 6 is based on the jaw fragment SMP-SMU 
61728. This contains a molar (Fig. 3, T), apparently the last, and 
the alveoli of the next anterior tooth. Type 6 resembles Kermackia 
texana in size and general proportions, but the metaconid of Type 6 
is much smaller, no higher than the paraconid. As in Kermackia, 
the crista obliqua runs to the metaconid. As the crest continues up 
the posterior surface of the metaconid, it is much weaker and nearly 
vertical. The talonid basin is widely open on the lingual side as in 
Kermackia, but the entoconid is almost absent, and an additional 
cusp is present on the crista obliqua. The top of the hypoconid has 
been broken off. Two trigonids from Greenwood Canyon, PM 
1065 and PM 1045, have the same structure. This form is suffi- 
ciently distinctive to deserve a name, which I propose formally 
as follows: 

Trinititherium, gen. nov. 

Diagnosis. The lower molars differ from those of Kermackia as 
follows: (1) metaconid and paraconid subequal; (2) an accessory 
cusp present on the crista obliqua; (3) the crest on the posterior 
surface of the metaconid weaker and more vertical; (4) entoconid 
rudimentary. Type species: 

Trinititherium slaughteri, sp. nov. 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 11 

Diagnosis. This is the only known species of Trinitiiherium. 

Holotype, SMP-SMU 61728, from Butler Farm. Illustrated by 
Slaughter (1965, Fig. 4, and 1971. PI. 1). 

W. A. Clemens (personal communication) has raised the question 
whether Triniiitherium might represent the posterior molar of Ker- 
mackia. This would imply a reduction of the metaconid towards 
the posterior end of the series. Such a process occurs as an aspect 
of carnassial specialization in Cenozoic carnivorous mammals (But- 
ler 1947). The metaconid is also reduced in Deltatheridiidae and 
Stagodontidae in the late Cretaceous. In all these, there is evidence 
of a sectorial specialization of the paraconid which is not shown by 
Trinititherium. The metaconid is similar in height to the paraconid 
in Kielantherium and probably in Aegialodon. In these genera, 
however, the talonid is shorter and narrower than in Trinititherium 
and Kermackia. 

TurnbuU (1971) grouped together Types 1, 2, and 5 as Holo- 
clemensia and Types 3, 4, and 6 as Pappotherium. He included in 
Holoclemensia the specimens of Type 2 that are here regarded as 
Pappotherium. He did not consider in detail the morphological 
features that distinguish Type 2 from Types 1 and 5. He ascribed 
the differences to individual variation, to position in the series, and 
possibly to the presence of more than one species of Holoclemensia. 
The teeth that TurnbuU identified as Pappotherium include those 
of Kermackia and Trinititherium, together with Type 4, regarded 
here as the last molar of Holoclemensia. The criteria by which he 
distinguished Pappotherium. from Holoclemensia are those which 
in the present paper are used to distinguish Kermackia and Triniti- 
therium from Pappotherium and Holoclemensia. Kermackia had 
not been described when TurnbuU wrote his paper, and his suppo- 
sition, that all the tribosphenic molars in the collection could be 
referred to Pappotherium or Holoclemensia, was not unreasonable. 

There remains to be considered the lower jaw with four teeth 
(SMP-SMU 61992; Fig. 3, S), referred by Slaughter (1971) to Pap- 
potherium. Two teeth are simple, and two are molariform. The 
first molariform tooth is narrower than the second and has less 
elevated cusps. Slaughter says the first is less worn than the second 
molariform tooth and identifies it as a premolar. The existence of 
a molariform premolar at such an early date seems very unlikely. 
Both molariform teeth are much more worn than the two premo- 



12 BREVIORA No. 446 

lariform teeth, one of which is in process of eruption. Slaughter's 
statement (1971: 137) that X-rays showed no unerupted teeth ex- 
cludes the possibility that the first molariform tooth is a milk 
molar. The molariform teeth must be regarded as permanent 
molars. 

Ml is similar in size to that of Kermackia; M: is larger than M|, 
though it is smaller than the smallest specimen of Type 2 (PM 930). 
On both teeth, the tops of the trigonid cusps are broken off or 
worn, but the metaconid was evidently higher than the forwardly 
projecting paraconid. On both molars the talonid is as wide as the 
trigonid, and the hypoconid is buccally situated in relation to the 
hypoconulid. The entoconid is as high as the hypoconulid on M2, 
but lower on Mi. On both molars the anterior end of the crista 
obliqua has been obliterated through wear, and its relations to the 
trigonid are uncertain. However, the metaconid of M2 carries a 
strong posterior ridge that resembles Kermackia and differs from 
Type 2. This specimen cannot be placed in any of the species so 
far described, and because of its importance I propose a name for 
it as follows: 

Slaughteria, gen. nov. 

Diagnosis. M: differs from lower molars of Type 2, referred to 
Pappotheriuni. in that (1) the metaconid is higher than the para- 
conid, and (2) the metaconid has a posterior ridge, probably con- 
tinuous with the crista obliqua. M: differs from lower molars of 
Kermackia in that (1) the talonid is equal in width to the trigonid, 
and (2) the entoconid is larger than that of Kermackia. There is 
no metaconid on the lower premolars. Type species: 
Slaughteria eruptens, spec. nov. 

Diagnosis. This is the only known species of Slaughteria. 

Holotype, SMP-SMU 61992, from Butler Farm. Illustrated by 
Slaughter, 1971, PL 9. It is a mandibular fragment with two molars 
and two premolars, the anterior premolar in process of eruption. 

PREMOLARS 

The last premolar of Slaughteria is a two-rooted tooth with a 
well developed, one-cusped heel but no anterior cusp. The anterior 
premolar is smaller and seemingly has a single root. There are some 
isolated two-rooted premolars from Greenwood Canyon. PM 1247 
(Fig. 4, a) is larger than the last premolar of Slaughteria. It has a 



1978 



MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 



13 



small posterior heel and an anterior basal cusp which is displaced 
towards the lingual side, indicating that the tooth is from the lower 
jaw. PM 1098 (Fig. 4, b) is the posterior part of a lower premolar, 
probably a little larger than PM 1247 and with a larger, two-cusped 
heel. PM 1 136 (Fig. 4, c) is a much smaller tooth with no anterior 




Figure 4. Premolars and canine, a, PM 1247, lingual and crown views; b, PM 
1098, lingual view; c, PM 1136, ? buccal view; d, PM 931, buccal and crown views; 
e, PM 1 124; f, SMP-SMU 62399, lingual and crown views (from a cast); g, SMP- 
SMU 61947, lingual and crown views; h, SMP-SMU 61948, buccal and crown views. 



14 BREVIORA No. 446 

cusp or posterior heel. Its anterior profile is strongly convex, and 
there is a posterior crest which shows wear. None of the Texas 
specimens resembles the Wealden specimen that Clemens (1963, 
Fig. 10) described. The latter is a narrow tooth with divergent roots 
and a low crown, having the appearance of a milk molar. SMP- 
SMU 61730 and 61731 (Slaughter 1965, Fig. 5) are triconodont. 

The submolariform premolars that Slaughter (1968a, 1971) de- 
scribed are of greater interest. SMP-SMU 61947 (Fig. 4, g) is a 
lower premolar with a small talonid, a low, lingually placed para- 
conid, and a low metaconid. A crest on the buccal side from the 
talonid cusp continues up the posterobuccal surface of the proto- 
conid. In addition, there is a median talonid crest (=crista obliqua) 
which terminates at the base of the trigonid. SMP-SMU 62399 
(Fig. 4, resembles SMP-SMU 61947 in having a small metaconid, 
but it differs from that specimen in a number of details. The tooth 
is narrower and the protoconid is taller. The crista obliqua con- 
tinues up the posterior surface of the protoconid to meet the crest 
that connects the protoconid with the metaconid. As the paraconid 
of SMP-SMU 62399 has broken off, the length of the tooth can 
only be estimated, but it was probably a little shorter than SMP- 
SMU 61947 (length about 1.4 mm, compared with 1.55 mm). 

It is unlikely that either of these teeth is a penultimate premolar 
(conventionally called PO- Among late Cretaceous mammals a 
metaconid is known on Py only in Gypsonictops. In that genus P3 
also has a partly molariform talonid, and its possession of a meta- 
conid can be ascribed to the unusually high level of molarization 
of the tooth. Such molarization is almost certainly a specialization 
of Gypsonictops, unlikely to be present in ancestral placentals 
(Butler 1977). A metaconid occurs on the last premolar (P4) in 
several late Cretaceous genera: Zalamhclalestes. Protungulatum, 
Procerberus and Batodon. Both the Albian specimens are inter- 
preted as P4. The differences between them show that they belong 
to different species. 

SMP-SMU 61948 (Fig. 4, h) is a triangular upper premolar which 
Slaughter (1968a) compared with a placental P\ The lingual lobe 
was broken off, but a small protocone must have been present 
originally. The paracone is a tall cusp, similar in height to the 
protoconid of SMP-SMU 62399. There is no metacone. This tooth 
resembles P^ of Asiorvctes. but whether it is a P^ or a P"* is uncer- 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 15 

tain. In Asioryctes, as in Kennalestes and Zalambdalestes, the 
paracone of P^ is higher than that of ?•* and subequal to the proto- 
conid of P4. However, P^ may have been secondarily enlarged 
(Butler 1977). There is no metaconid on the lower premolars of 
Kennalestes and Asioryctes, and comparison with these genera may 
be misleading. In Cretaceous marsupials, such as Alphadon, the 
last upper premolar is the largest, and this is true also of Deltathe- 
ridium. 

PM 931 (Fig. 4, d) is an upper premolar which differs greatly 
from SMP-SMU 61948. It has a metacone on the shearing crest, 
but no protocone. This last cusp is represented only by an indis- 
tinct cingulum. The strongly developed shearing crest suggests that 
PM 931 is a last premolar that functioned against the trigonid of 
Ml. The absence of the protocone suggests that the corresponding 
lower premolar lacked a metaconid, and therefore that PM 931 did 
not occlude with either of the known complex lower premolars. It 
is too large to fit the lower dentition of Slaughteria. There are only 
two roots. The posterior one is flattened in cross section and ar- 
ranged obliquely, to support the lingual as well as the posterior 
apex of the crown. A similar arrangement is known in symmetro- 
dont molars, but PM 931 is much too large to be referred to Spa- 
lacotheroides. 

Probably some of the submolariform premolars belong to spe- 
cies of which the molars are already known. Only in Slaughteria 
have premolars been found in association with molars, and there 
the last premolar has no metaconid. In Slaughteria, the length of 
the last premolar is 65 per cent of that of M:. The two lower pre- 
molars with metaconids are as long as or longer than the molars 
of Kermackia and Trinititherium, and they are therefore probably 
too large to belong to these forms. Moreover in Kermackia and 
Trinititherium the crista obliqua on the molars runs directly to the 
metaconid. The possibility that the submolariform premolars be- 
long to Pappotherium or Holoclemensia must therefore be con- 
sidered. 

The crista obliqua of SMP-SMU 62399 resembles that of Type 
2 (supposedly Pappotherium) molars in meeting the protoconid- 
metaconid crest, whereas that of SMP-SMU 61947 fades out near 
the base of the trigonid as on Holoclemensia molars. In the late 
Cretaceous marsupials, Alphadon and Pediomys, the last lower 



16 BREVIORA No. 446 

premolar is about three-quarters as long as the longest molar, and 
in the placental Kennalestes the proportion is about 80 per cent. 
SMP-SMU 61947 is 80 per cent as long as PM 1005, interpreted 
as M2 of Holoclemensia; the somewhat shorter SMP-SMU 62399 
would bear about the same proportion to PM 965, the largest 
supposed Pappotherium molar. Size as well as structure is therefore 
consistent with the reference of SMP-SMU 61947 to Holoclemensia 
and SMP-SMU 62399 to Pappotherium. 

The upper premolar SMP-SMU 61948 would fit either of the two 
complex lower premolars. It possesses a strongly developed stylar 
cusp on the buccal edge, anterior to the metastyle. This cusp might 
be considered serially homologous with Style C of the molars. For 
this reason I refer SMP-SMU 61948, tentatively, to Holoclemensia 
as a P'' or P^. 

PM 931 (Fig. 4, d) is too small and its shearing crest too longi- 
tudinal to have functioned against the trigonid of PM 966, the sup- 
posed Ml of Holoclemensia. The absence of the protocone on the 
premolar suggests that this cusp was relatively small on the molars. 
Perhaps this tooth belongs to the same species as the last molar 
PM 1287 (Fig. 2, p) in which the protocone is small. 

The mandible PM 583 (Patterson 1956, Figs. 10, 11) contains 
alveoli for four two-rooted premolars. The posterior alveoli are of 
a size to fit Kermackia molars and are probably too large for pre- 
molars of that genus. The last pair of alveoli, however, could have 
held either of the premolars with metaconids, and the simple pre- 
molar PM 1247 would fit the alveoli for P3. There is a possibility 
that this mandible belongs to either Holoclemensia or Pappothe- 
ium. The remarkably large canine alveolus differentiates this speci- 
men from SMP-SMU 62400 (Slaughter 1971, PI. 10), a fragment 
containing a small, procumbent canine. PM 1124 (Fig. 4, e) is an 
upright two-rooted canine of small size, probably from the upper 
jaw. The mandible PM 583 contains alveoli for four lower incisors. 
Slaughter (1971, PL 10) described a rather large, spatulate upper 

incisor. 

DISCUSSION 

Number of Genera Represented 

The material from the Albian of Texas consists almost entirely 
of isolated teeth. Until more complete specimens are discovered, 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 17 

any interpretations or conclusions must remain highly speculative. 
Differences in the molars, premolars, canines and incisors indicate 
that a number of distinct taxa are present, at least on the generic 
level. In attempting to allocate the various sorts of teeth, I have 
avoided the introduction of additional taxa as far as possible. The 
minimum number of taxa necessary to include the known molars 
seems to be the following six: 

1. Holoclemensia, the largest form, is represented by several 
upper and lower molars; complex upper and lower premolars are 
allocated to this form. 

2. Pappotherium, a somewhat smaller form, is represented again 
by several upper and lower (Type 2) molars; a complex lower pre- 
molar is referred to it. 

Holoclemensia and Pappotherium are the most common mem- 
bers of the fauna, but Pappotherium seems to be relatively more 
common at Greenwood Canyon than at Butler Farm. The tooth- 
less mandible may belong to either genus. 

3. Slaughteria is represented by a single mandibular specimen, 
which is the only one to show associated molar and premolar teeth; 
it is smaller than Pappotherium. 

4. Kermackia is a small form known only by three lower molars. 

5. Trinititherium, similar in size to Kermackia, is known by one 
complete lower molar and two trigonids. The small upper molar 
SMP-SMU 62402, and the last molar PM 1075 may belong to either 
Kermackia or Trinititherium. 

6. The last upper molar PM 1287 and the upper premolar PM 
931 do not appear to belong to any of the foregoing and probably 
represent one or two additional species. 

The Family Pappotheriidae 

As interpreted here, Pappotherium and Holoclemensia are much 
alike in the organization of their molar series. Both have four 
molars, of which the second is the largest, and in the upper jaw, 
M^ is reduced posterobuccally. M' o{ Holoclemensia is more sym- 
metrical than in Pappotherium, and M^ less so, but such differences 
are to be found frequently in closely related genera of mammals. 

There is a wide stylar shelf in both genera. The trigonid is ele- 
vated, and it has a well developed, trenchant paraconid. The talo- 
nid is narrower than the trigonid. The lower molars have no cingu- 



18 BREVIORA No. 446 

lum except on the anterior surface of the protoconid. The last lower 
premolar has a metaconid and, at least in Holoclemensia, the last 
upper premolar has a protocone. 

The large Style C of Holoclemensia is represented in some speci- 
mens of Pappotherium by a minor elevation of the buccal marginal 
ridge. The metacone of Holoclemensia is larger in comparison with 
the paracone than in Pappotherium. The lower molars of Holo- 
clemensia have a more compressed trigonid, a higher metaconid, 
and a more buccally situated hypoconid. The crista obliqua in 
Holoclemensia fades out at the base of the trigonid, whereas in 
Pappotherium it continues up the trigonid wall. These differences 
seem insufficient to place the two genera in separate families, and 
I propose to include them both in the Pappotheriidae Slaughter 
1965. 

Relationship of Pappotheriidae to Marsupials and Placentals 

The Pappotheriidae agree with the primitive marsupial Alphadon 
in the possession of four molars and in the wide stylar shelf of the 
upper molars. Holoclemensia resembles Alphadon in having a 
distinct Style C, but in Alphadon this is smaller than the stylocone 
(Style B), and it varies in size between the species. In Alphadon M^ 
is the largest molar, and its posterobuccal apex and metastylar blade 
are not reduced. A further difference appears to be that the proto- 
cone shelf of Alphadon is better differentiated with more strongly 
developed conules. In Alphadon the metacone is as tall or taller 
than the paracone; Holoclemensia, with its larger metacone, ap- 
proaches Alphadon in this respect more closely than Pappotherium 
does. 

The lower molars of Pappotheriidae resemble Alphadon in the 
trenchant paraconid and in the connection of the crista obliqua to 
the middle of the posterior trigonid wall. However, in the marsu- 
pials the talonid is as wide as the trigonid, sometimes wider. The 
hypoconulid is in a lingual position, close to the entoconid, and a 
cingulum, arising from the hypoconulid, passes behind the hypo- 
conid. In pappotheriids the trigonid, especially in Holoclemensia, 
is more elevated. Holoclemensia also differs from Alphadon in 
that the trigonid is more compressed on M2-M4. The metaconid is 
higher than the paraconid, and the hypoconid is higher in compari- 
son with the entoconid on M2 and M3. Pappotherium (i.e. Type 2) 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 19 




Figure 5. Reconstructions of cheek teeth in occlusion. P. Pappotherium patter- 
soni; H, Holoclemensia texana; A, Alphadon marshi (a marsupial); C, Cimolesles 
incisus (a placental). 

Specimens used in the reconstruction of Pappotherium: M', PM 999; M^ PM 
894; M3 and M^ the holotype; P4, SMP-SMU 62399; M,, PM 930; M2, PM 965; 
Ms, PM 948. 

Specimens used in the reconstruction of Holoclemensia: P*, SMP-SMU 61948; 
M'. PM 1000; M2, PM 886; M'. the holotype; M^ the paratype; P4, SMP-SMU 
61947; M,, PM 966; M2, PM 1005; Mj, SMP-SMU 61727; M4, SMP-SMU 61726. 



20 BREVIORA No. 446 

agrees more with Alphadon in these characters, as was noted by 
Clemens (1966:16-17). 

Alhertatherium (Fox 1971), from the early Campanian, ap- 
proaches Holoclemensia in that Style C is higher than the stylocone, 
and the metaconid is somewhat higher than the paraconid. How- 
ever, the stylar shelf of Alhertatherium is narrower than in Holo- 
clemensia. The metacone and metastyle of M^ are larger, and the 
conules are better developed. On the lower molars, the trigonid is 
less compressed; the talonid wider; and the hypoconulid is lingual 
in position, and the entoconid is higher. As Fox (1971) noted, 
Alhertatherium does not link Alphadon with Holoclemensia. Style 
C is also enlarged in Aquiladelphis, an early Campanian pediomyid 
in which the stylocone is reduced to two small cusps. In other re- 
spects this form resembles Holoclemensia even less than Alherta- 
therium: for example, the crista obliqua meets the posterior wall of 
the protoconid. 

The primitive eutherian molar pattern is believed to be preserved 
best in the late Cretaceous Cimolestes and Kennalestes (Butler, 
1977). Holoclemensia and Pappotherium differ from these in the 
retention of the fourth molar, in the wide stylar shelf with better 
developed styles posterior to the stylocone, and in the correspond- 
ingly weaker development of the protocone shelf. Lower molars of 
Pappotheriidae agree with those of primitive placentals in the high 
trigonid and comparatively narrow talonid, in the connection of 
the crista obliqua to the middle of the trigonid wall, and in the ab- 
sence of the posterior cingulum. Holoclemensia, not Pappothe- 
rium, agrees with Cimolestes and Kennalestes in that the metaconid 
is higher than the paraconid. If the tooth has been correctly re- 
ferred, the possession of a premolar protocone would constitute 
another resemblance of Holoclemensia to placentals. However, 
there is no metaconid on P4 in Cimolestes or Kennalestes. 

Fox (1975) noted a resemblance in the outlines of the last two 
upper molars between Holoclemensia and the late Cretaceous Del- 
tatheroides, which has four molars. He concluded that Holocle- 
mensia must have four molars. Because the penultimate molar of 
Pappotherium shows less reduction in the metastyle. Fox inferred 
that only three molars were present in that genus. I find his argu- 
ment unconvincing; the relationship between the shapes of teeth 
and their numerical position is too inconstant for such deductions 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 21 

to be made. Late Cretaceous didelphids have four molars, and yet 
the metastyle of their penultimate molar is more developed than in 
Pappotherium. Deltatheroides, in which the metastyle of M^ is 
reduced, can be regarded as being in the process of losing its M'*, 
which is absent in Deltatheridium (Kielan-Jaworowska 1975); 
Holoclemensia might by analogy be considered to be evolving 
towards the three-molar state characteristic of placentals. 

It seems impossible to fit Holoclemensia and Pappotherium into 
a simple metatherian-eutherian framework. They have a mixture 
of characters which indicate that they belong to neither infraclass. 
Though Holoclemensia approaches primitive marsupials in some 
ways, Pappotherium is more like them in others; Holoclemensia 
also shows some resemblances to primitive placentals. No doubt 
some of the characters that pappotheriids share with marsupials or 
placentals are plesiomorphs inherited from a common ancestor, 
which may include the wide stylar shelf of primitive marsupials and 
the narrow talonid of primitive placentals. The fourth molar is 
probably another plesiomorphous character lost in placentals but 
retained in Deltatheridiidae (Butler & Kielan-Jaworowska, 1973; 
Kielan-Jaworowska, 1975) as well as in marsupials. Other shared 
characters may be due to parallel evolution, such as the enlarged 
Style C. The metaconid of P4 may also have evolved independently, 
for it occurs in some placentals but not in others. 

Placentals have been reported from Khovboor in Mongolia 
(Beliajeva et al., 1974; Dashzeveg, 1975) in a deposit that is be- 
lieved to be of Aptian age. The fauna, still undescribed, is said to 
contain forerunners of the much later Djadokhta fauna. If the 
'Aptian age of these specimens is correct, the divergence of placen- 
tals and marsupials must already have taken place at that time. The 
Albian Pappotheriidae would in that case be too late to be the 
common ancestors of the two infraclasses. Clemens (1971: 174) 
states that "The evidence . . . does not exclude the possibility that 
when the tribosphenic grade of dental evolution was attained an 
adaptive radiation occurred producing lineages in addition to those 
ancestral to marsupials and placentals." The Pappotheriidae may 
well have belonged to one such lineage. 

During much of the Cretaceous the land surface was more than 
usually fragmented by epicontinental seas (Lillegraven, 1974; Tel- 
ford, 1974). North America was divided, from middle Albian time 



22 BREVIORA No. 446 

onwards, into eastern and western land-masses by the Interior Sea- 
way. The northern part of the Atlantic was closed: the seaway 
between Greenland and Canada developed only in the Campanian 
(Martin, 1973). Europe was separated from Asia by the Turgai 
Strait. In mid-Cretaceous time, eastern North America and Europe 
thus formed a single continent from which the mammals from Texas 
are the only known representatives of their class. Following Lille- 
graven (1974) one may imagine a wide dispersal, in the early Cre- 
taceous, of an ancestral group with tribosphenic molars, of which 
Aegialodon, from the English Wealden, is the only known example. 
Subsequently isolated on different land-masses, the ancestral group 
gave rise to the placentals in Asia and the marsupials in western 
North America (or as Telford (1974) suggests, in South America). 
Possibly the Pappotheriidae represent another lineage that evolved 
on the Euramerican continent, but subsequently became extinct. 
A molar from the Campanian of Champ-Garimond, France (Le- 
doux et al., 1966; Butler, 1977) might conceivably be a pappotheriid 
derivative, but it is more advanced, and it might equally well be a 
placental immigrant from Asia. 

Kermackiidae, Fam. nov. 

Kermackia and Trinititherium are distinguished from Pappo- 
therium and Holoclemensia by the continuation of the crista ob- 
liqua to the tip of the metaconid, so that on the posterior surface 
of that cusp it forms a ridge (the distal metacristid of Fox, 1975). 
The talonid is narrow, its basin is widely open lingually, and the 
entoconid is small or rudimentary. These genera are at a lower level 
of molar evolution than the Pappotheriidae. In the latter the con- 
nection of the crista obliqua to the trigonid has shifted to a more 
buccal position, the talonid is wider, and the entoconid is larger. 
Dashzeveg (1975) noted the resemblance oi Kermackia to Aegialo- 
don and Kielantherium, which belong to the family Aegialodonti- 
dae Kermack (1957). However, these differ from Kermackia in 
having the talonid reduced in length as well as in width. Because 
of this difference I propose the family Kermackiidae to include 
Kermackia and Trinititherium. The Kermackiidae resemble the 
Aegialodontidae in retaining the distal metacristid but differ from 
them in that the talonid is equal in length to the trigonid. The 



1978 



MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 



23 



Aegialodontidae, Kermackiidae and Pappotheriidae represent three 
stages in the evolution of the talonid (Fig. 6). 

Slaughteria, which probably possesses a distal metacristid, has 
a wide talonid, and (on M2, but not on Mi) a fully developed en- 
toconid like the Pappotheriidae. It may represent another family, 
but, as some doubt remains over the details of the molar pattern, 
I prefer to place Slaughteria provisionally in the Kermackiidae. To 
judge from tlie relative size of the penultimate premolar, the num- 
ber of premolars may have been reduced. Slaughteria is the only 
Albian therian for which there is evidence of tooth replacement, 
and on this character. Slaughter (1971) placed it in the Eutheria. 
However, diphyodonty occurs in pantotheres (Butler &. Krebs, 
1973), and it must be considered a plesiomorphous character, re- 




Figure 6. Left lower molars in lingual view, to illustrate possible evolutionary 
radiation in the early Cretaceous, a, Peramus (Pantotheria); b, Aegialodon; cKiel- 
antherium (Aegialodontidae); c, Kermackia; d, Trinititherium (Kermackiidae); f, 
Deltatheridium (Deltatheridiidae); g, Holoclemensia: h, Pappotherium (Pappo- 
theriidae); i, Alphadon (Metatheria); j, Champ-Garimond molar; k, Kennalestes 
(Eutheria). 



24 BREVIORA No. 446 

tained in placentals and lost (except for the last premolar) in mar- 
supials. 

CLASSIFICATION ABOVE THE FAMILY LEVEL 

A name is required for an infraclass to include those therians 
that have a functional protocone and have advanced beyond the 
pantotherian stage, but nevertheless cannot be regarded either 
as marsupials or placentals. Turnbull (1971) proposed a cohort 
Tribosphenata, equal in rank with his Marsupiata and Placentata, 
including all within the infraclass Eutheria. The Tribosphenata 
comprised two orders: (1) Tribosphena, containing Pappothehum, 
Holoclemensia, Aegialodon and Peramus, and (2) Zalambdodonta, 
consisting of a number of placentals and marsupials with zalamb- 
dodont molar teeth. Since zalambdodont molar patterns have been 
derived, more than once, from dilambdodont ("euthemorphic") pat- 
terns (Butler 1972), I consider the order Zalambdodonta to be arti- 
ficial. 

I propose the name Tribotheria for an infraclass standing between 
the Pantotheria, on the one hand, and the Metatheria and Eutheria 
on the other. The Tribotheria differ from the Pantotheria in having 
reached the tribosphenic stage of molar evolution by the acquisition 
of a protocone that functioned against the talonid. The infraclasses 
Metatheria and Eutheria originated from members of the Tribo- 
theria, but there were other, less diversified lineages, such as the 
Deltatheridiidae and Pappotheriidae, which may be left in the Tri- 
botheria. TurnbulPs order Tribosphena was similar in conception, 
but it included Peramus. which lacks a functional protocone. 

For Tribotheria with a distal metacristid I propose an order 
Aegialodontia. This corresponds to stage IIA in the scheme of 
molar evolution tabulated by Fox (1975, Table 1). The Aegialo- 
dontia includes the families Aegialodontidae and Kermackiidae. 
The late Cretaceous Deltatheridiidae may also be placed here. Their 
lower molars show a considerable resemblance to Kielantherium, 
and it seems likely that they have an aegialodontid ancestry, from 
which they have retained the small talonid. Potamotelses (Fox 
1975) may represent another family in the Aegialodontia. For the 
Pappotheriidae, which have lost the distal metacristid, and are at 
a stage of evolution comparable to the primitive marsupials and 
placentals, I propose a new order Pappotherida. 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 25 

The classification proposed may be tabulated: 
Subclass Theria 

Infraclass Pantotheria 
Infraclass Tribotheria 
Order Aegialodontia 

Family Aegialodontidae (Aegialodon, Kielantherium) 
Family Kermackiidae {Kermackia. Trinititherium, ? Slaugh- 
ter ia) 
Family Deltatheridiidae {Deltatheroides, Deltatheridium) 
Family ? (Potamotelses) 
Order Pappotherida 

Family Pappotheriidae {Pappotherium, Holoclemensia) 
Infraclass Metatheria 
Infraclass Eutheria 

SUMMARY 

1. At least six genera of mammals with tribosphenic molars are 
represented in the collection, but most specimens can be referred 
to Pappotherium and Holoclemensia. 

2. There are four types of upper molar of Holoclemensia pat- 
tern, and four of Pappotherium pattern, showing that both these 
genera had four molars. 

3. Of the six types of lower molar distinguished by Patterson 
and Slaughter, Types 1, 4, and 5 are referred to Holoclemensia 
and Type 2 (except for PM 966) to Pappotherium. 

4. Lower molars of Type 3 are Kermackia, and those of Type 6 
are named Trinititherium slaughteri gen. et sp. nov. 

5. The two lower premolars with metaconids are referred to 
Holoclemensia and Pappotherium respectively, and the upper pre- 
molar with a protocone is referred to Holoclemensia. 

6. The mandible SMP-SMU 61992, which Slaughter (1971) re- 
ferred to Pappotherium, has two molars and two premolars. It is 
named Slaughteria eruptens gen. et sp. nov. It is the only specimen 
that provides evidence of tooth replacement. 

7. Pappotherium and Holoclemensia are united in the family 
Pappotheriidae. They are neither marsupials nor placentals, but 
represent a separate line of evolution. 

8. Kermackia and Trinititherium are placed in a new family 
Kermackiidae. They are at a more primitive stage of molar evo- 



26 BREVIORA No. 446 

lution than the Pappotheriidae. Slaughteria is provisionally placed 
in the Kermackiidae. 

9. A new infraclass Tribotheria is proposed to include mammals 
with tribosphenic molars that are not classifiable as marsupials or 
placentals. This infraclass includes two new orders, Aegialodontia 
and Pappotheria. 

ACKNOWLEDGMENTS 

I thank W. Turnbull and B. Patterson for assistance with the 
examination of the specimens from Greenwood Canyon. B. Slaugh- 
ter very kindly allowed me to study material from Butler Farm at 
the time unpublished, and provided casts. B. Patterson, Z. Kielan- 
Jaworowska and W. A. Clemens read draft manuscripts, and I am 
much indebted to them for helpful suggestions. 

LITERATURE CITED 

Beliajeva, E. I., B. A. Trofimov, and V. J. Reshetov. 1974. (General Stages 
in evolution of late Mesozoic and early Tertiary mammalian fauna in Central 
Asia). Fauna i biostratigrafia Mesozoa i Kainozoa Mongolii. Trudy Sovm. 
Sov.-Mong. Pal. Exp. Moscow, 1: 19-45. 

Butler, P. M. 1947. The evolution of carnassial dentitions in the Mammalia. 
Proc. Zool. Soc. London, 116, Part II: 198-220. 

Butler, P. M. 1972. The problem of insectivore classification. M Joysey, K. A. 
and T. S. Kemp (Eds.), Studies in Vertebrate Evolution. Edinburgh: Oliver & 
Boyd: 253-265. 

Butler, P. M. 1977. Evolutionary radiation of the cheek teeth of Cretaceous 
placentals. Acta palaeont. Polon., 22: 241-271. 

Butler, P. M. and Z. Kielan-Jaworowska. 1973. Is Deltatheridium a mar- 
supial? Nature, 254: 105-106. 

Butler, P. M. and B. Krebs. 1973. A pantotherian milk dentition. Palaont. 
Zschr., 47: 256-258. 

Clemens, W. A. 1963. Wealden mammalian fossils. Palaeontology, 6: 55-69. 

Clemens, W. A. 1966. Fossil mammals of the type Lance Formation, Wyoming. 
Part II Marsupialia. Univ. CaUf. Publ. Geol. Sci. 62: 1 122. 

Clemens, W. A. 1971. Mammalian evolution in the Cretaceous. In Kermack, 
D. M. and K. A. Kermack (Eds.), Early Mammals. Zool. J. Linn. Soc, 50, 
Suppl. 1: 165-180. 

Crompton, a. W. 1971. The origin of the tribosphenic molar. In Kermack, D. 
M. and K. A. Kermack (Eds.), Early Mammals. Zool. J. Linn. Soc, 50, Suppl. 
1: 131-143. 

Dashzeveg, D. 1975. New primitive therian from the early Cretaceous of Mon- 
golia. Nature, 256: 402-403. 



1978 MAMMALIAN TEETH OF TRIBOSPHENIC PATTERN 27 

Fox, R. C. 1971. Marsupial mammals from the early Campanian Milk River 
Formation, Alberta, Canada. In Kermack, D. M. and K. A. Kermack (Eds.), 
Early Mammals. Zool. J. Linn. Soc, 50, Suppl. 1: 145-164. 

Fox, R. C. 1975. Molar structure and function in the early Cretaceous mammal 
Pappotherium: evolutionary implications for Mesozoic Theria. Canad. J. Earth 
Sci., 12: 412-442. 

HoFFSTETTER, R. 1970. L'histoirc biogeographique des Marsupiaux et la dichot- 
omic Marsupiaux-Placentaires. C. R. Acad. Sci., Paris, Ser. D, 271: 388-391. 

HoFFSTETTER, R. 1972. Donnccs et hypotheses concernant I'origine et I'histoire 
biogeographique des Marsupiaux. C. R. Acad. Sci., Paris, Ser. D, 274: 2635- 
2638. 

Kerm.-^ck, K. a. 1967. The Aegialodontidae — a new family of Cretaceous mam- 
mals. Proc. geol. Soc. London, 1640: 146. 

Kielan-Jaworowska, Z. 1975. Evolution of the therian mammals in the late 
Cretaceous of Asia. Part L Deltatheridiidae. Palaeont. Polon., 33: 103-132. 

Ledoux, J.-C, J.-L. Hartenberger, J. Michalix, J. Sudre, and L. Thaler. 
1966. Decouverte d'un mammifere dans le Cretace Superieur a dinosaures de 
Champ-Garimond pres de Fons (Card). C. R. Acad. Sci., Paris, Ser. D, 262: 
1925-1928. 

Lillegraven, J. A. 1969. Latest Cretaceous mammals of upper part of Edmon- 
ton Formation of Alberta, Canada, and review of marsupial-placental dichot- 
omy in mammalian evolution. Univ. Kansas paleont. Contr., 50: 1 122. 

Lillegraven, J. A. 1974. Biogeographical considerations of the marsupial-pla- 
cental dichotomy. Ann. Rev. Ecol, Syst., 5: 263-283. 

Martin, R. 1973. Cretaceous — early Tertiary rift basin of Baffin Bay — con- 
tinental drift without sea-floor spreading. In Pitcher, M. G. (Ed.), Arctic Geol- 
ogy. Amer. Assoc. Petrol. Geol. Mem. 19: 500-505. 

Patterson, B. 1956. Early Cretaceous mammals and the evolution of mamma- 
lian molar teeth. Fieldiana: Geol., 13: 1 105. 

Slaughter, B. H. 1965. A therian from the lower Cretaceous (Albian) of Texas. 
Postilla, 93: 1-18. 

Slaughter, B. H. 1968a. Earliest known eutherian mammals and the evolution 
of premolar occlusion. Texas J. Sci., 20: 3-12. 

Slaughter, B. H. 1968b. Earliest known marsupials. Science, N. Y., 162: 254- 
255. 

Slaughter, B. H. 1968c. Holoclemensia instead of Clemensia. Science, N. Y., 
162: 1306. 

Slaughter, B. H. 1971. Mid-Cretaceous (Albian) therians of the Butler Farm 
local fauna, Texas. In Kermack, D. M. and K. A. Kermack (Eds.), Early Mam- 
mals. Zool. J. Linn. Soc, 50, Suppl. 1: 131-143. 

Telford, R. H. 1974. Marsupials and the new paleogeography. In Ross, C. A. 
(Ed.), Paleogeographic Provinces and Provinciality. Soc. Economic Paleontol- 
ogists and Mineralogists, Sp. Publ. 21: 109-126. 
Turnbull, W. D 1971. The Trinity therians: their bearing on evolution in mar- 
supials and other therians. In Dahlberg, A. A. (Ed.), Dental Morphology and 
Evolution: 151-179. Chicago: Univ. Chicago Press. 



v..i500': 



MAR 1 8 1985 

■ lARVARD 



B R E V I 0"R""A 

Museiiim of Comparative Zoology 



us ISSN 0006 9698 



Cambridge, Mass. 20 December 1978 Number 447 

FOOD SELECTION BY BEAVERS: SAMPLING BEHAVIOR 

Stephen H. Jenkins' 

Abstract. Beavers in central Massachusetts frequently remove small pieces of 
bark (less than 25 cm") from standing trees. After removing a sample, the beavers 
may abandon these trees. At two ponds studied by the author in fall 1973, there 
were significant differences in species composition of trees sampled, felled, and 
available near the ponds. If beavers are physiologically capable of measuring the 
nutritional value of small pieces of bark, then sampling may be an economical way 
for them to assess spatial and temporal variation in relative nutritional value of 
different tree species. 

INTRODUCTION 

During a recent study of food selection by beavers (Castor can- 
adensis) at two ponds in central Massachusetts (Jenkins, 1974), I 
observed numerous trees from which the beavers had removed 
small patches of bark (often less than 25 cm-). Without using them 
further, the beavers then abandoned these trees. During September 
and October 1973, beavers at Blue Heron Cove cut into or partially 
stripped the bark from the lower trunks of 46 trees without com- 
pletely felling them. At Tamplin Road Pond, there were 36 such 
trees. The beavers removed less than 25 cm- of bark from 25 of the 
46 trees at Blue Heron Cove and the same amount from 17 of the 
36 trees at Tamplin Road Pond. 

Chabreck (1958) reported that beavers in Louisiana stripped the 
bark from the lower trunks of many trees without felling them. 
Aldous (1938) and many others have noted that beavers may not 
use all the bark of the trees they fell. The present study is the first 
to describe and to consider the possible adaptive significance in the 
beavers' pattern of removing small pieces of bark (less than 25 cm^) 

'Department of Biology, University of Nevada, Reno, Nevada 89557 



2 BREVIORA No. 447 

from standing trees. Secondly, the study compares the beavers' 
patterns of selection of species for sampling with their patterns of 
selection of species for more complete utilization. 

For econamy of expression, I will call the beavers' incomplete 
use of trees (described above) sampling. I define light sampling as 
use of less than 25 cm^ of bark near the base of a tree without fell- 
ing it and heavy sampling as cutting into the wood of a tree and/ or 
stripping more than 25 cm- of bark near its base without felling it. 
Several authors (Rozin, 1969; Freeland and Janzen, 1974; Westoby, 
1974) have attached functional connotations to the word sampling 
in their discussions of optimal foraging. I wish to avoid a specific 
functional connotation in using the word. Instead, I present six 
alternative hypotheses which could account for beavers' incomplete 
use of trees. The first of these hypotheses argues that beavers' 
sampling might not be of direct adaptive value but is rather an 
incidental byproduct of the foraging process. The remaining five 
suggest five different ways in which sampling might function as an 
adaptive component of a beaver's foraging strategy. My data are 
insufficient to discriminate completely among these hypotheses; I 
present them to establish that sampling by beavers is a phenomenon 
worth further study, especially in the context of recent developments 
in optimal foraging theory (Krebs et al., 1977; Pyke et al., 1977). 



RESULTS AND DISCUSSION 

Table 1 shows the generic distributions of trees sampled, felled, 
and available at the major tree cutting sites of Blue Heron Cove 
and Tamplin Road Pond between 1 September 1973 and 6 Novem- 
ber 1973 (Blue Heron Cove) or 2 November 1973 (Tamplin Road 
Pond). The following hypotheses might explain beavers' tree sam- 
pling behavior. 

(1) A beaver may be interrupted while starting to fell a tree. 
Sampled trees simply represent those abandoned because of such 
interruption. If the probability of interruption is independent of 
tree genus, and if the time to remove less than 25 cm^ of bark from 
a tree is less than the time to fell even a small tree (but one more 
than 2.5 cm diameter, the lower size limit considered in this study), 
then this hypothesis predicts that the generic distributions of lightly 
sampled and felled trees should be the same. (Heavily sampled 



1978 



FOOD SELECTION BY BEAVERS 



Table 1 

Generic Distributions of Trees Sampled. Felled, and Available at 
Blue Heron Cove and Tamplin Road Pond. Fall 1973 





Lightly 


Heavilv 








Sampled* 


Sampled** 


Felled 


.Available+ 


- 




Blue H 


eron Cove 




Oak (Quercus spp.) 


1 1 (44%) 


16 (76%) 


31 (51%) 


63 (25%) 


Maple {Acer spp.) 


4(16%) 


1 ( 5%) 


14 (23%) 


97 (38%) 


Birch (Betula spp.) 


9 (36%) 


3(14%) 


1 ( 2%) 


41 (16%) 


Others 


I ( 4%) 


1 ( 5%) 


15 (25%) 


53 (21%) 


Totals 


25 


21 


61 


254 






Tamplin 


Road Pond 




Oak {Quercus spp.) 


6(35%) 


7 (37%) 


2 ( 6%) 


15(12%) 


Maple {Acer spp.) 


4(24%) 


9 (47%) 


12(34%) 


59 (45%) 


Cherry {Prunus spp.) 


5 (29%) 


3(16%) 


15(43%) 


18(14%) 


Others 


2(12%) 


( 0%) 


6(17%) 


38 (29%) 


Totals 


17 


19 


35 


130 



*Less than 25 cm^ of bark removed. 

**Wood cut and or more than 25 cm- of bark removed. 

+ Based on randomly placed. 78.5 m% circular quadrats. See Jenkins (1975) for 

details. 

trees are excluded because it may take less time to fell a small tree 
than to sample heavily a large tree, and a difference in the generic 
distributions of sampled and felled trees might arise from differ- 
ences in the available diameter distributions of various genera.) At 
Blue Heron Cove, generic distributions of lightly sampled and felled 
trees are significantly different (x' = 15.0, 2 df, p < 0.00 1, only oak, 
maple, and birch considered because few trees of other genera were 
sampled). At Tamplin Road Pond, these distributions are not quite 
significantly different (x' - 5.2, 2 df, 0.05 < p < 0.10, only oak, 
maple, and cherry considered). This hypothesis may account for 
the existence of some sampled trees. However, it does not explain 
the marked difference in generic distributions of felled and sampled 
trees at Blue Hercn Cove (Table 1). 



4 BREVIORA No. 447 

(2) Beavers cannot identify trees taxonomically without tasting 
their bark, so sampling is necessary for selection of preferred tree 
genera. If this is true, the generic distribution of trees felled or 
sampled (the sum of columns 1, 2, and 3 in Table 1) should match 
that of trees available. The significant differences between these 
distributions at both ponds (Blue Heron Cove: x' — 50.4, 3 df, p< 
0.001; TampHn Road Pond: x' = 31.3, 3 df, p < 0.001) cast doubt 
on this hypothesis. 

(3) Sampling is used to avoid ingestion of potential toxins and 
demonstrates cautiousness with novel food items (Freeland and 
Janzen, 1974). This hypothesis suggests that unfamiliar food types 
will be sampled more than familiar ones. Comparison of lightly 
sampled and felled trees at both Blue Heron Cove and Tamplin 
Road Pond refutes this hypothesis. In particular, oak is the genus 
most often sampled as well as felled at Blue Heron Cove, and cherry 
is most often felled and also frequently sampled at Tamplin Road 
Pond. Cautiousness in using new, potentially toxic foods may lead 
to sampling behavior in some species, but it does not explain these 
instances of sampling behavior by beavers. 

(4) Sampling satisfies a requirement for certain vitamins or 
minerals, needed in small quantities and available only in some 
tree species. This hypothesis fails to explain the extensive sampling 
of maples and oaks at Blue Heron Cove and maples and cherries 
at Tamplin Road Pond. Many trees of these species are felled. 
The felled trees should supply any micronutrients which may be 
uniquely available in these species, unless the felled trees have con- 
centrations of particular micronutrients several hundredfold less 
than sampled trees of the same species. 

(5) Sampling represents practice at tree selection by young ani- 
mals. I have no data on the amount of sampling done by individual 
members of the two colonies. However, young animals could prac- 
tice tree selection on branches and logs brought to the edge of the 
pond by adults with less risk of predation than if the young animals 
sampled standing trees at distances as great as 50 m from shore. 
Therefore, it is unlikely that sampling would evolve solely to serve 
this function. 

(6) Sampling is used by beavers to assess possible spatial and 
temporal variation in the nutritional quality of alternative foods. 
Unlike some of the others, this hypothesis cannot be tested by 



1978 FOOD SELECTION BY BEAVERS 5 

comparing generic distributions of trees sampled, felled, and avail- 
able. I present evidence elsewhere that beavers select different 
genera of trees for cutting at different sites around a pond (Jenkins, 
1975), and different genera in different years at the same site (Jen- 
kins, in preparation). There is evidence of spatial (Cowan et al., 
1950) and temporal (Kramer and Koslowski, 1960) variation in 
nutritional value of some tree species. It seems reasonable to sug- 
gest that thre spatial and temporal variation in beavers' tree selec- 
tion is related to spatial and temporal variation in relative nutri- 
tional value of different tree species. Sampling may be a mechanism 
for achieving an optimal diet in the face of such variation. 

CONCLUSIONS 

Two kinds of data, both difficult to collect, are needed to prove 
more conclusively which of the above hypotheses are incorrect. 
First, direct observations of beavers in the act of sampling would 
help decide the validity of hypotheses 1 and 5. Unfortunately, bea- 
vers are primarily nocturnal, so special methods (e.g., radio telem- 
etry, use of night vision instruments) would be needed to assist in 
making such observations. Data might be obtained on sampling 
during twilight hours without such assistance. However, such data 
probably would be inconclusive. For example, failure to observe 
sampling by young animals during twilight would certainly not 
prove that young animals don't sample; they might in fact be active 
samplers, but only at night. Second, data on toxin and nutrient 
content of bark of individual trees would help decide the validity 
of hypotheses 3, 4, and 6. To get adequate data of this sort would 
require analyzing a large number of samples for numerous nutrients 
and possible toxins (since both intraspecific and interspecific, as 
well as spatial and temporal variations in these attributes of tree 
bark are important components of these hypotheses). In addition, 
some of the toxins probably are unidentified or incompletely char- 
acterized at the present time. 

In this note, I show that beavers are highly selective in sampling 
trees, but that they choose genera in different proportions than in 
felling trees. I suggest that sampling may be an economical way 
for beavers to assess spatial and temporal variation in relative nu- 
tritional value of different tree species, assuming they are capable 
of measuring the nutritional value of small pieces of bark. The 



6 BREVIORA No. 447 

physiological and behavioral mechanisms by which sampling func- 
tions are important subjects for future research. 

ACKNOWLEDGMENTS 

I thank R. Fagen, H. Hodgdon, and T. Schoener for critically 
reviewing this paper, and I thank K. Jenkins for help in collecting 
the data. 

LITERATURE CITED 

Aldous, S. a. 1938. Beaver food utilization studies. J. Wildl. Manage., 2: 215- 
222. 

Chabreck, R. H. 1958. Beaver-forest relationships in St. Tammany Parish, 
Louisiana. J. Wildl. Manage., 22: 104-107. 

Cowan, 1. McT., W. S. Hoar, and J. Hatter. 1950. The effect of forest suc- 
cession upon the quantity and upon the nutritive values of woody plants used 
as food by moose. Can. J. Res., Sect. D, Zool. Sci., 28: 249 271. 

Freeland, W. J., AND D. H. Janzen. 1974. Strategies in herbivory by mammals: 
the role of plant secondary compounds. Amer. Nat., 108: 269-289. 

Jenkins, S. H. 1974. Food selection by beavers. Unpublished Ph.D. dissertation. 
Harvard Univ., Cambridge, Ma., 165 pp. 

. 1975. Food selection by beavers: a multidimensional contingency table 

analysis. Oecologia (Berl.), 21: 157-173. 

. 1978. Seasonal and year-to-year differences in food selection by beavers. 



In preparation. 
Kramer, P. J., AND T. T. KosLOWSKi. 1960. Physiology of trees. McGraw-Hill, 

New York, 642 pp. 
Krebs, J. R., T. Ericksen, M. I. Webber, and E. L. Charnov. 1977. Optimal 

prey selection in the Great Tit (Parus major). Anim. Behav., 25: 30 38. 
Pyke, G. H., H. R. Pulliam, and E. L. Charnov. 1977. Optimal foraging: a 

selective review of theory and tests. Quart. Rev. Biol., 52: 137-154. 
RoziN, P. 1969. Adaptive food sampling patterns in vitamin deficient rats. J. 

Comp. Physiol. Psychol., 69: 126-132. 
Westoby, M. 1974. An analysis of diet selection by large generaUst herbivores. 

Amer. Nat., 108: 290-304. 



'^^'S. CCMP. ZOOL 
MAP 1 n .Q 

B R E V I R Af 

Museiiiii of Comparative Zoology 

us ISSN 0006 9698 

Cambridge, Mass. 20 December 1978 Number 448 



SYSTEMATIC NOTES ON THE LOONS (GAVIIDAE: AVES) 
Robert W. Storer' 

Abstract. The following discussion documents the treatment of the loons for 
the revision of Volume 1 of Peters' Check-list of Birds of the World. The family is 
thought to be most closely related to the Charadriiformes. The Cretaceous genera 
Enaliurnis and Lonchodytes are thought to be convergent with rather than related to 
the loons {contra Brodkorb). Gavia immer and G. adamsii are considered full species 
but members of the same superspecies, while pacifica is considered conspecific with 
arctica. The subspecies Gavia siellata squantata. G. arctica suschkini. and G. immer 
elasson are not recognized. The sequence of forms followed is: stellaia. arctica 
pacifica. a. arctica. a. viridigularis. immer. adamsii. 

I. INTRODUCTION 

The purposes of this paper are to present reasons for decisions 
made in preparing the section on the Gaviidae for the revision 
of Volume 1 of Peters' Check-list of Birds of the World and to 
present some preliminary information gained while examining and 
measuring specimens of loons preparatory to making an analysis of 
geographic variation within the family. To date, I have examined 
over 2200 study skins of loons but still need to study material in 
several important collections before the final analysis can be made. 

For a group of only four species, there have been many diver- 
gences of opinion on such matters as the systematic position of the 
family, the sequence of species within the genus, the specific status 
of some of the forms, and the validity of several of the subspecies. 

'Museum of Zoology 

The University of Michigan 

Ann Arbor, Michigan 48109 



2 BREVIORA No. 448 

II. SYSTEMATIC POSITION OF THE LOONS 

There is general agreement that the living loons belong to the 
single genus Gavia. This, with one or two genera known only as 
fossils, constitutes the family Gaviidae. Several fossil genera re- 
ferred to families of uncertain systematic position have also been 
included in the order Gaviiformes, which otherwise contains only 
the Gaviidae. These fossils are discussed below. 

The varied opinions on the systematic position of the living loons 
within the class Aves have been well summarized by Sibley and 
Ahlquist (1972: 53-58). On the basis of fossil evidence (Storer, 1956) 
and studies of egg-white proteins (Sibley and Ahlquist, he. cit.), 
loons are believed most closely related to the gulls, auks, and 
shorebirds. Ideally, they should be placed next to or near the 
Charadriiformes, but this is impractical as only the first volume of 
Peters' check-list is being revised at present. 



III. SEQUENCE OF SPECIES 

In presenting a linear sequence of species, it is customary to start 
with the most primitive and end with the most advanced. In the case 
of the loons, there is fossil evidence bearing on the matter of 
primitiveness. 

The known fossil record of these foot-propelled diving birds goes 
back to the Upper Eocene of England {Colymboides anglicus) and 
the Aquitanian deposits (late Oligocene or early Miocene) of France 
(C minutus) and probably the Oligocene of North America (Gavi- 
ella pusilla). The Lower Cretaceous genus Enaliornis and the Upper 
Cretaceous genus Lonchodytes have been considered as families of 
the same order (Gaviiformes) as the loons by Brodkorb (1963a: 
220-221; 1963b: 56-60). However, the material of these genera is 
fragmentary, and the pieces of leg bones resemble those of loons in 
showing adaptations for swimming and diving but in other respects 
are quite different from them. The distal end of the carpometa- 
carpus of Lonchodytes pterygius, as shown in Brodkorb's figure 
(1963b: 59), differs so markedly in the relative sizes, angles, and 
positions of the distal articulations from those of the loons as to cast 
considerable doubt in my mind on the alleged phylogenetic relation- 
ship between Lonchodytes and Gavia. 



1978 SYSTEMATIC NOTES ON THE LOONS 3 

If, as I am convinced, these Cretaceous genera are convergent 
with the loons and if we accept Brodkorb's comment that the 
position of the little-known Upper Paleocene genus Eupterornis is 
"tentative," the earliest known loons {Colymhoides and Gaviella) 
were smaller than later ones. The best-known species (C minutus) 
was about the size of a teal, and, judging from the size of its wings, a 
strong flier (Storer, 1956: 423). The earlier species (C. anglicus) is 
poorly knojwn but was somewhat larger and apparently had rela- 
tively shorter wings than C minutus. It may have "represented a 
specialised offshoot from a more generalised gaviid stem, rather 
than a direct ancestral form of either the later Gavia species or C 
minutus" (Harrison, 1976: 66). The Oligocene species, Gaviella 
pusilla, was also smaller than the species of Gavia. Known only 
from the proximal end of a carpometacarpus, Gaviella differs from 
other loons in several respects and is probably not closely related to 
them (Wetmore, 1940: 30). Thus the limited evidence from fossils 
indicates that Gavia evolved from smaller birds which were less well 
adapted for diving and had relatively larger wings. While the 
modern genus Gavia contains several Pliocene species, studies of 
these species have not as yet provided information regarding the 
relative antiquity or phylogenetic relationships among the Recent 
species. 

The Red-throated Loon, Gavia stellata, is the most distinct 
species in both its winter and nuptial plumages, and for this reason 
it should be placed at one end of the sequence. The square white 
spots on the upperparts and the well-developed stripes and iri- 
descence on the head and neck of the other three species can be 
considered advanced characters. G. stellata is the smallest of the 
living loons and also has the lowest wing loading. To estimate this, I 
divided the weight of the bird by the square of the length (arc) of the 
folded wing. The means of this index for ten specimens of each of 
the species are as follows: stellata 2.00, a. pacifica 2.25, immer 3.03, 
and adamsii 3.17 grams/cm-. Although detailed comparative stud- 
ies have not been made, stellata is said to be able to take off from 
water more easily than the other loons and alone among them can 
take off from land (Palmer, 1962: 59). As pointed out above, small 
size and greater flying ability may be considered primitive charac- 
ters in loons, so I have placed stellata first on the basis of its 
relatively simple nuptial plumage, small size and good flying ability. 



4 BREVIORA No. 448 

As members of a superspecies, adamsii and immer should be placed 
next to each other. Arctica has been known to hybridize with immer 
(Hunter and Dennis, 1972; Robertson and Fraker, 1974) and 
therefore these two species should be adjacent on the list. Assuming 
the largest species to be the most advanced and lacking other strong 
evidence to the contrary, I prefer the sequence arctica, immer, 
adamsii to the reverse for these three species. If one considers the 
forms of arctica to represent a broken rassenkreis, as I do, the 
sequence should be either pacifica, arctica, viridigularis or the 
reverse. I prefer starting with the smaller, better-known form, and 
thus the sequence I have adopted is: stellata, arctica pacifica, a. 
arctica, a. viridigularis, immer, adamsii. 

IV. SPECIES PROBLEMS 

The Common and Yellow-billed Loons, G. immer and G. 
adamsii. — The breeding ranges of these forms were presumably 
separated during the last glaciation — that of immer lying south of 
the glaciers and that of adamsii in Yukon-Bering Sea refugia to the 
north (Rand, 1948: 317-318). Of the principal differences between 
the species, the color of the bill and of the iridescence on the head 
and the head patterns may have been selected for as reproductive 
isolating mechanisms; all are located where major specific differ- 
ences among other groups of diving birds such as penguins and 
grebes are found. On the other hand, evolution in bill shape in 
adamsii was presumably related to feeding. The upturned mandibles 
of this species and of stellata appear to be adaptations for bottom 
foraging. When birds are moving close over the bottom, the bill 
must be held below the body to catch bottom-living prey. In this 
position of the head, the lower part of the upturned mandible would 
lie nearly parallel to the bottom, making capture of prey in this 
situation easier than if the bill were shaped like those oi immer and 
arctica. The upturned bills o{ adamsii and stellata are probably an 
example of convergence, because there is no other reason to 
consider the former more like stellata than it is like either of the 
other two species. The difference in size between immer and adamsii 
might be accounted for by Bergman's rule, and like bill form, is 
probably not primarily related to reproductive isolating mechanisms. 

The effectiveness of the isolating mechanisms mentioned above is 
evident in the paucity or lack of known hybrids between the two 



1978 SYSTEMATIC NOTES ON THE LOONS 5 

species. There is an adult "female" in the Royal Ontario Museum 
(No. 76,360) found dead at Pt. Credit, Ontario on December 7, 
1957, which has been thought to be a hybrid between immer and 
adamsii (Godfrey, 1966: 11). In the recurved lower outline of the 
proximal segment of the mandibular rami (character 6 of Binford 
and Remsen, 1974: 115) this specimen resembles adamsii. The size 
of the remaining white squares on the back are within the range of 
those of adamsii and of males of immer, but not of females of 
immer. In size of wing, tarsus, and bill, the bird is well within the 
range of males of immer. In the other bill characters described by 
Binford and Remsen, the specimen is within the range of immer. 
While the possibility that this bird is an immer — adamsii hybrid 
cannot be completely discounted, I think it is more probably a male 
immer which was mis-sexed by the collector, who recorded nothing 
concerning the size or condition of the gonads. 

The status of pacifica. — The three forms of the Arctic or 
Black-throated Loon (arctica) complex {arctica, viridigularis, and 
pacifica) replace one another around the arctic except in Iceland 
and Greenland, where none occurs. Thus they might be thought of 
as forming a broken rassenkreis. The nominate form, arctica, ranges 
from the British Isles eastward across the northern parts of Eurasia 
and intergrades with viridigularis in Siberia. The latter form is 
found in western Siberia, largely south of the range oi pacifica, and 
in parts of western Alaska. The widespread North American form, 
pacifica, also breeds in northeastern Siberia. Portenko (1939) and 
Bailey (1948) have shown that the breeding ranges oi pacifica and 
viridigularis overlap in Anadyrland and western Alaska and re- 
ported no interbreeding. They concluded that the two were, there- 
fore distinct species, an opinion followed by Vaurie (1965). 

There are several reasons for doubting the specific status of 
pacifica. In the first place, those who have discussed the problem in 
the past have ignored one pertinent question: would arctica and 
pacifica interbreed if their ranges were to expand and meet? I think 
it likely that they would because the throat color is the same and the 
difference in size is less than that between paa/Zca and viridigularis. 
Secondly, there has been no thorough, detailed field study of 
pacifica and viridigularis in the area of overlap, and thirdly, there is 
evidence of interbreeding between the two forms. 

Of the two forms, viridigularis is larger in all measurements, has a 
green (not purple) sheen on the throat, and a darker gray nape. 



6 BREVIORA No. 448 

Green- or blue-throated individuals, otherwise indistinguishable 
from typically purple-throated individuals of pacifica are known to 
occur (Bailey, 1948: 140; Palmer, 1962: 45; Vaurie, 1965: 5), but 
whether this is a result of introgression with viridigularis or part of 
the normal range of variation oi pacifica is unclear. To date, I have 
measured and examined 12 examples of viridigularis and approxi- 
mately 270 o{ pacifica in breeding plummage. Of the latter, at least 
nine, or 3.3 percent have green or blue-green throats. All nine have 
measurements well within the range oi pacifica. In addition, a male 
from Savoonga, St. Lawrence Island, Alaska (Colo. Mus. Nat. Hist. 
No. 26,769) labelled "viridigularis" has measurements well within 
the range oi pacifica, although all above the means for males of that 
form. The nape is dark, like that of viridigularis, and it is either a 
very small example of that form or an intergrade. Two other green- 
throated birds may be intergrades: a male (Nat. Mus. Canada No. 
8,816) from Barter Island, Alaska, has a nape intermediate in color 
between those of the two forms, a long wing (3 1 8 mm. or nearly 1 .5 
standard deviations above the mean iov pacifica), a long tarsus (near 
the maximum for pacifica), but a small bill (below the mean for 
males oi pacifica); and an unsexed bird (Amer. Mus. Nat. Hist. No. 
348,959) from SE Victoria Island, Canada, has a light nape, long 
wing (321 mm.), a short tarsus (between the means for males and 
females of pacifica), a long bill (above the mean for males of 
pacifica) and a deeper bill than that oi any pacifica I have measured. 
Two purple-throated males with long wings (323 and 325 mm.), long 
tarsi, and bills somewhat above the mean for males oi pacifica, are 
also within the range of viridigularis in all these measurements and 
may be intergrades. While the sample of viridigularis which I have 
seen to date is too small to permit an accurate analysis of variation 
within that form, the presence of several probable intergrades 
suggests that interbreeding occurs between it and pacifica. 

V. SUBSPECIES PROBLEMS 

Gavia stellata squamata Portenko. — Vaurie (1965: 4) lists this 
subspecies as not well differentiated but warranting nomenclatural 
recognition, while Dement'ev and Gladkov (1968: 291) question 
its validity. I have examined 12 breeding adults from the range 
of this form from Franz Josef Land (3), Spitsbergen (7), and 
Bear Island (2), and I find that the plumage characters used to 



1978 SYSTEMATIC NOTES ON THE LOONS 7 

differentiate this form from nominate stellata (Vaurie, loc. cit.) are 
not consistent within the population, and as Dement'ev and Glad- 
kov (loc. cit.) pointed out, are found in some specimens from 
outside the range. On the evidence now available, I see no value in 
recognizing this subspecies. 

Gavia arctica suschkini Zarudny. — Although this race, based on 
migrant individuals from Russian Turkestan, was recognized by 
Peters (1931: 34), it is generally considered a synonym of the 
nominate race by recent authors (e.g. Dement'ev and Gladkov, 
1968: 297; Vaurie, 1965: 5). I have not seen the material on which 
this race was based and follow the above authors in not recognizing 
suschkini. 

Gavia immer elasson Bishop. — This subspecies was also recog- 
nized by Peters (1931: 35), but not by Vaurie (1965: 7) and many 
other recent authors. This species varies greatly in size, wing lengths 
of adults ranging at least from 287 to 411 mm., but I have not 
collected sufficient data to work out the pattern of geographic 
variation, which appears to be largely clinal. While realizing the 
possibility that there may be recognizable subspecies within this 
species, I prefer to consider immer monotypic at least until a 
thorough revision can be made. 

LITERATURE CITED 

Bailey, A. M. 1948. Birds of Arctic Alaska. Colorado Mus. Nat. Hist.. Popular 

Ser., 8.. 317 pp. 
BiNFORD, L. C, AND J. V. Remsen, Jr. 1974. Identification of the Yellow-billed 

Loon (Gavia adamsii). Western Birds, 5: III 126. 
Brodkorb, p. 1963a. Catalogue of fossil birds. Bull. Florida State Museum. 7: 

179-293. 
. 1963b. Birds from the Upper Cretaceous of Wyoming. Proc. Xlllth 

Internal. Ornithol. Congress, pp. 55-70. 
DEMENT'EV, G. P., AND N. A. Gladkov. 1968. Birds of the Soviet Union, Vol. 2. 

Israel Program for Scientific Translations, Jerusalem, xi + 553 pp. 
Godfrey, W. E. 1966. The birds of Canada. Nat. Mus. Canada Bull., 203. 428 pp. 
Harrison, C. J. O. 1976. The wing proportions of the Eocene diver Co/i/H/'o/V/ei 

anglicus. Bull. Brit. Ornithol. Club, 96: 64-65. 
Hunter, E. N., and R. H. Dennis. 1972. Hybrid Great Northern Diver X Black- 
throated Diver in Wester Ross. Scottish Birds, 7: 89-91. 
Palmer, R. S. 1962. Handbook of North American birds. Vol. 1. Yale Univ. 

Press, ix + 567 pp. 
Peters, J. L. 1931. Check-list of birds of the world. Vol. 1. Harvard Univ. Press. 

xviii + 345 pp. 



8 BREVIORA No. 448 

PoRTENKO, L. A. 1939. Inst. Polyar. Zemled. Zhivotn 6, Fauna Anadyr. Kraya, 

Ptitsy, 2. pp. 144-152. 
Rand, A. L. 1948. Glaciation, an isolating factor in speciation. Evolution, 2: 

314-321. 
Robertson, 1., and M. Fraker. 1974. Apparent hybridization between a Com- 
mon Loon and an Arctic Loon. Canadian Field-Nat., 88: 367. 
Sibley, C. G., and J. E. Ahlquist. 1972. A comparative study of the egg white 

proteins of non-passerine birds. Peabody Mus. Nat. Hist. Bull. 39. vi + 276 pp., 

37 figs. 
Storer, R. W. 1956. The fossil loon, Colymboides minutus. Condor,58: 413-426. 
Vaurie, C. 1965. The birds of the Palearctic fauna. Non-Passeriformes. H. F. & 

G. Witherby. xx + 763 pp. 
Wetmore, a. 1940. Fossil bird remains from Tertiary deposits in the United 

States. J. Morph., 66: 25-37. 



''^l;s. comp. zoou 

L/DRARY 

MAR 3 8 1985 

'HARVARD 

B R E V I 0"1R' A 

Museum of C()iu])arative Zoology 



us ISSN 0006 9698 



Cambridge. Mass. 21 Februar y 1979 Number 449 

SOUTH AMERICAN ANGLES: THE SPECIES GROl PS. 
2. THE PROBOSCIS ANGLES (ANOLIS LAEVIS GROUP). 

Ernest E. Williams' 

Abstract. The Anolis laevls group is known from three species represented by 
only nine specimens, all males, from four localities. All are distinguished by a soft 
multi-scaled proboscis and form a graded series from Peruvian A. laevis (proboscis 
minimally developed) through Brazilian A. phyllorhinus (proboscis of moderate size) 
to Ecuadorian A. proboscis (proboscis very long). Color in life and habitat are 
known only for A. phyllorhinus. The proboscis is interpreted as primarily an 
intraspecific social signal increasing the virtual size of the animals. 

The most readily recognizable — at least in males — of all South 
American Anolis are the three species that have a nasal appendage 
or proboscis. The group may be described as follows: 
Anolis laevis species group 

Type species: A. laevis Cope 1876. 

Definition: (Many possibly useful details are not determinable in 
A. laevis and hence are not mentioned here.) Alpha anoles of 
moderate size (60 97 mm snout-vent length) distinguished by a soft 
multi-scaled nasal appendage projecting forward above the rostral 
scale. Four to 10 scales across the snout between the second 
canthals. Supraorbital semicircles in contact or separated by as 
many as three scales. Loreal rows two to five. Interparietal of 
moderate to large size in contact with the surpaorbital semicircles or 
separated by as many as three scales. Supralabials in contact with 
suboculars. Middorsal scales uniform or the median row raised into 
a crest of triangular scales. Tail crest single or double. Lamellae 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 



2 BREVIORA No. 449 

under fourth toes ranging between at least 17 and 26. Dewlap and 
postanals large in males. Females unknown. 

Distribution: Widely disjunct: (1) trans-Andean Ecuador; (2) 
southern tributaries of the Amazon in central Amazonia; (3) 
western Amazonia. 

INCLUDED SPECIES 

Anolis laevis Cope 1876 

(Fig. 1) 

Scytomycterus laevis Cope, Jour. Acad. Nat. Sci. Phila., NS 8: 165. 

Holotype. ANSP 11368, collected by Prof. James Orton. 

Type locality. "Between Moyabamba and Balsa Puerto on the 

river Huallaga in eastern Peru." 

Additional references: Boulenger, 1885: 56 (referred to the genus 
Anolis and placed between A. tigrmus and A. punctatus); Burt and 
Burt, 1933: 17 (listing only); Barbour, 1934; 154 (placed along with 
A. tigrinus as synonym of /I. transversalis; both synonymies quite 
mistaken); Williams, 1965: 6-13 (discussed as member of the 
punctatus group sensu lato)\ Peters and Donoso-Barros, 1970: 57 
(citation only); Malnate, 1971: 358 (listing of type in the Philadel- 
phia Academy). 

Anolis phyllorhinus Myers and Carvalho 1945 
(Figs. 2 and 3) 
Anolis phyllorhinus Myers and Carvalho, Bull. Mus. Nac. NS 
No. 43: 2. 

Holotype. MN (Rio de Janeiro) 1804, adult male collected by 
Alexandre Parko on June 14, 1943. 

Type locality. "Borba, lower Rio Madeira, State of Amazonas, 
Brasil." 

Additional references: Williams, 1965: 8-13 (discussed as member 
of the punctatus group sensu lato, first mention of the second 
known specimen MZUSP 7118, Jacareacanga, Rio Tapajoz, Para, 
Brasil'); Peters and Donoso Barros, 1970; 63 (citation only). 
Anolis proboscis Peters and Orces 1956 
(Figs. 4, 5, and 6) 
Anolis proboscis Peters and Orces, Breviora No. 62: 2. 



'Note that the region between the Madeira and Tapajoz is one of Haffer's (1974, 
p. 70) "distribution centers" for endemic Cis-Andean birds. 



1979 



ANOLIS LAEVIS SPECIES GROUP 




4 BREVIORA No. 449 

Holotype. MCZ (Cambridge) 54300, a mature male collected by 
Antonio Proano during April 27-29, 1953. 

Type locality. "Neighborhood of Cunuco. a small town at 1200 
meters elevation, five kilometers northwest of Mindo, on the south 
bank of the Rio Mindo, a northern tributary of the upper Rio 
Blanco, in Pichincha Province, Ecuador." (The five additional 
specimens — USNM 207671-73, in the National Museum of 
Natural History, Washington, D.C.; IPN 761 1, 7612, in the collec- 
tion of Gustavo Orces V in Quito, Ecuador — are from Mindo or 
"region of Mindo.") 

Additional references: Williams, 1965: 8 13 (discussed as member 
of the punctatus group sensu lato); Peters, 1967: 13, 17 (key and 
citation); Peters and Donoso Barros, 1970: 63 (citation only). 

Ecological information. None except for A. phyllorhinus Myers 
and Carvalho: "Mr. Parko collected the holotype of Anolis phyl- 
lorhinus in the capoeira (second growth or low jungle) that sur- 
rounds the town of Borba. This locality is in the Amazonian 
lowlands on the lower Rio Madeira, about 90 miles (airline) south- 
southeast of the city of Manaus. The lizard was caught on a low tree 
while Mr. Parko was collecting butterflies. . . ." 

Distinguishing characters of the species. A. laevis differs from the 
two other species in the small number of scales between the second 
canthals (4, rather than 9 or 10), the low number of loreal rows (2, 
rather than 3 to 5) and in having a very rudimentary proboscis. A. 
proboscis has a crested dorsum and tail and a very long proboscis, 
while A. phyllorhinus lacks such crests and in fact has two rows of 
scales dorsally on the tail and a shorter proboscis. Table 1 lists the 
standard scale characters for each species. 

Size. Snout-vent length: laevis 60 mm, phyllorhinus 73 mm, 
proboscis 74 mm. Tail: laevis 79 mm, phyllorhinus 153 mm, 
proboscis 97 mm. Hind limb length: laevis 39 mm, phyllorhinus 
49 mm, proboscis 42 mm. 

Color. Laevis: Cope reported the preserved specimen as follows: 
"Color above dark gray, below pigmented white (in spirits). The two 
colors are abruptly defined between the orbit and there are brown 
spots behind the axilla. Tail distinctly annulate." 

Phyllorhinus — Myers and Carvalho: "Color in alcohol (formalin- 
fixed) plain, dull, bluish gray above, lighter beneath, with no sign of 
a color pattern anywhere save on the toes and undersides of the legs. 



1979 



ANOLIS LAEVIS SPECIES GROUP 




a. 



BREVIORA 



No. 449 



Table 1. Scale counts and characters. 



lac vis 

soft protuberance 

covered with ? 

small scales 



N = I 



phvlldihinus 

leaf-like 

laterally 

compressed with 

granular scales 

N = 2 



proboscis 

leaf-like 

laterally 

compressed with 

elongate scales 

N = 6 



scales 




between second 




canthals 


4 (Cope 


scales between 




semicircles 





scales between 




interparietal 




and semicircles 





loreal rows 


2 


rows between 




suboculars and 




supralabials 





labials to 




center of eye 


9 


sublabials in 




contact with 




infralabials 


? 


median gular 




scales in contact 




with mental 




between sublabials 


7 


middorsal rows 




enlarged 






tail crest 



double row, 
no crest 



10 



1-2 

4-5 



6-7 



5-6 



9 10 
1-3 

1-3 

3-5 



9-12 



2-3 



double row, 
no crest 



(crest of tri- 
angular scales) 

crest present 



lamellae 4th toe 



25 26 



17 21 



1979 



ANOLIS LAEVIS SPECIES GROUP 
I 




iV-v^-t 



8 BREVIORA No. 449 

The undersurfaces of the legs and arms bear a fine, Ught mottUng, 
perhaps better described as a bluish reticulation on a yellowish 
ground color. The undersurfaces of the digital expansions are dark. 
The superior surfaces of the toes and fingers (but not of the feet and 
hands) are light, with bold, blackish crossbands. The tail gradually 
becomes brownish toward the end. A color sketch made by Mr. 
Parko from life shows the creature bright blue-green, the dewlap 
yellowish, with traces of pink or red on the toes, the top of the 
snout, and the end of the rostral appendage." Mr. Parko's notes say 
the beast immediately changed from bright blue-green to brown 
when captured, but changed back to green on preservation. 

Proboscis — Peters and Orces: "Dorsal ground color (in alcohol) 
a dull gunmetal blue, with irregular, parallel, horizontal black 
streaks along vertebral margin. These streaks vaguely line up and 
form a pair of lateral bands between the limbs. Dark brownish- 
black spot at shoulder. Limbs and lateral surfaces spotted with light 
yellowish white; limbs vaguely barred with darker blue; skin in 
interstices between scales of dorsum of digits light, giving appear- 
ance of banding on fingers and toes. Dorsum of head unicolor, as 
back; temporal region and lips lighter, with faint reddish or purplish 
tinge and a marked light spot over ear opening. Ventral surfaces 
generally lighter, with vague reddish tints on chest and chin; belly 
stippled with light spots. Venter of limbs heavily spotted with white 
proximally, becoming totally white on the foot. Tail alternately 
barred with gunmetal blue and blackish bands approximately equal 
in width." 

TAXONOMY AND NOMENCLATURE 

The single recognition character — the proboscis — differs 
considerably in the three known species, which are abundantly 
distinct. The extent of the differences might reasonably raise the 
issue of parallel evolution, but the proboscis is so singular a feature 
that it seems subjectively more probable that the known species are 
the relicts of a once much larger radiation. It is also very possible 
that there are more species still to be discovered, perhaps with quite 
restricted distributions. 

Jt is clear that the first adumbrations of a proboscis are found in 
A. putu lulus and some of its relatives. In these the rostral scale and 
rostra] area are produced to overhang the lower jaw. In laevis there 



1979 



ANOLIS LAEVIS SPECIES GROUP 




10 HRl MORA No. 449 

appears to be a soft protuberance which involves the smaller post- 
rostral scales. In A. phvllorhinus and A. proboscis the rostral 
remains at the same level of development as in laevis but the area 
abo\e the rostral is expanded into an impressive nose leaf, longest in 
A. prohoscis. 

The distinction between the proboscis and punctatus group is 
thus perhaps somewhat artificial. The morphological sequence is, 
however, clear and leads from proboscis at one extreme back to 
pufuiaius at the other. I find it convenient to make the distinction at 
the point at which the scales and tissues above the rostral are 
involved in the swelling. Since the relation of this area to the rostral 
is very similar in the three species, it provides greater plausibility for 
the contention that this is a single lineage. It is not, on the other 
hand, clear that A. punctatus is directly related to the species that 
mimic it by means of the presence of a projecting rostral in the male. 
However, any decision on this point is necessarily tentative, and I 
emphasize convenience rather than certainty of phyletic affinity as a 
basis for recognition of this group. 

Nothing is known of the function of the proboscis (see below) and 
all nine specimens of this group thus far collected have been males. 
This has raised (e.g. Peters and Orces, 1956) the issue that the 
proboscis anoles might be the males of species already described on 
the basis of females. However, there are few described species still 
known only from females. Also the other external characters of each 
of the proboscis anoles are such that it would require a degree of 
sexual dimorphism beyond anything known in lizards to make any 
of the three proboscis species possibly conspecific with any other 
described anoles. In the case of A. punctatus and the other species 
with projecting rostrals, this and the presence of a dewlap, enlarged 
postanal scales and sometimes minor differences in color distinguish 
the males. Scale counts and other aspects of morphology do not 
significantly differ between the sexes. 

Only A. proboscis is known from series (the type, three newly 
collected specimens given to the United States National Museum by 
Orces, and two in the Orces collection in Quito). A. phvllorhinus is 
represented by only two specimens, the type and a badly preserved 
second specimen. A. laevis is still known only from the type, which 
is now in extremely poor condition. 



1979 



ANOLIS LAEVIS SPECIES GROUP 



11 




12 BREVIORA No. 449 

EVOLUTIONARY ASPECTS 

Three species, nine specimens, four localities and one note on 
habitat are the sum of our objective data on the proboscis anoles. 
They provide little opening for discussion or for evolutionary 
speculation. 

It is possible, however, to place this minimal amount of data 
against a background of the much greater knowledge of anoles as a 
whole. It is a peculiarity of the proboscis anoles that in no other way 
are they known to be different or indeed especially interesting. The 
single aspect that appears striking is the proboscis itself. 

Other bizarre structures are known in anoles. The dewlap would 
be bizarre were it less common and less familiar. High tail fins 
supported by heightened caudal neural spines occur in a number of 
distantly related species. Casque heads — heads enlarged by crests 
and ridges — occur in several Anolis and in the related anole genera 
Phenacosaurus and Chaniaeleolis. All these structures — dewlap, 
tail fin, casque head and proboscis — have the effect of increasing 
the apparent size of the animals that bear them. In the case of the 
dewlap, there is evidence of use in agonistic encounters, and 
evidence (Rand 1967) that larger animals usually win in such 
encounters. 

Trivers (1976) has shown the importance of sexual selection for 
size in one species of Anolis; he has inferred the generality of the 
rule. There are, however, aspects of increase in real size that make 
the course of evolution complex. A real increase in size with its 
implied increase in weight can carry with it functional restrictions 
that compel changes in the life style of the species that exercises this 
option. Sometimes such changes are realized within the ontogeny of 
a single species. Sometimes we see them as the alternative strategies 
utilized by sympatric species (Schoener, 1970; Williams, 1972). 
Virtual changes in size — unreal but visually effective — are still 
another alternative. Virtual size has the advantage of introducing 
few if any functional problems. 

I have called attention to elongation of the head in many crown 
anoles (Williams, 1965). It is probable that this initially has a (still 
undefined) functional explanation. However, there are side effects: 
the change of head shape probably has its species-recognition 
aspects and the size of the head has its agonistic advantage aspects. 
Yet if these latter aspects have too much effect, the bony structure of 



1979 



ANOI.IS 1 AEVIS SPECIES GROUP 



13 




14 BREVIORA No. 449 

the head is elongated beyond its functional optimum. There will 
therefore again be a premium on virtual rather than real change of 
head shape. We see the beginning of such a phenomenon in Anolis 
punctatus and in some related species where (in males) only the 
rostral scale is enlarged. A. laevis carries this process a slight step 
further. A. phyllorhinus and A. proboscis have gone much further. 

It is significant that the probosces in A. phyllorhinus and A. 
proboscis are not similar in detail. One would expect this if there 
were functional aspects to the structure per se. On the contrary, it 
appears a similar visual effect may readily be achieved by only 
broadly comparable means. This corresponds with species recog- 
nition devices and intra-species display where there is no require- 
ment that structures be similar; rather the requirement is that they 
be different. (In view of the wide allopatry of the known proboscis 
anoles, there could be no selection for striking difference now. but 
perhaps it has existed in the past.) 

Within the Iguanidae, the basiliscines would seem to be quite 
parallel cases. Head casques — very different in structure — are 
present in all three genera; in Basiliscus, dorsal and tail fins are 
present, different and wonderfully conspicuous in three species, very 
reduced in a fourth. (Once a bizarre structure has been adopted as 
an intra- and interspecies signal, an option open to a new member of 
a group is to abandon the structure: absence of a signal is itself a 
signal.) The absence of a dewlap is a strategy adopted by two species 
in the very complex anole fauna of Cuba; the dewlap is consistently 
retained in the simplest faunas and is reduced only in the fauna 
almost as locally complex as Cuba, that of Hispaniola (Williams 
and Rand, 1977). 

Outside the Iguanidae, a number of species in the closely related 
family Agamidae show parallels to the proboscis anoles: Cerato- 
phora (Ceylon, three species, see M. Smith, 1935 and Taylor, 1953); 
Harpesaurus (Java, Sumatra, Nias, five species fide Wermuth, 
1967); Lyriocephalus (Ceylon, one species, Taylor, 1953) and Co- 
photis (Ceylon and Sumatra, De Roois, 1915). The probosces in 
these are all soft structures but considerably more variable in shape, 
number and squamation than those of the proboscis anoles. (Figs. 
7 9 show the differences within the genus Ceratophora.) For cha- 
meleons Rand (1961) has already discussed the function of the 
ornamentation that is so remarkably developed in the montane 



1979 



ANOLIS LAEVIS SPECIES GROUP 



15 




16 BRFVIORA No. 449 

species of Africa. In the horned chameleons, however, the ornament 
is a solid structure. In this case, the ornament may serve a physical 
function in aggression, as the horns do. 

Still further afield, Hopson (1975) has discussed the singular 
crests of hadrosaurian dinosaurs as cranial display structures. His 
paper (see also Molnar. 1977) also cites the parallels with mam- 
malian horns and has references to the extensive literature. 

Of course, alternative explanations are available for structures 
such as probosces in Ano/is, e.g. camouflage. However, such other 
functions, if they exist, do not disprove the display function of 
Anolis probosces or basiliscine crests. The value of a structure for 
more than one function is a familiar phenomenon and will, not 
surprisingly, be sustained by natural selection. 

ACKNOWLEDGMENTS 

The extant world material of proboscis anoles has been made 
available to me by the late James A. Peters, W. R. Heyer and 
Ronald Crombie of the National Museum of Natural History, by 
Gustavo Orces-V of the Escuela Polytecnica Nacional, Quito, 
Ecuador, by Edmond V. Malnate of the Academy of Natural 
Sciences, Philadelphia and by P. E. Vanzolini of the Museum of 
Zoology of the University of Sao Paulo. To all of them I am 
indebted for their help and kindness. I have discussed the function 
of the probosces with A. S. Rand. Robert Bleiweiss called my 
attention to the refugium between the Rios Tapajoz and Madeira 
cited by Haffer. Laszlo Meszoly has provided the illustrations of the 
proboscis anoles and of their counterparts in the agamid genus 
Ceratophora. This study has been supported in part by National 
Science Foundation grant GB 3773 IX. 

LITERATURE CITED 

Barboi R. T. 1934. The Anoles II. The mainland species from Mexico southward. 

Bull. Mus. Comp. Zool. 77: 119 155. 
Boui ENGER, G. A. 1885. Catalogue of the lizards in the British Museum (Natural 

History), 2nd ed., London, xiii + 492 pp. 
Bi Ri, C. E. AM) M. D. Bi'Ri. 1933. A preliminary checklist of the lizards of 

South America. Trans. Acad. Sci. St. Louis 28: 1-104. 
Cope, E. D. 1876. Report on the reptiles brought by Prof. James Orton from the 

Middle and Upper Amazon and western Peru. Jour. Acad. Nat. Sci. Phila. (2) 8: 

159 188. 



1979 



ANOLIS LAEVIS SPECIES GROUP 



17 




18 BREVIORA No. 449 

De Roou, N. 1915. The reptiles of the Indo-Australian Archipelago. I. Lacertilia, 

Chelonia, Emydosauria. E. J. Brill, Leiden: i-.\iv + 384 pp. 
Haffer, J. 1974. Avian speciation in tropical South America. Publ. Nuttall 

Ornith. Club No. 14. Cambridge. England, viii -•- 389 pp. 
Hopsov. J. A. 1975. The e\olution of cranial display structures in hadrosaurian 

dinosaurs. Paleobiol. 1: 21-43. 
M AL.NATE, E. V. 1971. A catalog of primary types in the herpetological collections 

of the Academy of Natural Sciences, Philadelphia (ANSP). Proc. Acad. Nat. 

Sci. Phila. 123: 345-375. 
Moi.NAR, R. E. 1977. Analogies in the evolution of combat and display structures 

in ornithopods and ungulates. Evol. Theory 3: 165-190. 
M'l [ Rs. G. S. AND A. Carvai HO. 1945. A strange new leaf-nosed lizard of the 

genus Anolis from Amazonia. Bol. Mus. Nac. Rio de Janeiro N.S. 43: 1-14. 
Peters, J. A. 1967. The lizards of Ecuador, a checklist and key. Proc. U. S. Nat. 

Mus. 119: 1-49. 
Peters, J. A. and R. Donoso-Barros. 1970. Catalogue of the Neotropical 

Squamata. Part 11. Lizards and amphisbaenians. Bull. U. S. Nat. Mus. 297: viii + 

293 pp. 
Peters, J. A. and G. Orces-V. 1956. A third leaf-nosed species of the lizard 

genus Anolis from South America. Breviora No. 62: 1-8. 
Rand. A. S. 1961. A suggested function of the ornamentation of East African 

chameleons. Copeia 1961: 411 414. 
. 1967. Ecology and social organization in the iguanid lizard, Anolis 

lineatopus. Proc. U. S. Nat. Mus. 122: I 79. 
ScHOENER. T. W. 1970. Size patterns in West Indian Anolis lizards. II Correla- 
tions with the sizes of particular sympatric species — displacement and 

convergence. Amer. Natur. 104: 99 103. 
Smith. M. 1935. The Fauna of British India, including Ceylon and Burma 

Reptilia and Amphibia. IL Sauria. Taylor and Francis. London: i-.xiii + 440 pp. 
Taylor, E. D. 1953. A review of the lizards of Ceylon. Univ. Kans. Sci. Bull. 35: 

1525-1585. 
Trivers, R. L. 1976. Se.vual selection and resource-accruing abilities in Anolis 

garmani. Evol. 30: 253-269. 
Wermhth. H. 1967. Liste der rezenten Amphibien und Reptilien. Agamidae. Das 

Tierreich 86: i-.\iv + 127 pp. 
WiEi lAMS. E. E. 1965. South American anoles (Sauria, iguanidae): two new 

species of the punctaius group. Breviora No. 235: 1-15. 
. 1972. The origin of faunas. Evolution of lizard congeners in a complex 

island fauna: a trial analysis. Evol. Biol. 6: 47 90. 
Wii I lAMs, E. E. AND A. S. Rand. 1977. Species recognition, dewlap function. 

and faunal size. Amer. Zool. 17: 261 270. 



1979 



ANOLIS LAEVIS SPECIES GROUP 



19 




MUS. COM P. ZOOU 
LIORARY 

MAR 1 8 1985 

HARVARD 



B R E V I 0^'E""A 

Museum of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 21 February 1979 Number 450 

A NEW SPECIES OF MOENKHAUS/A FROM THE 

MATO GROSSO REGION OF BRAZIL 

(PISCES: CHARACIDAE) 

William L. Fink' 

ABSiR\t 1. A new species of the characid fish genus XUn'nklwitsia is described trom 
the Arinos River of the Mate Grosso region of Brazil. A brief discussion of some ot 
the characters used in analyses of characid relationships is included. 

INTRODUCTION 

The fish described here as new presents some interesting problems 
regarding current classification criteria for characids. These prob- 
lems arise primarily because the system now in use, which dates 
from the work of Eigenmann (1917). is obsolescent and no longer 
able to incorporate the diversity of the fishes it was intended to 
classify. Most of the characters which are used to "define" genera 
and higher categories have been found to be inadequate to express 
hypotheses about relationships within the family. The bases of the 
subfamilies Cheirodontinae and Tetragonopterinae have been re- 
cently reviewed by several authors (see Fink and Weitzman. 1974) 
and this critical reappraisal is being carried on in other areas within 
the Characidae. 

The characters which cause problems in hypothesizing the rela- 
tions of the fish herein described are lateral line length and color 
pattern. The former character is of major importance in the 
"Eigenmann" system while the latter has been virtually ignored by 
characid systematists. Both of these characters are discussed below 
after the description. 



'Museum of Comparative Zoology, Harvard lini\ersity. Cambridge. Mass. 02138. 



2 BREVIORA No. 450 

The methods of taking morphometric and meristic data used in 
this analysis are described in Fink and Weitzman ( 1974). Propor- 
tions are given as percent of standard length (SL). Paratypes have 
been deposited in the following institutions: Academy of Natural 
Sciences of Philadelphia (ANSP), British Museum of Natural 
History (BMNH), California Academy of Sciences, San Francisco 
(CAS), National Museum of Natural History, Washington, D.C. 
(LJSNM), Museum of Comparative Zoology, Cambridge, Mass. 
( MCZ), Museu de Zoologia, Universidade de Sao Paulo ( MZUSP), 
and the Zoologisch Museum, Universiteit van Amsterdam (ZMA). 

hh>enkhausia phaconoia, new species 
Figures 1, 2 

Holotype: MZUSP 13793; Brazil, Mato Grosso, from the shores 

of an island "below Rio dos Peixes," Rio Arinos, 

Jurena-Tapajos drainage; collected by Harald Schultz, 

16 June 1962; 32.6 mm SL. 
Paratypes: ANSP 135924, same data as holotype, one specimen 

26.7 mm SL. 

BMNH 1978.9. 12.1, same data as holotype, one speci- 
men 28.8 mm SL. 

CAS 41707, same data as holotype. one specimen 29.1 

mm S.L. 

FMNH 83856, same data as holotype, one specimen 

25.6 mm SL. 

MCZ 54087, same data as holotype, three specimens 

27.5-29.9 mm SL. 

MZUSP 13794, same data as holotype, one specimen 

19.5 mm SL. 

MZUSP 13795, same data as holot\pe, one specimen 

20.6 mm SL. 

USNM 218798, same data as holotype, twenty speci- 
mens 16.1 36.4 mm SL. 

ZMA 115.273, same data as holotype, one specimen 
24.1 mm SL. 

Di'scriplion 

See Tables 1 and 2 for variable meristic and morphometric data. 
Body elongate, compressed laterally. Predorsal body profile slightly 



1979 



NEW SPECIES OF MOENKHAUSIA 




4 BREVIORA No. 450 

convex with a slight concavity at nape. Dorsal body profile abruptly 
angles ventrally along dorsal-fin base, then becomes straight or 
slightly convex to adipose fin where profile dips ventrally a small 
amount, then continues as a nearly straight line to dorsal procurrent 
caudal-fin rays. Ventral profile gently rounded to anus; steepest 
inclination ventral to jaws. Ventral profile protrudes ventrally its 
greatest distance just anterior to pelvic-fin insertions. Body profile 
along anal fin-base nearly straight, or slightly convex; at anal-fin 
termination, profile nearly straight or slightly concave to procurrent 
caudal-fin rays. 

Maxillary with 3-7 teeth; in specimens below 21 25 mm SL, teeth 
mostly conical; specimens in that size range usually with dorsalmost 
tooth tricuspid (rarely with 4 cusps); specimens above that range 
usually with tricuspid maxillary teeth. 

Premaxillary outer row teeth, 2-5, larger fish usually with larger 
tooth number; in smaller specimens teeth usually conical; in speci- 
mens above about 24 mm SL, teeth usually tricuspid. 

Premaxillary inner row teeth, 5-6, each tooth usually smaller 
than tooth medial to it. Ontogenetic changes include replacement of 
earlier teeth by teeth with a greater number of cusps until adult 
condition reached. Smaller specimens (about 16-19 mm SL) usually 
have medial three teeth with 3 cusps, with lateral teeth having I or 2 
cusps, giving a tooth cusp formula of 3331 1 or 33321. In the size 
range 22-25 mm SL there is a variable pattern: medial tooth 3 4 
cusped, second tooth 3-5 cusped, third tooth 3 5 cusped, fourth 
tooth with 3 cusps, teeth lateral to those with 1-2 cusps. In adults, 
inner row premaxillary tooth formula usually 45533 or 45433. 

Dentary teeth show ontogenetic trends similar to those exhibited 
by inner row premaxillary teeth, replacement teeth usually with 
more cusps. Small specimens usually have a formula 3331 +(6 10 
very small conical teeth); adults usually have tooth formulae 
ranging from 4333 +(6-10) to 5353 +(6 10); second tooth from 
symphysis is offset anteriorly in adults and rarely has more than 3 
cusps. 

Fontanels moderate, that pari anterior to the epiphyseal bar 
slightly more than two-thirds as long as that part posterior to bar. 
Gill rakers moderate. Infraorbital bones well ossified, infraorbital 3 
wide, with a narrow naked area behind and below. 



1979 NEW SPECIES OF MO£A'A:///l(y'5//l 




^jlW^^^^ 



Figure 2. Dentition of Moenkhaiisia phaeonota, paratype, USNM 218798, 
28.2 mm SL. 



6 BREVIORA No. 450 

Scales moderately large, thin, cycloid with concentric circuli and 
about 2 7 radii on the exposed posterior field. Lateral line with a 
slight ventral curve along body, complete, with scales perforated to 
end of caudal base. Scales above lateral line 5; scales below lateral 
line 4. Scale sheath of about 4-7 scales along anterior anal-fin base. 
Scales extend over about 1/5 to 1/4 of caudal-fin rays. Small 
accessory scales present ventral to pelvic-fin insertion. 

Dorsal-fin origin anterior to anal-fin origin, posterior to pelvic fin 
insertion, nearer eye than caudal-fin base. Third or fourth ray of 
dorsal fin longest with posterior rays shorter, forming a straight or 
slightly convex posterior margin to fin. First unbranched ray of anal 
fin usually not visible externally; last unbranched anal-fin ray and 
first through fourth or fifth branched rays longer than posterior 
rays, forming a slightly concave fin margin. Pectoral fin low on 
body, pointed or slightly rounded, reaching to or beyond pelvic-fin 
insertion. Pelvic fin with distal tip reaching from just anterior to just 
posterior to anal-fin origin; i 7 rays. Caudal fin with 10 9 principal 
caudal-fin rays; fin forked, not split to base. Dorsal and ventral 
caudal-fin lobes equal in size. Dorsal and ventral procurrent caudal- 
fin rays are equal in size and extend cephalad same distance. All fin 
rays without hooks. 

Color in alcohol. — Ground color light brown. Nape, top of 
head, and jaws covered with numerous small br9wn melanophores. 
Back with a reticulate pattern of dark brown melanophores. A 
broad stripe almost devoid of melanophores extends from just 
above and behind eye along side of body, terminating ventral to 
dorsal procurrent caudal-fin rays. Below pale stripe is a broad dark 
stripe consisting of numerous melanophores extending from cheek 
and opercle behind eye, across body side and onto caudal peduncle 
and posteriorly onto about six of middle caudal-fin rays. Immedi- 
ately posterior to opercle on second through fourth lateral scales, 
melanophores are extremely dense, forming a large humeral spot 
about two-thirds diameter of eye. Posteriorly, above anal fin and on 
caudal peduncle, melanophores very dense and tend to concentrate 
along junctions of myomeres. Immediately dorsal to anal fin, 
melanophores abruptly become less dense, leaving a pale stripe at 
anal-fin base which is continuous with pale color of belly. Dorsal fin 
with melanophores concentrated on interradial membrane between 
first and fifth branched rays. Pectoral and pelvic fins with a few 



1979 NEW SPECIES OF A/O£A'A7//10'5//4 



Table 1 . Measurements of Moenkhausia phaeonota 



Character 


N 


Range 


X 


s 


Holotype 


Standard Length (SL) 


31 


16.1-36.4 


25.8 


5.670 


32.6 


% SL 












Greatest Depth 


29 


25.6-35.5 


30.2 


2.302 


29.1 


Eye-Dorsal Fin Origin 


31 


34.7-38.2 


36.7 


0.890 


36.2 


Dorsal-Fin Origin — 


31 


50.5-56.2 


53.3 


1.307 


52.5 


Base of Caudal Peduncle 












Snout-Dorsal Fin Origin 


31 


50.3-53.4 


51.6 


0.737 


50.6 


Snout-Pectoral Fin Origin 


31 


24.7-29.8 


26.4 


1.017 


25.5 


Snout-Pelvic Fin Origin 


31 


42.4-47.2 


45.3 


1.164 


44.8 


Snout-Anal Fin Origin 


31 


55.7-60.7 


58.8 


1.157 


59.2 


Dorsal Fin Length 


25 


26.0-31.7 


28.1 


1.461 


30.7 


Pectoral Fin Length 


30 


18.5-24.6 


21.9 


1.448 


19.3 


Pelvic Fin Length 


31 


10.8-16.4 


14,6 


1.217 


14.1 


Head Length 


31 


19.4-28.6 


25.2 


2.141 


24.5 


Eye Diameter 


31 


9.6-13.8 


11.3 


1 .038 


10.1 


Snout Length 


31 


6.2- 7.8 


6.9 


0.384 


6.4 


Bony Interorbital 


31 


7.5- 9.0 


8.3 


0.354 


8.0 


Caudal Peduncle Depth 


31 


8.3-10.7 


9.5 


0.659 


9.8 


Caudal Peduncle Length 


31 


10.4-14.3 


12.6 


l.OIl 


11.3 



8 BREVIORA No. 450 

scattered melanophores on interradial membranes. Caudal fin with 
melanophores along edges of all rays, in addition to the extremely 
dark middle caudal rays mentioned above. Anal fin with melano- 
phores loosely scattered on interradial membranes, more concen- 
trated between anterior rays and especially along outer margin of 
fin. Adipose fin with many small melanophores. 

Etymology. — From phaios, meaning dusky brown and the suffix 
— nota, meaning having the attribute or quality of. The name refers 
to the color pattern of the fish. 

DISCUSSION 

This species, according to Eigenmann (1917), belongs in the genus 
Moenkhausia, which is diagnosed by the following combination of 
characters: complete lateral line, partially scaled caudal fin, five 
inner row premaxillary teeth and a naked area between the third 
infraorbital and the lower limb of the preopercle. However, in color 
pattern, preserved specimens are extremely similar to several species 
of Hyphessobrycon. A large number of Moenkhausia species were 
examined durng this study in an attempt to find a potential sister 
group for M. phaeonota (see Comparative Material Examined) but 
no currently recognized species was considered to have any derived 
characters in common with it. In Eigenmann's (1917) key to the 
species of Moenkhausia, those that are included in the couplet 
"depth usually more than 2.75 in length" are most similar to M. 
phaeonota in general body shape but they have marked color 
differences and in most cases meristic count differences. Specimens 
identified as M. dichroura (Kner) by Eigenmann (USNM 86792) 
approach M. phaeonota in both anal-fin ray number and lateral 
scale number, but are very different in color pattern, in being 
heavier bodied, and in having a larger third infraorbital. Of the 
more recently described species, M. takesi Gery is phenetically more 
similar to M. phaeonota than other members of the genus. Both are 
fairly similar in body and head shape, although M. takesi is 
"stouter." Moenkhausia phaeonota has more predorsal scales (1 1-13 
versus 9), more lateral scales (36-38, usually 36, versus 33), more 
anal-fin rays (iii, 22-25, usually 22 or 23, versus iv, 18-19) [The 
holotype of M. takesi has four unbranched anal-fin rays rather than 
three as reported by Gery (1964)], and fewer gill rakers (17-21, 
usually 18, versus 20-21). The color pattern of M. takesi in life and 



1979 r<EW SPEC\ES OF MOENKHA US/A 



Table 2. Meristic counts of Moenkhausia phaeonota 

Character N Range x s Holotype 



8.9 0.250 



23.1 0.854 23 



10.7 0.599 



7.0 0.000 



35.5 1.635 37 



Branched 


Dorsal 




8 


9 








Fin Rays 


^ 


31 


2 


29 








Branched 
Fin Rays 


.Anal 


31 


22 
7 


23 
18 


24 


25 
3 




Branched 


Pectoral 




9 


10 


11 


12 




Fin Rays 




31 


1 


9 


20 


1 




Branched 


Pelvic 




7 










Fin Rays 




31 


31 










Lateral 






33 


34 


35 


36 


37 


Scales 




11 


2 


1 


2 


1 


4 


Perforated 




34 


35 


36 


37 


38 


Lateral Scales 


9 


2 


1 


2 


3 


1 



36.0 1.414 37 

Scales Around 12 13 14 

Caudal Peduncle 8 ~1 H ~6 13.6 0.744 14 

Predorsal \^ ]1 ^ ^_1 

Scales 26 2 18 ^ ^ 11.2 0.710 13 

Gill r7 |8 ]9 20 2] 

Rakers 13 2 9 ^ ^ ~[ 18.2 0.987 17 

Precaudal 13 14 

Vertebrae 31 TT 20 13.6 0.486 14 

Caudal 23 24 25 

Vertebrae 31 Ts 72 H 23.5 0.568 23 

Total 37 38 

Vertebrae 31 28 ^ 37.1 0.301 37 



10 BREVIORA No. 450 

in preserved specimens consists primarily of dense black melano- 
phores dorsal to a light lateral stripe and includes a dark area on the 
caudal peduncle and the area dorsal to the anal fin. The black band 
in M. phaeonola extends from the middle caudal-fin rays onto the 
base of the tail and continues to the posterior border of the opercle. 

In summary, M. phaeonola seems to share no unique characters 
with any known species of Moenkhausia and is rather different 
phenetically from the only Moenkhausia species which looks even 
slightly similar [M. takesi). Inasmuch as there is questionable 
phylogenetic reality to Moenkhausia, it is useful to look beyond the 
bounds of that genus to another group with which M. phaeonola 
seems to share a number of similarities. This leads us to a group of 
species within Hyphessobrycon, the members of which share a 
specialized color pattern. This group includes H. metae Eigenmann 
and Henn, H. loretoensis Ladiges, H. peruvianus Ladiges, and 
perhaps H. agulha Fowler and H. herbertaxelrodi Gery. All of these 
species have a color pattern in preserved specimens which includes a 
band of dark brown or black chromatophores which generally 
extends along the lateral body from the eye to the caudal fin base. 
The area of greatest concentration of chromatophores is usually 
dorsal to the anal-fin base, and there is often a secondary concentra- 
tion in the "humeral" area behind the opercular aperture. In life, the 
lateral band is dark brown to black, and dorsal to it is a bright 
lateral stripe, usually gold, goldish-red or creamy white. In addition, 
in live specimens there is generally some red coloration on the 
caudal and dorsal fins, and in some species, dorsally on the eye. 
Preserved specimens of Moenkhausia phaeonola have a color 
pattern nearly indistinguishable from H. meiae of Colombia, and 
differ from that species primarily in meristic counts including: 
number of lateral scales (33-37, compared with about 33 in melae), 
number of branched anal-fin rays (22 25, compared with 17 19 in 
metae), and number of vertebrae (37-38, compared with 33 in 
melae). 

In view of this striking resemblance to these members oi Hyphes- 
sobrycon, I would propose a relationship of M. phaeonola with 
those species. That group is currently being revised by myself and S. 
H. Weitzman and M. phaeonola will be considered in that study, 
after material available to us has been further examined and 
analyzed. In the interim, the species described herein will be referred 



1979 NEW SPECIES OF iWC>£.VA'///lL'5//l II 

to the genus Moenkhausia, where it belongs according to the 
diagnostic characters of that series. A revision of this group of 
characids must include a redefinition of certain generic limits and a 
full review of these fishes, something which I am not yet prepared to 
present. 

COMPARATIVE MATERIAL EXAMINED 

Hyphessohrycon agulha Fowler, holotype, ANSP 39232, Brazil, 

Rio Madiera, about 200 miles E of Long W62°20'. Coll. E. A. 

Smith, 1912. 
Hyphessohrycon herbertaxelrodi Gery, holotype, USNM 196089, 

Brazil, Mato Grosso, Rio Paraguay Basin, Coxim on the Rio 

Tacquary. Coll. K. H. Stegemann, 1960. 
Hyphessohrycon loretoensis Ladiges, MCZ 52154, Colombia, Rio 

Loreto Yacu, in small creeks about 1 km from river mouth (Rio 

Loreto Yacii is a trib. of Rio Maranon), approx. 03°30'S, 

070°10'W. Coll. Alfonso Doaz, Dec. 1976. 
Hyphessohrycon metae Eigenmann and Henn, holotype, CAS (lU) 

13421, Colombia, Barizora, Rio Meta, Coll. M. Gonzales. 
Hyphessohrycon metae Eigenmann and Henn, paratypes, CAS (lU) 

13422, same data as holotype. 

Moenkhausia hrowni Eigenmann, holotype, FMNH 52733, British 
Guiana: Arnataima. Coll. C. H. Eigenmann, 1908. 

Moenkhausia collettii Steindachner. USNM 66228, British Gui- 
ana: Erukin Creek, lower Potaro River. Coll. C. H. Eigen- 
mann, 1908. Identified by Eigenmann. 

Moenkhausia copei (Steindachner), USNM 66230, British Gui- 
ana: Rockstone, Essequibo River. Coll. C. H. Eigenmann, 
1908. Identified by Eigenmann. 

Moenkhausia cotinho Eigenmann, USNM 66244, British Guiana: 
Tukeit, lower Potaro River. Coll. C. H. Eigenmann, 1908. 
Moenkhausia dichroura (Kner), USNM 86792, Bolivia: Rurrena- 
baque. Coll. N. E. Pearson, Oct. 1921. 

Moenkhausia eigenmanni Gery, holotype, USNM 198640. Colom- 
bia: Rio Marracacias into upper Rio Meta, at Restrepo. Coll. 
Ross Socolof, June, 1963. 

Moenkhausia lepidura gracilima Eigenmann, holotype, USNM 
120279, Brazil: Serpa. Coll. Thayer Expd. 1866. 



12 BREVIORA No. 450 

Moenkhausia takesi Gery, holotype, USNM 198136. Brazil: Rio 
Guana basin, near Belem do Para. Coll. Takese, 1963. 

Moenkhausia rohertsi Gery. holotype. USNM 200427. Peru: upper 
Amazon, near Iquitos. Coll. J. Roberts. 1963. 

ACKNOWLEDGEMENTS 

I wish to thank Sara Fink and Stanley H. Weitzman for editorial 
comments on the manuscript. James E. Bohlke and W. E. Esch- 
meyer generously loaned specimens of Hyphessohrycun meiae. 

Literature Cited 

EiGENMANN, C. H. 1917, The American Characidae. I. Mem. Mus. Comp. Zool., 

Harvard 43(1): 102 pp. 
Fink, W. L. and S. H. Weiizman. 1974. The so-called cheirodontin characids of 

Central America, with descriptions of two new species (Pisces: Characidae). 

Smithsonian Contrib. Zool. 172: 46 pp. 
Gery, J, 1964. Two new tetras from the lower Amazon basin. Trop. Fish Hobb. 

12(7): 13 15 and 59 69. 



B R E V f'W-W*^ 

MAR 1 8 1985 
Museum of Comparati¥#xvZjWlogy 

us ISSN 0006 9698 

Cambridge, Mass. 21 February 1979 Number 451 

A NEW SPECIES OF CYBOTOID ANGLE 
(SAURIA, IGUANIDAE) FROM HISPANIOLA 

Albert Schwartz' 

Absiract. Anolis strahmi. a new species of clivicolous cybotoid anole from 
Hispaniola, occurs both north and south of the Sierra de Baoruco, the mountain 
range associated with the Dominican Peninsula de Barahona. Two subspecies are 
recognized. Additionally, a new subspecies of Anolis lon^itlhialls Noble is named 
from the Peninsula; nominate A. I. longitihialis occurs on Isla Beata off the southern 
tip of the peninsula. Details of distribution and known ecology of the two species are 
given, and comparisons of the two species (and their respective subspecies) are made. 
A hypothetical evolutionary and geographical sequence relating these two species to 
parent A. wholes has been postulated. 

INTRODUCTION 

Anolis cyhotes Cope is the first-named of a series of related taxa 
of Hispaniolan anoles. A. cyhotes. sensu strict o, is a widespread 
mesophilic species of moderate size that occurs in wooded to rather 
open situations throughout much of Hispaniola. Although the 
species prefers mesic habitats, it does not completely shun situations 
which are xeric. In extremely arid areas, the species is often confined 
to oases or other shaded enclaves. It reaches elevations in excess of 
about 1,525 m, depending upon which named upland populations 
one accepts as subspecies of .4. cyhotes (see Schwartz and Thomas, 
1975:77). The species is also known from several of the Hispaniolan 
satellite islands: He de la Gonave. Isla Catalina. Isla Saona, Ile-a- 
Vache. lie de la Tortue. and He Grande Cayemite. Only on the first 
of these has an endemic subspecies been named. The present paper 



'Miami-Dade Commurity College North, Miami. Fla. 33167. 



2 BREVIORA No. 451 

does not attempt to deal with those named populations (armouri 
Cochran, duri.s Barbour, haeiianus Garman) that are or have been 
associated with A. cyhoies nomenclaturally. 

Three other species form the cybotes complex. A. shrevei (Coch- 
ran, 1939) is restricted to the uplands of the Dominican Cordillera 
Central; this has been recently affirmed as a distinct species by 
Williams (1975). A xerophilic ally of /4. cybotes is A. whitenmni 
Williams. This species centers in the Cul de Sac-Valle de Neiba 
plain, which today encompasses the area between the north and 
south paleoislands (Williams, 1961). east into the Llanos de Azua. 
A. w'hitemani and A. cybotes occur macrosympatrically in this 
region; the former occupies xeric situations whereas the latter 
occupies oases and riverine woods — shaded, mesic situations. 
There are also records of A. whitemani from the northwestern 
Haitian Presqu'ile du Nord Quest (Mole St. Nicholas) and near 
Gonaives. It is likely that these "isolated records" are one end of a 
linear coastal range from the Cul de Sac Plain along the Golfe de la 
Gonave onto the xeric northwestern peninsula, but records are 
absent to affirm this continuity. A. whitemani is also known from 
the xeric Monte Cristi region in northwestern Repiiblica Domini- 
cana. Most recently, Williams (1975) named A. marcanoi from the 
southern xeric slopes and associated plains of the Sierra de Ocoa, a 
southern affiliate of the Cordillera Central. A. marcanoi is sym- 
patric (and even rarely syntopic) with A. cybotes. Williams has 
given very pertinent details of this association, and Webster (1975) 
affirmed species-rank for A. marcanoi by electrophoresis. These two 
species differ slightly meristically, but in living animals the dewlap 
color and the head and body pattern are distinctive and dichoto- 
mous. 

Anolis loni^itihialis was described from Isla Beata, off the south- 
ern tip of the Peninsula de Barahona. Cochran (1934) first 
combined A. longitibialis with A. cybotes: the species are indeed 
very similar in general appearance and scutellation. It would seem 
quite logical that wide-ranging ^4. cybotes would have a local insular 
population on Isla Beata, despite the absence o{ A. cybotes from the 
Peninsula de Barahona itself, at least as far as Cochran's data ( 1941 ) 
suggested. 

In 1969, Richard Thomas collected a single large female cybotoid 
on the Peninsula de Barahona. This lizard was so much larger than 



1979 NEW SPECIES OF CYBOTOID ANGLE 3 

female A. c\ botes from the adjacent foothills of the Sierra de 
Baoruco that suspicion was aroused that it might represent A. c. 
loniiltlhlai'l.s. Comparisons were made, and it was determined this 
lizard was indeed much more similar in size and ventral scutellation 
to Beata lizards than to A. cyhotes from the nearby mountains. In 
1971, at a locality 9.6 km N of Pedernales, I collected cyhutes and 
longitihialis at the same locality; the habitat was grassy and shrubby 
pasture adjacent to limestone cliffs and their talus. Since Thomas's 
lizard was secured sleeping adjacent to a cave in limestone cliffs, it 
seemed a possibility that on the mainland longitihialis was limited to 
this habitat, whereas cyhotes preferred more shaded and less 
extreme habitats. There was no doubt, however, that we were 
dealing with two distinct species, A. cyhotes and A. longitihialis, 
which have limited sympatry and syntopy on the Peninsula de 
Barahona, and of which one {A. longitihialis) occurs also on Isla 
Beata. 

In June 1974, Fred G. Thompson of the Florida State Museum 
collected in the Repiiblica Dominicana on the north face of the 
Sierra de Baoruco near the Dominico-Haitian border, just northeast 
of the border post of El Aguacate. He courteously lent me his 
material from that expedition. Two anoles from 2 km NE of EI 
Aguacate are extremely distinctive; both are males, one is very large, 
and each has its head more attenuate with a different profile and is 
much less "jowly" than A. cyhotes. Most pertinently, the dewlaps 
were mustard-colored when I examined them 4 months later. 
Although this is not the dewlap color in life of this population, 
nevertheless, the retention of such deep and distinctive colors after 
preservation suggests the dewlaps are differently and brightly 
colored in life, in contrast to the pale colors of A. cyhotes. 

In December 1974, Michael H. Strahm and I made an attempt to 
reach Thompson's locality; however, the road to the north face of 
the Sierra de Baoruco from La Florida was in disrepair and our 
vehicle was unable to reach the high elevations (900 m) needed to 
secure the lizards. Dr. Thomson had advised me that the lizards 
were common on rocky and exposed roadcuts, but we never reached 
such a situation. In the summer of 1975 we were successful in 
reaching El Agucate from the north, where the road has high to 
moderate artificial "cliffs" along its eastern margin as it ascends the 
mountains. We finally found a population of Thompson's lizards 



4 BREVIORA No. 451 

and secured a series on three visits. My earlier contentions proved 
correct: 1) the dewlaps are very brightly colored, and 2) the lizards 
are large and are confined to creviced roadcuts in this area. Their 
status as a species distinct from A. cyhotes was confirmed by taking 
typical A. cyhoies on fenceposts across the road (5m) from the cliffs 
which harbored the new lizards. Thus we seemed to have an easily 
solvable problem: a derivative oi A. cyhotes which is very special- 
ized in habitat (cliff faces) occurring on the north face of the Sierra 
de Baoruco at elevations of about 900 m. The two species differ 
grossly in adult size, head shape, and dewlap color as well as in 
habitat; there was no question that they are distinct and sympatric. 

Also during the summer of 1975. Ernest E. Williams and William 
E. Haas, while collecting in the Peninsula de Barahona lowlands, 
encountered large cybotoids in a shaded and fairly mesic ravine near 
sea level at a locality 5 km SE and 2.9 3.0 km N of Pedernales. 
Superficial examination of their specimens in the field, and subse- 
quent collection of a short series taken there by Thomas Wiewandt 
and Diderot Gicca, all served to confirm that these lizards were very 
comparable to the series from high elevations on the north face of 
this same mountain range. The habitat of the new population near 
Pedernales is very like that of the El Aguacate specimens — a 
limestone ravine or wash that is much creviced; however, vegeta- 
tionally this ravine is much more shaded and mesic than the bare 
and exposed road cut near El Aguacate. 

In 1975 Richard Thomas secured a fine series oi A. longitihialis at 
a locality 17 km NW of Oviedo that represents the easternmost 
record for the species. This locality is a wooded but xeric ravine with 
creviced limestone walls, that lies on the eastern edge of the north- 
south limestone ridge which bisects the Peninsula de Barahona. 
There are specimens of A. longitihialis from west (Pedernales) and 
east (17 km NE of Oviedo) of the ravine which harbors the 
differently-colored dewlapped population discovered by Williams 
and Haas. Both species in this region show a predilection for 
limestone cliffs; they and the north slope lizards show remarkable 
agility and familiarity with their cliffs and are completely "at home" 
upon them. Other anoles (/4. cyhotes. A. hrevirostris Bocourt) occur 
on cliffs in this general region but rarely seek refuge in crevices; 
rather, when pressed, they drop to the ground to hide. This is not 
true of either the new species or of A. longitihialis both of which 



1979 NEW SPECIES OF CYBOTOID ANGLE 5 

almost invariably seek refuge in crevices (see additional comments 
below). I have ne\er taken the new species at night which suggests 
that the lizards sleep in crevices. A. loni^ifihla/is, however, has been 
secured while asleep on shrubs and low trees adjacent to cliffs. This 
implies that the two species differ in sleeping sites, although they 
may forage on the same vertical rocky faces. 

Williams (1975:7-8) made the following comment in his "justifi- 
cation" for the naming of A. marcanoi despite weak meristic 
differences between that species and A. cyhoies: "It becomes more 
obvious that, in addition to those species in which museum ta.x- 
onomists rejoice because they are very distinct in terms of the 
characters conventionally studied, there are in many groups valid 
biological species only imperfectly separable on museum characters, 
if at all. This phenomenon is only interesting in terms of the history 
of museums, not of biology. ... It will not be necessary in the near 
future to defend or specially comment on cases like that here 
described." The same comments, to some extent, apply here. I have 
no doubt that the peculiarly isolated populations herein described as 
a new species differ biologically from A. longitibialis. Except for 
some size and modal differences in scale counts, without preknowl- 
edge of the dewlap colors as a re-enforcing mechanism, one might 
be easily led into misinterpreting the situation. Verifying that A. 
longitibialis is distinct from A. cy botes required the chance en- 
counter of the two species syntopically after the lapse of two years. 
A. longitibialis geographically surrounds two of the three known 
populations of the new species. Differences between this new species 
and A. longitibialis are not subtle in life, but assigning long- 
preserved and discolored museum specimens to one or the other 
species is and will continue to be difficult. This fact, however, 
should not be a deterrent from naming a new taxon. Two biological 
entities (species) exist. It is the duty of the systematist to recognize 
this fact nomenclaturally and to present whatever data he may have 
to support his contention. 

Before proceeding to the new species, it is pertinent to examine 
variation in A. longitibialis. Noble's (1923) description is detailed 
but lacks data on some features which are important when com- 
paring the Beata population with the Peninsula de Barahona 
population. These two samples differ from each other and from the 
new species about to be named in a number of ways; the differences 



6 BREVIORA No. 451 

in the former case are those which are accepted as being of 
subspecific rank. Accordingly, I first define A. lougitihialis in those 
terms which will differentiate it from the new species, and then 
name a new mainland subspecies of A. longitihialis. 

Anolis longitihialis Noble 

Anolis longitihialis Noble, 1923, Amer. Mus. Novitates, 64:4. 

Type locality: Isla Beata, Republica Dominicana. 

Holotype: AMNH 24329. 

Definition. A cybotoid anole characterized by small size (males to 
72 mm, females to 59 mm), supraorbital semicircles occasionally not 
in contact, modally I 1 scales between the supraorbital semicircles 
and the interparietal, smaller median dorsal scales (35 to 57 in snout- 
ear distance), subocular scales often in contact with supralabial 
scales, usually 4 or 5 postrostral scales, few fourth toe lamellae 
(15-21), dewlap pale yellow to pale orange, throats of adult males 
almost always immaculate and rarely streaked. 

Anolis longitihialis specuuni. new subspecies 

Holotype: MCZ 132,370, adult male, 17 km NW of Oviedo 
Nuevo, Pedernales Province, Republica, 183 m, 7-8 August 1975, 
Richard Thomas, coll. Original number RT 3,461. 

Paratypes: All paratypes are from Republica Dominicana, Ped- 
ernales Province. ASFS V26,898-90K RT 3,462, LSUMZ 29,541 
47, same data as holotype; ASFS V16,728, 6.4 km SE Pedernales, 
17 May 1969, J. B. Strong: ASFS V21.531, 8 km N, 2 km E of Cabo 
Rojo, 1 August 1969, R. Thomas; ASFS V29, 766-68, 7 km N, 20.0 
km SE of Cabo Rojo, 183 m, 14 August 1971, A. Schwartz; ASFS 
V30,073-74, 7 km N, 20.0 km SE of Cabo Rojo, 183 m, 22 August 
1971, D. C. Fowler; ASFS V42,235, 17 km NE of Oviedo Nuevo, 
183 m, 12 August 1975, M. H. Strahm; ASFS V30.086-87, 7 km N, 
17.6 km SE of Cabo Rojo, 152 m, 22 August 1971. A. Schwartz, B. 
R. Sheplan; ASFS V30,114, 9.6 km N of Pedernales, 244 m, 23 
August 1971, A. Schwartz; ASFS V4I,914, 7 km N, 2 km SE of 
Cabo Rojo, 29 July 1975, M. H. Strahm; UF FSM 21.567-68, I km 
SW of Las Mercedes, 380 m, 21 March 1974, R. Franz; MCZ 
146,848, 1.2 km E of cave near intersection Cabo Rojo-Pedernales 
roads, 21 July 1975, W. E. Haas; MCZ 146,849, 7 km N. less than 



1979 NEW SPECIES OF CYBOTOID ANGLE 7 

1.2 km E of intersection of Cabo Rojo-Pedernales roads. 21 July 
1975. W. E. Haas; MCZ 128,319, MCZ 128,342-43, 2 km E of turn 
to Cabo Rojo. 16 July 1971, T. P. Webster, R. B. Huey; MCZ 
143,483. Cabo Rojo, behind police station, 5 July 1974, P. E. Hertz, 
R. B. Huey; MCZ 151,864-78, Cabo Rojo, behind laundry, 1-3 
October 1976, W. E. Haas; MCZ 132,378 80, 7 km N, 1.2 km SE of 
Cabo Rojo, 2 October 1976, N. Atkins, W. E. Haas; MCZ 151.902, 
7 km N, 1 .2 km E of Cabo Rojo. 2 October 1976. W. E. Haas; MCZ 
I32.381-82,'17.6 km NW of Oviedo Nuevo, 2 October 1976, W. E. 
Haas; MCZ 151,828-48. 17.6 km NW of Oviedo Nuevo, 2-3 
October 1976, W. E. Haas. 

Associated specimen. MCZ 58,419. 30 km NW of Oviedo, 
Pedernales Province, Republica Dominicana. 

Definition. A subspecies characterized by the combination of 
large size (males to 72 mm. females to 59 mm. snout-vent length), 
larger dorsal scales (35-52). larger ventral scales (31-52). usually 4 
of 5 postrostral scales, dorsum brown with 4 transverse dumbbells 
and often 1 pair of sacral blotches, throats of females almost always 
longitudinally streaked with dark brown, very rarely so in adult 
males; dewlaps very pale orange to very dull yellow or dirty yellow. 

Description. An adult male with a snout-vent length of 71 mm 
and femur length of 23 mm (measured as proposed by Ruibal and 
Williams, 1961:214); snout scales at second canthal scales 7. vertical 
loreal rows 4. supraorbital semicircles in contact, scales between 
semicircles and interparietal 1 1 (counted as proposed by Schwartz, 
1968:260); subocular scales in contact with supralabial scales bilat- 
erally; 5 enlarged scales in supraocular discs, 7 postmentals, 4 
postrostrals, 3 3 canthals, 19 fourth toe lamellae on phalanges II 
+ HI, 49 median dorsal scales and 43 median ventral scales in 
snout-ear distance. In life, dorsum brown with 4 darker brown 
dorsal dumbbells (narrow middorsally). the anterior pair the largest 
and forming a pair of blotches, somewhat faded along their anterior 
borders, the posterior 3 very fragmented and barely discernible as 
dumbbells; a dark but centrally very flattened postocular U, the 
arms very short and touching the upper eyelids; temples, neck, and 
remainder of dorsum with scattered longitudinal dark dashes or 
small spots; a pale narrow subocular crescent; lores and infraorbital 
area pale and without dark mottling; \enter white, chin and throat 
immaculate; upper surfaces of fore- and hindlimbs concolor with 



8 BREVIORA No. 451 

dorsum and with a very few scattered dark brown dots and without 
defined crossbands; dewlap dull yellow. 

Variation. The series of 76 A. I. specuum is composed of 38 
males and 38 females. MCZ 151,828 is the largest male with a 
snout vent length of 72 mm; the largest female (ASFS V2I,531) has 
a snout-vent length of 59 mm. The smallest lizard is a female (RT 
3,460) with a snout vent length of 37 mm. Femora measurements 
(in mm.) of all males are 14.2-24.4 (x = 21.1) and are 11.8-19.2 
( 16.5) in all females. Snout scales between the second canthals vary 
between 6 and 10 (.\ = 7.4; M,, = 7-44% of the specimens); loreal 
rows are 4 to 6 ( M„ = 5-53%); supraorbital semicircles are in contact 
in 69 specimens and are separated by one row of small scales in five 
lizards. Scales between the supraorbital semicircles and the inter- 
parietal scale are modally 1/ 1 (40 specimens); other counts include 
0/0 (2), 0/1 (3), 1/2 (12), 2/2 (13). Dorsal scales in snout ear 
distance are 35-52 (x = 44.3) and ventrals in the same distance are 
3 1 52 (x = 38.5). The subocular scales may or may not be in contact 
with the supralabial scales; bilateral separation between these scales 
occurs in 44 specimens, unilateral contact occurs in six, and bilateral 
contact in 16; thus, there is contact at least unilaterally between the 
suboculars and supralabials in 33% of the lizards. Enlarged scales in 
the supraocular discs vary between five and 1 1 (x = 7.0; Mo = 7 - 
27%); this count is difficult to take since there may be many small 
(but not granular or tiny) scales along the periphery of the disc that 
might be counted if one desired. Postmental scales are 2 to 9 (x = 
6.3; M„ - 41%); postrostral scales are 3 to 5 (x = 4.2; M,, = 4 or 
5 36% in each category). Canthals are usually 3/3 but three lizards 
have 4/4 canthal scales and one has 3/4. Fourth toe lamellae on 
phalanges II + III are 15 21 (x = 17.6); femur/ snout-vent length 
ratio X 100 is 28.7-36.3 (x = 32.5) in males, 28.8-35.2 (33.2) in 
females. 

Color notes in life on both males and females indicate that the 
dorsal ground color is brown with darker brown markings. There is 
often a white subocular semicircle and a gray lateral stripe which 
may be absent or may be suggested by the remnants of dark dorsal 
and ventral outlining. In males there are usually four dumbbell- 
shaped figures with their narrowest portions lying across the midline 
of the back and followed by a pair of dark sacral blotches; of the 
dumbbells, the first is the best defined and largest and is less clearly 



1979 NEW SPECIES OF CYBOTOID ANGLE 9 

delimited anteriorly than posteriorly as in the holotype. Occasional 
adult males (MCZ 128,342) are virtually patternless above, without 
traces of the dumbbells and with only some longitudinal dashes and 
dark dots remaining. Another smaller male (ASFS VI6,728) has 
only the finest transverse indications of the dumbbells, whereas in a 
comparably sized male (ASFS V30,074) the dumbbells are large and 
form prominent butterfly-shaped markings across the back. The 
truncate pogtocular U described for the holotype is present in most 
males, but may be very reduced, only indicated, or absent. A fine, 
vertically diagonal dark line often courses across the temporal 
region toward the dorsal midline and may even occasionally (MCZ 
128,342) form a complete nuchal V or vague W. The throats of 
males are variable; young specimens have them streaked with dark 
brown while full adults have them most often immaculate. An 
exceptional male (RT 3,462) has streaking remnants on the throat at 
a snout vent length of 71 mm. 

Females have the same dorsal pattern variation as do males. The 
dumbbells are, when present, conspicuous, dark, and well devel- 
oped. Some females (ASFS V30,086) have the dorsal pattern much 
reduced or even absent and the postocular truncate U is more often 
only indicated rather than present and well developed. One female 
(ASFS V41,914) was described in life as having an orangish 
middorsal streak with dark brown dorsal flecking between the 
dumbbells. The throats of females are variable — many are 
longitudinally streaked with dark brown whereas others have the 
throats immaculate white. Therefore there seems to be no correla- 
tion of size and throat streaking. 

Male dewlaps have been described as orange, pale orange, dirty 
yellow, or dull yellow (the latter two conditions at the type-locality). 
A male (ASFS V30,074, snout-vent length 54 mm) was recorded as 
having the dewlap orange (PI. 9K5; all color designations from 
Maerz and Paul, 1950); a topotypical male (ASFS V42,235; snout- 
vent length 67 mm) had the dewlap dull yellow (PI. 1216) shortly 
after death. The vestigial dewlaps in females were recorded as pale 
orange to orange. 

Comparisons. A. I. longifihialls from Isla Beata (Schwartz and 
Thomas, 1975:89, incorrectly assigned A. longitihialis to Isla Alto 
Velo whence the species is unknown) differ from A. I. specuui}i in 
several ways. The Beata subspecies is smaller (males to 67 mm. 



10 BREVIORA No. 451 

females to 57 mm snout vent lengths), the dorsal scales (39 57) and 
ventral scales (29 51) are smaller, and there are modally four 
postrostrals (46'"f). 

The series of A. I. loni^liihialis consists of 30 males and 13 
females. The dorsal coloration in life is grayish tan to gray-brown 
and the color closely matches the cliffs upon which the species lives 
on Isla Beata. In males, the lower sides are greenish and in this sex 
the pattern consists of a series of longitudinal fine dark brown lines, 
dashes, or dots; there is no indication of dark brown dumbbells. The 
truncate occipital U is barely indicated at best and is usually absent; 
the fine diagonal temporal line is present but faint or fragmented. 
All these features, plus dorsal ground color, differentiate A. I. 
longirihialis from A. I. specuutn. Throats of males are almost 
always immaculate; seven males show some vague streaking or dark 
scribbling on the throats. Females are much like the males dorsally, 
except that there may be a more clear indication of the dumbbells 
(AMNH 41,431) than in males. Female throats always show some 
indication of dark streaking or scribbling, but this is seldom bold 
and prominent. Noble (1923:4) noted that A. longitihialis from Isla 
Beata had the dorsum with "a fine penciling of dark brown." He 
also commented that the color (in alcohol) was pale chocolate- 
brown, finely marked with a number of narrow lines of dark brown. 
Some of the lizards which were available to Noble have been 
examined by me and they still retain these features. 

One peculiarity of Noble's description is his comments on dewlap 
color. He stated (1923:4) that the dewlap was "bright" and that it 
"oddly enough, sometimes retains its color in preservative." This 
latter is a peculiarity of this species, of A. marcauoi, and of the one 
next to be described. Most Antillean anoles with orange, red, or 
yellow dewlaps lose the pigments shortly after preservation; but this 
seems not to be the case in these cybotoids. Despite Noble's 
comments on dewlap color and its persistence in A. longitihialis, 
nowhere does he mention what are the characteristics of the bright 
dewlap color except to say that it is "yellowish." Thomas's ( 1964) field 
notes state that in males the dewlap is dirty yellow anteriorly to 
orange on the posterior three-quarters of the dewlap. It appears that 
the dewlap color in the two subspecies of A. longitihialis is fairly 
comparable. 



1979 NEW SPECIES OF CYBOTOID ANGLE U 

Remarks. There is one specimen (MCZ 58,419, from 30 km NW 
of Oviedo. Pedernales Province) of whose status I am uncertain. 
This is an adult male 67 mm in snout vent length. It differs from 
both A. I. specuum and the new species described below in that it 
has a complete dark nuchal band, three complete dark dorsal bands 
between the limbs, and a pair of sacral blotches that almost form a 
fourth complete band. There are no distinctive head markings and 
the specimen is somewhat faded (collected in 1958). If it is an A. I. 
specuum, it is remarkably well patterned dorsally; the throat is now 
immaculate. The small ventral scales assure that it is not A. cybotes. 
The locality places it within the distribution of the former taxon but 
I refrain from assigning it to /I. /. specuum since it differs in pattern 
details as noted above. I suspect that it is an exceptionally well 
patterned male A. I. specuum, but the cautions noted in the 
introduction are re-enforced here: without knowledge of habitat or 
color and pattern in life, one is strongly handicapped in confidently 
assigning older specimens to any of these taxa. 

The name specuum is from the Latin for "of the crevices or caves" 
and alludes to the crevice-dwelling habits of this subspecies. The 
word specuum is genitive plural. 

Specimens examined. A. I. longitihialis. Repiiblica Dominicana, 
Isla Beata. just E of Punta Beata (ASFS V17.2I5 19); no other 
locality (ASFS V2,772-80, USNM 83,878, USNM 83,880, AMNH 
41.415. AMNH 41,422, AMNH 41,424-32, AMNH 52,449-51, 
MCZ 17,686, MCZ 31,774, MCZ 37,480-82, UP FSM 21,572-78). 

The new species referred to in the introduction is composed of two 
populations which differ from each other. Michael H. Strahm 
collected most specimens of the northern population. Accordingly, 1 
associate his name with this new species, which I call: 

Anolis strahmi. new species 

Definition. A cybotoid anole characterized by large size (males to 
79 mm, females to 64 mm snout-vent lengths), supraorbital semi- 
circles always in contact, modally 2 2 scales between the supra- 
orbital semicircles and the interparietal, larger median dorsal scales 
(32 to 51 in snout-ear distance), subocular scales always separated 
from supralabial scales by one row of scales, usually three post- 



12 BREVIORA No. 451 

rostral scales, more fourth toe lamellae (17-24), dewlaps deep 
orange to deep orange-brown, throats of males marked, or without 
dark flecking or scribbling, by population. 

Head. Moderately massive, length from snout to posterior bor- 
der of eye much shorter than either femur or tibia. Head scales 
mostly smooth in males and very weakly carinate in females. Five to 
nine scales across snout at level of second canthal scales. A very 
shallow frontal depression and a deeper parietal depression. Nares 
in front of and above canthal ridge composed of three or four scales. 
Anterior nasal scale in contact with rostral. Snout distinctly pointed 
when viewed from above and with a distinctive profile due to the 
raised nares. 

Supraorbital semicircles in contact and are separated from the 
supraocular discs by 2 rows of granules. Supraocular discs consist of 
about 4 to 1 1 enlarged smooth to very weakly keeled (males) or 
keeled (females) scales separated by about 4 rows of scales and 
granules from the scales of the supraciliary rows. Two elongate 
supraciliaries are continued posteriorly by a double row of mod- 
erately enlarged scales. Canthus distinct, the first canthal scale the 
largest. Loreal rows five to nine, the lower rows larger and more 
regular. Supratemporal area scales are granular, grading rather 
abruptly into larger scales surrounding the interparietal depression. 
Interparietal is about the same size as ear opening and is separated 
from the supraorbital semicircles by one to three scales (modally 
two) or rarely unilaterally in contact. 

Subocular scales are almost always separated from supralabials 
by one row of scales (very rarely in contact, very rarely by two rows 
of scales), anteriorly grading into loreals, posteriorly grading into 
large scales at the corner of the mouth. Usually six supralabials to 
center of eye. 

Mentals are equally broad and long and almost equilateral, and 
are in contact posteriorly with one to ten small elongate postmental 
scales. Infralabials are broadly rectangular anteriorly, narrow pos- 
teriorly, in contact with three large tetrahedronal sublabials. Throat 
scales are small, granular, not keeled, and the anterior ones are 
elongate. 

Trunk. Middorsal scales are in two or four abruptly enlarged 
rows, about four times as large as flank scales and 32-51 in snout- 
ear distance. Ventrals are small, about 1.5 times as large as 



1979 NEW SPECIES OF CYBOTOID ANGLE 13 

middorsal series, cycloid, smooth, and 31 to 61 in snout-ear 
distance. Two postanal scales are enlarged in males. 

CJular fan. Very large; scales smooth, those along margin about 
twice as large as ventrals. 

Limbs and digits. Hand and foot scales are smooth. Between 14 
and 24 scales are under phalanges II + III of fourth toe. Largest 
scales of arm are smooth to very weakly unicarinate, those of leg 
smooth and both those of arm and leg are larger than ventrals. 

Tail. Slightly laterally compressed, each verticil surmounted by 
three sharply keeled scales and ventrally by three pairs of unicari- 
nate slightly smaller scales. 

Anolis strahnii strahnii. new subspecies 

Holutype: MCZ 132,371. an aduh male, from 3 km NE of El 
Aguacate, Independencia Province, 854 m, Republica Dominicana, 
taken on 19 July 1975 by Michael H. Strahm. Original number 
ASFS V4 1,729. 

Paratypes. All paratypes are from Independencia Province, Re- 
publica Dominicana. ASFS V4I, 730-34, same data as holotype; 
ASFS V41, 284-94, same locality as holotype, 10 July 1975, M. H. 
Strahm; ASFS V28,453, ASFS V41, 308-09, same locality as holo- 
type. 14 July 1975; ASFS V44,991-94, same locality as holotype. 22 
December 1976. A. Schw-rtz. W. B. Southerland; UF FSM 21.565- 
66, 2 km NE of El Aguacate. 900 m, 30 June 1974. F. G. Thompson. 

Definition. A subspecies of .4. strahnii characterized by modally 
eight scales in the supraorbital discs, dorsum is pale gray and is at 
best flecked with dark gray and often unpatterned, throat is usually 
unpatterned in both sexes but occasionally has some vague scrib- 
bling, and dewlap is very deep orange to orange-brown. 

Description. An adult male with a snout-vent length of 78 mm 
and femur length of 25.5 mm; six snout scales at second canthal 
scales, seven vertical loreal rows, supraorbital semicircles in contact, 
scales between semicircles and interparietal 2/2; subocular scales 
separated by one row of scales from supralabials bilaterally; 1 1 
scales in supraocular discs, six postmentals, three postrostrals, 3 3 
canthals. 20 fourth toe lamellae on phalanges II + 111. 41 median 
dorsal scales and 48 median ventral scales in snout-ear distance. In 
life, dorsum is pale gray with a few scattered darker gray dots, most 
prominent above the forelimb insertions; remnants of a postocular 



14 BREVIORA No. 451 

truncate U are present, its very short arms abutting against the 
upper e\elids; supra- and infralabials contrastingly spotted with 
pale and dark gray; a very pale blue-gray subocular crescent; venter 
pale grayish, throat immaculate and without pattern; limbs and tail 
without any crossbands or other prominent markings; dewlap 
(shortly after death) deep orange (PI. 4A11). 

lariation. The series of 26 A. s. strahnii consists of nine males 
and 17 females. The holotype and another male (UF/FSM 21.566) 
are the largest males with snout vent lengths of 78 mm; the largest 
female (ASFS V41.288) has a snout-vent length of 64 mm. The 
smallest lizard (ASFS V44.994) is a female with a snout-vent length 
of 42 mm. Femora measurements (in mm.) in all males are 15.7-26.4 
(.X = 21.5) and are 13.0-21.5 (18.9) in all females. Snout scales 
between the second canthals vary between five and eight (x = 7.0; 
M, ,= 7 43*7 of the specimens); loreal rows are five to seven (Mo = 
5-52%); supraorbital semicircles are in contact in all specimens. 
Scales between the supraorbital semicircles and the interparietal 
scale are modally 2/2 (14 specimens), with other counts of 1/0 (1), 
1/ 1 (2). 1 2 (4), and 2 3 (1). Dorsal scales in snout-ear distance are 
34-48 (x = 39.3) and ventrals in the same distance are 32 61 (x = 
41.2). The subocular scales are separated from the supralabial scales 
by one row of scales bilaterally in all but two lizards which have 
either two rows of scales or are in contact unilaterally. Enlarged 
scales in the supraocular discs vary between five and 12 (x = 8.0; M„ 
= 8-36%); the same precautions made in the account of A. I. 
specuuin apply here. Postmental scales are four to seven (x = 5.3; 
Mo = 6-36%;); postrostral scales are three to five (.x = 3.5; M,, = 
3-58%). Canthal scales are always 3 3. Fourth toe lamellae on 
phalanges II + III are 16-21 (x = 18.6); femur snout vent length 
ratio X 100 is 31.7-34.3 (x = 33.3) in males and 30.5-36.1 (32.9) in 
females. 

Color notes on both males and females in life indicate the dorsal 
ground color is gray, remarkably similar to the color of the rocks on 
which the lizards live. Both sexes are patterned similarly dorsally; 
there are no dumbbells or flank stripes present (although the 
position of the latter is occasionally indicated by vague elongate 
dashes along its putative upper and lower margins). The dorsum is 
more or less randomly dotted or marked with scattered dashes 
aligned longitudinally; young males have vague dumbells which are 



1979 NEW SPECIES OF CYBOTOID ANOIE 15 

SO reduced as to be only faint transverse lines hollowed laterally — 
I.e., they are mere vestiges of what presumably are basically 
dumbbells. The general aspect of the males and females is oi a faint 
and randomly dotted or lineate pattern, but many adults and 
subadults completely lack any sort of dark body pattern elements. 
The postocular truncate U is represented by its outlines and these 
may be fragmented and very obscure. The same is true of the 
diagonal terhporal line. The venter is dark gray to yellowish gray in 
females. The throat is marked with some confused scribbling, 
oriented longitudinally in one adult male (ASFS V41,285) and in 
one subadult male (UF/FSM 21,565; snout-vent length 54 mm). 
Females generally have gray throats without markings, but one 
adult (ASFS V4 1,290; snout vent length 62 mm) has some vague 
darker scribbling and two other females have remnants of similar 
markings only very faintly shown on the posterior portion of the 
throat. In life, the limbs are crossbanded with pale and dark gray, 
but these markings are not obvious on the preserved specimens. A 
distinctive feature of males (and somewhat less so of females) is the 
spotted supra- and infralabials. A very pale blue-gray subocular 
crescent is present and moderately conspicuous in life. 

The dewlaps in males are very deep orange to orange-brown and 
these colors are especially rich and almost velvety in texture. Color 
designations are PI. 13JI0 for one living male and PI. 1 1 II I and PI. 
4AII for two recently dead males. Females have the same basic 
intensity of coloration on their vestigial dewlaps. 

Comparisons. Comparisons of A. s. strahmi with the subspecies 
on the south side of the Sierra de Baoruco as well as with A. I. 
specuum will be withheld until the former is described below. 

Remarks. I have commented previously on Dr. Thompson's 
observations on the habitat o{ A. s. strahmi. Our own observations 
amply confirm his. On 10 July 1975 we searched several kilometers 
of the road below the post of El Aguacate and saw only very 
occasional lizards on the gray rocks of the roadcut and did not 
observe any on adjacent shrubs or herbs. In much of this distance, 
the roadcut is rather densely covered with vines, small shrubs, and 
herbs, but elsewhere it is almost bare with only scattered vegeta- 
tional cover. We finally encountered .4. .v. strahmi in some abun- 
dance along an exposed section of roadcut which was about 30 m in 
length. The roadcut here was almost bare of plants with many 



16 BREVIORA No. 451 

crevices and solution holes. Between 1330 and 1600 hrs this section 
of roadcut was shaded (due to the direction of its face rather than to 
any arborescent or herbaceous cover) and the lizards were seen in 
their retreats, their heads or foreparts extended outside the crevices 
in an alert manner. They were not easily alarmed, relying upon the 
extremely cryptic coloration of their gray dorsa agreeing very 
closely with the hue of the roadcut face. Two lizards were seen 
"sunning" themselves (in the shade) vertically, head down, on 
completely exposed rocky surfaces; one was observed clinging to the 
roof of a small solution hole. In addition to the 1 1 specimens 
collected during this 2.5 hour period, at least eight others were seen 
but not collected. They allowed close (1 m or less) approach of the 
collector before becoming alarmed and retreating into crevices or 
solution holes. Recovery time was brief; on several occasions, as we 
walked slowly down this 30 m stretch of road, we noted refuges of 
specific lizards. Upon returning a maximum of 10 minutes later the 
lizards had already reappeared at the crevice mouths (which were 
their individual retreats) or on adjacent rock faces. On subsequent 
visits we observed lizards in the same precise places as previously. 
Their agility and assurance on the rock faces were impressive. 

On our third visit to the type locality on 19 July 1975, six lizards 
were collected between 1720 and 1830 hrs in the same stretch of 
roadcut. The adult male holotype was taken just below a slightly 
more covered section. About three or four other anoles were seen 
but escaped capture. Although the cliff face was completely in shade 
and the air was cool (but not cold) the rock faces were still warm to 
the touch. 

On 22 December 1976, W. B. Southerland and I visited the type 
locality once again. Our arrival was at 1 200 hr and we saw very few 
lizards. The roadcut was in shade and the shade increased until 1500 
hr so that maximum insolation had occurred in the early morning 
prior to our arrival. However, the rocks were still warm to the 
touch. As the afternoon progressed, more A. s. strahmi appeared, so 
that about 12 were seen. Some large males, once disturbed, did not 
reappear on the roadcut face. Four specimens were taken: a male on 
an open rock face, a female on a dead stem 8 cm from the base of the 
cliff and two other females in or near crevices or cavities in the 
roadcut face. The female noted here on a dead stem is the only 
individual we observed or collected not on the rocks. 



1979 NEW SPECIES OF CYBOTOID ANGLE 17 

Above and below the road at the type locality are steep slopes. An 
outstanding feature of the immediate region is the eroded limestone 
terraces and jumbled rocks and boulders, all sparsely covered with 
herbs and scattered shrubs and with only occasional large trees. The 
general aspect of the area is rather bleak. 

Two other observations at the type locality are pertinent. We 
encountered an active Vromacer cateshyi Schlegel foraging on the 
roadcut face- occupied by A. s. strahmi. These snakes eat lizards but 
are rarely seen in this habitat type and prefer trees and shrubs. In a 
crevice at this site we also encountered a young Epicrates striatus 
Fischer; when young, these snakes eat lizards. It is not unlikely that 
both snakes were associated with this particular section of roadcut 
because of the ready supply of food (A. strahmi). Both snakes are 
diligent predators in that they search for prey by penetrating 
crevices, holes, cavities, and other possible hiding places. Elongate 
U. catesbyi is especially well adapted for this foraging pattern 
during the day and E. striatus forages similarly at night. 

Anolis strahmi abditus, new subspecies 

Holotype: MCZ 146,827, an adult male, from dirt road to Las 
Mercedes, 2.9 km from intersection (= 5 km SE, 2.9 km N of 
Pedernales), Pedernales Province, Republica Dominicana, 19-20 
July 1975 by William E. Haas collector. Original number MCZ F- 
29,006. 

Paratypes. All paratypes are from Pedernales Province, Repub- 
lica Dominicana. MCZ 146,920, MCZ 146,828 47, same data as 
holotype; ASFS V41,908 12, same locality as holotype, 29 July 
1975, M. H. Strahm; UF/FSM 34,423 27, 5 km SW, 2.5-3.0 km N 
of Pedernales, 8 August 1975, T. Wiewandt and D. Gicca; MCZ 
151,879-901. 5 km SE, 2.9 km N of Pedernales, 30 September and 1 
October 1976, W. E. Haas; MCZ 132,383, 5 km SE, 2.9 km N of 
Pedernales, 30 September 1976, N. Atkins and W. E. Haas; MCZ 
151.849-54, MCZ 151,857-63, between 15 and 16 km N of Cabo 
Rojo, Alcoa road. 3 October 1976, W. E. Haas. 

Definition. .A subspecies oi A. strahmi characterized by modally 
six or seven scales in the supraorbital discs; dorsum tan to grayish 
tan, at times marked with small brown to reddish blotches; lateral 
flank stripe tan to whitish present, dorsum with transverse dumb- 



18 BREVIORA No. 451 

bells or hourglasses in both sexes; throat flecked or scribbled in both 
sexes with dark brown and dewlap deep orange. 

Description. An adult male with a snout-vent length of 78 mm 
and femur length of 26 mm; six snout scales at second canthals, 
supraorbital semicircles in contact, scales between semicircles and 
interparietal 2/ 2, subocular scales separated by one row of scales 
from supralabials bilaterally; seven scales in supraocular disc, six 
postmentals, three postrostrals, 3 3 canthals, 19 fourth toe lamellae 
on phalanges II + III, 35 median dorsal scales and 40 median ventral 
scales in snout-ear distance. As preserved, the dorsum is grayish 
brown, with four dark brown dumbbell remnants, all much hol- 
lowed or with their lateral expanded ends virtually missing and an 
additional fifth dumbbell remnant postsacrally; a dull interocular 
bar and a truncate postocular U, both hollowed; lores, supra-, infra-, 
and sublabials mottled dark and pale brown; sides of body with 
elongate dark brown linate fragments, with two of the lowermost 
fragment-series in part outlining the tlank stripe; limbs marbled or 
mottled dark brown and brown, the only clear transverse dark bar 
on the crus; both fingers and toes with a moderately clear dark 
brown crossband; venter grayish to tan, chin and throat dotted with 
dark brown, sides of abdomen flecked with grayish brown; dewlap 
pinkish 6 months after preservation. 

Variation. The series of 68 A. s. abditiLS consists of 33 males and 
35 females. The largest male (ASFS V41,911) has a snout-vent 
length of 79 mm and the largest female (MCZ 146,838) a snout-vent 
length of 63 mm. The smallest lizard (UF/FSM 34,425, a female) 
has a snout vent length of 39 mm. Femora measurements (in mm.) 
in all males are 16.0 27.4 (x = 21.7) and are 13.2-20.8 (18.1) in all 
females. Snout scales between the second canthals vary between five 
and nine (x = 6.9; Mo = 7-52% of the specimens); loreal rows are 
four to seven (Mo = 5-62%); supraorbital semicircles are in contact 
in all specimens. Scales between the supraorbital semicircles and the 
interparietal scale are modally 2/2 (35 specimens), with other counts 
of 1/ 1 (9), 1/2 (5), 2/3 (6), and 3/3 (8). Dorsal scales in snout-ear 
distance are 32-51 (x = 39.8) and ventrals in the same distance are 
31-53 (x = 40.2). The subocular scales are separated from the 
supralabial scales by one row of scales in all but two specimens; 
these lizards have these scales in contact either bilaterally or 



1979 NEW SPECIES OF CYBOTOID ANGLE 19 

unilaterally. Enlarged scales in the supraocular discs vary between 
four and 13 (x = 7.3; M,, = 6 or 7 27% in each case). Postmental 
scales are four to 10 (x= 7.3; M,, = 6 33%); postrostral scales are 
three to five (x = 3.7; Mo = 3-43%). Canthal scales are most often 
3 3, but one lizard has 2/2 and another 3 4 scales in this position. 
Fourth toe lamellae on phalanges 11 + III are 17-24 (x = 19.0); 
femur snout-vent length ratio XlOO is 28.7-38.4 (x= 33.5) in males 
and 30.5 3(1.3 (33.4) in females. 

Color notes state that in both males and females in life the 
dorsum is tan to grayish tan with five narrow dumbbell or hourglass 
remnants; these dorsal pattern elements end above the flank stripe 
which is either tan or white. Despite its prominence in life, the flank 
stripe is outlined only by dark brown line fragments, including the 
remnants of the dorsal transverse dumbbells. A dark brown inter- 
ocular stripe and a postocular truncated U are both present and may 
be either prominent or vague; the U may be most conspicuous at its 
posterolateral "corners" on the superior temporal region. Some 
males (ASFS V4I,910 12, MCZ 146,829) show a progressive degen- 
eration of the dorsal pattern into a series of more or less random 
dumbbell-fragments and associated longitudinal lines or dots which 
are brown to reddish. In each of these lizards, the peculiar dorsal 
pattern is obviously a derivative of the customary dorsal dumbbells. 
Two females (ASFS V41,909, MCZ 146,838) are comparable to the 
above mentioned males in diminution of the dorsal pattern. In both 
sexes, the throat is marked with some sort of pattern; this varies 
from marbling or scribbling to longitudinal dark lines which is more 
common in females than males. The intensity of the throat markings 
is variable. The lores and supra-, infra-, and sublabials are contrast- 
ingly marked with dark and pale and the lip markings extend 
ventrally to give rise to the throat pattern. The male dewlaps are 
deep orange in life (PI. 12F9); two dead males had comparable 
dewlaps (PI. 12F9, P1.12H8). The deep orange color of the dewlap 
center is slightly richer along the margin. 

Comparisons. A. s. abditus requires comparison with the taxa A. 
s. strahmi and^. /. specuum. Anolis cyhotes also occurs sympatri- 
cally with both subspecies oi A. strahmi. but it is easily distinguish- 
able in dewlap color (pale yellow, pale pink, almost white), smaller 
snout-vent length, and much larger ventral scales. Anolis hreviros- 



20 BREVIORA No. 451 

tris. a small gray dorsoventrally compressed species with a pale 
yellow dewlap and a paramedian double row of snout scales, also is 
syntopic with A. s. ahditus. 

The two subspecies of A. strahmi are very similar, although they 
are altitudinally and geographically separated. The dewlaps in both 
are deep and rich orange (to orange-brown in A. s. strahmi) and 
reach the same size in both sexes. A. s. strahmi differs from A. s. 
abditus in that the former has modally eight scales in the supra- 
orbital discs versus six or seven in A. s. ahditus. Dorsal coloration in 
A. s. strahmi is pale gray versus tan to grayish tan in A. s. abditus. 
The dorsum is at best flecked and often unpatterned in A. s. strahmi 
males versus relatively prominent dumbbell or hourglass remnants 
in both sexes of /4. .s. abditus. Finally, throats in both sexes of /4. s. 
strahtni are rarely marked versus throats in both sexes o{ A. s. 
abditus which are marked with flecking, dotting, scribbling, or lines. 
The dorsal color and pattern, as well as the throat markings, are the 
most distinctive characteristics separating the two subspecies. 

A. s. abditus has not as yet been taken sympatrically or syntopi- 
cally with A. I. specuum (see Fig. 1); it is unlikely that these two 
species will be taken together since both are clivicolous. Though 
they seem to be ecological equivalents, it is possible that at some 
locality on the Peninsula de Barahona they occur together. The two 
species should be easily differentiable, if they are sympatric. The 
dewlap in A. s. abditus is deep orange, whereas that oi A. I. specuum 
is dull or dirty yellow to pale orange. A. s. abditus reaches a larger 
size (males to 79 mm, females to 63 mm) than does A. I. specuum 
(males to 72 mm, females to 59 mm). Although there are three to 
five postrostral scales in both taxa, the mode in A. s. abditus is three 
and m A. I. specuum is four or five. Two other scale relationships 
will aid in differentiating the two species. \nA. I. specuum. there are 
more often 0/0 1/ 1 scales between the supraorbital semicircles and 
the interparietal (64% of the specimens) whereas in A. s. abditus 
there are more often 1/2 3/3 (79%) scales in this position. A. I. 
specuui7i more often (33%) has the subocular scales in contact with 
the supralabials than does A. s. abditus (3%). Although none of 
these scale counts or relationships is absolute, in combination they 
serve to differentiate the two species. 

There are some striking resemblances between A. I. specuum and 
A. s. abditus in color and pattern. The former is brown with darker 



1979 



NEW SPECIES OF CYBOTOID ANGLE 



21 



,.„,..»a 




Figure 1. Map of the Peninsula de Barahona, Repiiblica Dominicana. Hispan- 
ioia, and Isla Beata. Pertinent population centers are indicated b\ labeled squares. 
Known distributions are shown for the anoles A. longiiihialis (triangles) and A. 
sirahmi (circles). The distributions are further designated for A. I. longiiihialis (solid 
triangle), A. I. specuum (hollow triangles), .-1. v. sirahmi (solid circles), and A. s. 
ahdiiiis (hollow circles). 



22 BREVIORA No. 451 

brown markings whereas the latter is tan to grayish tan with brown 
markings. Both are prominentK marked dorsally with 4 dumbbells; 
however, in A. I. spccuuni. the first dumbbell is well developed 
laterally and open anteriorl\ to \ield a large almost ocellus-like 
spot. The dewlap colors of the two species are extremely distinctive 
since no A. I. specuuni is known that has the deep orange dewlap of 
A. s. ahditus. One final feature serves to distinguish the two species 
in this area. A. I. specuum males most often have the throat 
immaculate as adults (one adult male is known with a streaked 
throat) whereas the throat is invariably patterned in A. s. ahditus 
males. All A. s. ahditus females have marked throats whereas the 
throats of A. I. specuum females are variable — from immaculate to 
streaked with dark brown. 

Remarks. There are only two localities for A. s. ahditus. The first 
is a ravine (the type locality) between the Pedernales-Oviedo road 
and the village of Las Mercedes. At about 2.5 km N of the main 
road, the dirt road to Las Mercedes passes (for about 1 km) through 
a shaded and broad ravine with reddish rocky natural cliffs on both 
sides. These cliffs are creviced and have solution holes, but are 
separated from the road itself by low talus slopes which extend some 
five to ten meters between the cliffs and the roadway. The flora of 
the ravine stands in strong contrast to the Acacia-cactus desert 
which surrounds it; the ravine is distinctly more mesic, with trees 
and an understory of grasses, herbs and BryophyUum. At the time 
of our 1975 visit the outside desert was exceptionally parched 
whereas the ravine was modestly luxuriant and moist. The ravine is 
almost at sealevel but at its northern end the road begins the ascent 
of the lower front ranges of the Sierra de Baoruco on which the 
village of Las Mercedes lies. The second locality is on the Alcoa 
road between Cabo Rojo and Aceitillar; I have not visited the site 
but William E. Haas (in litt.. 22 October 1976) wrote that it is "up a 
path just past km 16 on the Alcoa truck road, and the habitat is 
somewhat different [from the type locality] and the physiography a 
bit more so." 

At the time of our 29 July 1975 visit to the type locality, the 
lizards were scarce at 1300 hr. The sun was almost directly overhead 
and the ravine was hot although fairly well shaded. As the afternoon 
progressed, the lengthening shadows of the western wall began to fill 
the ravine and bv 1530 hr the lizards were moderatelv abundant. 



1979 NEW SPECIES OF CYBOFOID ANOLE 23 

Thev inhabited crevices and solution holes in the cliff faces (much 
after the fashion of A. s. strahmi). One male was seen foraging at 
1630 hr on a shrub adjacent (1 m) to the cliff and had a large 
caterpillar in its mouth when collected. Foraging thus involves 
securing food elsewhere than directly on the cliff face itself. A very 
small juvenile was dislodged from the basal rosette of a moderately 
sized dead sisal plant (Agave) on the cliff face but escaped without 
being collected. One behavorial incident is worthy of report: we 
pursued a large male along the cliff face and the lizard finally leaped 
to an isolated large (2 m diameter) boulder, ran across its top at full 
tilt and literally launched itself off the far side of the boulder onto 
the ground. Several minutes later the same lizard was collected 
within a few centimeters of the boulder's base, immobile in the 
grassy and herbaceous ground cover. Whether its immobility was 
traumatic or whether it was "hiding" in a very unusual situation is 
unknown. This is one of only two instances when A. strahmi was 
secured in any situation other than a clivicolous one (the other 
exception is the female A. s. strahmi on the dead stem near the cliff 
base at El Aguacate). As with A. s. strahmi, A. s. abditus is 
impressively "at home" on its cliff faces. It is very agile, and its 
camouflage, tendency to remain immobile when approached — 
relying upon its cryptic coloration, and its prompt recovery time, all 
are extremely similar to those features in the behavior of A. s. 
strahmi. 

DISCUSSION 

It seems obvious that A. longitihiaHs and A. strahmi form a 
compact and closely related duo of cybotoid anoles whose distribu- 
tion centers upon the Peninsula de Barahona. Although I am 
convinced that they represent two separate species rather than 
subspecies, the latter interpretation is not to be lightly dismissed. 
There are no absolute meristic characters which separate the two, 
and they are similar in many features, not the least of which is their 
predilection for cliffs and vertical rockv surfaces to which they are 
unquestionably adapted. If, however, A. lotigitibialis and A. strahmi 
are conspecific. the peninsular distribution of the latter is peculiar in 
that it seems to be in enclaves surrounded on both east and west by 
(or interdigitating with) A. longitihiaHs. This arrangement can be 



24 BREVIORA No. 451 

interpreted as a mosaic, where one species (/I. longitihialis) inhabits 
open and xeric cliff faces and the other (A. sirahmi) inhabits shaded, 
more mesic, and less rigorous cliffs. The dewlap colors of the two 
species are very distinctive and since dewlap color in eye-minded 
anoles is so important in species recognition and territorial defense, 
the very fact of the striking color differences of dewlaps in these two 
taxa strongly suggests that they are indeed species. The situation 
here is not so complex as that between A. cyhotes and A. marcanoi 
(Williams, 1975): in this case two unquestioned species without 
strong meristic differences but with distinctive and contrasting 
dewlap colors and different body patterns are sympatric and even 
rarely syntopic. In these two species, however, there appear to be 
subtle differences in habitat preference, and as Williams (1975:9) 
pointed out the "balance of power" between them in any particular 
area may be tenuous. The same situation may exist in the geo- 
graphical and ecological relationships between A. longitihialis and 
A. strahmi on the peninsula but presently the differences seem more 
overt. 

More puzzling is the occurrence of two subspecies oi A. strahmi 
on the north and south slopes of the Sierra de Baoruco. This is 
coupled with the striking difference in elevations involved (near sea 
level, and between 854 and 900 m). The habitats of the two 
subspecies are comparable — generally creviced cliff faces or 
roadcuts which can be regarded as cliff-face surrogates. But the 
similarity ceases with this description. The ravine near Pedernales, 
although it transects an otherwise hostile and xeric area, has 
relatively luxuriant vegetation and is well shaded. The El Aguacate 
roadcut is open and exposed to direct sunlight (the same is true for 
localities where A. longitihialis has been taken on the peninsula 
itself) and the lizards seem to shun those sections of the roadcut that 
are heavily vegetated and overgrown. Even sections of the roadcut 
that are exposed and open often lack or have minimal populations 
of A. strahmi. The area is quite cool in the afternoon and cold at 
night. One explanation for the apparently precise niche occupied by 
A. s. strahmi may well be that, at such high elevations, insolation of 
cliff faces and lack of vegetational cover will determine the diurnal 
rhythm of these lizards. Cliffs, exposed or not, facing in directions 
where the sunlight will not reach them until rather late in the 
morning are unsuitable because of lack of time exposed to the 



1979 NEW SPECIES OF CYBOTOID ANGLE 25 

warming effect of the sun. In contrast, vegetationally covered cliffs, 
even those oriented to achieve maximum early insolation, are too 
protected by vegetation to allow the lizards a sufficient period of 
activity during each day. 

The disjunct nature of the two subspecies of A. strahmi may be 
more an artifact than a reality. There are few suitable cliff faces 
readily attainable by road in this entire region. It seems likely that 
A. sfrahmi ifihabits suitable habitats around the general periphery 
of the Sierra de Baoruco. Cliffs and roadcuts on the peninsula along 
the eastern edge of the mountains between Barahona and Enriquillo 
were searched casually. These cliffs are often at sea level but are well 
shaded and generally face east. We found both A. cybotes and A. 
hrevirostris here but not A. strahmi or A. longitibialis. Such 
negative evidence is questionable at best, although we did not find 
A. cybotes syntopically on cliffs or roadcuts with either of the 
clivicolous species. However, A. cybotes does occur sympatrically 
(but not on cliffs) with A. I. specuum and both subspecies of A. 
strahmi. It is interesting to note that in those places where A. 
cybotes inhabits cliffs and roadcuts it is much less adapted for such 
habitats than are the two clivicolous species. When pressed A. 
cybotes is more likely to leave the cliff face and jump to the ground 
to seek refuge under adjacent ground cover than are A. longitibialis 
or A. strahmi which seek sanctuary in crevices or solution holes. 

Geographically it seems likely that there are more or less discon- 
tinuous populations oi A. strahmi scattered around the periphery of 
the Sierra de Baoruco (but not across its summit) and the lower 
eastern regions of the adjacent Haitian Massif de la Selle. Popula- 
tions should be looked for on the bare roadcut-cliffs between Fond 
Parisien and Soliette and on the northern face of the La Selle in 
Haiti near the Dominico-Haitian border south of Fond Verrettes. 
Although these cliffs seem suitable (more for A. s. abditus than for 
A. s. strahmi), it may be that an appropriate habitat niche occurs in 
this region which will accommodate the latter subspecies. If A. s. 
strahmi did occur there at suitable elevations on the northern face of 
the Massif de la Selle, it may now be extirpated due to extreme 
modification of habitat (the region has been denuded of much 
vegetational cover). 

A. longitibialis. on the other hand, presents a more familiar 
pattern. It is a Barahona Entrapment xerophile derived from 



26 BREVIORA No. 451 

primarily mesophilic A. cybotes with which, under favored circum- 
stances, it is still sympatric. From the peninsula it has invaded Isla 
Beata and differentiated on the subspecific level. A phyletic se- 
quence between these three species might well be: widely distributed, 
ecologically tolerant but primarily mesophilic A. cybotes with a pale 
dewlap -^ xeric and cliff adapted A. longitibialis with a pale dewlap 
— clivicolous and more mesic adapted A. sirahmi with a vividly 
colored dewlap limited to disjunct enclaves where its ecological 
requirements are rather precisely met. 

Another possibility that should not be overlooked is that the four 
taxa involved (longitlbiall.s, speciium, strahmi, abditus) should be 
regarded as separate species. Certainly they are isolated from each 
other {longitibialis is insular; specuum occurs on the Peninsula de 
Barahona; strahmi and abditus occur to the north and south of the 
Sierra de Baoruco massif, respectively). Regarding them as species 
has much to recommend it, but I have taken the more conservative 
course that no external characteristics are sufficiently obvious to 
make one seriously consider that there are four species involved 
rather than two. However, skeletal or electrophoretic evidence 
might well prove differently. 

ACKNOWLEDGMENTS 

I wish to thank the following curators for allowing me to study 
material in their collections: Richard G. Zweifel, American Museum 
of Natural History (AMNH), Douglas A. Rossman, Museum of 
Zoology, Louisiana State University (LSUMZ), Ernest E. Williams. 
Museum of Comparative Zoology, Harvard University (MCZ), 
Richard Thomas, University of Puerto Rico (RT), Walter A. 
Auffenberg, University of Florida, Florida State Museum (UF/ 
FSM), George R. Zug and Ronald L Crombie, National Museum of 
Natural History (USNM). Specimens collected by myself and 
parties are in the Albert Schwartz Field Series (ASFS). Much 
pertinent material in the MCZ has been collected under National 
Science Foundation grant GB-3773IX and preceding grants to 
Ernest E. Williams, and some specimens in the ASFS have been 
collected under National Science Foundation grants GB-7977 and 
B-023603. In the field I have had the competent assistance of the 
following men, without whose help much of the material for the 
present problem would not be available: Danny C. Fowler, David 



1979 NEW SPECIES OF CYBOTOID ANOLE 27 

C. Leber, James A. Rodgers, Jr., W. Barry Southerland, Michael H. 
Strahm, James B. Strong, Bruce R. Sheplan, and Richard Thomas. 
Most especially I am grateful to Fred G. Thompson tor allowing me 
to examine the first specimens of A. strahnii. 

LITERATURE CITED 

Cochran. D^ M. 1934. Herpetological collections made in Hispaniola by the 
Liowcma expedition, 1934. Occ, Papers Boston Soc. Nat. Hist. 8: 163-188. 

. 1939. Diagnoses of three new lizards and a frog from the Dominican 

Republic. Pioc. New England Zool. Club 18: 1-3. 

1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mas. 177; vii + 



398 pp. 
Mafrz. a., AM) M. R. Pmi. 1950. A dictionary of color. McGraw-Hill Book 

Co. Inc., N.Y., vii + pp. 1 23 and 137 208, 56 pis. 
Noble. G. K. 1923. Four new lizards from Beata Island. Dominican Republic. 

Amer. Mus. Novitates 64: 1-5. 
Ri iBAi . R., AM) E. E. Wii I i-xMs. 1961. The ta,\onomy of the Anolis honmlechis 

comple.x of Cuba. Bull. Mus. Comp. Zool. 125(8): 211 246. 
ScHvvARiz. A. 1968. Geographic variation in Anolis clistichus Cope ( Lacertilia, 

Iguanidae) in the Bahama Islands and Hispaniola. Bull. Mus. Comp. Zool. 

137(2): 255-309. 
, AND R. Thomas. 1975. A check-list of West Indian amphibians and 

reptiles. Carnegie Mus. Natur. Hist. Spec. Publ. 1: 216 pp. 
Webster, T. P. 1975. An electrophoretic comparison of the Hispaniolan lizards 

Anolis cyhoies and A. marcanoi. Breviora, No. 431: 1-8. 
Williams, E. E. 1961. Notes on Hispaniolan herpetology. 3. The evolution and 

relationships of the Anolis semilineaius group. Breviora, No. 138: 1-8. 
. 1975. Anolis marcanoi new species: sibling to Anolis cyhotcs: description 

and field evidence. Bre\iora No. 430: 1-9. 



/zr 



MU3. CCMP. ZOOU 
LIPRARY 



MAR 1 8 1985 

R E V I l-i- 

iiseiiiii of Comparative Zoology 

us ISSN 0006-9698 



Cambridge, Mass. 21 February 1979 



Number 452 



LIZARDS OF THE SCELOPORVS ORCUTTI COMPLEX 

OF THE 
CAPE REGION OF BAJA CALIFORNIA 

William P.Hall'^ and Hobart M. Smith' 

Abstract. Sceloporus orcuiii licki as hitherto defined is shown to embrace two 
cryptic species, neither of which is subspecific in relation to the allopatric 5. orcutli: 
an arboreal species, 5'. licki, and a petricole, 5'. hunsakeri sp.n., from 3 mi. E of San 
Bartolo (holotype MVZ 73.570); both are restricted to the Cape region of Baja 
California. All three species, S. orcuiti. S. licki and 5. hunsakeri have identical 2n = 
34 (.\Y d ) karyotype patterns, but differ in numerous other respects. In these three 
species, the most succinctly diagnostic character states are: presence of a dorsolateral 
light stripe in S. licki only, 59 or fewer dorsal scales plus femoral pores in most 
(96% of 27) 5. orcutti. and 61 or more in most S. hunsakeri (98% of 61). 



INTRODUCTION 

Field work in 1969 indicated that the Cape Region of Baja 
California (the area south of the Isthmus of La Paz) is inhabited by 
three distinct species of Sceloporus which are broadly sympatric and 
frequently syntopic with one another (Figs. 1, 2). Two of the three 
species, Sceloporus licki Van Denburgh and a species we describe 
here, are sufficiently similar to one another and to .S. orcutti 
Stejneger that Smith (1939) considered them variants of a single 
race, S. orcutti licki. However, HalTs discovery in 1969 that the licki 
variant was arboreal and the other petricolous prompted reexam- 
ination of material from Baja California Sur. Specimens of the 
unnamed taxon have been available in museums since 1890. 



'Department of Environmental. Population and 

Organismic Biology, University of Colorado 

Boulder, Colorado 80309 
-Current Address: Department of Genetics 

University of Melbourne, 

Parkville, Victoria 3052, Australia 



BREVIORA 



No. 452 




Pacific 
Ocean 



O Sceloporus orcutti 
• Sceloporus n.sp. 
A Sceloporus licki 



Fig. 1. Distribution of the three species of the Sceloporus orcutli complex in 
Baja California. 



1979 



LIZARDS OF THE SCELOPORVS ORCUTTl 



California 




Pacific 
Ocean 



O Sceloporus orcutt. 
9 Sceloporus n sp 
/^; Sceloporus Itcki 







30 Kn 



SAN JOSE 
DEL CABO 



CABO DE SAN LUCAS 



Fig. 2. Distribution of Sceloporus himsakeri. licki and orcutii on or near the 
Cape region of baja California. 



4 BREVIORA No. 452 

Comparative material relevant to this problem has been exam- 
ined from most of the major museums in the country. We are 
indebted to the curators and directors of these institutions. Cited 
specimens are referenced to collections by the following acronyms: 
CAS. California Academy of Sciences; CUM. University of Colo- 
rado Museum; LACM, Los Angeles County Museum; LMK, L. M. 
Klauber pers. coll.; LSJU, Leland Stanford Jr. Univ.; MCZ, 
Museum of Comparative Zoology, Harvard University; MVZ, 
Museum of Vertebrate Zoology, University of California, Berkeley; 
SDSNH, San Diego Society of Natural History Museum; UMMZ, 
University of Michigan Museum of Zoology; and USNM, U. S. 
National Museum of Natural History. 

We name the newly recognized species for Dr. Don Hunsaker H, 
in recognition of his pioneering studies in the comparative ethology 
of the genus Sceloporus and for stimulating karyological study of its 
species by Hall. 

SceU)ponLs hunsakeri. new species 

Holoivpe: MVZ 73,570, adult male, tail complete, intact although 
fractured in two places in distal half, not regenerated, from 3 mi. E 
of San Bartolo ±500 ft.. Feb. 17, 1960. R. G. Crippen, field no. 277. 

Paratvpes: Thirty-three, including eight topot\pes, two of which 
were taken on the same date as the holotype ( MVZ 73.572, 73,575) 
and six on March 14, I960 (MVZ 73,579-84); two from 5 mi. SW of 
San Jose del Cabo, July 6, 1970, Richard L. Holland (CUM 
45,880-1); two from Migriiio, 33 mi. SSE of Todos Santos, July 5, 

1969, RLH (CUM 40,779 80); one from '/: mi. N of Migrifio, July 8, 

1970, RLH (CUM 45,882); one from 1 V2 mi. N of Migrifio, July 9, 
1970, RLH (CUM 45,883); two from 7 km WNW of Buenavista, 100 
m, Aug. 3, 1969, W. P. Hall (MCZ R 1, 222,41 3); one from bridge at 
Km 105.32 on Mex. Hwy. 1, Aug. 3, 1969. W. P. Hall (MCZ 
R122,242); one from Cabo San Lucas. Aug. 4. 1919. J. R. Slevin 
(MCZ 15,565); one from 2 mi. NW of Cabo San Lucas. P. Elias, M. 
Merrill, and J. Lazell. Jan. 29. 1972 (MCZ R13I.704); one from 
Cabo San Lucas. Elias. Merrill and Lazell. Jan. 29. 1972 (MCZ 
R 131. 706); 10 from 6.4 mi. E of San Lucas. Playa Publica. T. Paul 
Maslin. Apr. 8. 1974 (CUM 51.401 10); and three from 6.5 mi. E of 
San Lucas. 300 ft.. T. Paul Maslin, Apr. 9, 1974 (data from last 13 
not included in tabulations herein). All of the type material is from 



1979 LIZARDS OF THE SCELOPORUS ORCITTI 5 

the Cape Region south and east of the Isthmus of La Paz in the 
territory of Baja CaHfornia Sur. Mexico. 

Other specimens examined. A total of 67, all from the Cape 
Region of Baja California Sur. Mexico, was examined only by Hall: 
Agua Caliente, 800 ft. (CAS 46,790-1, MVZ 1 1,709-1 1); 2.7 mi. W 
of Agua Caliente, El Chorro (LACM 18,880 2); Rancho Buena 
Vista (LACM 18,883): 4 mi. SE of Rancho Buena Vista, Cerro 
Agua Amargo, 23° 36' N, 109° 37' W (MVZ 50,070); Boca de la 
Sierra (CAS 91,457, LACM 18,879. SDSNH 30,188-9); Cabo San 
Lucas (CAS 16.532-3. 46,800-6, 46,808, SDSNH 32,903); 0.3 mi. 
SW of Cabo San Lucas (CAS 91.446); 7.7 mi. NE of Cabo San 
Lucas (CAS 9 1 ,3 19-20); El Triunfo (CAS 46,779, 46,823. 46,826-8); 
1.3 mi. N of El Triunfo (CAS 91,476); La Paz (CAS 46.775. 
SDSNH 17.696-8); 10 mi S of La Paz (SDSNH 30,191 3); 12.4 mi. 
E of La Paz on road to Las Cruces (CAS 91,255): 12.9 mi. E of La 
Paz. Arroyo de los Pozos (CAS 91,367); 16 mi. E of La Paz 
(SDSNH 30,186); 19 mi. N of La Paz [N of Pichilinque Bay] (CAS 
91,093); 19.5 mi. SE of La Paz, on road to Los Planes (CAS 
91.105-6); 20 air mi. SE of La Paz. 1 mi. SW of La Trinchera (CAS 
91.974-6); 21 mi. ESE of La Paz, 18.5 mi. from jct. road to Los 
Planes and La Paz airport road (CAS 91,222); Los Frailes (SDSNH 
32,920); Bahia Los Frailes (CAS 15.739); 7 mi WNW of Los Planes 
near Lat. 24° (MVZ 73.587-8); Ojos Negros (USNM 37.675-6; San 
Antonio (CAS 46.830); San Bartolo (CAS 46.787, SDSNH 38,075); 
1.1 mi. SE of San Bartolo (CAS 91,384-6); 3 mi. E of San Bartolo 
(CAS 90,972 5); 15 mi. S of San Bartolo (SDSNH 52,991). An 
additional 12, all young, were examined by both authors: seven 
topotypes. five of which were taken on the same date as the holotype 
(MVZ 73,571. 73.573-4. 73.576. and 73.578) and two on March'l4. 
1960 (MVZ 73.585-6); one from 2 mi. NW of Cabo San Lucas. 
Elias. Merrill and Lazell. Jan. 29, 1972 (MCZ R13 1,703); one from 
Cabo San Lucas, Elias, Merrill and Lazell, Jan. 29, 1972 (MCZ 
131,705); and three from Boca de la Sierra, 2 mi. N of Miraflores. 
April 17, 1977, R. Seib (MVZ 144,800-2). 

Diagnosis. A small (to 86 mm (5,71 9 ) member of the spinosus 
group of Sceloporus (as of Smith, 1939; Smith and Taylor, 1950), 
with supraoculars in a single row (undivided) rear two in contact 
with median head scales, no dorsal nuchal collar, dorsal scales 
33-37 (X 35.1); femoral pores 12-18 (15.9); posterior supraocular 
usually (94';r) in contact at least narrowly with a superciliary scale; 



6 BREVIORA No. 452 

zero to 6 (1.9) supernumerary granules between supraoculars and 
superciliaries (i.e., granules in excess of a single row or parts 
thereof); internasals two only, rarely varying in number or position; 
subnasal usually (65.2%) present; preocular usually (72.3%) hori- 
zontally divided; scales between postparietals often (45.5%) four or 
five; number of single scales between subocular and supralabials 
often (64.6%) 2-4; preanal escutcheon scales 7 19; no evidence of 
continuous light lateral stripe in shoulder region; dark bars on 
throat of females and young usually not including a pair of central 
parallel lines; a central dark blue patch on throat in adult males; 
black patch on shoulder never containing a central light spot. The 
species is a petricole and has a 2n = 34 karyotype. 

Description of holotype. Four symmetrical postrostrals, laterals 
about 1 '/4 times as large as medial, squarish pair; a single lorilabial 
row contacting rostral between postrostrals and supralabials; no 
subnasal; two canthals; a large loreal separating 2nd canthal from 
lorilabials; latter in two rows posterior to middle of 1st canthal, 
except for interruption by one or two scales between rear of 
subocular and supralabials; latter 4 4 to below middle of eye; 
preocular divided; one enlarged, strongly keeled postocular in 
contact with subocular, upper postocular apparently destroyed by 
abrasion, others small, similar to temporals. 

Two large, symmetrical internasals narrowly contacting 1st can- 
thai, broadly contacting median pair of postrostrals and all three 
frontonasals; one small supranasal; two minute postnasals; median 
frontonasal somewhat larger than lateral frontonasals; two pre- 
frontals, each intermediate in size between lateral and medial 
frontonasals, broadly in contact medially; frontal divided, posterior 
section 2/3 size of anterior, narrowly contacting interparietal; one 
large frontoparietal on each side, 1/2-2/3 size of posterior section 
of frontal; interparietal large, about as broad as long, widest 
posteriorly; a large parietal on each side, each about 3/5 size of 
interparietal; a well defined postparietal on each side, each over 
twice as wide as long, separated medially by 4 scales, narrowly 
contacting interparietal on one side only. 

Supraoculars 5 5, undivided, anterior three separated from me- 
dian head scales by a single row of small scales contacting posterior 
canthal, posterior two broadly in contact with parietal and inter- 



1979 LIZARDS OF THE SCELOPORiS ORCLTTI 7 

parietal; four strongly imbricate superciliaries as seen from above, 
two more subimposed under junction of 3rd and 4th dorsal 
superciliaries; one row of small scales extending from rear edge of 
1st superciliary (or very near it) posteriorly between all supraoculars 
and superciliaries except at posterior extremity, where 5th supra- 
ocular directly contacts (for about 3/5 of its length) rear supercili- 
ary; 2-2 small supernumerary scales (over and above one continu- 
ous row) in the area between supraoculars and superciliaries. 

Auricular' lobules 4-4 directly over ear opening, upper longest, 
next one broadest, lower one smallest and scarcely protruding over 
tympanum. 

Infralabials 5-5 to below middle of eye; three pairs of post- 
mentals, members of anterior pair in contact medially, those of 2nd 
separated by two scales, of 3rd by four; labiomental row separating 
postmentals and infralabials, except for one anterior scale in each 
series, allowing broad contact of 1st postmental and 1st infralabial; 
gulars all smooth and with a single apical notch, except below ear. 
where a faint keel and two notches separated by a median mucrone 
are present on a few scales. 

Dorsal scales strongly keeled, strongly mucronate. in vertebral 
region more acutely pointed and with fewer and shallower notches 
and shorter accessory m.ucrones than in more lateral regions, in 
obliquely converging rows, 33 in least count from interparietal to 
level of rear edge of thighs held at right angles to trunk. Ventrals 
smooth, with a single apical notch except at sides, where they blend 
with lateral scales. Femoral pores 16 17. the two series separated by 
six scales. Preanal scales similar to other ventrals. except for an 
escutcheon involving eight scales in two transverse series near 
posterior edge of preanal area; escutcheon scales thickened, smooth- 
edged, not notched, somewhat similar to femoral pore scales but 
the glandular (?) area spread over the entire exposed surface, not 
restricted to one area, and not so deeply penetrant. A deep lateral 
nuchal pocket, without scales; no axillary or postfemoral pocket. 

Scales oxi limbs are smaller in general than those on body, but 
otherwise grossly similar; scales on rear of thighs as large as scales 
preceding femoral pores, but mucronate, keeled and with accessory 
mucrones and notches; lamellar formula for fingers 9 12 16 17-12 
(9 13-16 17 12) and for toes 8-12-17-20-15 (8-12 17 20-14). 



8 BREVIORA No. 452 

Snout-vent 78 mm; tail 117 mm; snout-posterior margin of ear 
opening 18 mm; maximum width of head, at ear level, 17 mm; 
foreleg 32 mm; hind leg 49 mm; 4th toe from base of 5th, 20.5 mm. 

In preservatives, color more or less uniform dark tan above, with 
no markings except an iridescent bluish streak on some scales, 
especially along keel; foreleg with weak crossbands, more conspicu- 
ous distally; head somewhat lighter than trunk; tail with dim darker 
brown bands, most distinct medially. 

Melanization of ventral surfaces nearly but not quite complete, 
with a light chin blending with a darker infralabial pigmentation 
crossed by a few narrow diagonal light lines; central and entire rear 
area of throat black with dark blue lateral highlights; black on 
throat extending onto anterior chest area, but light areas remaining 
in central lateral chest areas at level of axillae; throat color 
extending dorsad as a shoulder patch, light margined (one scale 
wide) posteriorly, entering lateral nuchal pocket anteriorly but not 
passing beyond it; light posterior margin crossing arm insertion; no 
light areas within shoulder patch. Sides of belly dark mauve, 
blending with a broad black median border shot with dark iri- 
descent blue areas; lateral belly patches not fully confluent medially 
or with gular patch, although limited continuity exists in both areas; 
black in groin continuous with belly patches but rear half of ventral 
surfaces of thighs largely light; preanal area entirely light. 

Variation. The 33 specimens in the type-series have 33 37 dorsals 
(33, five; 34, six; 35, five; 36. thirteen; 37, four) with a mean of 35.1. 
The range in the counts taken by Hall for the "other specimens" is 
considerably greater (range 31-41; 31, one; 33, four; 34, three; 35, 
nine; 36. eleven; 37, eighteen; 38, ten; 39, seven; 41, one) and the 
mean 36.3 is also greater. The counts by Hall were taken at various 
times under less controlled conditions and include several partially 
decornified specimens which are difficult to count accurately be- 
cause the underlying scales are very soft, easily distorted and 
difficult to discern. Specimens reexamined by Smith with few 
exceptions have 13 fewer dorsals than Hall originally counted. 
Nevertheless we regard older counts that cannot now be rechecked 
as useful indicators of a somewhat greater range than is represented 
in the relatively small type-series. 

The femoral pore counts taken by both authors are essentially 
identical and our pooled results yield a range of variation in 169 



1979 I 1/ AKDS OK IHE SCELOPORLS ORCLTTI 9 

counts from 12 to 18(12, one; 13. five; 14, sixteen; 15, thirty-five; 16, 
fifty-eight; 17, forty-three; 18, eleven) with a mean of 15.9. There is 
no significant difference between counts in the two sexes (males, N 
= 98. 13-18, X 15.89; females N= 58, 12-18, X 15.84; 13 counts not 
allocated to sex, 13 17, X 15.84). 

Head scale characters were recorded only for the 33 individuals of 
the t\pe-series. In 66 counts on the contact of rear supraocular with 
a superciliary, only 4 (1.6''x) had the rear supraoculars separated 
from the superciliaries (the first supraocular is in broad contact with 
the first superciliar\' in all). The supernumerary scales var\' from 1 to 
6 (0, nine; 1 . seventeen; 2. twenty-one; 3. ten; 4. four; 5. three; 6, one) 
with a mean of 1 .9. 

The variation in other head scales, including very small scales, is 
exceptionally limited. The scales in the internasal area are especially 
constant relative to the variation occurring in most Sceloporus. All 
scales are as described for the holotype except: separate postrostrals 
four in 30, two in 1, five in 1, median pair fused with the supranasals 
on each side in 1, one pair of large internasals in 32, three in 1; 
internasals contacting anterior canthal on both sides in 21. sep- 
arated by a separate small scale on both sides of two, on one side in 
four, by contact of a postnasal and lateral frontonasal on both sides 
of three, on one side in one, and in one specimen by a postnasal- 
lateral frontonasal contact on one side and by a separate small scale 
on the other. The internasals always contact the postrostrals 
(although the central two postrostrals are fused with the supranasals 
in one specimen) and frontonasals, so that always there are two 
scales in a straight median (or closely paramedian) line between 
median frontonasal and rostral. Subnasal present on both sides in 
20, on one side in three, on neither side in 10; preocular horizontally 
divided on both sides in 21, on one side in four, on neither side in 
seven (indeterminate on one side in one, divided on other); post- 
nasals 2-2 in nine, 2-3 in nine, 3-3 in 14, 4-4 in one; scales between 
postparietals two in two, three in 16. four in 11. five in four; 
postparietals contact interparietal on both sides in 29, on one side in 
three, on neither side in one; two enlarged, keeled postoculars in all; 
number of single scales contacting both subocular and supralabials 
4 (59 counts: 0, three; 1, twenty; 2, twenty-three; 3, seventeen; 4, 
two); 1 St canthal failing to contact lorilabials only on one side of one 
(lower edge split off as a separate scale); two frontoparietals on one 



10 BREVIORA No. 452 

side in five, all others with 1-1; scales of 1st pair of chinshelds 
separated by one scale in one, of 2nd pair by three scales in two, and 
by two in 31, of 3rd pair by four (14), five (18), or six (1) scales; 
frontal divided into three scales in one, into four in one (in the latter, 
the anterior of three transverse divisions is split into right and left 
components). 

All mature males possess a clearly defined preanal escutcheon, 
each scale of which contains a light colored flaky tissue surrounded 
by a narrow border of cornified black pigmented tissue. Scales in 
the center of the escutcheon are more strongly modified than those 
towards the margin. The structure is also discernible in females and 
young by scale thickness and the lack of an apical notch, but the 
modified scales can be reliably counted only in the adult males. In 

1 1 such individuals from the type-series, counts range from 8 to 14 
(8, two; 9, two; 10, three; 1 1, two; 13, one; 14, one) with a mean 9.3. 
In 33 comparable specimens examined only by Hall, the range is 7 
to 19 with a mean of 12.5 (see Comparisons for details). All counts 
are more or less subjective, as frequently several scales are only 
partially modified to various degrees. 

In live, fully melanistic males (68 mm s-v and larger, as recorded 
in color photographs taken in the lab, within a week of capture, 
when the animals were at or near optimum temperature), the central 
gular, chest, preanal. shoulder and shank areas are black, the lateral 
belly patches distinctly navy blue; four to five iridescent green 
diagonal stripes one to two scales wide divide the blue belly patches 
into darker ventral chevrons, each about three scales wide. The 
underside of the tail is abruptly lighter than the trunk and only 
faintly dusky. The underside of the foreleg, chin and infralabial areas 
are darker, and some evidence of narrow light bars is present on the 
lower lips and chin. Dorsally, the basic body color is an iridescent 
greenish bronze to light metallic green which grades to a slaty, 
metallic blue or purple along middorsum from between shoulders to 
over the tail base. This darker area is about seven to eight scales 
wide at the widest and grades into the lighter ground ct)lor rather 
than being sharply differentiated from it. In no specimen is there a 
light central spot in the shoulder patch. 

Adult females and subadult males are light brown to dark 
greenish brown or slate above, with scattered scales exhibiting an 



1979 LIZARDS OF THE SCELOPORUS ORCUTTI 11 

iridescent bluish apex; lips, tail and limbs barred, more strongly on 
distal parts. In both sexes, 3 4 dark bars separated by lighter stripes 
radiate from the eye on sides of the head. A black shoulder patch is 
always present, light-bordered both anteriorly and posteriorly in the 
young. In older animals of both sexes the anterior light border 
becomes obscured by encroachment of black, whereas the posterior 
border remains intact and distinct in all specimens. Typically the 
upper and lower extremities of the anterior border persist for a time 
as isolated light spots, and in due time only the lower persists, 
sometimes completely enclosed by black. In mature adults even that 
disappears completely. In no developmental stage is there ever a 
central light spot within the shoulder patch. Very small individuals 
show about 1 1 transverse rows of small, obscure, dark brown spots, 
larger and more distinct paravcrtebrally. on the trunk; the series 
continues onto the tail. This pattern soon fades with maturity, 
although the row of spots at shoulder level becomes briefly accentu- 
ated and weakly collarlike, before it becomes obscured by the 
general darkening of the dorsal surfaces; adult females may retain 
vestiges of the collarlike marking but it is completely indiscernible in 
adult males. 

On ventral surfaces, the dark gular bars are bluish, and faint 
bluish belly patches shading laterally to tan are visible in mature 
females. The medial borders of the belly patches are comparatively 
dark but still considerably fainter than the gular bars. The patches 
are separated by about five to seven scale rows along the midline. 

In all young specimens the throat, chin and lower lips are whitish 
but prominently barred, the dark lines twice as wide as the 
intervening spaces and converging toward the rear center of the 
throat. These markings persist throughout life, except that in 
mature males they are extensively obscured by the black area 
expanding from the rear center of the throat. The throat bars 
seldom form parallel lines on midthroat. 

One adult male S. huusakeri from the paratypic series had a 
karyot\pe (MCZ Karyology catalog no. Y 14,683. main catalog no. 
MCZ R 122.241 ) of 34 chromosomes in the typical primitive pattern 
for sceloporines (Paull et al. 1976). 5". licki and 5". onuiti have the 
same chromosome number (Cole 1970, Hall 1973. in prep.). The 
karyotyped male -S. hunsakeri was collected while actively involved 



12 BREVIORA No. 452 

in territorial defense and was maintained in the lab with a female 5". 
hunsakeri and several S. licki for seven weeks before being sacri- 
ficed for karyology. The male was frequently seen courting the 
female throughout this time. He paid no attention to the S. licki 
housed in the same cage. When karyotyped, his testes still showed 
active spermatogenesis, with high frequencies of spermatogonial 
mitoses, meiotic prophases, second metaphases and maturing 
spermatids. We conclude from this that the male was sexually active 
at least from the period of collection, 3 Aug. 1969, through 24 Sept., 
when it was karyotyped. 

Ecologically, 5". hunsakeri appears to be a strict petricole and 
extremely wary. Many were observed 2 3 August 1969, when Hall 
was collecting in the Cape region of Baja California, but except for 
the two male S. hunsakeri collected while actively displaying to one 
another and a female trapped under a small bridge, all took alarm at 
a distance many times the 2 m length of the noose pole. In most 
cases, just at the sight of the collector's vehicle, they escaped into 
deep crevices or under massive boulders. None were observed in 
alluvial or wooded areas lacking massive rock outcroppings. Essen- 
tially all collection localities recorded for S. hunsakeri are rocky. 

Comparisons 

Past consideration of 5". hcki as a subspecies of S. orcutti was a 
product of I) the similarities oi S. hunsakeri and S. hcki in size and 
squamation; 2) the close parallelism of 5'. hunsakeri and S. orcutti in 
ecology and coloration; and 3) the near parapatry of 5". orcutti and 
S. hunsakeri. To establish S. hunsakeri and reestablish S. hcki as 
valid species we must show: 1) that the differences between 5". 
hunsakeri and S. hcki are sufficient to rule out the possibility that 
these are not simply different morphs of a polymorphic population 
(as Smith [1939] originally believed); and 2) that the differences 
between S. hunsakeri and S. orcutti are sufficient to indicate the 
probable existence of intrinsic barriers to gene flow between the two 
populations. 

S. hunsakeri vs. S. hcki. Although hcki and hunsakeri are 
essentially identical in body and head proportions, in femoral pore 
and body scale counts, and in most cranial scale characters, other 
differences are of a degree and nature that cannot be accounted for 
by allelic differences at a small number of loci. Most of the 



1979 



LIZARDS OF THE SCELOPORL'S ORCUTTI 



13 



differences are summarized in Table 1. The most constant differ- 
ences in scutellation pertain to the relation of the posterior supra- 
ocular to the superciliaries ( Fig. 3), irregularities of internasals ( Fig. 
4) and the size of the escutcheon (Fig. 7). In S. hunsakeri the 
modified escutcheon scales are in two or more transverse rows, 
whereas in 5. licki usually only one scale row is involved, with three 
to five scales fully modified and a couple more partially so (i.e., at 
least half of the scale surface modified). A few additional scales, not 
counted in the tabulated totals, may show slight modifications of 
the scale tip. Though in S. hunsakeri more scales are involved in the 
secretion, not all are as sharply differentiated from the surrounding 
normal scales. The developmental field controlling differentiation 
appears to have a much smoother gradient so that the scales at the 
anterior and lateral peripheries of the escutcheon are all only 
partially modified. Thus the counts of these scales for both S. 
hunsakeri and S. orcutti, which have similar developmental patterns 
for this character, are not precise. Some females and juveniles show 





hig. 3. Contrast between 5. licki (left, MVZ 144,803) and 5'. hunsakeri (right, 
MCZ R 122,241) in number of supernumerary scales between superciliaries and 
supraoculars, and in contact of posterior supraocular with superciliaries (actual field 
length is 8.6 and 9.1 mm respectively). In 144,803, rear supraocular and superciliary 
are completely separated by small scales and five supernumeraries. In R 122,241, rear 
supraocular contacts rear superciliary, and two supernumeraries present. 



14 



BREVIORA 



No. 452 



^ 



^ 



■r- E 



i^ ^ ~ 



^ ir :rr 



sC, ^ „ 



^ W-) 



^ -d 



.S — "^ 



S XJ _- > 



33 E « 



^ r. a 



-z ^ ^ 



+ 2 



w o 



^ t; .±; 



^ CA 



1979 LIZARDS OF THE SCELOPORUS ORCUTTI 15 



u [^ -o 



5 I ^ ^ is 



S 2 S S 2 






if 2 '-' - 
.T! n iS c 



2 «J ^ t: >■ ? '2 
i; c — Bj — :i — ; 



s ^ 



2 s 



■n o 



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16 



BREVIORA 



No. 452 



some degree of modification of the preanal scales but never to the 
extent seen in the adult males. The appearance of the escutcheon 
strongly suggests an organ specialized for the secretion of a 
substance to be rubbed against the substrate as the vent is wiped 
across it. Since fully developed escutcheons are only found in 
reproductively mature males, the secretion may have a pheromonal 
function possibly involved in maintaining reproductive isolation. 
The scale differences between the two species, though not con- 
spicuous, are diverse and distinctive. However, differences are 
strikingly apparent in color pattern and appear to be completely 
diagnostic in living and preserved material (Fig. 5). Color observa- 
tions are based on the entire series of preserved specimens examined 
by Hall, on color transparencies of live specimens of both sexes and 
on the preserved material available to both authors. One set of 




Fig. 4. Contrast between 5'. licki (left. MVZ 144,803) and S. hun.sakcri {ngbl. 
MVZ 144,800) in structure of internasal area; actual field length 1 1.5 mm in each (in 
median line). In 144,803 three posterior internasals, two primary internasals, latter 
separated from first canthal on each side by tiny scale posterior to an upper(left) and 
2nd (right) postnasal; the four postrostrals irregular; suprana.sal on each side 
relatively large, frontoparietals in contact. In 144.800, primary internasals the only 
internasals present, separated from first canthal on each side b> contact of upper 
postnasal and lateral frontonasal; this symmetry is typical. 



1979 



LIZARDS OF THE SCELOPORi'S ORCUTTI 



17 





Fig. 5. Contrast between young male (39.5 mm s-v) 5. licki (left, MVZ 73,577) 
and young male (35.5 mm s-v) S. hunsakeri (right, MCZ R13 1,703) in pattern of 
throat bars: actual field length 16.0 and 16.5 mm respectively. Parallel paramedian 
lines pronounced in 73,577, visible throughout life in both se.xes since males lack a 
central rear gular patch; in R 1 3 1 ,703 the lines are largely oblique, scarcely parallel at 
any point, and are obscured in adult and subadult males by a rear central gular patch. 



differences associated with overall ground coloration and patterning 
appears to be correlated with ecological differences between the 
species. The remaining differences all relate to specific body areas 
presumably used in social signaling and species recognition. 

The body color below the dorsolateral light stripe also shows some 
patterning of slightly lighter vertically elongated spots in both sexes 
of 5'. licki. In the field, differences in basic ground coloration and 
the conspicuously visible dorsolateral stripes of 5". licki allow the 
collector to distinguish between these two species from distances of 
many meters. These differences are correlated with the basic colors 
of the preferred substrates frequented by each species. The relatively 
dull and uniform pattern of 5". hunsakeri is adaptive to its habitat on 
unpatterned rocks, whereas the 5". licki pattern is equally as 
disruptive as the light and shadow on or under the trees that 
constitute the habitat for that species. 

Other color differences are all associated with body structures 
presumably involved in social signaling and/or species recognition. 



18 



BREVIORA 



No. 452 



The dark shoulder patch seems hkely to serve as an orientation 
mark for lateral displays (Fig. 6). In S. himsakeri it may be set off in 
front by a light line or series of spots in incompletely melanized 
individuals. Faint ventral patches are visible in females of both 
species, but unlike the contrast in males, they are more intense 
medially, with the median dark edges separated by several scale 
rows. 

Although karyotypes of 5. hunsakeri, S. licki and S. orcutti are 
identical (Cole 1970, Hall 1973, in prep.), there is evidence that the 
reproductive seasons of 5". licki and 5". hunsakeri differ. Two licki 
males were karyotyped, one six days after capture 6 August (MCZ 
original number Y18,801 = MCZ 136,188), the other about seven 
weeks after capture (MCZ original no. Y14,689 = MCZ R122,233). 
Both showed completely regressed testes, with the seminiferous 
tubules collapsed to a few mm"* within the loose and shriveled 
tunica, only a few mitoses and no hint of sperm or any stage of 
meiosis. On the other hand, the 5". hunsakeri male karyotyped after 
7 weeks of captivity still showed very active spermatogenesis and 





Fig. 6. Contrast between young male (39.5 mm s-v) .V. /;< A/ (lett.MCZ 144,803) 
and a subadult male (61.5 mm s-\) S. hunsakeri (right, MCZ 73,575) in relation of 
dark shoulder patch to a central light spot; actual field length 5.8 and 12.0 mm 
respectively. A light dot, conspicuous m life hut small here, within dark spot in S. 
licki, absent in 5'. hunsakeri. 



1979 



LIZARDS OF THE SCELOPORUS ORCUTTI 



19 




Sceloporus licki 

Paratype 
CAS 1423 



5 

\ I I I I I m m 




Sceloporus n. sp. 
CAS 46828 

Fig. 7. Contrast between adult male .S. //cA/deft, CAS 1,423) and adult male 5. 
hunsaken (right, CAS 46,829) showing development of preanal escutcheon scales. In 
5. hunsakeri. last row of preanals folded under and are not visible in illustration. 



20 BREVIORA No. 452 

courtship behavior. The 5'. licki individuals held in the same cage 
with the S. hunsakeri, though apparently in good health, showed 
very little social or courtship behavior. Although this sample size is 
small, these observations along with the visual and pheromonal 
differences noted above, suggest these two species may have quite 
different reproductive seasons. 

Therefore, although these two species are similar in most charac- 
ters of proportion and squamation, they show many significant 
differences in other aspects of their biology, and we believe they 
should be considered genetically independent and ecologically 
differentiated sympatric species. Based only on these color pattern 
differences, even the oldest of the preserved specimens may be easily 
allocated to one or the other species. 

5". hunsakeri vs. S. orcutti. It is evident that S. hunsakeri and S. 
orcutti occupy similar ecological niches characteristic of strict 
petricoles. They also have many aspects of coloration in common 
(e.g., see Smith, 1939). There is no evidence that their ranges 
contact, except for one 5". orcuiii and four 6". hunsakeri with locality 
data of "La Paz" (which may only be a collecting headquarters). The 
Isthmus of La Paz is an alluvial outwash from the mountains to the 
south and is presumably of fairly recent origin. According to 
topographic maps of the area (Comision Intersecretarial Coordina- 
dora del Levantamiento de la Carta Geografica de la Republica 
Mexicana, 1:500,000 scale, quadrangles I2R-VII, I2R-VIII, and 
12Q-II), the maximum elevation of the divide across the Isthmus 
between the Pacific and Gulf slopes is less than 200 m. The depicted 
200 m contour indicates that at least 20 km separates any area of the 
Cape over 200 m from any area of the Peninsula proper over 200 m. 
Except for the questionable specimens from "La Paz", there are no 
locality records for either 5'. orcutti or 5". hunsakeri from this 
lowland. Papenfuss (person, commun.) has collected extensively in 
this area and he does not recall having observed "orcutti" or any 
potential S. orcutti habitat. Presumably S. orcutti and S. hunsakeri 
are dichopatric, although their ranges are close to each other 
(specimens with reliable localities are separated by about 40 km). In 
the absence of sympatry to assure that reproductive isolation exists, 
other criteria must be examined to determine the taxonomic status 
of these two isolated populations. 



1979 LIZARDS OF THE SCELOPORLS ORCUTTI 21 

S. hunsakeri and 5". orcuiti differ conspicuously in morphology, 
unlike 5". hunsakeri and 5'. licki. A considerable size difference and 
diagnostic scale characters are obvious from comparisons of Baja 
California Sur materials (Table I). Northern 5". orcutti reach an 
even larger size than this small sample indicates. 

Head scales differ in percentages of given character states but not 
categorically in any character. The most constant distinction we 
discovered li^s in complete separation of the rear supraocular from 
the rear superciliary in most (85%) 5'. orcutti and contact in most 
(94%) 5'. hunsakeri. Secondly, the internasal scales are quite irregu- 
lar in number and or position in S. orcutti, but are seldom 
unsymmetrical in S. hunsakeri. Thirdly, the major pair of inter- 
nasals contacts the first canthals on neither side in ten 5". orcutti and 
on 47 of 66 sides (71.2%) in 5". hunsakeri. Lesser distinctions are 
noted in Table 1. 

Finally, there is little overlap in either dorsal scale or total 
femoral pore counts for the two species. Dorsal scales for 28 
southern peninsular S. orcutti examined by Hall range from 29 to 34 
(X[±SEM] 31.77 ± 1.295) while 81 S. hunsakeri v?ingQirom7>\ to 43 
(X 38.13 ± 1.884). The hypothesis of equivalency is rejected at the 
.005 level using the ttest with 32 d.f. Total femoral pore counts 
range from 22 to 27 (X 23.83 ± 1 J81) in 28 5. orcutti, whereas 84 S. 
hunsakeri range from 25 to 36 (X 31.56 ± 2.282) with equivalency 
rejected at the .005 level. A discriminant function can be generated 
from these data (Fig. 8) for 27 S. orcutti and 81 S. hunsakeri by 
adding the total femoral pore number to the dorsal scale count for 
each individual. This function separates all but two of the individ- 
uals examined. All S. orcutti except one have 59 or fewer dorsal 
scales plus femoral pores, whereas all but one 5. hunsakeri have 61 
or more. The mean value of the discriminant for 19 5'. hunsakeri 
with localities within 32 km of La Paz (i.e., the Isthmus of La Paz) is 
68.23 ± 2.96 compared with 68.03 ± 2.93 for all 81 5. hunsakeri. The 
four individuals from "La Paz" range from 69 to 71 with a mean of 
70.0. Two 5. orcutti, one from "La Paz" and one from 20 mi. W of 
La Paz are respectively 54 and 53, at the opposite end of the range of 
variation in S. orcutti from the values (60-74) for 5". hunsakeri. 

Color differences between the two species are completely diag- 
nostic, insofar as we can compare them. S. hunsakeri does not 



22 



BREVIORA 



No. 452 



N 
O 



00 -"^ 

II C^ ^ " 

CM • -CM 
II CD CO 11 




Fig. 8. Comparison of discriminant functions for S. hunsakeri and .9. orcutii 
(most of the latter from Baja California Sur). S. orcuiti specimens are indicated by 
circles, 5. hunsakeri by squares. Solid symbols indicate specimens from "La Paz"; 
symbols containing an X indicate specimens with collection localities within 20 miles 
of La Paz. 



1979 LIZARDS OF THE SCELOPORVS ORCUTTI 23 

possess the distinctive, narrow diagonal crossbars on both dorsal 
and ventral surfaces so conspicuous in northern S. orcutti. The 
barring, extensively obscured in adult S. orcutti, persists at least on 
the sides of the body in the largest specimens. Other differences are 
noted in Table 1. Southern S. orcutti appear to maintain these 
distinctions, although we do not now have substantial recent 
material to support this contention. 

Clearly these data do not provide evidence for past or present 
introgression between the distinctly larger, larger-scaled 5". orcutti 
and the smaller, smaller-scaled ^. hunsakeri. To conclude this 
comparison we are left with the problem of assessing the taxonomic 
level of difference between populations which are and have been 
geographically separated. Our experience with the genus leads us to 
regard the observed differences as a strong indicator that the two 
taxa would be genetically isolated if they w^re in contact either 
through parapatry or sympatry, and therefore that S. hunsakeri 
should be recognized as a full species, rather than as a subspecies. 

S. licki vs. S. orcutti. S. licki would never have been closely 
compared with 5". orcutti if it had not been for the confusion of S. 
licki and >S. hunsakeri from the very outset (see next paragraph), 
perpetuated by Smith (1939). Early workers referred or compared 
material representing S. licki to S. undulatus consohrinus appar- 
ently because of similarities in pattern observed also by Hall. 
Yarrow (1883:61) and Belding (1887:98) so identified materials 
taken in 1882 at "La Paz" by Belding, and Van Denburgh's ( 1895) 
primary comparison of 5". licki was with 5". consohrinus. That 
association does not now seem rational since the closest relatives of 
S. licki certainly occur in more nearly adjacent territories. 

General comments on the S. orcutti complex 

A single paratype of 5". licki was recorded in Van Denburgh's 
description (1895:114) from San Jose del Cabo. taken Sept. 16, 
1890, by W. E. Bryant. Given the probable lack of 5". licki habitat 
there and the verified presence of 5". hunsakeri in later collections, 
the Bryant specimen was probably the earliest acquisition of 5'. 
hunsakeri. Van Denburgh also recorded a dark central gular patch 
in one male of his Miraflores series (1895:112; 1922:357); that 



24 BREVIORA No. 452 

specimen must likewise represent S. hunsakeri, since later collec- 
tions from Miraflores confirm that both S. hunsakeri and 5". licki 
are present in the vicinity. 

Unfortunately, the California Academy of Sciences collection 
(including the S. licki type and all but three paratypes distributed to 
other museums) was destroyed in the San Francisco earthquake and 
fire. Nevertheless, Van Denburgh's type-description referred to the 
striped form, which he clearly differentiated from 5". orcuiti by its 
smaller and more sharply keeled ("rougher") dorsal scales, from 5'. 
magister and 5". zosteromus by smaller and more mucronate scales, 
and from all of these species by color pattern. Each of the three 
surviving licki paratypes is clearly the striped morph. Two paratypes 
entered in the Stanford University collection at the time of the fire 
have since been returned to the California Academy of Science. One 
of these was designated by Smith (1939) as a neotype under its 
Stanford University tag number (neotype: CAS 1423 = LSJU 
2987a; when seen by Hall, both tags were still on the specimen). The 
other specimen from this series is CAS 1426 = LSJU 2988. The third 
paratype is from the Miraflores series, and is in the U.S. National 
Museum of Natural History (USNM 23,749). 

Stejneger (1893) described 5'. orcutti from USNM 16,330, from 
Milquatay Valley, San Diego County, California, collected by C. R. 
Orcutt. In reference to the type-locality it may be of interest to 
record that the bottle containing the holotype also contains a rather 
fragile note dated Aug. 14, 1934, signed by L. M. Klauber. We quote 
it here to insure against loss of the information it contains: "Orcutt 
once told me that the 'Milquatay' referred to was the flat just east of 
Campo, San Diego Co., Calif. Milquatay is said to mean wheat field 
in Indian, and almost any flat was known as Milquatay. The town 
of Guatay (or Quatay) is not the type locality of orcutti." 

The preanal escutcheon of S. hunsakeri and S. Ucki discussed in 
the preceding account occurs in 5". orcutti and several other species 
of the S. spinosus group as well. The development of the escutcheon 
in S. orcutti is very similar to the pattern seen in S. hunsakeri. 
However, its histology, functions, distribution in other species, and 
general taxonomic value remain to be explored. 

Dr. George Gorman and his students at UCLA have recently 
begun a survey of the electrophoretic variability of the Baja 
California Sceloporus. They inform us (person, commun.) that 



1979 LIZARDS OF THE SCELOPORUS ORCUTTI 25 

genetic distances between representatives oi S. orcuiti, S. Ilcki, and 
S. hunsakeri are all consistent with our separation of these popula- 
tions at the specific level. They also report that there appears to be 
electrophoretic evidence suggesting that even S. onutti as we have 
restricted its definition may be polytypic. However, we see no 
supportive external morphological evidence. 

Given the many taxonomic problems involved in classifying 
allopatric populations, we do not here make any formal attempt to 
allocate 5". orcuiti complex populations on the Gulf of California 
islands to any of the three peninsular species. This cannot be done 
definitively until information on the biochemical genetics of both 
mainland and insular populations is more complete, although 
external morphology and coloration will yield preliminary indica- 
tions. Prior to such examination, we suggest that the Espiritu Santo 
and Ballena island populations may be provisionally classified with 
5'. hunsakeri (although some differences in coloration are evident), 
and that populations from the more northern islands be grouped 
with S. orcutti. 



Literature Cited 

Belding, L. 1887. Reptiles of the Cape region of Lower California. W. Am. 
Sclent. 3(24): 97 99. 

Cole, C. J. 1970. Karyotypes and evolution oi i\\t spinosus group of lizards in the 
genus Sceloponts. Amer. Mus. Novitates (2431): 1-47. 

Hai I , W. P. 1973. Comparative population cytogenetics, speciation and evolu- 
tion in the iguanid lizard genus Sceloporus. Ph.D. Thesis, Harvard Univ. 

Hall, W. P. and R. K. Selander. 1973. Hybridization of karyotypically 
differentiated populations in the Sceloporus grammicus complex (Iguanidae). 
Evolution 27: 226-242. 

Paull, D., E. E. Wii llxms and W. P. H \i i . 1976. Lizard karyotypes from the 
Galapagos Islands: Chromosomes in phylogeny and evolution. Bre\iora No. 
441: I-3I. 

Smlih. H M. 1939. The Mexican and Central American lizards of the genus 
Sceloporus. Zool. Ser. Field Mus. Nat. Hist. 26: 1-397. 

Smiih, H. M. and K. R. Larsen. 1975. A new species of the /()/7»(;.vi/.v group of 
the lizard genus Sceloporus. Copeia 1975: 47 50. 

Smiiu. H. M. \\I) .1. D. Lv\c H. 1967. A new cryptic lizard (Iguanidae: Scelopo- 
rus) with comments on other reptiles from Oaxaca, Mexico. Herpetologica 23: 
18-29. 

Smith. H. M. AM) ,A. H.Swiizk'i. 1974. Another cryptic associate of the lizard 
Sceloporus fonuosus in Guerrero, Mexico. J. Herpetol. 8: 297-303. 



26 BREVIORA No. 452 

Smith, H. M. and E. H. Tavior. 1950. An annotated checklist and key to the 

reptiles of Mexico exclusive ot the snakes. Bull. U.S. Nat. Mus. No. 199: 1-253. 
Stejneger, L. H. 1893. Annotated list of the reptiles and batrachians collected by 

the Death Valley Expedition in 1891, with descriptions of new species. N. Amer. 

Fauna 7: 159 228. 
Van DtNBiRCiH, J. 1895. A review of the herpetology of Lower California. Parti. 

— Reptiles. Proc. California Acad. Sci. (2)5: 77-163. 
Van Denbi'RGH, J. 1922. The reptiles of western North America. Part 1. Lizards. 

Occ. Pap. California Acad. Sci. 10: 1-611. 
Yarrow, H. C. 1883. Check list of North American Reptilia and Batrachia, with 

a catalogue of specimens in the LInited States National Museum. Bull. U.S. Nat. 

Mus. (24): 1 249 (1882). 



APPENDIX 

ADDITIONAL SPECIMENS EXAMINED 

(Ail from Baja California Sur, Mexico, unless otherwise noted.) 

Sceloporus Ikki. Seoivpe: Sierra San Lazaro (CAS 1.423 = LSJU 2,987). 
Paratypes: Sierra San Lazaro (CAS 1,426 = LSJU 2.988); Mirafiores (USNM 
23,749). Other specimens: Boca de la Sierra (LMK-SDSNH 30,187); 2 mi. NNW of 
Cerro San Antonio (SDSNH 53,136 43); El Sauce ( = E1 Sauz). 4000 ft.. Victoria Mts. 
(MVZ 11,702 08); Guamuchil Rancho (CAS 46,809); La Burrera (SDSNH 45.062- 
64, 53.132-35); Arroyo Palmellar, 6 mi. ENE of Rancho La Burrera (LACM 
34,592-93); "La Paz" (USNM 53,392); 40 mi. S of La Pa/, 2 mi. above El Valle 
Perdido (CAS 90,544, 90,558); Miraflores (USNM 64,472); nr. km 70. Mex. Hwy. 1, 
8 km SE of San Antonio (MCZ 136.188); San Bartolo (CAS 46.780 1, 46,783 86; 
UMMZ 56,044; MVZ 144,809-13); 1.1 mi. SE of San Bartolo (CAS 91,383); 1.5 mi. 
E of San Bartolo, ±500 ft. (MVZ 73, 589); 3 mi. E of San Bartolo, ±500 ft. (MVZ 
73,570). 

Sceloporus orcutti. Holotype: California: San Diego Co., Milquatay Valley. 2500- 
2600 ft. elevation (USNM 16,330). Paraivpes: Same locality (USNM 16,238, 29, 
-33. -34). Other specimens: 15 mi. S of Canipole (LMK-SDSNH); 43 mi. N of 
Canipole (LMK-SDSNH); Rancho Chenque (SDSNH 51.044 46); Coyote Bay, 13 
mi. SE of Mulege (MVZ 37,318); 12 mi. E of El Arco (SDSNH 17.471); La Paz 
(LMK-SDSNH 38,339); 20.0 mi. W of La Paz, ±500 ft. (MVZ 73.590); San Ignacio 
(MVZ 10.653, -54, -56, 13,597. -99; LMK-SDSNH 4.053. 54; UMMZ 76,482[2], 
80,900); Mision Santa Gertrudis [Baja California Norte] (SDSNH 17,529 36). 



MAR 1 8 1985 

n DIVERSITY. 

B R E V I R A 

Museium of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 21 February 1979 Number 453 

THE LARGE PALAEOTRAGINE GIRAFFID, 

PALAEOTRAGUS GERMAIN/, FROM 

LATE MIOCENE DEPOSITS OF 

LOTHAGAM HILL, KENYA 

C. S. Churcher' 

Abstract. An isolated and slightly damaged left M' constitutes the first record of 
the species P. germaini in East Africa. This specimen matches in size and develop- 
ment of the buccal ribs and styles the upper molars of P. germaini from Oued el 
Hammam, Algeria. 

An isolated and damaged left upper molar, probably M' (KNM- 
LT 414, field no. 138/67K), was collected by Dr. V. J. Maglio from 
Lothagam Hill, Kenya, in 1967. The tooth was found at Lothagam-I 
near the base of Member B (Patterson et al. 1970), and is deposited 
in the collections of the International Louis Leakey Memorial 
Institute for African Prehistory, Kenya National Museums, Nairobi. 

The specimen (Fig. 1) is very slightly worn on the occlusal surface 
and derives from an immature individual. Minor damage has 
removed the mesial face of the crown, including the mesial half of 
the protocone, but the paracone is essentially entire. Slight damage 
to the distobuccal corner of the metacone has not destroyed the 
shape of the cusp or of the distobuccal wing of the hypoloph. Minor 
fragments of enamel are absent from the buccal surfaces of the 
apices of the paracone, metacone, and metastyle. The surface of the 
enamel is typically giraffid in its overall rugosity and thickness. The 
crown is tilted lingual on its roots in the anteroposterior (parasagit- 
tal) plane, and the roots tip distad. The mesial pair of roots is more 
damaged than the distal pair, all four roots are separate and are 



'Royal Ontario Museum and Department of Zoology, University of Toronto, 
Toronto, Ontario M5S lAl, Canada. 



BREVIORA 



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1979 PALAEOTRAGUS GERMAINI 3 

round or oval in section. The exception is the distobuccal or 
metacone root which is grooved on its distal surface and crescentic 
in section. 

The cusps are typically giraffid with tapered protocone and 
hypocone set buccally from the lingual cingulum. The paracone and 
metacone similarly are set lingually towards the mid-line of the 
crown. The buccolingual diameter over the mesial cusps (paracone 
and protocohe) is greater than that over the distal cusps (metacone 
and hypocone). There is a well-developed median rib on the buccal 
surface of the metacone and a small pU is present distally in the 
postfossette on the wall of the hypoloph. A similarly located and 
formed pli, which is broken away, appears to have been present on 
the protoloph. 

The specimen is attributed to Palaeotragus germaini Arambourg 
1959 because of its similar size (Table 1 ) and because it matches well 
with Arambourg's (1959, Pis. XIV. XV) illustrations of first and 
second upper molars of that species (Fig. 2, A & B) from Oued el 
Hammam. The specimen resembles P. germaini in 1) the presence of 
strongly developed parastyles and mesostyles and 2) a strong rib on 
the buccal surface of the metacone of M', in the feeble junction of 
the paraloph and metaloph, and in the overlap of the mesostyle 
mesiobuccally to the distal end of the paraloph. Arambourg's 
specimens possess interlophar endostyles or pillars between the 
bases of the protocones and hypocones of the first and second 
molars. The Lothagam-1 specimen lacks any such endostyle. How- 
ever, this structure is not always present in modern Giraffa 
camelopardalis (Fig. 2, F), Okapia johnstoni, or in fossil P. 
primaevus (Fig. 2, C; Churcher 1970, 1977). Therefore it cannot be 
considered a reliable diagnostic character. The specimen resembles 
P. primaevus in the presence of the metacone rib, and in the 
presence of plis on the buccal walls of the protocone and metacone. 
The specimen differs in that the ribs are more developed and the plis 
finer and directed mesiad rather than buccad as in P. germaini (Fig. 
2, A & C). 

The Lothagam tooth expresses the other characters ascribed to P. 
germaini by .Arambourg (1959, 100-101), but many of these char- 
acters (e.g.* strong cingula or deep fossettes) are common to 
Giraffidae and are probably not diagnostic for species or genus. 

The dimensions of the Lothagam P. germaini tooth fall within the 
ranges reported by Arambourg (1959) for Mi's of P. germaini from 



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1979 PALAEOTRAGVS GERMAIN! 5 

Oued el Hammam and marginally on or below the ranges for M-'s 
(Table 1 ). In all instances, the measurements of the Lothagam tooth 
are greater than any reported for P. primaevus from Fort Ternan 
(Churcher. 1970), or for Samotherium sineme (Fig. 2, E) or S. 
neumayri reported by Bohlin ( 1926) and Arambourg (1959) respec- 
tively. {S. sinense is probably a junior synonym of S. neuniavri 
according^ to Erdbrink. 1976). The Lothagam specimen is assigned 
to Palaeotragus germaini on the bases of both size and morphology. 

Palaeotragus germaini was originally described by Arambourg 
(1959) from Oued el Hammam or Bou Hanifa, Algeria. Churcher 
(1978) assigned the large palaeotragines from Beni Mallal, Morocco 
(Lavocat 1961), Bled ed Douarah, Tunisia (Robinson and Black 
1974), and possibly Smendou, Algeria (Joleaud 1937) and Douaria, 
Tunisia (Roman and Solignac 1934) also to this taxon. These North 
African sites range from middle to late Miocene in age, with 
Smendou and Douaria extending into early Pliocene times. 

Arambourg (1959) described P. germaini as a large giraffid with 
elongate neck and legs, and with the forelimb slightly longer than 
the hind. The radius is longer than the tibia and the femur is short, 
the metapodials are subequal, long, slender and laterally compressed 
with subrectangular sections. Arambourg considered that P. ger- 
maini'% upper molars were characterized by strong buccal walls, 
interlophar endostyles, metacones separated from paracones, and 
dep fossettes that are usually open lingually. He considered the teeth 
to be strongly brachydont, but the tooth from Lothagam is not 
more brachydont than teeth of similar size from modern G. 
camelopanlalis nor do his illustrations show teeth thaf are more 
brachydont than that from Lothagam. The ossicones were consid- 
ered to be long, supraorbitally placed, subtriangular in section and 
oriented posterodorsally over the middle region of the skull. The 
single ossicone known from Oued el Hamman is proportionately 
smaller than those of Samotherium and is similar to those of other 
species of Palaeotragus. 

Arambourg described the Oued el Hammal skeletal material as 
larger than that of other Palaeotragus species, similar in robustness 
to that of Samotherium. and intermediate in size between that of 
Samotherium and Giraffa. He considered that P. germaini tx\\\h\\.td 
a parallel evolution separate from Giraffa or Samotherium. and its 
lineage would thus be separate from those of the Giraffinae or 



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1979 PALAEOTRAGi'S GERM AIM 7 

Sivatheriinae and would represent the more progressive and larger 
Palaeotraginae (Churcher 1978, Fig. 9). Arambourg assigned the 
Qued el Hammam giraffid to Palaeotragus on the proportions of 
the limbs, especially the elements of the hind limb. However, the 
characters of the molar teeth also place the taxon within the genus 
Palaeotragus rather than any other genus of the Giraffidae. 

P. gernujini was known previously only from North African sites 
of middle to late Miocene age, or possibly early Pliocene age. The 
Lothagam-1 late Miocene tooth represents the first East African 
record of the species. Some confusion may e.xist between some of 
the reported occurrences oi Samoiherium in Africa and P. germaini, 
especially when dental material is absent. The cervical vertebrae and 
two crushed ossicones from Fort Ternan that were assigned to 
Samotherium africanum by Churcher (1970) may in fact represent 
P. germaini or a form ancestral to it, since these ossicones are not 
excessively large and the single ossicone from Oued el Hammam 
cannot be considered necessarily typical. The identification of 
Samoiherium by Aguirre and Leakey (1974) from the Nakali Tuffs 
of the Ngorora Formation, Kenya, suggests that both genera were 
present in East Africa during the late early to late Miocene. Their 
determination is founded on an isloated first or second upper molar 
(Fig. 2, D) which possesses a strong parastyle and mesostyle, a 
strong paracone rib, and reduced cingulum and metacone rib. which 
are all features of Samoiherium. 

Acknowledgments 

I wish to thank Charles L. Smart for sending the tooth to me and 
suggesting that I publish a description of it; Professor Bryan 
Patterson for allowing me to include mention of the specimen in my 
1978 review of the African Giraffidae and for suggestions about this 
report; and Mary Lynn Richardson for advice and constructive 
criticism. The work was supported by grant A-1704 from the 
National Research Council of Canada. The Museum's expeditions 
to Kenya were supported by National Science Foundation grants 
GP-1188 and GA-425. 

References 

Agi IRRE, E. and p. LtAKE'i. 1974. Nakali, nueva fauna de Hipparion del Rift 
Vallev de Kenya. Estud. Geol. 30: 219 227. 



8 BREVIORA No. 453 

Aramboi RCi. C. 1959. Vertebres continentaux de Miocene superieurde I'Afrique 

du Nord. Mem. Ser\. Carte Geol. Algerie, n.s.. No. 4: 1-161. 
BoHi IN, B. 1926. Die Familie Giraffidae. Palaeont. Sinica, Ser. C, 4 (1): 1-178. 
Cm RCHfR, C. S. 1970. Two new Upper Miocene giraffids from Fort Ternan, 

Kenya, East Africa: Palaeoiragus primaevus n. sp. and Samotherium africanum 

n. sp. Fossil Vert. Africa 2: 1-109. 
. 1978. Giraffidae, Chapter 25. In Evolution of Mammals in Africa, V. J. 

Maglio and H. B. S. Cooke (eds.). Harvard Univ. Press; Cambridge, Mass., 

pp. 509-536. 
Erdbrink, D. p. B. 1977. On the distribution in time and space of three giraffid 

genera with Turolian representatives at Maragheh in N.W. Iran. Proc. Koning. 

Nederl. Akad. Wetensch.. Ser. B. 80 (5): 337-355. 
JoLEALiD, L. 1937. Remarques sur les giraffides fossiles d'Afrique. Mammalia 1: 

85-96. 
Lavocat, R. 1961. Le gisement de vertebres de Beni Mellal (Maroc). Etude 

systematique de la faune de mammiferes et conclusions generales. Notes Mem. 

Serv. Geol. Maroc 155: 29-94, 109-145. 
Patterson, B., A. K. Behrensmeyer and W. D. Sill. 1970. Geology and fauna 

of a new Pliocene locality in northwest Kenya. Nature 226: 918-921. 
Robinson, P. and C. C. Black. 1969. Note preliminaire sur les vertebres fossiles 

du Vindobonien (formation Beglia), du Bled Dourah, Gouvernorat de Gafsa, 

Tunisia. Notes Serv. Geol. Tunisia 31: 67-70. 
Roman, F. and M. Solignac. 1934. Decouverte d'un gisement des mammiferes 

pontiens a Douaria (Tunisia Septentrionale). C. R. Acad. Sci. Paris 199: 1649- 

1650. 



sg' 



L.iL5n/-\r\ I 



MAR 1 8 1985 

REV I»©«.R A 

useum of Comparative Zoology 



us ISSN 0006-9698 



Cambridgf. Mass. Jriv31, 1979 



Number 454 



EVOLUTION OF LIFE HISTORIES: 

A COMPARISON OF ANOLIS LIZARDS 

FROM MATCHED ISLAND AND MAINLAND HABITATS 

Robin M. Andrews' 



Abstract. Faunal comparisons between islands in the West Indies and the 
South and Central American mainland suggest that Anulis lizards play very different 
trophic roles in these two regions. Island anoles, with few predators, appear to be 
food limited relative to the heavily preyed upon mainland species. Theoretical 
considerations of species living under different competitive regimes suggest that 
island and mainland anoles should have evolved major differences in their life 
histories. 

In order to evaluate the evolution of life history in the genus Anoli.s, I consider the 
degree of food limitation for anoles in comparable island and mainland habitats, the 
life history consequences of different selective regimes, and the correlation between 
observed life histories and theoretical expectations. Food availability on island and 
mainland sites was assessed from determinations of the standing crop of arthropods 
relative to the standing crop of anoles. A more direct assessment was accomplished 
by evaluating growth rates of /4«o//.y juveniles. Both comparisons indicate that island 
anoles are food limited relative to mainland counterparts. Island anoles exhibit 
delayed maturity, long adult survivorship, large adult size, high sexual dimorphism 
ratios, and probably low reproductive effort in comparison with mainland anoles of 
similar size. For the most part, observed Ufe histories fit the predictions based on 
current life history theory. However, for a given population, maximum body size, 
sexual dimorphism ratios, and offspring size are relatively invariant in expression. 
On the other hand, growth rate, rate of egg production, and age at sexual maturity 
vary as a function of food availability. 

INTRODUCTION 

In this paper I evaluate the effects of the competitive milieu on 
evolution of life history in the lizard genus Anolis. Various observa- 
tions suggest the island and mainland members of this large group 



'Department of Biology, Virginia Polytechnic Institute and State University, 
Blacksburg, VA 24061. 



2 BREVIORA No. 454 

differ in the way that their populations are regulated. Anoles of the 
South and Central American mainland have numerous competitors 
for their insect prey and also have many predators; these anoles play 
minor roles in the middle levels of their complex food webs (Rand 
and Humphrey, 1968). On the other hand, anoles of the West Indies 
have few competitors or predators. There are fewer species of birds 
on these islands than in comparable mainland habitats and many 
types of insectivorous and lizard-eating birds are totally absent. 
These missing species include motmots, antbirds, woodhewers, 
ovenbirds, and puffbirds (Bond, 1971). Moreover, West Indian bird 
faunas do not exhibit density increases as a result of fewer species; 
density of insectivorous birds is lower in the West Indies than in 
comparable mainland habitats (Recher, 1970; Terborgh, pers. 
comm.). At a given West Indian site there tend to be fewer species of 
frogs (possible competitors for small insect prey) and snakes 
(possible predators on lizards) than at a comparable mainland site. 
The number of lizard species is also often lower than on the 
mainland (Rand, 1961; Schwartz and Thomas, 1975). Army ants 
which compete with anoles on the mainland have never reached the 
West Indies. It is this poverty of potential competitors that allows 
Williams (1972) to conclude that competition among West Indian 
anoles is, for all practical purposes, intrageneric. 

Trophic level differences suggest that island anoles, with few 
predators, may be food limited relative to the heavily preyed upon 
mainland species (Slobodkin et al. 1967; Wiegert and Owen, 1971). 
Recent theory predicts that such differences in mode of population 
regulation might have profound effects on Anolis life histories 
(MacArthur and Wilson, 1967; Gadgil and Bossert, 1970; Gadgil 
and Solbrig, 1972; Pianka, 1970, 1972). 

The effect of the competitive milieu on the allocation and 
utilization of energy result in the predictions listed in Table 1. 
Heightened competition in saturated environments will either 1) 
reduce the overall amount of energy that an organism can gather per 
unit time or 2) force changes in the allocation of time and energy 
devoted to conflicting demands, or both will occur (Pianka, 1972). 
The consequences of this are important. The absolute amount of 
energy that can be used for reproduction is reduced. Thus a 
relatively low reproductive effort is predicted for populations in 
saturated environments. Moreover, selection will usually favor 



1979 A COMPARISON OF ANOLIS LIZARDS 3 

Table I. Predictions of relative differences in life history between island and 
mainland A/iolis lizards. 

ISLAND ANOLES: MAINLAND ANOLES: 

Food Limited Not Food Limited 

1. Reproductive effort Low High 

2. Sexual maturity Late Early 

3. Relative size of offspring Large Small 

4. Size Large Small 

5. Sexual dimorphism in size High Low 



individuals with attributes that make them successful in competitive 
situations. In Anolis. selection for large size can be expected to be 
particularly efficacious because larger individuals are usually domi- 
nant over smaller individuals in competition for space, which 
provides an advantage for obtaining food and mates (Rand, 1967; 
Trivers, 1976). Sexual dimorphism in size is a means by which 
competition for food between male and female Anolis is reduced 
(Schoener, 1967). High sexual dimorphism ratios are expected when 
competition is intense. Since juveniles fare poorly in competition 
with established adults, selection should favor long adult Ufe with 
reproduction occurring in more than one breeding season (Murphy. 
1968). Tinkle et al. (1970) predict that lizards living in saturated 
environments will have relatively large eggs and hatchlings because 
of the relationship between size and competitive ability. In sum, 
island anoles should be K-selected in the sense of MacArthur and 
Wilson (1967). 

On the other hand, if mainland anoles are generally limited by 
their predators, they should be r-selected relative to island anoles 
that are limited by food resources (Pianka, 1970. 1972). Because the 
risk of death is high for all age classes of mainland anoles, fitness is 
maximized by high reproductive effort. Attributes that maximize 
present reproduction will be favored by selection because the chance 
of future reproduction is small. Because age at first reproduction 
and size are closely and positively related (Fenchel, 1974), selection 
for early reproduction (which is the most effective way to increase r) 
will parallel selection for small adult body size. 

In order to evaluate the evolution of life history in the genus 
Anolis, I have considered three main questions: Are island anoles 
food limited relative to mainland anoles? If so, what have been the 



4 BREVIORA No. 454 

life history consequences of the different selective regimes? Do the 
observed life histories conform to theoretical expectations? 

To evaluate the hypothesis that island anoles are food limited 
relative to mainland anoles, I first compare the standing crop 
biomass of Anolis predators and their arthropod prey in com- 
parable island and mainland habitats. A lower ratio of prey to 
predator biomass on islands than on the mainland would provide 
indirect evidence supporting the hypothesis. More direct evidence is 
sought in comparisons which involve the relative allocation of 
energy to feeding and social interactions as opposed to growth and 
reproductive effort. The hypothesis will also be tested with data on 
population turnover. Higher individual persistence in island versus 
mainland populations would provide evidence that predation pres- 
sure is lower in island than in mainland habitats. Next, various life 
history attributes of island and mainland anoles are contrasted to 
determine if island and mainland anoles differ in body size, age at 
sexual maturity, sexual dimorphism ratios, reproductive effort, and 
size of offspring. Finally, I discuss the evolution of life histories in 
Anolis. 

MATERIALS AND METHODS 

Comparisons of island and mainland Anolis are based on data 
that I collected from cacao plantations on Dominica (an island) 
during August 1971, March 1972, and July 1972 and in Costa Rica 
(mainland) during April and May 1971, August 1971, and February 
1972. Cacao plantations were chosen because different localities 
have similar climates; cacao requires rainforest climates with high, 
evenly distributed rainfall. In such environments seasonal differ- 
ences in lizard reproduction are minimal (Licht and Gorman, 1970; 
Sexton et al., 1971) facilitating comparisons of different sites. 
Moreover, the closed canopy of a mature cacao plantation and the 
shade required by the trees themselves make the understory of a 
cacao plantation similar to that of an undisturbed rainforest 
(Leston, 1970). 

Sites and anoles stuilied 

The island site was located on the windward side of Dominica at 
Londonderry Estate 2 km N of the Melville Hall Airport at an 
elevation of about 15 m. Weather stations in this area frequently 



1979 A COMPARISON OF ANOLIS LIZARDS 5 

average over 2500 mm of rain a year and even during the relatively 
dry period, mid-January to mid-March, rainfall is usually more 
than 100 mm a month (Beard. 1949; Hodge, 1954). Rainfall at 
Melville Hall Estate for August 1971 was 268 mm. for March 1972 
was 265 mm. and for July 1972 was 108 mm (Christopher Maximae, 
pers. comm.). 

Dominica has a single species of anole. Anolis oculaius. On the 
basis of geographical variation in size and color pattern four 
subspecies have been described (Lazell. 1972). A. uculatus winsioni 
is found in lowland habitats on the windward side of the island. 
Subsequent references to A. oculatus will apply to this subspecies. 
Anolis oculatus are common in such diverse habitats as beach 
strand and undisturbed forest but appear to be most abundant in 
man-made "forests" such as cacao and orange groves. In cacao 
plantations A. oculatus occupy all structural components of the 
habitat, i.e. individuals are seen on the ground, on trunks, and in the 
canopy. 

The mainland study site was located in the Atlantic lowlands of 
Costa Rica about 45 km W of Limon (Limon Prov.) on Finca La 
Lola, the cacao experimental station of the Institute Interamericano 
de Ciencias Agricolas (IICA, Turrilaba) at an elevation of 33 m. 
Average annual rainfall at La Lola for the years 1949-1967 was 3501 
mm (Soria et al.. 1969). There are two relatively dry periods a year 
(February-March and September), but rainfall rarely falls below 
100 mm a month. Rainfall at La Lola for April 1971 was 266 mm. 
for May 1971 was 111 mm. for August 1971 was 106 mm. and for 
February 1972 was 332 mm (Alfredo Paredes. pers. comm.). 

Although seven species of anoles are recorded from nearby 
forests, only two are commonly found in the cacao plantations 
(Talbot. 1977). Of these. A. limifrons is by far the most abundant, 
being found in all localities where shade is present. In the cacao, 
individuals are usually found less than 2 m above the ground but 
seldom on the ground itself. In contrast, the terrestrial A. humilis is 
restricted to older cacao stands where understor\' vegetation is 
sparse. 

Mark-recapture studies 

Mark-recapture studies, conducted during three visits over about 
a year, were made in mature cacao plantations characterized by a 
continuous canopy and sparse unaerstory. In Dominica, the 625 m- 



6 BREVIORA No. 454 

study site included 43 regularly spaced cacao trees and six large 
breadfruit trees (Fig. 1). In Costa Rica, the 652 m- study site (in 
E.xperimental Section 19 of Finca La Lola) included 45 regularly 
spaced cacao trees and two large Hura crepitans (Fig. 2). Insecti- 
cides or other chemical treatments were not used on the study sites 
during this study or for many years previously. 

During census periods, anoles were captured by hand and 
information on size, sex. individual identification number, and 
location were recorded in the field. Anoles were released shortly 
after capture where they were caught. Snout-vent length (SVL) was 
measured to the nearest mm for each capture and during some 
census periods weight (W) was determined to the nearest one-tenth 
of a gram (Pesola scales). Sex of adults and all but the smallest 
juveniles was readily distinguishable by the presence of the large 
dewlap in males only. Additionally, male A. oculatus of all sizes 
have enlarged post-anal scales which females lack. Individuals were 
identified by clipping the terminal phalange of toes in unique 
combinations involving not more than one toe. 

Population size was estimated from mark-recapture data. ¥ox A. 
oculatus, I estimated population size with the Peterson method using 
a normal approximation to place 95% confidence intervals around 
the mean (Seber, 1973). Census data for these estimates were 
collected on 17 20 March and 30 March-1 April, 1972. For A. 
limifrons. population size was estimated with a regression procedure 
(Marten, 1970) using data of five census periods during April and 
May 1971. 

Adult survivorship on the study sites was determined from the 
proportion of marked adults collected during the two site visits after 
the initial marking period. 

Diet analyses 

Anoles were collected- for stomach-content analysis in cacao 
plantations adjacent to and similar in vegetation structure to the 
mark-recapture sites. On Dominica, this collection was made on 
two days in August 1971, and in Costa Rica, on six days in early 
May 1971 . Collections were made in late afternoon on rainless days 
so that all individuals had an equal time to feed. Individuals were 



-These collections have been deposited in the Museum of Comparative Zoology, 
Harvard University. 



1979 



A COMPARISON OF ANOLIS LIZARDS 



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BREVIORA 



No. 454 




1979 A COMPARISON OF ANOLIS LIZARDS 9 

collected by size in proportion to their occurrence in the popula- 
tions. Specimens were killed within an hour of capture and pre- 
served in alcohol. 

Stomach contents were analyzed using the following techniques. 
The length of each item found in the stomach proper was measured 
and the average width and depth were estimated to the nearest 0.1 
mm with an ocular micrometer. The volume of each arthropod was 
determined by multiplying these three dimensions. Arthropods were 
assigned to one of 15 taxonomic categories (Table 2). 

Anolis body size and reproductive condition 

For each lizard in the above collections SVL and head length 
(HL) were measured. Head length was measured as the distance 



Table 2. Composition of arthropod collections made by sweeping understory 
vegetation on the Dominica and Costa Rica cacao sites. Values are percentages of 
total numbers in each collection and do not include collembola and mites. 





DOMINICA 


COSTA 


RICA 




cacao 


forest 


cacao 


forest 




foliage 


foliage 


foliage 


foliage 


Lepidoptera larvae 


0.2 


0.0 


1.9 


0.3 


Lepidoptera adults 


0.3 


2.7 


5,8 


0.7 


Orthoptera 


1.9 


0.5 


2.3 


4.1 


Blattidae 


0.1 


0.0 


0.0 


1.1 


Hymenoptera: Formicidae 


40.8 


52.5 


10.3 


21.4 


Hymenoptera: Other 


10.7 


3.1 


18.5 


12.1 


Homoptera 


5.5 


5.8 


10.6 


6.4 


Hemiptera 


1.5 


0.0 


1.8 


0.7 


Coleoptera adults 


1.2 


2.3 


13,3 


11.2 


Diptera adults 


32.3 


20,6 


29.3 


21.7 


Holometabolous larvae 


0.1 


0.0 


O.I 


0.1 


(except Lepidoptera) 










Araneida 


4.5 


11.8 


5.5 


6.3 


Isopoda 


0.0 


0.0 


0.1 


2.4 


Isoptera 


0.3 


0.0 


0.0 


9.6 


Miscellaneous 


0.5 


0.7 


0.5 


1,9 


Sweeps 


700 


450 


250 


350 


Total arthropods 


3522 


3180 


1952 


1219 


No. ^ 5 mm 1000 sweeps 


243 


44 


972 


526 



10 BREVIORA No. 454 

from the anterior edge of the ear opening to the tip of the snout. The 
sex of each specimen was confirmed by dissection and the repro- 
ductive condition of females was indexed by the number of oviducal 
eggs and the diameter of the largest ovarian follicle. 

The relation of head length and weight to SVL was determined by 
a least squares analysis. A single regression equation was appropri- 
ate for each of the three species because separate analyses for males 
and females did not alter the equation significantly (p > 0.05). 

Lizards were classified as adult male, subadult male, adult female, 
and juvenile. Females were considered adult (sexually mature) at 
SVLs not less than that of the smallest female with oviducal eggs. 
For species in which males are considerably larger than females, 
subadult males are equal in range of SVL to adult females (after 
Schoener, 1968). Subadult males are small, sexually mature males 
(Licht and Gorman. 1970; Sexton et al., 1971) that appear to be 
excluded from the breeding population by the larger adult males 
(Rand, 1967; Andrews. 1971a) and thus form a separate class. 
Juveniles, defined as immature individuals, were sometimes divided 
into two groups: juvenile I and juvenile II, with smaller and larger 
individuals, respectively. 

The total live weight of anoles on each of the study areas was 
estimated using the following technique. I divided the A. limifrons 
collected in May 1971 and the A. oculaius captured in March 1972 
into SVL categories separated by 1 mm increments. The appropriate 
regression equation was used to convert SVL to weight. Weight was 
then multiplied by the number of individuals in each SVL category 
and the results were summed within classes to give the total weight 
per class. These values were adjusted for the estimated number of 
individuals per class on the study areas. 



Behavioral studies 

Information on foraging behavior was collected through a series 
of standard 30-minute observation periods (e.g. focal-animal sam- 
pling, Altmann, 1975). Foraging behavior was typified by adult 
females because foraging is more clearly defined in females than in 
(adult) males (Andrews, 1971a). On Dominica, observations were 
made between 6 and 18 August 1971 on the capture-recapture area. 
On the Costa Rica site, observations were made between 29 April 



1979 A COMPARISON OF ANOLIS LIZARDS 11 

and 10 May 1971 on both the capture-recapture area and the 
collection area. 

Characteristics of arthropod communities 

Arthropod abundance, size distribution and taxonomic composi- 
tion on the cacao study sites and also in undisturbed forests nearby 
were determined from sweep samples of understory foliage. Sweep 
samples provide representative collections of the kinds of prey eaten 
by anoles that forage in this stratum (Andrews. 1971b). The limita- 
tions and assets of sweep sampling as an ecological technique are 
discussed in previous literature (Southwood. 1966; Janzen and 
Schoener, 1968; Janzen, 1973a). 

On the cacao sites, sweep collections for arthropods were made 
within two days of the collections of anoles and were taken on the 
same areas as the anole collections. The forest site on Dominica 
(Palmist Ridge) was located 5 km SW of the cacao site at an 
elevation of about 275 m. Rainfall is approximately 4000 mm y. 
This ridge top forest is extensive, with a high (max. 44 m) and fairly 
open canopy. The understory is sparse enough to walk through 
easily at most points. This site is further described by Soriano-Ressy 
et al. (1970). Sweep samples were taken in this forest in March 1972. 
The forest site in Costa Rica was located 2 km from Finca La Lola 
near the road between Limon and Siquirres and at a similar 
elevation to the cacao site. The forest area was relatively small 
(perhaps less than 1 km-) and was surrounded by cacao and areas 
recently clear cut. Trees had been selectively removed from the 
forest and therefore in some places the understory vegetation was 
dense enough to prohibit easy access. Sweep samples were taken in 
this forest on 29 February 1972. 

Methods used for sweep sampling are described by Janzen 
(1973a). The total number of sweeps on each site ranged from 250 to 
700 with sets of 50 separated for statistical purposes. Arthropods 
were manually sorted from the leaves and detritus immediately after 
collection and then preserved in alcohol. Sampling was conducted 
only during sunny or intermittently cloudy conditions and at the 
same time of day (1000-1200 hours) on both sites. The same 
methods of taxonomic categorization and measurement of length 
were used for these arthropods as for the arthropods taken from the 
AnoUs stomachs. However, dry weight in each length class was 



12 BREVIORA No. 454 

estimated using regression equations (Sage, 1974, Appendix Table 
2) rather than from a volumetric determination. The following 
relationship for adult insects was used for most length classes: 

Ln Y = -7.761 + 0.3498X — 0.003931X2 
where Y = dry weight in grams and X = length in mm. For 
Lepidoptera larvae and Orthoptera of 15 mm and greater, I used 
equations specific for these taxa. 

RESULTS AND DISCUSSION 

Prey Availability: Size. Abundance, and Taxa 

Size-frequency distributions of arthropods collected in cacao 
plantations show that large individuals were poorly represented in 
the island sample ( Fig. 3). Only 4.8% of all individuals were 5 mm or 
greater in length on the Dominica site in contrast with 12.8% on the 
Costa Rica site (p ^ .001, Kolmogorov-Smirnov two-tailed test). 
This contrast between sites is accentuated when dry weight in each 
arthropod length class is plotted (Fig. 4) and further illustrated by 
summing total dry weights per 1000 sweeps, 6.4 g for Dominica and 
17.8 g for Costa Rica. Moreover, significantly fewer arthropods 
were collected on the Dominica site than on the Costa Rica site 
(Table 2, p = 0.02, Mann-Whitney two-tailed test). The Dominica 
site averaged 247 arthropods per 50 sweeps and the Costa Rica site, 
391 arthropods per 50 sweeps. Thus the low abundance of arthro- 
pods on the Dominica site is related to, in part, the low numbers of 
large individuals. 

The collection of arthropods from forest understories on Domin- 
ica had significantly fewer large arthropods than did the collection 
from the Costa Rica site (Table 2, p^ .001. Kolmogorov-Smirnov 
two-tailed test). The island site also had a lower dry weight biomass 
of arthropods per 1000 sweeps than did the mainland site, 6.5 gas 
opposed to 8.1 g, respectively. However, the Dominica forest 
collection had significantly more individuals per 50 sweeps than did 
the Costa Rica forest collection (p = .02, Mann-Whitney U test, 
two-tailed). This is the result of large numbers of one very small 
myrmicine ant in the Dominica collection. 

The taxonomic composition of the arthropod collections on the 
two sites is contrasted in Table 2. Formicidae dominate the island 
collections (41-53%') with the one myrmicine species contributing 
respectively 27.4 and 46.7% of all arthropods collected from the 



1979 

10,000 



A COMPARISON OF ANOLIS LIZARDS 



13 



5000- 



9- 1000 



on 
O 



O 
a 
o 



E 

D 

z 



500 



100 



50 



10 



5- 



"T 1 1 1 1 1 1 r 



-I 1 r 



I ' 



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I 
I 
I 



I" 



"I r r 



12 



T r- 



n 



16 



20 



24 



28 



Arthropod Length (mm) 



Fig. 3. Size-frequency distributions for arthropods collected in understory foli- 
age in cacao plantations in Dominica ( — ) and La Lola, Costa Rica ( — ). Arthro- 
pods of 28 mm or more in length are not shown (2 in Dominica and 1 in Costa Rica). 



cacao and forest foliage. In contrast, Formicidae contributed only 
10-219f of all arthropods from the mainland site. 

In general, large arthropods are well represented in sweep sample 
collections from mainland rainforests. On the Costa Rica site, 526 
arthropods 5 mm or greater in length were collected per 1000 sweeps 
and this comprised 15.1% of all arthropods. Comparable figures are 



14 



BREVIORA 



No. 454 





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Arthropod Length (mm) 

Fig. 4. Dry-weight — frequency distributions for arthropods collected in under- 
story foliage in cacao plantations. Conventions as in the preceding figure. 



available for two other lowland rainforests in Costa Rica. In wet 
and dry season collections, 1 (Andrews, 1971b) found 510-525 
arthropods 5 mm or greater in length per 1000 sweeps and these 
comprised 22.4-31.6% of all arthropods. In a dry season collection, 
Janzen and Schoener (1968) found 201 arthropods 5 mm or greater 
in length per 1000 sweeps and this comprised 25.2% of all arthro- 
pods. In comparison, very few large arthropods were collected on 
Dominica, the one island locality for which comparable data are 
available. I collected only 44 individuals 5 mm or more in length per 
1000 sweeps and only 0.6% of all arthropods collected were in this 
size category (Table 2). 



1979 A COMPARISON OF ANOLIS LIZARDS 15 

More extensive comparisons of island and mainland arthropod 
communities have been made in second growth habitats. Allen et al. 
(1973) and Janz.en (1973a. b) sweep sampled comparable sites on 
Puerto Rico and nearby smaller islands and on the mainland in 
Costa Rica. Both studies found a significantly lower abundance of 
arthropods in the island than in the mainland collections. 

Population Densities 

Anoles were two to three times more abundant on the island than 
on the mainland study site (Table 3). The total population size of /I. 
oculatus was estimated at 299 363 individuals and of .4. limifrons, 
123 individuals. I assume that gains and losses to the adult 
populations during the census intervals were small because of their 
fidelity to established territories (Rand, 1967; Ruibal and Phili- 
bosian, 1974a; Andrews and Rand, unpub. mainland data) and 
because mortality would be slight during the census periods. The 
Peterson estimate for A. oculatus iwvenWes is probably reasonable 
because the census period spanned only two weeks. However, the 
regression technique used for A. limifrons adults gave unrealistically 
high values for A. limifrons juveniles. This was the result of the large 



Table 3. Population composition, density, and biomass of A. oculatus on the 
Dominica study site and A. limifrons on the Costa Rica study site. Estimates for 
each class are followed by 95% confidence limits. Asterisked values for juveniles 
indicate the total number captured during the census period. Two totals for A. 
oculatus are given, one for each estimate of juvenile numbers. 

A. oculatus A. limifrons 

Site area (m-) 625 652 

Number of Individuals 
Juveniles 

Subadult males 
Adult females 
Adult males 
Total adults 

Total numbers 
Total biomass (g) 



170 (138-241) 


— 


106* 


46* 


44 (22-oc) 


— 


ill (80-198) 


— 


38 (28-65) 


— 


— 


77 (72-82) 


299-363 


123 


1056-1121 


96 



16 BREVIORA No. 454 

number of unmarked individuals that were captured during the 
seven week census period. The census period was long enough to 
allow recruitment from eggs. Moreover, because of a lower site 
fidelity of juveniles than adults (Rand, 1967), appreciable numbers 
of unmarked individuals may have moved into the study area and 
marked individuals moved away from it. A conservative estimate of 
juvenile population si/e is the total number of juveniles captured on 
each site during the census period (Table 3). Only 22 A. hunilll.s were 
encountered on the mark-recapture area; the inclusion of its num- 
bers would increase the number of anoles on the mainland site very 
little. 

For similar habitats, the commonest species on islands character- 
istically have higher population densities than the commonest 
mainland species (Table 4). In non-forested and disturbed habitats, 
some island species become extremely abundant. For example, 
densities of 1400 and 2000 individuals per 1000 m- are recorded for 
the grass anoles, A. poncensis and ,-1. pulchellus. on Puerto Rico. 
Population densities of more than 400 individuals per 1000 m- are 
typical of trunk-ground anoles in disturbed habitats (e.g. A. ocu- 
latus, A. lineatopus, A. acutus, A. sagrei). The island species listed 
with relatively low densities share habitats with species of equal or 
greater abundance. On Bimini, although A. distichus. A. carolinen- 
sis, and A. angusticeps are generally uncommon compared to A. 
sagrei, the range of combined densities on five sites where all four 
species were present was 197 935 individuals per 1000 m'(Schoener 
and Schoener, unpub. data). Anolis lineatopus and A. opalinus had 
a combined density of 312 individuals per 1000 m- on Jamaica. 

In contrast to the island examples, the highest densities recorded 
for anoles in disturbed mainland habitats are all less than 200 
individuals per 1000 m-. On Barro Colorado Island in Panama, A. 
auratus, a mainland counterpart of the West Indian grass anoles, 
had an estimated population density of 137 individuals per 1000 m- 
(Andrews, unpub. data) even when population densities were 
unusually high (A. S. Rand, pers. comm.). 

Anole densities are lower in forests than in more open and 
disturbed habitats on both islands and on the mainland. On Abaco, 
Schoener and Schoener (unpub. data) estimate population densities 
for A. sagrei of 167 235 individuals/ 1000 m' on forest sites and 
439 969 individuals 1000 m- on secondary sites over a two year 
period. Fleming (pers. comm.) found only six A. limifrons per 1000 



1979 A COMPARISON OF ANOLIS LIZARDS 17 

m- in forest plots in the Atlantic lowlands of Costa Rica, in contrast 
to my estimate of 189 individuals 1000 m- in cacao plantations. 
Greater densities of some anoles in disturbed habitats compared 
with forested habitats has also been noted in Mexico by Henderson 
and Fitch (1975). 

Anolis density estimates on islands and on the mainland may be 
even more disparate than indicated in Table 4. Sparse populations 
are usually'not chosen for ecological study because of difficulty in 
obtaining sufficient data to accurately estimate population size. 
Since Anolis densities are generally lower on the mainland, main- 
land site selection is more likely to be biased towards high density 
situations. 

Population densities have not been determined for any South 
American anoles. Work by Rand and Humphrey ( 1968), Fitch et al. 
(1976) and Duellman (in press) indicate that the most conspicuous 
characteristic of Anolis communities in this region is the extremely 
low densities of all species. An on-going study (Miyata. 1977) is 
being conducted, in part, in disturbed habitats where some species 
are encountered fairly frequently. It is obvious that low densities 
have generally discouraged ecological research on anoles in South 
America. 

Diets 

The island cacao site is characterized by low arthropod abun- 
dance, arthropod size distributions skewed towards small sizes, high 
frequencies of ants and a high density of Anolis lizards relative to 
the mainland site. These differences in the prey community should 
be reflected in the diets of the island and mainland cacao anoles. 

Daily food intake of the cacao anoles was assessed from the 
quantities of food material found in their stomachs (Fig. 5). The two 
mainland species consistently had greater volumes of prey in their 
stomachs per gram body weight than did the island species. The 
differences were the greatest for the adult females of A. limifrons 
and A. humilis which had about twice the prey volume as did the 
similarly sized juveniles of A. oculatus. The greatest similarities 
were for adult males of the two mainland species which had mean 
prey volumes comparable to those of similarly sized .4. oculatus. 

Metabolic rate, and thus energy requirements, are a function of 
temperature in lizards (Bennett and Dawson, 1976). Since A. 
oculatus and A. limifrons seem to have similar thermal regimes, 



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A COMPARISON OF ANOLIS LIZARDS 



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BREVIORA 



No. 454 



100 



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10 



Weight (grams) 

Fig. 5. Mean volumes of prey per individual for A. oculatus. A. limifrons. and 
A. humilis expressed as a function of body weight. Head lengths converted to weight 
equivalents with equations in Table 10. 

their energy requirements should not differ on this basis. Ruibal and 
PhiUbosian (1970) measured mid-day body temperatures for A. 
oculatus in a cacao plantation near my study site. Lizards had a 
mean body temperature of 29.8° C for a range in air temperature of 
28.6 29.8° C. A. R. Kiester and I measured mid-day body tempera- 
tures for A. limifrons along a path at La Lola and at the forest site in 



1979 A COMPARISON OF ANOLIS LIZARDS 21 

May 1971. Body temperatures at these sites should bracket those in 
the cacao site because they had somewhat more open and more 
closed canopies respectively. Sixteen adult A. Utnifrons along the 
path had a mean body temperature of 30.1°C (S. E. = 0.44) with a 
range in air temperature of 28. 0-32.0° C. In the forest, six adults had 
a mean body temperature of 29.3° C (S.E. = 0.21 ) with a range in air 
temperature of 28.4 30.3° C. Mid-day body temperatures of both A. 
oculaius a-nd A. linufrons averaged about 30° C. 

In some qualitative aspects the prey size utilization curves for the 
three cacao anoles are similar (Figs. 6 8). The distributions by prey 
numbers are all strongly skewed towards the smaller prey sizes, i.e. 
the items eaten most frequently by all species were small. Such 
distributions seem to be typical for anoles (Schoener, 1968; Sexton 
et al., 1972) and probably reflect log-normal distributions of size in 
insect communities (Schoener and Janzen, 1968). The distributions 
by prey volume are more normal in appearance although still 
skewed towards small prey sizes for some lizard classes. Arthropod 
length is related to volume (or weight) by a power function 
(Schoener, 1969a). Therefore, large items, although eaten relatively 
infrequently, contribute substantially to the volume of prey eaten 
and thus are more important than frequency would indicate. 

Larger individuals of all species tended to take larger prey than 
small individuals. This was particularly clear for A. oculatus where 
differences were significant for all such comparisons (p ^ .01, 
Kolmogorov-Smirnov two-tailed test, Siegal 1956). Females tended 
to take larger prey than equally sized sub-adult males but differ- 
ences were not significant (K-S two-tailed test, p > 0.05). 

For A. limifrons. the classes listed in order of decreasing prey size 
are: adult female, juvenile II, adult males and juvenile I (p^ 0.05 for 
all pairwise comparisons, K-S two-tailed tests). Thus, males took 
relatively small prey for their size. For A. humilis, males and 
females took prey of similar sizes but significantly larger prey than 
did the juvenile classes. 

Although qualitatively similar, the prey size utilization curves of 
the island and mainland anoles differ markedly when absolute prey 
sizes are considered. The sampled population of ,4. oculaius took 
significantly smaller prey than did the A. limifrons population (p ^ 
.001, K-S two-tailed test). This difference is also illustrated from 
comparisons of lizard size classes (Fig. 9). The largest males were 



22 



BREVIORA 



No. 454 



ADULT MALE ( N ^ 28 ) 

Prey vol ( mm5)/,ndividuol : 228 I 
Prey no /individual - 490 



SilBAfHJLT MAL f (N; 10) 
Prey vol ( mm-') /individual 
Prey no /individual = 61 6 



ADULT FEMALE (N^ 32) 

m')/individuol 
Prey no /individual = 60 4 




JUVENILE II (N = 28) 

Prey vol (mm3)/individuol =49 2 
Prey no /individual = 52 6 



JUVENILE I (N= 20) 

Prey vol ( mm')/individiiol - 12 
Prey no /individual - 31 H 



12 16 20 

PREY LENGTH (mm) 



Fig. 6. Prey utilization curves by the fraction of prey numbers in each prey- 
length class ( — ) and the fraction of prey volume in each prey-length class ( — ) for 
A. oculaiiis. Arrows indicate mean prey sizes for the volume distribution. 



1979 



A COMPARISON OF ANOLIS LIZARDS 



23 



ADULT MALE (N:49) 

Prey vol { mm^) / individual =22 9 
Prey no /individual =38 



ADULT FEMALE (N = 65l 

Prey vol ( mm^) /individual = 58. 1 
Prey no /individual =4,0 




JUVENILE I I {N = 6I) 

Prey vol ( mm')/indi viduol = 20.2 
Prey no. /indi vidual = 4.0 



JUVENILE I (N=33) 

Prey vol ( mm^) individual =6.5 
Prey no /individual =5.4 



12 16 20 

PREY LENGTH (mm) 



Fig. 7. Prey utilization curves for A. limifruns. Conventions the same as for the 
preceding figure. 



24 



BREVIORA 



No. 454 



ADULT MALE (N=I8) 

Prey vol ( mm')/ i ndividual = 300 
Prey no /indi vidual - 2 9 



ADULT FEMALE ( N= 19) 

Prey vol (mm') / indi vidual = 66 4 
Prey no /individual = 75 




JUVENILE II (N= 12) 

Prey vol ( mm^) / individual = 16 5 
Prey no /individual =63 



JUVENILE I ( N = 24) 

Prey vol ( mm') /indi viduol = 7.8 
Prey no /individual =5.5 



Fig. 8. 



8 12 16 20 

PREY LENGTH (mm) 

Prey utilization curves lor A. hunnlis. Conventions the same as Figure 6. 



1979 



A COMPARISON OF ANOLIS LIZARDS 



25 



the only A. uculatus that averaged larger prey than any A. limifrons 
and this was a result of three large orthopterans eaten by one 
individual which strongly influenced the mean. Mean prey sizes for 
A. hiiniilis are not plotted but are very similar to those of A. 
limifrons (compare class means in Figs. 7 and 8). However, in the 
comparison of the prey size distributions for the entire collection of 
both species, A. limifrons was found to take significantly larger prey 
than did A. humilis (p^ .001, K-S two-tailed test). Associated with 
the differences in prey sizes were the numbers of prey found per 
anole individual — 32-62 for A. oculatus as opposed to 4-5 for A. 
limifrons and 3-8 for A. humilis. 

The three species also differed in the taxonomic composition of 
their diets. For A. oculatus, the most frequently eaten arthropods 



Limifrons, 




J I L. 



4 5 6 7 8 9 10 II 12 13 14 15 16 17 18 19 20 
HEAD LENGTH (mm) 

Fig. 9. Mean prey length for the volume distribution (see Fig, 6) as a function of 
head length for A. oculatus and A. limifrons. 



26 



BREVIORA 



No. 454 



were Formicidae (ants) which contributed 44 57% of all items 
(Table 5). Formicidae. Isoptera, and miscellaneous (mostly psocids, 
collembola, and mites) arthropods were the most important con- 
tributors to prey volume although decreasingly so for the larger 
anoles with contributions of 77, 61 , 58, and 39% for juveniles, adult 
females, subadult males, and adult males, respectively. For A. 
limifrons, no one taxon was outstanding in frequency but Lepidop- 
tera larvae were the most important contributor to volume for all 
classes (27 44%) while Homoptera, spiders, and Orthoptera were of 
lesser importance (Table 6). For A. hiimilis, frequency of various 
ta.xa was more consistent, with holometaboulous larvae (mostly 
Diptera and Coleoptera), spiders, miscellaneous, and Isopoda being 
consistently highly ranked (Table 7). Again the most important 
contributor to volume was Lepidoptera larvae (25-28%). 

Stomach content analyses show that the diet of the island A. 
oculaius is made up of numerous small insects, primarily ants, and 



Table 5. Dietary composition of A. oculaius on the Dominica site in August 
1971. Values shown are the decimal fraction of total numbers and volume (mm') 
for each class. 









Adult 


Subadult 


Ad 


ult 




Juvenile 


Female 


Male 


Me 


le 




No. 


Vol, 


No, 


Vol. 


No. 


Vol. 


No. 


Vol. 


Lepidoptera L. 


.001 


.011 


.003 


.002 


.002 


.005 


.012 


.088 


Lepidoptera Ad. 


— 


— 


.001 


.006 


— 


— 


.001 


.001 


Orthoptera 


.002 


.020 


.004 


,035 


.006 


.025 


.005 


.300 


Blattidae 


.002 


.023 


.005 


,041 


.002 


.006 


.001 


.002 


Hymenoptera: 


















Formicidae 


,460 


,256 


.502 


,244 


.570 


.224 


.444 


.206 


Hymenoptera: Other 


.036 


,011 


.008 


,001 


.010 


.005 


.009 


.042 


Homoptera 


.006 


,006 


,006 


,010 


.005 


.007 


.005 


.001 


Hemiptera 


,006 


.004 


.004 


.004 


.002 


.001 


.001 


.001 


Coleoptera Ad. 


,084 


,039 


.067 


,058 


,063 


.009 


.191 


.102 


Diptera Ad. 


,052 


.080 


.015 


,144 


,014 


,356 


.008 


.067 


Holometabolous L. 


.012 


,020 


.019 


,006 


,013 


,005 


,005 


.002 


Araneida 


,033 


,019 


.012 


,087 


,014 


,005 


,015 


.005 


Isopoda 


— 


— 


— 


— 


— 


— 


— 


— 


Isoptera 


.077 


.377 


.113 


,210 


.119 


,274 


.180 


.126 


Misc. 


.229 


,135 


.242 


.153 


.181 


.078 


.124 


.056 


Total N 


2110 


1934 


616 


1373 


Total Volume 


1633.8 


3656.0 


1415.2 


6407.6 



1979 



A COMPARISON OF ANOLIS LIZARDS 



27 



Table 6. Dietary composition of A. limifrons on Costa Rica site in May 1971. 
Values shown are the decimal fraction of total numbers and volume (mm') for 
each class. 









Ad 


ult 


Ad 


ult 




Juvenile 


Female 


Ma 


lie 




No. 


Vol. 


No. 


Vol. 


No. 


Vol. 


Lepidoptera L. 


.153 


.437 


.123 


.273 


.179 


.318 


Lepidoptera Ad. 


.014 


.017 


.054 


.097 


.043 


.038 


Orthoptera 


.002 


.004 


.042 


.125 


.027 


.039 


Blattidae 


.019 


.066 


.034 


.120 


.005 


.001 


Hymenoptera: Formicidae 


.074 


.008 


.069 


.008 


.109 


.033 


Hymenoptera: Other 


.108 


.030 


.061 


.011 


.038 


.023 


Homoptera 


.108 


.129 


.088 


.076 


.147 


.090 


Hemiptera 


.026 


.005 


.019 


.004 


.005 


.003 


Coleoptera Ad. 


.053 


.030 


.161 


.078 


.125 


.084 


Diptera Ad. 


.110 


.056 


.050 


.024 


.125 


.049 


Holometabolous L. 


.026 


.030 


.004 


.001 






Araneida 


.165 


.099 


.119 


.056 


.087 


.230 


Isopoda 


.045 


.071 


.142 


.076 


.071 


.064 


Isoptera 


— 


— 


— 


— 


— 


— 


Misc. 


.096 


.017 


.034 


.051 


.038 


.028 


Total N 


418 


258 


184 


Total Volume 


1445.4 


3776.1 


1122.0 



of the mainland A. limifrons of a few large insects, primarily 
Lepidopteran larvae. This dichotomy seems typical of most island 
and mainland anoles so far studied (Rand. 1967; Schoener. 1967, 
1968; Schoener and Gorman, 1968; Andrews, 1971a, b; Sexton et 
al., 1972; Lister, 1976; Scott et al., 1976). If the frequency of prey by 
length is considered, it seems generally true that the majority of the 
prey of island anoles (population or class) are less than 5 mm in 
length and the majority of the prey of mainland anoles are 5 mm in 
length or more. Ants are conspicuous in the diets of island anoles 
only. A "giant" anole, A. cuvieri of Puerto Rico, seems to be an 
exception (Rand and Andrews, 1975), but generalizations discussed 
here may not apply to such a large species. This problem is discussed 
in the Overview Section. 

Although intraspecific comparisons indicate that head and body 
dimensions of anoles are related to prey size (Schoener, 1968; 
Schoener and Gorman, 1968), comparisons between species, and 



28 



BREVIORA 



No. 454 



Table 7. Dietary composition of ^. humilis on the Costa Rica site in May 1971. 
Values shown are the decimal fraction of total numbers and volume (mm') for 
each class. 











Ad 


uh 


Adult 






Juvenile 


Female 


Ma 


le 






No. 


Vol. 


No. 


Vol. 


No. 


Vol. 


Lepidoptera L. 




.091 


.281 


.085 


.276 


.132 


.254 


Lepidoptera Ad. 




.014 


.009 


.028 


.090 


.019 


.070 


Orthoptera 




.039 


.136 


.021 


.030 


.076 


.189 


Blattidae 




.005 


.021 


.007 


.007 


.019 


.036 


Hymenoptera: F( 


Drmicidae 


.082 


.032 


.042 


.008 


.094 


.039 


Hymenoptera: Other 


.039 


.012 


— 


— 


— 


— 


Homoptera 




.029 


.036 


.042 


.087 


.057 


.037 


Hemiptera 




.048. 


.014 


— 


— 


— 


— 


Coleoptera Ad. 




.048 


.028 


.014 


.006 


.132 


.042 


Diptera Ad. 




.043 


.020 


.063 


.135 


.019 


.001 


Holometabolous 


L. 


.173 


.207 


.507 


.196 


.132 


.058 


Araneida 




.149 


.039 


.092 


.026 


.094 


.089 


Isopoda 




.101 


.142 


.070 


.137 


.113 


.127 


Isoptera 




— 


— 


— 


— 


— 


— 


Misc. 




.139 


.028 


.028 


.001 


.113 


.058 


Total N 




208 


142 


53 


1 


Total Volume 




386.2 


1261.6 


540.8 



particularly between species that occupy different habitats, show 
that lizard size alone is a poor indicator of prey size. For example, 
most sizes of A. limifrons eat larger prey than do A. oculatus even 
though A. oculatus is by far the larger species. Moreover, A. 
oculatus has a relatively longer head and greater weight per unit 
SVL than A. limifrons. There is apparently no direct functional 
relationship between the size and body proportions of anoles and 
the size of prey eaten. For A. oculatus. the linear relationship 
between lizard body size and mean prey size may result from 
interference competition where larger individuals monopolize the 
largest prey through social dominance over smaller individuals. 

Anolis activity patterns 

Both diets and population densities suggest that island anoles 
may have to spend more time and energy foraging and interacting 
with other individuals than do mainland anoles. If turnover rates 
and caloric values of stomach contents are comparable, then for 



1979 A COMPARISON OF ANOLIS LIZARDS 29 

similar energy requirements, the utilization of small prey will 
necessitate more active foraging by island than by mainland anoles. 
This prediction is borne out by results of observations on adult 
females on the cacao sites (Table 8). A. oculatus individuals were 
considerably more active than A. litnifrons during foraging as 
judged by the number of shifts in position and the number of 
movements that were directed towards potential prey. A. oculatus 
made about twice the number of major shifts in position (greater 
than one body length) than did A. limifrons. A. oculatus was 
observed to attack prey about five times more frequently. The 
greater activity of A. oculatus was associated with a relatively 
shorter prey attack distance with 70% of the 60 prey capture 
attempts made at 10 cm or less as opposed to only 25% of the 8 such 
attempts for A. liniifrons. Although A. oculatus females fed more 
on the ground than did A. limifrons females (85%- versus 38% of 
attacks), the success rate of both species was similar; about 80% of 
prey capture attempts resulted in capture. 

Where anole densities on islands are greater than on the main- 
land, the activities and associated energetic costs of social interac- 
tions will be greater for island than for mainland anoles. Social 
interactions in reptile populations appear to increase as a function 
of population density (Brattstrom, 1974). Presumably, increased 
density is associated with increased levels of interactions between 
males and females as well as increased levels of aggression between 
individuals of the same sex. Ruibal and Philibosian ( 1974b) found a 
linear increase in aggressive encounters with density for both male 
and female A. acutus. Brad Lister (pers. comm.) recorded the 



Table 8. Indices of foraging activity of adult female Anolis on the island and 
mainland cacao sites. 



Hours of observation 
Position shifts/ h 
Prey capture attempts/h 
% successful attempts 
Mean prey distance (cm) 
% captures on ground 



oculatus 


A. limifrons 


18 


12 


9 


6 


12.1 


5.2 


6.7 


1.3 


82 


75 


19 


30 


85 


38 



30 BREVIORA No. 454 

activities of adult male A. sagrei in areas with densities of 13 and 50 
individuals per 231 m- on Abaco Island in the Bahamas. He found 
that this appro.ximately four-fold increase in male density was 
associated with a 15-20 fold increase per anole in such socially 
relevant activities as chases and displays (headbobs and dewlaps). 
High activity rates of island anoles may only be possible 
because of the relaxation of predation in island environments. With 
low risk of predation, island anoles maximize fitness by moving 
frequently to capture prey and to engage in social interactions with 
neighbors. In contrast, mainland anoles live in environments where 
a moving lizard has a relatively high probability of attracting the 
attention of a predator (Andrews, 1979). Mainland anoles maximize 
fitness by feeding infrequently on large, high yield items. 

Observations by Toft (1976) address this point. Diurnal members 
of frog communities in tropical rainforests exhibit one of two 
foraging patterns. The poisonous dendrobatids and some bufonids 
are active foragers, moving almost incessantly and eating very small 
items such as ants. In contrast, the palatable frogs (mostly lepto- 
dactylids) are sit-and-wait predators on relatively large arthropods. 
Their foraging behavior is much like that of the arboreal sit-and- 
wait anoles. The conspicuous behavior of dendrobatids and bufo- 
nids may be possible only because they are distasteful to most 
predators. 
Survivorship 

Survivorship of adult A. oculatus on the island site was consider- 
ably greater than of adult A. limifrons on the mainland site (Table 
9). Nine months after the original census of the A. limifrons 
population, none of the males that were present on the site in May 
were recaptured and only one (6.6% of females captured) female was 
recaptured. In contrast, a high proportion of the originally marked 
A. oculatus were recaptured in following censuses. After 7 and II 
months respectively, 32.6% and 25.0% of males and 62.9%- and 
53.5% of females were recaptured. 

Although there are few long-term population studies on anoles, 
the evidence at hand suggests that adult survivorship is greater for 
island than for mainland anoles. Annual adult survivorship of A. 
acutus on St. Croix ranged from 27 40% for females and 0-19% for 
males (Ruibal and Philibosian, 1974a). Schoener and Schoener 
(1978) found annual survivorships up to 20 50% for some classes of 



1979 



A COMPARISON OF ANOLIS LIZARDS 



31 



A. sagrei, A. distichus, and A. angusiiceps on Bimini and Andros. 
Several populations oi A. opalinus also had many adult individuals 
that lived for more than one year (T. A. Jenssen, pers. comm.). On 
the other hand, both capture-recapture data and seasonal changes in 
population composition suggest complete annual turnover of adults 
for mainland A. polylepis (Andrews, 1971b). A. limifrons in Costa 
Rica (Table 9; Fitch, 1973b), A. limifrons in Panama (Andrews and 
Rand, in prep.; Sexton et al., 1963). A. nehulosus (Jenssen, 1970), 
A. cupreus (Fitch, 1973a: Fleming and Hooker, 1975). and other 
mainland anoles (Fitch, 1973b). 
Size Relationships 

The island and mainland cacao anoles are markedly different in 
size (Table 10). A. oculatiis is large and exhibits a pronounced 
sexual dimorphism in adult maximum size. Males reached 79 mm 
SVL or about 1 1 g and females 63 mm SVL or about 5 g. In 
contrast. A. limifrons and A. humilis are small with slight sexual 
dimorphism in size. Female A. limifrons reached 41 mm SVL and 
female A. humilis 40 mm SVL at weights of about 1.3 g and males 
were a few mm less in SVL. This difference in size and in degree of 
sexual dimorphism is typical of the differences between island and 
mainland anoles. 

Sexual dimorphism ratios (SDR's) of Greater Antillean and 
mainland species are contrasted in Figure 10. Species size is indexed 
by maximum SVL of males and females (Williams. 1972). In this 
comparison, the number of sympatric species (as many as 6-7) are 
similar in both the island and mainland faunas. This comparison is 



Table 9. Survivorship of adult anoles on cacao sites. For A. 
included all individuals ^ 48 mm SVL. 



.ulatus this 



A. oculaius 



A. limifrons 



No. adults initially marked 

Time elapsed 

Total captured (% marked) 

Time elapsed 

Total captured (% marked) 



44(5 

85 9 
7 months 
46 $ (32.6) 
62 $ (62.9) 
1 1 months 
44(5(25.0) 
71 9 (53.5) 



23(5 

32 9 
3 months 
18 (5 (55.6) 
17 9 (47.1) 
9 months 
18(5 (0.0) 
15 9 (6.6) 



32 



BREVIORA 



No. 454 



^1 

■ 1) 



a. -^ 

o ^ 

-> c ^ 

J. — J « 

■w as c 



'« _ « 

.2 00 g" 

cd u c« 

u - ? 

N > O 



CO X 
Of c/^ 



<TD 



> > 

C/0 C/2 



^ ^ 



-J -J -J 
> > > 

c/5 00 c/5 



Csl ^^l Csl 



X K X 



OO «-! lO 



XXX 



1979 



A COMPARISON OF ANOLIS LIZARDS 



33 



a 
I/) 



o 

z 




Sexual Dimorphism Ratio 

Fig. 10. Se.xual dimorphism (male ma.ximum SVL female ma.ximum SVL) 
ratios for Greater Antillean (solid line) and mainland (South and Central America) 
(dashed line) Anolis. Data for the Greater Antillean species are from Schoener 
(unpublished measurement tables, only species with a sample size of 15 or more 
individuals were used). Data for mainland species are given in Table 12. 



also a conservative test of the prediction that island anoles will have 
higher SDR's than mainland anoles (Table 1) because of lower 
SDR's in the Greater Antilles than on the one- and two-species 
islands of the Lesser Antilles (Schoener, 1977). 

Island species are found to have significantly greater sexual 
dimorphism ratios than do mainland species (p ^ .001, 2X2 Chi- 
squared test, Siegal 1956). Fitch (1976) has also measured SDR's of 
mainland Anolis. Excluding those species for which I present data, 
the range of SDR's (0.80-1.36) and mean SDR (1.07) for the 
tropical anoles in his data set are very similar to mine (Fig. 10) and 
further support the prediction of relatively high sexual dimorphism 
ratios for island anoles. 



34 BREVIORA No. 454 

An even more relevant analysis is suggested by Schoener's 
(unpublished MS) demonstration that ecomorphs. which are species 
occupying similar structural habitats, are also similar in their sexual 
dimorphism ratios. Although 6-7 ecomorphs have been described 
(Rand and Williams, 1969), the present discussion is restricted to 
species whose structural habitat includes shrub level vegetation. 
Thus plotting the SDR's of bush-grass and trunk-ground anoles as a 
function of ma.ximum male SVL shows that island and mainland 
species form exclusive groups ( Fig. 1 1 ). For any given male size, the 
island bush-grass species have higher sexual dimorphism ratios than 
mainland counterparts. Furthermore, the island trunk-ground spe- 
cies have greater SDK's than all but one mainland trunk-ground 
species. Thus, even when similar ecomorphs are compared, the 
island anoles have greater SDK's than mainland species. 

Comparisons of size are difficult since there is a wide range of size 
of different species as indexed by maximum male size both on 
islands and on the mainland. In the Greater Antilles the range in 
species size increases with the number of species per island, and on 
Cuba, the most extreme case, the range is from 38 191 mm 
(Williams, 1972). Although Cuba has by far the largest Anolis. the 
range of species sizes in the Greater Antilles is generally similar to 
that for mainland species where the range is from 38 to about 150 
mm (Williams, pers. comm.). On the one- and two-species islands of 
the Lesser Antilles the range in species sizes is smaller, from 47- 128 
mm; extreme sizes are absent. Thus the range in species size is 
related to species richness; the greater range for Greater Antillean 
than for mainland anoles is possibly due to ecological opportunities 
that are not present on the mainland (Williams, 1972). 

To avoid this problem I compare the relative abundance of 
various sized species within Anolis communities on islands and on 
the mainland. 

Relatively large species are common and frequently dominant 
members of anole communities in the West Indies. On single species 
islands in the Lesser Antilles maximum male SVL's tend to range 
between 65 and 95 mm (Williams, 1972). On islands with two 
species, one tends to be larger and the other smaller than the average 
for solitary anoles (Williams, 1972). However, both members of 
these species pairs are frequently co-dominants in the habitats where 
they occur together (G. Gorman, pers. comm.). There are many 



1979 



A COMPARISON OF ANOLIS LIZARDS 



35 







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36 BREVIORA No. 454 

species in the Greater Antilles with maximum male sizes of 55 mm 
SVL or less (Schoener, unpub. measurement tables). Despite this, 
small bush-grass anoles and large trunk-ground anoles that are 
sympatric may have comparable densities (Schoener and Schoener, 
1971a, b; Jenssen, 1973) or large trunk-ground anoles may have 
greater densities than sympatric bush-grass anoles (Turner and Gist, 
1970; Rand. 1967). 

In contrast to the island situation, Anolis communities on the 
mainland are either dominated by small species or are comprised 
entirely of species with low population densities. According to Fitch 
et al. (1976): "Throughout Mexico and Central America one or 
another of the seven species mentioned [A. hwnilis, A. limifrons, A. 
cupreus, A. sericeus, A. nebulosus, A. subocularis, and A. tropi- 
donotus] tend to dominate habitats suitable for anoles". To his list I 
would add A. polylepis, the dominant in southwestern Costa Rica. 
All of these species are small, with males not exceeding 55 mm in 
maximum SVL. Furthermore, all species occupy shrub-level struc- 
tural habitats. Large congeners are rarely encountered relative to 
the small dominants. Examples of large species with low population 
densities are A. frenatus in Panama (Andrews and Rand, unpub- 
lished data), A. capita in Costa Rica (Andrews, 1971b), A. attenu- 
atus and A. woodi in Costa Rica (Fitch et al., 1976), and A. 
biporcatus and A. pentaprion in Costa Rica and Panama (Andrews, 
unpublished data; Fitch, unpublished data). 

In South America, the species most frequently encountered may 
be relatively large in size. It appears that the structure of Anolis 
communities in South America may be quite different than in 
Central America and Mexico, but because all species are infre- 
quently encountered, much more information is necessary before 
such a generalization is possible. 

Because of the high abundance of large species on islands, 
comparisons of population density (Table 4) provide a minimal 
contrast between Anolis community structure on islands and on the 
mainland. For example, the total live weight oi A. oculatus on the 
island study was estimated at 1094 g (including 173 juveniles) and 
the total live weight of A. limifrons on the mainland site at 96 g 
(Table 3). The 2 3 fold difference in numbers becomes a ten-fold 
difference in biomass. 



1979 A COMPARISON OF ANOLIS LIZARDS 37 

Reproduction (females) 

For lowland Anolis species, reproductive activity is associated 
with rainfall (Licht and Gorman, 1970). Judging from studies on 
sites with rainfall regimes similar to those of this study, reproduc- 
tion probably continues year round for A. oculatus and /I. limifrons. 
Possibly some dimunition occurs in the number of females with 
oviducal eggs during January-March (Licht and Gorman, 1970; 
Fitch, 1973t:). Data on reproductive condition was collected during 
times of the year that both A. oculatus and A. limifrons females 
should have been producing eggs (August 1971 and April-May 
1971, respectively). The majority of adult females on the cacao study 
sites were reproductive as measured by the presence of oviducal eggs 
(Table 11). A greater proportion of female A. limifrons were 
considered reproductive by this criterion than female .4. oculatus (p 
^ .01, 2X2 Chi-squared test, Siegal, 1956). Using reproductive 
condition as an index of the rate of egg production (Licht and 
Gorman, 1970), the data of Table 11 suggest that the rate of egg 
production by A. limifrons females might be greater than for the A. 
oculatus females. 

A general comparison of reproductive effort or proportion of 
energy budgets allocated to reproduction (Hirshfield and Tinkle, 



Table 11. Reproductive condition of female Anolis in study populations 
expressed as the number of individuals with oviducal eggs. 

Repro. class Number of % females 

(no. of eggs) Females with eggs 

A. oculatus 13 

Aug. 1971 1 22 63 

Dominica 2 

A. limifrons 6 

May 1971 1 53 91 

Costa Rica 2 6 

A. hum His 4 

May 1971 1 13 80 

Costa Rica 2 2 



38 BREVIORA No. 454 

1975) for island and mainland anoles is not now possible. Such 
comparisons require information on the rate at which eggs are 
produced during the breeding season as well as their caloric worth. 
Rates of egg production under field conditions are known for only 
two tropical species. A. aeneus on Grenada laid eggs at intervals of 
about 12 days (Stamps, 1975) and A. limifrons (both Panamanian 
and Costa Rican populations) at intervals of 7-8 days (Andrews and 
Rand, 1974). Although reproductive condition through the annual 
cycle has been determined for many anoles (Licht and Gorman, 
1970; Sexton etal, 1971; Andrews, 1971b; Gorman and Licht, 1974; 
Sexton and Brown, 1977), such data cannot be used to index the rate 
at which eggs are laid unless environmental conditions are com- 
parable. One confounding factor is that anoles retain eggs under dry 
conditions (Stamps, 1976). Thus, two oviducal eggs may indicate 
retention rather than high rates of production. A second influence is 
the year to year variation among species in fat accumulation during 
dry periods and its utilization for egg production during the 
breeding season (Licht and Gorman, 1970; Fleming and Hooker, 
1975). Fat storage would facilitate high rates of egg production over 
short periods. Both of the above factors make the interpretation of 
reproductive condition data difficult. Associated energy costs of re- 
production, such as territorial defence or preparing egg laying sites, 
are even more poorly known than the direct costs of eggs per se. 

OVERVIEW 

We are now in a position to evaluate the evolution of life histories 
in island and mainland Anolis. A critical portion of the information 
that has been reviewed concerns present-day conditions in these two 
environments. The expectation that island and mainland anoles will 
differ in life histories is based on supposed differences in the way 
that their populations are regulated. Therefore, I first consider the 
evidence that island anoles are food limited relative to mainland 
anoles. 

Determinations of arthropod and Anolis abundance suggest that 
food availability is less on islands than on the mainland. My 
Dominica site had a ten-fold greater live weight of anoles but about 
one-third the dry weight of arthropods than the Costa Rican site 
had. Although 1 measured standing crop, measurements of site 



1979 A COMPARISON OF ANOLIS LIZARDS 39 

productivity for arthropods would presumably enhance the differ- 
ence between sites. Assuming the rate of population turnover is 
roughly a function of individual size, production of small arthro- 
pods would be roughly comparable on both sites due to the similar 
representation of arthropods ^ 8 mm in length (Figs. 3 and 4). 
Production of large arthropods would be greater on the mainland 
site because such individuals were almost absent in the island 
collection. Limited arthropod data from other islands support my 
findings (Janzen, 1973b). More conclusively, the evidence for high 
anole densities on islands is well documented (Table 4) and would 
be strengthened by use of biomass rather than numbers of individu- 
als because of the relatively large size of common island anoles. 

The sizes and kinds of arthropods in anole diets suggest that 
island anoles eat more non-preferred items than do mainland 
anoles. Like many generalized predators, anoles prefer large items 
to small ones (Andrews, 1971b, unpub. data; Schoener, 1969a); the 
number of large prey eaten should be some function of their 
availability to a feeding lizard. Small items, particularly ants, are 
relatively more abundant in island than mainland habitats and more 
abundant in the diets of island than mainland anoles as well. The 
large numbers of prey in the stomachs of island anoles is a necessary 
consequence of small average prey sizes. Thus, a comparison of 
diets suggest that while mainland anoles have the "option" of 
feeding on large, high yield prey items, island anoles do not. 

The above comparisons constitute indirect evidence that anoles 
on islands are food limited relative to anoles on the mainland. In 
addition, comparisons of processes that are potentially energy 
limited would provide more direct evidence in support of this 
hypothesis. The growth of juveniles is such a process and is, in fact, 
considerably slower for island than mainland species. For com- 
parable lowland habitats, island juveniles grew at one-half to one- 
third the rate of mainland juveniles (Andrews, 1976). The low 
growth rates of island anoles may reflect low food availability or a 
relatively high allocation of energy to foraging and social activities 
or both. 

The data at hand suggest that island anoles are food limited 
relative to mainland anoles. The data on survivorship are in accord. 
Mainland anoles have greater population turnover than island 
anoles, presumably reflecting the greater predation intensity in 



40 BREVIORA No. 454 

mainland than island habitats. The hypothesized dichotomy in 
selective regimes for island and mainland anoles is strongly sup- 
ported. The consequences of such differences are predictable, at 
least in theory. I now review the life histories of island and mainland 
A no I is. 

First, is age at maturity later for island than mainland anoles? 
Island females were estimated to reach sexual maturity in 5 9 
months as opposed to 2-4 months for mainland females (Andrews, 
1976). Comparing species with similar female size, A. Uneatopus 
(Jamaica) becomes sexually mature in 156 days and A. polylepis 
(Costa Rica) in 100 days. However, in Anolis. size at sexual 
maturity is a linear function of maximum size (Andrews and Rand, 
1974). Thus, the later maturity of island than mainland anoles must 
be a function of their lower growth rates. Moreover the low growth 
rates of island anoles are facultative; with augmented food, rates of 
growth may be more than doubled (Andrews, 1976). This means 
that age of maturity in female Anolis on islands will vary with food 
availability. 

Two observations support the prediction that reproductive effort 
is lower for island than for mainland anoles. First, less energy is 
available for the growth of juveniles on islands than on the 
mainland. Energy may likewise limit reproduction for island fe- 
males. On the other hand, island females have the potential to lay 
eggs at rates comparable to those of mainland females (Andrews 
and Rand, 1974). Second, hatchling size is primarily a function of 
female size (Andrews and Rand, 1974). Island females do not have 
larger hatchlings than mainland females which may be related to 
constraints of the arboreal life styles of anoles. However, the slope 
of the linear function relating hatchling weight to female weight is 
considerably less than one. Therefore, proportionally, the caloric 
investment per offspring (ignoring associated costs) will be consid- 
erably less for large than for small species. For comparisons in 
which island species have larger females than mainland species, 
relative investment per offspring would be lower for island females 
even if rates of egg production in the two groups were the same. 

High SDR's are characteristic of West Indian Anolis. The male is 
larger in all cases. However, if sexual dimorphism is solely a 
function of the need to partition food by size (Schoener, 1967, 1968; 
Schoener and Gorman, 1968), it should not matter which sex is the 



1979 A COMPARISON OF ANOLIS LIZARDS 41 

larger. On the mainland, the situation is quite different. For 21 of 
the 59 populations measured by Fitch (1976) and myself, females 
were larger than males. 

One explanation for the larger size of males than females is sexual 
selection. Trivers (1972) points out that when females are clumped 
in space but dispersed in time by asynchronous breeding that one 
male can monopolize breeding. The more females potentially 
monopolized, the more intense will be male-male competition for 
this resource. These assertions are particularly applicable to Anolis 
because: 1) adult males are often associated with more than one 
female (Rand, 1967; Andrews 1971a), 2) females potentially mate 
just prior to the ovulation of each egg (Crews. 1973), and 3) male 
size is positively related to reproductive success (Trivers, 1972). 
Because of the potential for males to monopolize many females, 
sexual selection should be particularly effective in insular situations 
where Anolis densities may be high. 

In addition to sexual selection, factors that promote fitness in 
females may also explain the larger size of males than females in the 
West Indies. Because size at first reproduction is linearly related to 
maximum size (Andrews and Rand, 1974) for the same growth 
rates, the smaller the species size the sooner females can lay eggs. If 
we ignore other factors, early maturity should be advantageous 
because age at first maturity is such an important contributor to rmax 
(Cole, 1954). Furthermore, decrease in female size lowers metabolic 
requirements faster than it lowers offspring size. For example, 
reduction of female weight from 4 g to 2 g reduces standard energy 
requirements by about 40% but offspring weight by only about 20%. 
In highly competitive situations, it may be advantageous for females 
to have low energy demands and yet be able to produce relatively 
large offspring. Thus female size may be a compromise between low 
energy demands and early reproduction and the ability to produce 
competitive offspring. This might explain Schoener's ( 1969b) obser- 
vation that there is less variation in median female size than in 
median male size between islands differing in number of congeners. 

The prediction that mainland anoles will be smaller in size than 
island anoles is supported by community level comparisons; in 
mainland habitats common species are generally small and in island 
habitats common species are both small and large. I have argued 
that small size is one of a suite of r-selected life history attributes 



42 BREVIORA No. 454 

adaptive in mainland Anolis habitats where predator pressure is 
high. However, large species do occur in these habitats. The large 
mainland anoles appear to fall into one of two ecological categories, 
each minimizing the risk of predation in different ways. 

The very low densities of many large species such as A. capita and 
A. frenatus are associated with dietary specialization on large prey 
items. The mean length of 62 prey items eaten by 23 adult A. capita 
was 16 mm when calculated by prey numbers or 30 mm when 
calculated by prey volume (Andrews, 1971b). Adult A. frenatus eat 
similarly sized prey (Scott et al., 1976; Andrews, unpub. data). 
Individuals of 15 mm in length or more constitute less than 0.4% of 
arthropods taken by sweep sampling at lowland rainforest sites in 
Central America (Andrews, 1971b; Janzen and Schoener, 1968, 
Guapiles site). Therefore large anoles that specialize on rare re- 
sources must themselves be rare. Although small arthropods are 
much more abundant than large ones, large anoles are probably 
large prey specialists because the low foraging rates are adaptive in 
their rainforest habitats (Andrews, 1979). The tendency of the large 
trunk-ground species such as A. capita, A. frenatus, and A. woadi 
to "freeze" rather than to flee when disturbed (Fitch, pers. comm.; 
Andrews, 1971b) also suggests that they avoid detection by preda- 
tors by long periods of immobility and highly cryptic coloration. 

The mainland anoles that are both large and relatively abundant 
seem to be adapted to specialized habitats where predator pressure 
may be relatively low. The "aquatic" anoles have maximum male 
sizes ranging from 70-100 mm SVL (Table 12, Fitch 1976). Camp- 
bell (1973) found densities of A. paecilapus and A. lianatus up to 
100 individuals per 1000 m' in Panama. Escape from avian pred- 
ators may be relatively easy in the boulder strewn, stream side 
habitats where these anoles live. The "rock" anoles, with maximum 
male SVLs of about 80 mm, are locally abundant in Mexico (Fitch 
and Henderson, 1976). Here also, predator pressure may be low. 

In contrast, the overriding determinant of Ana/is life histories on 
islands is competition for food. Large size is one of the K-selected 
attributes which maximize fitness in situations where resources are 
limited. The contention by Case (1978) that the increased size of 
territorial lizards on islands is a function of a greater availability of 
food resources is not supported by the Analis data. Both insect 



1979 A COMPARISON OF ANOLIS LIZARDS 43 

Standing crops and the growth rates of juveniles in comparable 
island and mainland habitats indicate that the ratio of resource 
supply to resource demand by anole populations is relatively low on 
islands. Although food availability may provide one constraint to 
the maximization of body size in island habitats, it does not explain 
the relative increase in size of island anoles. 

The dichotomy in the life histories exhibited by island and 
mainland anoles generally conforms to the initial predictions (Table 
1). However, life history theory implies that life history attributes 
are invariant expressions of the genome. It is obvious from exam- 
ination of Anolis life history attributes that while some attributes 
are relatively constant in expression, others are extremely plastic. In 
Anolis, attributes such as maximum size, sexual dimorphism ratios, 
and offspring size have easily definable values for a given popula- 
tion. These indicate selection for fitness under environmental 
conditions that are "constant" across many generations. On the 
other hand, growth rate, age at sexual maturity, and rate of egg 
production are highly variable under experimental conditions and 
presumably in the field as well. There has been a recent recognition 
that such variability may be highly adaptive in heterogeneous or 
fluctuating environments (Hickman, 1975; Hirshfield and Tinkle, 
1975; Nichols et al., 1976). The ability of the food limited island 
anoles to exploit temporally or spatially abundant food by increas- 
ing rates of growth or egg production is not surprising considering 
the importance of early maturity and reproductive effort to individ- 
ual fitness. 

The life history model proposed here must be qualified in one 
important respect. It was developed by focusing on the major 
selective forces acting on island and mainland Anolis that live in 
lowland tropical environments. Although the available data support 
the predicted island-mainland dichotomy, only two anole eco- 
morphs, the bush-grass and the trunk-ground species, have con- 
tributed to these data. The anoles we know the least about, very 
large species, canopy species, and species that live in extreme 
environments, are also the most likely to prove exceptions to the 
model (Andrews, 1976). The trophic position of the very large anole 
species may very well differ from small species since many are 
frugivorous (Brach, 1976; Schoener, unpub. data) and some include 



44 



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46 BREVIORA No. 454 

other anoles in their diets (Rand and Andrews, 1975; O. J. Sexton, 
pers. comm.). Furthermore, the environment may place physiologi- 
cal constraints on anoles in some habitats that override the biotic 
factors considered here. As examples, A. onca, which occupies xeric 
coastal regions of Venezuela, has a very high SDR for a mainland 
anole (Table 12), and the low growth rates of A. tropidolepis in 
Costa Rica may be a function of low environmental temperatures in 
its cloud forest habitat (Fitch, 1972). Whether or not a distinct 
island-mainland dichotomy can be extended to such "atypical" 
species is yet to be resolved. 

ACKNOWLEDGMENTS 

This study was partially funded by NSF Grant Nos. B019801X 
and GB-37731X to E. E. Williams. Research was conducted during 
tenure as a Smithsonian postdoctoral fellow and a visiting research 
associateship at the Smithsonian Tropical Research Institute. I 
would like to thank G. Gorman. L. Houck, A. R. Kiester, E. G. 
Leigh, A. S. Rand, T. W. Schoener, E. E. Williams, and one 
anonymous reviewer for useful comments on the many versions of 
this paper. For assistance in Dominica, I am grateful to Christopher 
Maximae of the Forestry Office for information and logistic 
support and to Frobel Laville for permission to work on London- 
derry Estate. In Costa Rica, IICA also provided logistic support for 
field work. The kindness and assistance of Alfredo Paredes and 
Albertina Johnson at La Lola was most appreciated. 



Literature Cited 

Allan, J. D., L. W. Barnthouse, R. A. Prestbye, and D. R. Strong. 1973. On 
foliage arthropod communities of Puerto Rican second growth vegetation. 
Ecology, 54: 628-632. 

Altmann, J. 1975. Observational study of behavior: sampling methods. Be- 
haviour, 29: 227-267. 

Andrews, R. M. 1971a. Structural habitat and time budget of a tropicaMA;o//5 
lizard. Ecology, 52: 262-270. 

Andrews, R. M. 1971b. Food resource utilization in some tropical lizards. Ph.D. 
Thesis, Univ. of Kansas. Lawrence. 

Andrews, R. M. 1976. Growth rate in island and mainland anoline lizards. 
Copeia, 1976: 477-482. 

Andrews, R. M. 1977. Report, In E. E. Williams (ed.). The Third Anolis 
Newsletter. Museum of Comparative Zoology, Harvard University, Cambridge. 



1979 A COMPARISON OF ANOLIS LIZARDS 47 

Andrews, R. M. 1979, The lizard Coryiophanes cristatus: an extreme "sit-and- 

wait" predator. Biotropica, in press. 
Andrews, R. M. and A. S. Rand. 1974. Reproductive effort in anoline lizards. 

Ecology, 55: 1317-1327. 
Beard, J. S. 1949. The Natural Vegetation of the Windward and Leeward 

Islands. Oxford, Clarendon Press. 142 pp. 
Bennett, A. F. and W. R. Dawson. 1976. Metabolism. In Cans, C. (ed.). 

Biology of the Replilia. Vol. 5 (Physiology A), New York, Academic Press, 

pp. 127-223. 
Bond, James. 1961. Birds of the West Indies. Boston, Houghton Mifflin Co. 

256 pp. 
Brach, V. 1976. Habits and food of Anolis equestns in Florida. Copeia, 1976: 

.187-189. 
Brattstrom, B. H. 1974. The evolution of reptilian social behavior. Amer. 

Zool., 14: 35-49. 
Campbell, H. W. 1973. Ecological observations on Anolis lionotus and Anolis 

poecilopus (Reptilia, Sauria) in Panama. Amer. Mus. Novitates, 2516: 1-29. 
Case, T. J. 1975. Species numbers, density compensation, and colonizing ability 

of lizards on islands in the Gulf of California. Ecology, 56: 3-18. 
Case, T. J. 1978. A general explanation for insular body size trends in terrestrial 

vertebrates. Ecology, 59: 1-18. 
Cole, L. C. 1954. The population consequences of life history phenomena. 

Quart. Rev. Biol., 29: 103-137. 
Crews, David. 1973. Behavioral correlates to gonadal state in the lizard, Anolis 

carolinensis. Hormones and Behavior, 4: 307-313. 
Echelle, a. a., a. F. Echelle, and H. S. Fitch. 1971. A new anole from Costa 

Rica. Herpetologica, 27: 354-362. 
Fenchei , T. 1974. Intrinsic rate of natural increase: the relationship with body 

size. Oecologia, 14: 317-326. 
Fitch, H. S. 1972. Ecology of Anolis tropidolepis in Costa Rican cloud forest. 

Herpetologica, 28: 10-21. 
Fitch, H. S. 1973a. Observations on the population ecology of the Central 

American iguanid lizard, Anolis cupreus. Carib. Jour. Sci., 13: 215-230. 
Fitch, H. S. 1973b. Population structure and survivorship in some Costa Rican 

lizards. Occas. Papers Univ. Kans. Mus. Natur. Hist., 18: 1-41. 
Fitch, H. S. 1973c. A field study of Costa Rican lizards. Univ. Kans. Sci. Bull., 

50: 39-126. 
Fitch, H. S. 1976. Sexual size differences in the mainland anoles. Occas. Papers 

Univ. Kans. Mus. Natur. Hist., 50: 1-21. 
Fitch, H. S., A. A. Echelle, and A. F. Echelle. 1972. Variation in the Central 

American iguanid lizard, Anolis cupreus, with the description of a new sub- 
species. Occas. Papers Univ. Kans. Mus. Natur. Hist., 8: 1-20. 
Fitch, H. S., A. F. Echelle, and A. A. Echelle. 1976. Field observations on 

rare or little known mainland anoles. Univ. Kans. Sci. Bull., 51: 91-128. 
Fitch, H. S. and R. W. Henderson. 1976. A field study of the rock anoles 

(Reptilia, Lacertilia, Iguanidae) of southern Mexico. J. Herpetol., 10: 303-311. 
Fleming, T. H. and R. S. Hooker. 1975. Anolis cupreus: the response of a lizard 

to tropical seasonality. Ecology, 56: 1243-1261. 



48 BREVIORA No. 454 

Gorman. G. C. and R. Harwood. 1977. Notes on population density, vagility, 

and activity patterns of the Puerto Rican grass lizard, Anolis pulchellus (Rep- 

tilia, Lacertilia, Iguanidae). J. Herpetol.. 11: 363-368. 
Gorman, G. C. and P. Light. 1974. Seasonality in ovarian cycles among tropical 

Anolis lizards. Ecology, 55: 360 369. 
Gorman, G. C. and P. Light. 1975. Differences between the reproductive cycles 

of sympatric Anolis lizards on Trinidad. Copeia, 1975: 332-336. 
Henderson, R. W. and H. S. Fitgh. 1975. A comparative study of the structural 

and climatic habitats of Anolis sericeus (Reptilia: Iguanidae) and its syntopic 

congeners at four localities in southern Mexico. Herpetologica, 31: 459-471. 
Hickman, J. C. 1975. Environmental unpredictability and plastic energy alloca- 
tion strategies in the annual Polygonum cascadense (Polygonaceae). J. Ecology, 

63: 689-702. 
HiRSHFiELD, M. F. AND D. W. TiNKi E. 1975. Natural selection and the evolution 

of reproductive effort. Proc. Natl. Acad. Sci., 72: 2227-2231. 
Hodge, W. H. 1954. Flora of Dominica, B.W.E. Lloydia, 17: 1-238. 
HoLDRiDGE, L. R. 1967. Life Zone Ecology. Tropical Science Center, San Jose, 

Costa Rica, rev. ed. 124 pp. 
Jackson, J. F. 1973. Notes on the population biology oi Anolis tropidonotus in a 

Honduran highland pine forest. J. Herpetol., 7: 309-311. 
Janzen, D. H. 1973a. Sweep samples of tropical foliage insects: description of 

study sites, with data on species abundances and size distributions. Ecology, 54: 

659-686. 
Janzen, D. H. 1973b. Sweep samples of tropical foliage insects: effects of sea- 
sons, vegetation types, elevation, time of day, and insularity. Ecology, 54: 

687-702. 
Janzen, D. H. and T. W. Schoener. 1968. Differences in insect abundance and 

diversity between wetter and drier sites during a tropical dry season. Ecology, 

49: 96-110. 
Jenssen, T. A. 1970. The ethoecology of .4 a!o//5 A7£'/'i//o5(/5 (Sauria, Iguanidae). J. 

of Herpetol., 4: 1-38. 
Jenssen, T. A. 1973. Shift in the structural habitat of Anolis opalinus due to 

congeneric competition. Ecology, 54: 863-869. 
Lazeil, J. D., Jr. 1972. The Anoles (Sauria, Iguanidae) of the Lesser Antilles. 

Bull. Mus. Comp. Zool., 143: 1-115. 
Leston, D. 1970. Entomology of a cocoa farm. Ann. Rev. Entomol. 15: 237-294. 
LiCHT, P. AND G. C. Gorman. 1970. Reproductive and fat cycles in Caribbean 

Anolis lizards. Univ. Calif. Publ. m Zool., 95: 1-52. 
Lister, B. C. 1976. The nature of niche expansion in West Indian Anolis lizards 

I: Ecological consequences of reduced competition. Evol., 30: 659-676. 
MacArthur, R. H. and E. O. Wilson. 1967. The Theory of Island Biogeogra- 

phy. Princeton Univ. Press. 203 pp. 
MacArthur, R. H., J. M. Diamond, and J. R. Karr. 1972. Density compen- 
sation in island faunas. Ecology, 53: 330-342. 
Marten, G. G. 1970. A regression method for mark-recapture estimation of 

population size with unequal catchability. Ecology, 51: 291-295. 



1979 A COMPARISON OF ANOLIS LIZARDS 49 

Meyer, J. R. 1968. Distribution and variation of the Mexican lizard, Anolis 

barkeri Schmidt (Iguanidae), with a redescription of the species. Copeia, 1968: 

89-95. 
MiYATA, K. 1977. Report, In E. E. Williams (ed.). The Third Anolis Newsletter, 

Museum of Comparative Zoology, Harvard University, Cambridge. 
Murphy, G. 1. 1968. Pattern in life history and the environment. Amer. Natur., 

102: 391-403. 
Myers, C. W. 1971. Central American lizards related to Anolis pentaprion: Two 

new species from the Cordillera de Talamanca. Amer. Mus. Nov., No. 2471: 

1-40. 
Nichols, J. D., W. Conley, B. Batt, and A. R. Tipton. 1976. Temporally 

dynamic reproductive strategies and the concept of r- and K-selection. Amer. 

Natur., 110: 995-1005. 
Pianka, E. R. 1970. On r and K selection. Amer. Natur., 104: 592-597. 
PiANKA, E. R. 1972. r and K selection or b and d selection? Amer. Natur., 106: 

581-588. 
Rand, A. S. 1961. Ecology, behavior, and morphology of anoline lizards in 

Puerto Rico. Ph.D. Thesis, Harvard Univ., Cambridge, Mass. 
Rand, A. S. 1967. Ecology and social organization in the iguanid lizard Anolis 

lineatopus. Proc. U.S. Natl. Mus., 122: 1-79. 
Rand, A. S. and R. M. Andrews. 1975. Adult color dimorphism and juvenile 

pattern in Anolis cuvieri. J. Herpetol., 9: 257-260. 
Rand, A. S. and S. S. Humphrey. 1968. Interspecific competition in the tropical 

rain forest: ecological distribution among lizards at Belem, Para. Proc. U.S. 

Natl. Mus., 125: 1-17. 
Rand, A. S. and E. E. Williams. 1969. The anoles of La Palma: aspects of their 

ecological relationships. Breviora, 327: 1-19. 
Recher, H. F. 1970. Population density and seasonal changes of the avifauna in 

a tropical forest before and after gamma irradiation. In Odum, H. T. (ed.), A 

Tropical Rain Forest, Springfield, VA, US AFC: E69-E86. 
RuiBAL, R. AND R. Philibosian. 1970. Eurythermy and niche expansion in 

lizards. Copeia, 1970: 645-653. 
RuiBAL, R. AND R. Philibosian. 1974a. The population ecology of the lizard 

Anolis acutus. Ecology, 55: 525-537. 
RuiBAL, R. and R. Philibosian. 1974b. Aggression in the lizard Anolis acuius. 

Copeia, 1974: 349-357. 
Sage, R. D. 1974. The structure of lizard faunas: comparative biologies of 

lizards in two Argentina deserts. Ph.D. Thesis, Univ. of Texas, Austin. 
Schoener, T. W. 1967. The ecological significance of sexual dimorphism in size 

in the lizard Anolis conspersus. Science, 155: 474-477. 
Schoener, T. W. 1968. The Anolis lizards of Bimini: resource partitioning in a 

complex fauna. Ecology, 49: 704-726. 
Schoener, T. W. 1969a. Models of optimal size for solitary predators. Amer. 

Natur., 103: 277-313. 
Schoener, T. W. 1969b. Size patterns in West Indian Anolis lizards. I. Size and 

species diversity. Syst. Zool., 18: 386-401. 



50 BREVIORA No. 454 

ScHOENER, T. W. 1977. Competition and the niche. In Gans, C. and D. W. Tinkle 

(Eds.), Biology of the Reptilia, New York, Academic Press: pp. 35-136. 
ScHOENER, T. W. AND G. C. GoRMAN. 1968. Somc niche differences in three 

Lesser Antillean lizards of the genus Anolis. Ecology, 49: 819-830. 
SCHOENER, T. W. AND D. H. Janzen. 1968. Notes on environmental determinants 

of tropical versus temperate insect size patterns. Amer. Natur., 102: 207-224. 
SCHOENER, T. W. and A. ScHOENER. 1971a. Structural habitats of West Indian 

Anolis lizards. II. Puerto Rico Uplands. Breviora, 375: 1-39. 
Schoener, T. W. and a. Schoener. 1971b. Structural habitats of West Indian 

Anolis lizards. I. Lowland Jamaica. Breviora, 368: 1-53. 
Schoener, T. W. and A. Schoener. 1978. Inverse correlation of survival in 

lizards with island size and avifaunal richness. Nature, 274: 685-687. 
Schwartz, A. and R. Thomas. 1975. Checklist of West Indian amphibians and 

reptiles. Carnegie Museum of Natural History, Special Publication No. 1, 

216 pp. 
Scott, N. J., D. E. Wilson, C. Jones, AND R. M. Andrews. 1976. The choice of 

perch dimensions by Anolis lizards. J. Herpetol. 10: 75-84. 
Seber, G. a. F. 1973. The estimation of animal abundance and related param- 
eters. London, Griffin. 506 pp. 
Sexton, O. J. 1967. Population changes in a tropical lizard Anolis limifrons on 

Barro Colorado Island, Panama Canal Zone. Copeia, 1967: 219-222. 
Sexton, O. J., J. Bauman, and E. Ortleb. 1972. Seasonal food habits of Anolis 

limifrons. Ecology, 53: 182-186. 
Sexton, O. J. and K. M. Brown. 1977. The reproductive cycle of an iguanid 

lizard Anolis sagrei, from Belize. J. Natur. Hist., II: 241-250. 
Sexton, O. J., E. P. Ortleb, L. M. Hathaway, R. E. Ballinger, and P. Licht. 

1971. Reproductive cycles of three species of anoline lizards from the Isthmus 

of Panama. Ecology, 52: 201-215. 
SlEGAL, S. 1956. Nonparametric statistics. McGraw-Hill. 312 pp. 
Slobodkin, L. G., F. E. Smith, and N. G. Hairston. 1967. Regulation in ter- 
restrial ecosystems, and the implied balance of nature. Amer. Natur., 101: 

109-124. 
SoRiA, J., O. EsQUivEL, L. A. Paredes, and R. DlalZ-Romeu. 1969. Finca La 

Lola. Cacao, 14: 1 42. 
Soriano-Ressy, M., a. P. Desmarais, and J. W. Perez. 1970. A comparison of 

environments of rain forests in Dominica, British West Indies, and Puerto Rico. 

In Odum, H, T. (Ed.), A Tropical Rain Forest, Springfield, VA, US AEC: 

B329-B346. 
SouTHWOOD, T. R. E. 1966. Ecological Methods With Particular Reference to 

the Study of Insect Populations. London, Methuen. 391 pp. 
Stamps, J. A. 1975. Courtship patterns, estrus periods and reproductive condi- 
tion in a lizard, Anolis aeneus. Physiology and Behavior, 14: 531-535. 
Stamps, J. A. 1976. Egg retention, rainfall and egg laying in a tropical lizard, 

Anolis aeneus. Copeia, 1976: 759-764. 
Talbot, J. J. 1977. Habitat selection in two tropical anoline lizards. Herpeto- 

logica. 33: 114-123. 



1979 A COMPARISON OF ANOLIS LIZARDS 51 

Tinkle, D. H.. H. M. Wilbur, andS. G. Tilley. 1970. Evolutionary strategies in 

lizard reproduction. Evol., 24: 55-74. 
Toft, C. A. 1976. Partitioning of food in a community of tropical frogs. Ph.D. 

Thesis, Princeton Univ., New Jersey. 
Trivers, R. L. 1972. Parental investment and sexual selection. In Campbell, B. 

(Ed.), Sexual Selection and the Descent of Man, Chicago, Aldine Publishing 

Co.: pp. 136-179. 
Trivers, R. L. 1976. Sexuai selection and resource-accruing abilities in Anolis 

garmani. Evol., 30: 253-269. 
Turner, F. B. -wn C. S. Gist. 1970. Observations of lizards and tree frogs in an 

irradiated Puerto Rican forest. In Odum, H. T. (ed.), A Tropical Rain Forest, 

Springfield, VA, US AEC: E25-49. 
Vanzolini, p. E. and E. E. Williams. 1970. South American anoles: The 

geographic differentiation and evolution of the Anolis chrysolepis species group 

(Sauria, Iguanidae). Arquivos de Zoologia, S. Paulo, 19: 125-298. 
Wiegert, R. G. and D. F. Owen. 1971. Trophic structure, available resources, 

and population density in terrestrial vs. aquatic ecosystems. J. Theoret. Biol., 

30: 69-81. 
Williams, E. E. 1972. The origin of faunas. Evolution of lizard congeners in a 

complex island fauna: A trial analysis. Evol. Biol., 6: 47-90. 



MUS. CO MP. ZOOL 
LIBRARY 



\n 



<^ "■.•"'■' 



B R E V I ,Q.B A 



useiioi of Comparative Zoology 



us ISSN 0006 9698 



Cambridge, Mass. July 31, 1979 Number 455 

A NEW PALEOCENE PALAEANODONT AND THE 
ORIGIN OF THE METACHEIROMYIDAE (MAMMALIA) 

Kenneth D. Rose' 

Abstract. A new genus and species of palaeanodont, Propalaeanodon schaffi. 
from the late Paleocene (late Tiffanian) Polecat Bench Formation of the Bighorn 
Basin, Wyoming, is the oldest and most primitive known metacheiromyid. Pro- 
palaeanodon foreshadows Palaeanodon in the reduction of its postcanine teeth. It 
resembles epoicotheriids in the presence of postcanines along the entire length of the 
horizontal ramus, and resembles the slightly older Amelotabes. in particular, in the 
retention of seven postcanines. Propalaeanodon thus provides additional evidence 
for uniting the Metacheiromyidae and the Epoicotheriidae in the suborder Palaean- 
odonta, and it strongly suggests that the Metacheiromyidae evolved from the 
Epoicotheriidae, probably during the Tiffanian. 

Two humeri from the same stratigraphic level as the holotype are tentatively 
referred to Propalaeanodon. They possess specializations typical of Eocene palaean- 
odonts, including a prominent deltopectoral crest and a large supinator crest. These 
features indicate that Propalaeanodon was, already in the late Tiffanian, a highly 
specialized fossorial animal. 

The systematic position of the Palaeanodonta is uncertain. They have been 
considered to be related to the orders Xenarthra or Pholidota or both by all recent 
students and, indeed, they are the only known fossil group that evolved specializa- 
tions similar to those in these two orders. Although it is possible that palaeanodonts 
were an early radiation of probably myrmecophagous, fossorial mammals merely 
convergent to these orders, the evidence now available enhances the probability that 
the three groups share a special relationship. 

INTRODUCTION 

The edentate-like members of the rare mammalian suborder 
Palaeanodonta have long been considered early relatives of the 
Xenarthra, the Pholidota, or both. As now known, they are 
restricted to the earlier Tertiary of western North America. Palaean- 
odonts are classified in two families, the Epoicotheriidae and the 



'Museum of Paleontology, University of Michigan, Ann Arbor, Michigan 48109. 



2 BREVIORA No. 455 

Metacheiromyidae. To date, nine genera and fifteen species have 
been recognized, several of which were described quite recently 
(West, 1973; Rose et al., 1977; Rose, 1978). They range in age from 
Tiffanian (late Paleocene) to Orellan (middle Oligocene). 

Recently the dentary of a primitive new palaeanodont was 
discovered by Charles Schaff, Museum of Comparative Zoology, in 
late Tiffanian rocks in the northern Bighorn Basin, Wyoming. The 
oldest and most primitive metacheiromyid, it sheds new light on the 
origin of this peculiar family and further documents the unity of the 
Palaeanodonta. Two palaeanodont humeri, collected much earlier 
from the same stratigraphic interval, probably represent the same 
new species. 

Abbreviations of institutional names cited herein are as follows: 

AC Pratt Museum, Amherst College, Amherst, Massachusetts. 

AMNH American Museum of Natural History, New York. 

MCZ Museum of Comparative Zoology, Harvard University, 
Cambridge, Massachusetts. 

PU Princeton University Museum, Princeton, New Jersey. 

UM University of Michigan Museum of Paleontology, Ann 
Arbor, Michigan. 



SYSTEMATIC PALEONTOLOGY 

Suborder PALAEANODONTA Matthew, 1918 

Family METACHEIROMYIDAE Wortman, 1903 

PROPALAEANODON, gen. nov. 

Type species. Propalaeauodon schaffi, sp. nov. 

Included species. Type only. 

Distribution. Late Paleocene (late Tiffanian) oi the Bighorn 
Basin, Wyoming. 

Diagnosis. Only metacheiromyid with seven postcanine teeth that 
extend to the back of the horizontal ramus. All postcanines single- 
rooted except P4, which is two-rooted. Postcanines small, peglike, 
separated by short diastemata, and with rounded crowns, in these 
features resembling Palaeanodon; but crowns higher and more 
pointed than in Palaeanodon. Dentary slender, slightly smaller than 
in Palaeanodon ignavus; deepest beneath first postcanine (Pi), 
shallowest beneath last postcanine (M3), as in Palaeanodon. Medial 
buttress prominent but less so than in Palaeanodon. 



1979 



PALEOCENE PALAEANODONT 



Etymology. In allusion to its resemblance to, lower stratigraphic 
level than, and probable relationship to Palaeanodon. 

PROPALAEANODON SCHAFFI, sp. nov. 
Fig. lA and B, 2A, and 3B 

Holotype: MCZ 20122, left dentary with ?:_,, alveoli for Pi, P4, 
Mi_3, and posterior margin of canine alveolus. 

Hypodigm. Holotype and tentatively PU nos. 13928, 13929. 

Horizon and Locality. Latest Paleocene (late Tiffanian), "Silver 
Coulee beds" of Polecat Bench Formation (of Jepsen, 1940), Jepsen 
Valley Quarry, NW1/4, Sec. 27, T.57N., R.IOOW., Park County, 
Wyoming. 





Figure J. Lateral (A) and dorsal (B) views of holotype of Propalaeanodon 
schaffi. MCZ 20122, left dentary with second and third postcanines (P:_3). Dorsal 
view of left dentary of Palaeanodon sp. (C), showing extended edentulous part of 
ramus behind fifth postcanine; reconstructed from UM 63611 and 66243. 



BREVIORA 



No. 455 




Figure 2. Medial views of left dentaries of palaeanodonts. (A) Prupalaeanodon 
schaffi, holotype, MCZ 20122. (B) Palaeanodon sp., reconstruction based on UM 
63611 and 66243. (C) Ameloiabes simpsoni, holotype, PU 14855 (right dentary, 
reversed). 



Diagnosis. Only known species of the genus; measurements in 
Table 1. 

Etymology. For Charles R. Schaff, expert collector and prepara- 
tor, who discovered the holotype. 

Description. Only two teeth are preserved in the holotype, 
although the dentary is nearly complete and provides much in- 
formation about the mandibular morphology of Propalaeanodon 
schaffi. The horizontal ramus is intact as far forward as the 
posterior margin of the canine alveolus. This alveolar margin 
reveals that the canine was large, as in other palaeanodonts, and its 



1979 



PALEOCENE PALAEANODONT 



root robust, a feature also indicated by thickening of the dentary 
anteriorly. 

The dentary contained seven postcanine teeth, presumably four 
premolars and three molars, which were greatly modified from the 
primitive eutherian condition. Seven is a greater number of teeth 
than in any other metacheiromyid, indeed more than in any other 
palaeanodont except Amelotabes, which also has seven (Rose, 
1978). Alocodontulum, in which the lower dentition is unknown, 
has seven upper postcanines (Rose et al., 1977, 1978), and thus 
probably had seven lowers as well. The postcanines of Propalaean- 
odon occur along the entire length of the horizontal ramus, as in 




Figure 3. Left humeri of palaeanodonts. Scale is 10 mm. (A) Early Eocene 
metacheiromyid Palaeanodon ignavus, based on AMNH 15088 and 16832; after 
Matthew (1918), but with corrected orientation of proximal end. (B) IPropalaeano- 
don schaffi, PU 13928. (C) Early Eocene epoicotheriid Pentapassalus woodi. part of 
holotype, AC 2766. Note that this species is referable to Pentapassalus (Rose, 1978), 
not to Palaeanodon (Guthrie, 1967; Emry, 1970). The supinator crests in (A) and 
(C) are incomplete and have not been reconstructed because of the variable develop- 
ment of this crest in different taxa of palaeanodonts; they were in any case larger 
than depicted. 



BREVIORA 



No. 455 



TABLE 1 

Dimensions of the holotype of Prupalaeanodon schaffi, MCZ 20122. 
(Measurements are in mm to nearest .05 mm) 









Length 


Breadth 


Pi alveolus 






1.35 


0.85 


P: alveolus 






damaged 




P: crown 






1.25 


1.00 


Pi alveolus 






2.40 


1.15 


Pi crown 






1.80 


1.10 


P4 alveolus 






2.80 


1.10 


Ml alveolus 






2.00 


1.15 


M2 alveolus 






2.00 


1.15 


Ml alveolus 






1.60 


1.00 


Pi-M, 






17.10 (approx.) 




Depth of mandible, lingually 


at 


P,: 


6.10 (approx.) 




Depth of mandible, lingually 


at 


M, 


: 4.80 





epoicotheriids but not in later metacheiromyids, in which the back 
of the horizontal ramus is edentulous. All teeth in Propalaeanodon 
are single-rooted except P4, which has two roots (probably a 
primitive feature). Judging from the alveoli. Mi and M2, and 
possibly P3, had single roots that were faintly bilobate. The roots of 
all postcanines were tapered and closed, as in Palaeanodon. The 
epoicotheriids Amelotabes (Fig. 2C) and Tubulodon are similar in 
having a two-rooted P4, but in these taxa several other postcanines 
are also two-rooted. In Pentapassalus Mi has a bilobate root but P4 
is single-rooted (Gazin, 1952). The postcanines in Palaeanodon are 
all single-rooted (Fig, IC, 2B). 

The crowns of P: and Pi are preserved. P: has become dislodged 
from its alveolus as a result of damage to the front of the jaw and 
appears higher than P3. Both teeth are peglike with rounded crowns 
apparently devoid of enamel. In these characters they closely 
resemble the postcanines of Palaeanodon (Fig. IC, 23 and Rose, 
1978), although they are slightly larger, particularly longer, higher 
crowned, and less rounded. As in Palaeanodon they show no cusp 
pattern on the crowns, nor is there any evidence of direct occlusion 
with the upper teeth. Occlusal wear is discernible in molars and even 
in premolars of some epoicotheriids. The alveolus of P4 indicates 



1979 PALEOCENE PALAEANODONT 7 

that it was the largest cheek tooth, a feature in common with 
Anielotahes but in contrast to Palaeanodon. There is a slight size 
reduction from P4 to My. 

All postcanines are separated by short diastemata. At the alveolar 
margin, the alveoli of P2 4 slightly exceed the size of the roots they 
contain; this probably obtained for the molars as well. These traits 
are characteristic also of Palaeanodon, some advanced epoicoth- 
eriids, and many toothed xenarthrans. 

The ascending ramus and condyle of Propalaeanodon resemble 
those of Pentapassalus pearcei and Metacheiromys tatusia (Gazin, 
1952; Simpson, 1931). The coronoid process is slightly smaller than 
in contemporary proteutherians {sensu Butler, 1972) and rises at an 
angle of about 110° to the ventral border of the mandible. The 
condyle, which is narrower transversely than in most toothed 
xenarthrans and contemporary proteutherians, is situated relatively 
low but above the level of the tooth row. Its convex articular surface 
is directed dorsally and backward, as in P. pearcei. The mandibular 
foramen is also relatively low, being situated on the posterior pro- 
jection of the tooth row. The internal mandibular groove extends 
from just anterior to this foramen to a point beneath Mi. 

As in all metacheiromyids, there is a prominent medial buttress, 
forming anteriorly a low, narrow ridge lingual to the molars. The 
dorsal surface of the buttress is roughened, particularly behind M3. 
On the external surface of the mandible the masseteric fossa, though 
rather poorly defined, is deeper than in most other palaeanodonts. 
The fossa is bounded in front by a crest formed by the thickened 
anterior border of the coronoid. A more rounded ridge running 
forward from the condyle forms its lower limit. The horizontal 
ramus is rather slender and shallow, deeper in front than in back, as 
in Palaeanodon. A small mental foramen is present beneath P3. 

Two palaeanodont humeri from Princeton Quarry are probably 
referable to Propalaeanodon schaffi (Princeton Quarry is about Vi 
mile northwest of Jepsen Valley Quarry and at about the same 
stratigraphic level). They are the only other palaeanodont specimens 
from the same stratigraphic interval as the holotype. PU 13928, a 
complete left humerus (Fig. 3B), and PU 13929, a distal end of a left 
humerus, are of appropriate size for Propalaeanodon. They are 
smaller than the humeri of Palaeanodon ignavus (Fig. 3 A) and 
Pentapassalus woodi (Fig. 3C and Rose, 1978) and slightly larger 



BREVIORA No. 455 



TABLE 2 



Dimensions (mm.) of humeri in certain palaeanodonts. 
(Known humeri of Palaeanodon are too incomplete to allow accurate measurements.) 



S E 







£ < 

1 S 
3 ^ 


Pentapassa 
pearcei (fro 
Gazin, 195: 


1 ^ 


length 

length of deltopectoral crest 
(from proximal end of 
greater tuberosity) 


39.3 
26.3 


74.0 

54.7 


37.0 
28.0 


47.1 
35.5 



length of supinator crest 19.7 38.8 22.0 20.8a 

(from distal end of humerus) 

greatest transverse breadth 19.8 34.0 21.7 22.6 

of distal end of humerus 

a = approximate 

than the humeri of Pentapassalus pearcei (Gazin, 1952). However, 
all these specimens are essentially similar in structure (see Table 2 
for comparative dimensions). 

The deltopectoral crest is broad and well developed in PU 13928. 
It may be relatively slightly longer than in Palaeanodon, but this is 
not certain because the known humeri of the latter are fragmentary 
and poorly preserved (AMNH nos. 15088, 16831, 16832). The crest 
is relatively shorter than in Pentapassalus and Metacheiromys (see 
Simpson, 1931: 340-342). Its distal end is developed into an 
anteromedially directed prominence for the insertion of the pecto- 
ralis major. Proximolaterally the crest gives rise to a low process for 
the insertion of the deltoid. A less well-defined crest extends distally 
from the lesser tuberosity almost to the entepicondylar foramen and 
bears a small teres tubercle midway along the shaft, on which the 
teres major inserted, as in Metacheiromys dasypus (Simpson, 1931). 

The supinator crest is somewhat smaller than in Pentapassalus 
pearcei and Metacheiromys dasypus and lacks the hooklike, dor- 
sally projecting flange characteristic of the latter two species. 



1979 PALEOCENE PALAEANODONT 9 

Nonetheless, this crest is very large and served as the site of origin 
for the powerful supinator muscle and carpal extensors, and 
probably the brachioradialis. The median epicondyle (the point of 
origin of the pronator teres and the carpal flexors) is also large, 
although smaller than in Palaeanodon and Pentapassalus. These 
features of the humerus indicate an animal highly adapted for 
fossorial habits (Reed, 1954; Hildebrand, 1974). Although the 
development of the deltopectoral crest in palaeanodonts is (as 
Emry, 1970, asserts) more like that of manids than of armadillos, 
the development of the supinator crest is more extreme, even in PU 
13928, than in either manids or xenarthrans. 

DISCUSSION 

Propalaeanodon is the second known Paleocene palaeanodont 
and the only known Paleocene metacheiromyid. A number of 
resemblances to epoicotheriids were described above — e.g., seven 
postcanines, teeth present at the back of the horizontal ramus, two- 
rooted P4 — but these are primitive features of palaeanodonts. 
Propalaeanodon resembles Palaeanodon (Fig. 1 and 2) in the 
manner of reduction of its postcanines to peglike teeth (all but one 
single-rooted) with rounded crowns that apparently lack enamel. 
Like Palaeanodon, its postcanine teeth are housed in tapered alveoU 
that are larger than the roots and separated by diastemata. Al- 
though some of these features also evolved in advanced epoicothe- 
riids (e.g., Bridgerian Tetrapassalus mckennai, Chadronian-?Orel- 
lan Xenocranium pileohvale: Simpson, 1959; Colbert, 1942), the 
total suite of features evolved much earlier in the metacheiromyids. 
Thus when metacheiromyids had already achieved these traits, 
contemporary epoicotheriids, as now understood, retained rela- 
tively unreduced teeth with enamel and with very short or no 
diastemata between them. Tetrapassalus and Xenocranium are 
more specialized than Propalaeanodon in having fewer teeth and a 
shortened dentary, but they are more primitive (and typically 
epoicotheriid) in showing occlusal wear on the teeth. 

As metacheiromyids reduced the number of postcanine teeth, 
they retained a long dentary, hence leaving the back of the 
horizontal ramus edentulous. The trend in the Epoicotheriidae was 
toward shortening the dentary as teeth were lost. The antiquity and 
primitive grade of Propalaeanodon results in resemblances to both 



10 BREVIORA No. 455 

metacheiromyids and certain epoicotheriids and highlights the 
extent of parallehsm that has occurred within the Palaeanodonta. 

Propalaeanodon is one of those rare fossils that is morphologi- 
cally intermediate between two well defined families, and it strength- 
ens the alliance of the Epoicotheriidae and the Metacheiromyidae in 
the suborder Palaeanodonta. Because of its very early acquisition of 
advanced traits found also in Palaeanodon and its stratigraphic and 
geographic proximity to Palaeanodon, I believe Propalaeanodon is 
best regarded as a primitive metacheiromyid, probably lying in or 
very near the ancestry of Palaeanodon. Its very late Paleocene age 
makes derivation of the Metacheiromyidae from an epoicotheriid 
similar to early late Tiffanian Amelotabes highly probable. More- 
over, the primitive aspect of Propalaeanodon suggests that meta- 
cheiromyids did not become distinct from epoicotheriids before the 
Tiffanian. 

The humeri tentatively referred to Propalaeanodon possess spe- 
cializations characteristic of fossorial mammals (see Simpson, 1931 
and Rose, 1978 for additional functional interpretations). They are 
of interest in demonstrating that the humerus was already highly 
specialized in this late Paleocene palaeanodont. The deltopectoral 
crest is very similar to that in Palaeanodon. The supinator crest may 
have been slightly larger than in Palaeanodon, but no specimen of 
Palaeanodon preserves a complete supinator crest. 



AFFINITIES OF THE PALAEANODONTA 

Resemblances of palaeanodonts to Xenarthra and Pholidota have 
been enumerated by several authors (Matthew, 1918; Simpson, 
1931; Emry, 1970). Two questions arise from these comparisons: To 
what extent are these features demonstrably synapomorphic with 
one or the other of these orders? To what extent do they reflect 
parallelism or convergence? 

Detailed comparisons led Matthew (1918) to conclude that the 
Palaeanodonta are definitely related to the Xenarthra and probably 
to the Pholidota as well. He found no morphologic obstacles to 
direct descent of both orders from Palaeanodon, but for geographic 
and faunal reasons he deemed this origin of Xenarthra unlikely. He 
speculated that "early Tertiary faunas of South America . . . must be 
derived apparently from some late Cretacic fauna, unknown to us 



1979 PALEOCENE PALAEANODONT 11 

but presumably inhabiting some part of North America" (Matthew, 
1918: 653). Simpson (1931) detailed many characters to support 
alliance of the Metacheiromyidae and the Xenarthra, but he 
concluded that they are not so closely related to the Pholidota. 
Nearly four decades later, Emry (1970) critically reviewed Simpson's 
criteria, presenting arguments that he believed vitiated them. In 
Emry's view, the Metacheiromyidae were directly ancestral to the 
Manidae, and he included both in the order Pholidota. He rejected 
any special relationship between palaeanodonts and Xenarthra. 
Emry therefore discontinued formal recognition of the suborder 
Palaeanodonta, questioning whether epoicotheriid-metacheiromyid 
ties were really closer than metacheiromyid-manid relationship. 

A primary concern of these and other authors has been that 
known palaeanodonts were too specialized too early in the record to 
be direct ancestors of xenarthrans, since contemporary and dif- 
ferently specialized xenarthrans were known from South America 
(e.g., the early Eocene dasypodid Utaetus). In particular, tooth 
reduction was regarded as "prematurely specialized." The discovery 
of a primitive epoicotheriid (Rose, 1978) and now a primitive 
metacheiromyid, both from the late Paleocene and both with much 
less reduced dentitions, may obviate this objection. Meager evidence 
attests to the presence of dasypodids in the Riochican of Patagonia 
(Simpson, 1948), and these are possibly contemporaneous with the 
Paleocene palaeanodonts. However, while the age of the Riochican 
is usually given as late Paleocene, its precise correlation with North 
American land mammal ages is uncertain (Simpson, 1978). At least 
part of the Riochican is considered by some authors to be younger 
than late Paleocene, i.e. equivalent to the Clarkforkian and early 
Wasatchian (e.g., Marshall et al., 1977). It is known that the teeth of 
early xenarthrans had enamel and developed gabled wear surfaces 
(Simpson, 1932). Therefore Propalaeanodon itself, whose post- 
canine teeth apparently had already lost enamel and did not develop 
wear facets, may have been too specialized to be directly ancestral to 
the Xenarthra (but not Pholidota). These objections do not apply to 
Amelotabes, and nothing now known about it precludes it from 
possible direct ancestry to Xenarthra or Pholidota. Some post- 
cranial features, such as the extreme development of the supinator 
crest in palaeanodont humeri, are more specialized than in any 
known xenarthrans or pholidotans. However, some members of 
both groups possess an expanded, though smaller, supinator crest. 



12 BREVIORA No. 455 

If known palaeanodonts are ancestral to either Xenarthra or 
Pholidota, secondary reduction of the supinator crest must have 
occurred. This would not be required, however, if either or both 
orders were derived from more primitive palaeanodonts in which 
the supinator crest was less developed. 

Paleogeographic evidence must also be considered. There was no 
land connection between North and South America in the late 
Cretaceous or in the Paleocene (Freeland and Dietz, 1971; Malfait 
and Dinkelman, 1972). Therefore if palaeanodonts gave rise to 
xenarthrans, they could only have reached South America by 
rafting (Patterson and Pascual, 1972). In view of the specializations 
of metacheiromyids and the probability that Metacheiromyidae did 
not evolve before the Tiffanian, their resemblances to xenarthrans 
are more likely the result of convergence than of true affinity. 
Among known fossil forms, epoicotheriids are the most likely 
candidates for xenarthran ancestors, although this is probably 
dependent on the existence of earlier representatives than are now 
known. Derivation of manids from early palaeanodonts is possible 
and may be supported by the presence of an early manid, Patri- 
omanis, in the Chadronian of North America. However, contem- 
porary (or possibly older) manids have long been known from the 
Quercy Phosphorites of Europe (e.g. von Koenigswald, 1969) and a 
middle Eocene manid was recently reported from Messel, Germany 
(Storch, 1978). A direct land connection between North America 
and Europe existed until the end of Sparnacian (early Eocene) time 
(McKenna, 1972), hence it was presumably possible for manids to 
disperse in either direction before the middle Eocene. If manids 
evolved from palaeanodonts, it is most parsimonious to postulate 
North America as the place of origin, with subsequent dispersal to 
Europe. 

There can be little question that palaeanodonts were the closest 
ecological analogues of xenarthrans and pholidotans in the early 
Tertiary. It may be that palaeanodonts represent an early independ- 
ent radiation convergent to these orders. Evidence presented here 
and in other recent studies, however, strengthens the possibility of 
relationship among these groups. Once again it appears reasonable, 
as Matthew (1918: 655) wrote sixty years ago, "that there is, after 
all, a real affinity between the Pholidota and Xenarthra." 



1979 PALEOCENE PALAEANODONT 13 

ACKNOWLEDGMENTS 

I am greatly indebted to Dr. Parish A. Jenkins, Jr. and Mr. 
Charles R. Schaff (MCZ) for permission to describe the holotype, 
and to Drs. Donald Baird (PU) and Frederick Szalay (Hunter 
College) for permission to study the humeri from Princeton Quarry. 
To Mr. William Amaral (MCZ), I extend my gratitude for prepar- 
ing the holotype. I thank also Drs. Walter Coombs (AC), Robert J. 
Emry (National Museum), Philip D. Gingerich (UM), and Malcolm 
C. McKenna ( AMNH) for access to specimens used in this research, 
and Dr. George Gaylord Simpson for calling my attention to the 
recent article by Dr. Gerhard Storch. The figures were skillfully 
drawn by Ms. Karen Payne. Finally, I am grateful to Drs. Emry and 
Gingerich, Mr. David Krause and Prof. Bryan Patterson for their 
careful critical reading of the manuscript and suggestions for its 
improvement. 

REFERENCES 

Butler, P. M. 1972. The problem of insectivore classification. In K. A. Joysey 

and T. S. Kemp (eds.). Studies in Vertebrate Evolution. Edinburgh: Oliver and 

Boyd, p. 253-265. 
Colbert, E. H. 1942. An edentate from the Oligocene of Wyoming. Notulae 

Naturae, No. 109: 1-16. 
Emry, R. J. 1970. A North American Oligocene pangolin and other additions to 

the Pholidota. Bull. Amer. Mus. Nat. Hist. 142: 455-510. 
Freeland, G. L., and R. S. Dietz. 1971. Plate tectonic evolution of Caribbean 

— Gulf of Mexico region. Nature 232: 20-23. 
Gazin, C. L. 1952. The Lower Eocene Knight Formation of western Wyoming 

and its mammalian faunas. Smithson. Misc. Coll. 117: 1-82. 
Guthrie, D. A. 1967. The mammalian fauna of the Lysite Member, Wind River 

Formation, (Early Eocene) of Wyoming. Mem. S. Calif. Acad. Sci. 5: 1-53. 
HiLDEBRAND, M. 1974. Analysis of vertebrate structure. New York: John Wiley 

and Sons, 710 pp. 
Jepsen, G. L. 1940. Paleocene faunas of the Polecat Bench Formation, Park 

County, Wyoming. Part I. Proc. Am. Phil. Soc. 83: 217-341. 
Malfait, B. T., and M. G. Dinkelman. 1972. Circum— Caribbean tectonic and 

igneous activity and the evolution of the Caribbean plate. Geol. Soc. Amer. Bull. 

83: 251-272. 
Marshall, L. G., R. Pascual, G. H. Curtis, and R. E. Drake. 1977. South 

American geochronology: radiometric time scale for middle to late Tertiary 

mammal-bearing horizons in Patagonia. Science 195: 1325-1328. 



14 BREVIORA No. 455 

Matthew, W. D. 1918. Part V. Insectivora (cont.), Glires, Edentata. In W. D. 

Matthew and W. Granger, A revision of the Lower Eocene Wasatch and Wind 

River faunas. Bull. Amer. Mus. Nat. Hist. 38: 565-657. 
McKenna, M. C. 1972. Was Europe connected directly to North America prior 

to the middle Eocene? Evol. Biol. 6: 179-188. 
Patterson, B., and R. Pascual. 1972. The fossil mammal fauna of South 

America. In A. Keast, F. C. Erk, and B. Glass (eds.). Evolution, Mammals, and 

Southern Continents. Albany: State University of New York Press, p. 247-309. 
Reed, C. A. 1954. Some fossorial mammals from the Tertiary of western North 

America. J. Paleontol. 28: 102-111. 
Rose, K. D. 1978. A new Paleocene epoicotheriid (Mammalia), with comments 

on the Palaeanodonta. J. Paleontol. 52: 658-674. 
Rose, K. D., T. M. Bown, and E. L. Simons. 1977. An unusual new mammal 

from the early Eocene of Wyoming. Postilla, No. 172: 1-10. 
. 1978. Alocodontulum. a new name for Alocodon Rose, Bown and 

Simons, 1977, mm Thulborn 1973. J. Paleontol. 52: 1162. 
Simpson, G. G. 1931. Metacheirumys and the Edentata. Bull. Amer. Mus. Nat. 

Hist. 59: 295-381. 
. 1932. Enamel on the teeth of an Eocene edentate. Amer. Mus. Novitates, 

No. 567: 1-4. 
. 1948. The beginning of the age of mammals in South America. Bull. 

Amer. Mus. Nat. Hist. 91: 1-232. 
1959. A new middle Eocene edentate from Wyoming. Amer. Mus. Novi- 
tates, No. 1950: 18. 
1978. Earlv mammals in South America: Fact, controversy, and mystery. 



Proc. Am. Phil. Soc. 122: 318-328. 
Storch, G. 1978. Em Schuppentier aus der Grube Messel— zur Palaobiologie 

eines mitteleo/.anen Maniden. Natur und Mus. 108: 301-307. 
VON KoENKiSWM 1). W. 1969. Die Maniden(Pholidota, Mamm.)deseuropaischen 

Tertiars. Mitt. Bayer. Staatssamml. Palaont. hist. Geol. 9: 61-71. 
West, R. M 1973. .\n early middle Eocene epoicotheriid (Mammalia) from 

southwestern Wvomum. J. Paleontol. 47: 929-931. 



MAR 1 8 mS 

B R E V I O-I-A 

Miiseimi of Comj^arative Zoology 



lis ISSN ()()()6 96^ 



Cambridge, Mass. July 31, 1979 Number 456 



DESCRIPTION OF A NEW HAWAIIAN GOBIID FISH 
OF THE GENUS TRIMMA 



Phillip S. Lobel' 



Abstract. A new gobiid fish species, Trimma taylori from Hawaii is described. A 
key to the nine nominal species of Trimma and a discussion of the Hawaiian gobies are 
inckided. The genus Trimma is reviewed and a preliminary assessment it presented of 
the possible relationships of Trimma. The nominal species of Trimma as now- 
recognized may eventually be seen to comprise more than one group. 

INTRODUCTION 

Gobiid fishes are among the least known of the tropical marine 
fauna due to their generally miniscule size and secretive habits. 
Despite the difficuhy of collecting and identifying such small reef 
fishes, a great number of species (over 1 500) and genera (ca. 550) have 
been described (E. Lachner, pers. comm.). This high species diversity 
of gobiids rivals and exceeds that of most other fish families. 
However, the full extent of goby species diversity is far from known. 
For example, Trimma presently contains eight nominal species but 
an additional 25 to 30 remain to be described (D. Hoese, pers. 
comm.). The primary task of gobiid systematists at this time remains 
the study of species identification and distribution. The new goby 
described herein is the first Trimma to be found in Hawaiian waters. 
This species brings the total number of Hawaiian fishes to 682 (see 
Randall 1976). 



'Museum ot Comparative Zoology, Harvard University. Cambridge. Massachu- 
setts 02138. 



BREVIORA 



No. 456 



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1979 GOBIID FISH OF THE GENUS TRIMMA 3 

MATERIALS AND METHODS 

All specimens were measured with dial calipers. Proportional 
measurements were made by micrometer in a dissecting microscope 
and with calipers. Proportions are expressed as percent of standard 
length (SL) in millimeters (Table 1). Fish were collected with the 
anesthetic quinaldine mixed with 95% isopropyl alcohol. Specimens 
of T. taylori were captured in a fine meshed net and immediately 
placed into vials to prevent damage. Color photographs were taken 
as records of live coloration. Fifty specimens have been deposited as 
representative types of the species. 

Abbreviations of institutional names cited herein are as follows: 
AMS. Australian Museum, Sydney; BPBM. Bernice P. Bishop 
Museum, Honolulu; BMNH, British Museum of Natural History, 
London; CAS, California Academy of Sciences, San Francisco; 
MCZ, Museum of Comparative Zoology, Harvard University; 
USNM, National Museum of Natural History, Washington, D.C. 

Comparative material examined: Trimma caesiura, holotype, 
USNM 51772; USNM 156727 to 156738 (25+ specimens); USNM 
218362. T. tevegae. paratypes, USNM 203437 (9). T. eviotops, 
holotype, USNM 116169; paratypes, USNM 116170 (3); paratype, 
MCZ 37267 ( 1 ). X-ray photographs of these species are on file at the 
USNM and BPBM. I have also examined species of related genera 
deposited at the USNM, MCZ, AMS, and BPBM. 



Trimma Taylori sp. nov. 
Fig. 1; Table 1. 

Holotype: BPBM 19919, 15.5 mm SL, taken in a caveat 44 meters 
depth off the leeward coast of Oahu, Hawaii, near Makaha Beach, 
July 12, 1975, P. S. Lobel. 

Paratypes: BPBM 19920. 15 mm SL, locality same as holotype, 
July 20, 1975, P. S. Lobel (specimen photographed in color and 
figure 1). MCZ 51685, 14 mm SL, collected with holotype. BPBM 
19921, 2 specimens each 14 mm SL; collected with BPBM 19920 
(cleared and stained). MCZ 51686, 2 specimens, 13 15 mm SL; 
collected with BPBM 19920. MCZ 51687, five specimens, 12-13.5 
mm SL; collected with holotype. USNM 215323. 9 specimens, 6 13 
mm SL; same location as holotype, Aug. 13, 1975, P. S. Lobel. AMS 
118686-001, eight specimens, 6-13 mm SL; same location as 



BREVIORA 



No. 456 




Figure 1. Trimnia lavlori. male 15 mm. s.l. Paratype BPBM 19920. 

holotype, Aug. 13. 1975, P. S. Lobel. CAS 35450, six specimens, 
11 13 mm SL; collected with BPBM 19920. BPBM 19922, eight 
specimens, 6 14.5 mm SL; same location as holotype, Aug. 13, 1975, 
P. S. Lobel. BMNH 1976.4.6.1-7, seven specimens, 6 13 mm SL; 
same location as holotype. Aug. 13, 1975, P. S. Lobel. 

Diagnosis. This species can be easily distinguished from all other 
Trimma spp. by a combination of characters: Dorsal VI L 10, anal L 
10, and pectoral 14; second spine of the first dorsal fin filamentous, 
longest, extending past the midpoint of the second dorsal fin when 
depressed. 

Description. Morphometric characters of specimens are presented 
in Table I . Tooth shape and pattern are illustrated in Figure 2. Head 
length 25% of standard length. Eye diameter about 40% head 
length. First dorsal fin originates one third the standard length 
distance from snout. Mouth slanted about 45° upward, corner of 
mouth in line with midpoint between forward margin of eye and 
pupil. Eye to snout distance approximately one-half eye diameter. 
Body scaled except for head and operculum, but including nape. 
Scales in longitudinal series from opercule to hypural base, 22-23. 
Seven scales from the origin of the anal to the first dorsal. Middorsal 
area from first dorsal spine to head with three scales. Gill raker count 
of three specimens: 3 1-12, 3 1-13, and3-l-14. Five lobes on 



1979 



GOBIID FISH OF THE GENUS TRIMMA 



pseudobranch. Fin ray formulae: Dorsal VI-1. 10. anal I. 10. pelvic 
15, and pectoral 14. Vertebral count 25, not including urostyle. 
Dorsal fins separate. Second spine of first dorsal fin elongated, 
extending over second dorsal fin. Posteriormost ray of second dorsal 
fin and of anal fin split to base but counted as one. Pelvic fins 
connected at base but not united by frenum. Additional characters 
are the same as described for the genus (see Cohen and Davis 1969). 

Color in life. Overall color pale straw yellow with darker yellow 
pigment spots bordering base of each scale. Immediately posterior to 
the eye, orange spot almost equal in area to eye. Dorsal portion of this 
orange spot speckled with small melanophores. Head dorsally 
speckled with numerous orange and black chromatophores. Dorsal 
and anal fins with broad deep orange band at base and with thin band 
of black pigmentation above it; fins overall light orange, with a 
diffuse dark edge. Caudal fin similar in color to dorsal and anal fins 
but without deep orange band. Pelvic and pectoral fins yellow- 
orange. 




Figure 2. Trinuna taylori. detail ol mouth and teeth, S.E.M. photo by E. Seling. 



BREVIORA 



No. 456 







MALE 




FEMALE 

Figure 3. Trimnia laylori. sexual dimorphism of genital papillae. Drawn from 
MCZ 51686. Paratypes by Karsten Hartel. 

Color in alcohol. All colors vanish, body pale. Pigment spots on 
dorsum of head visible after one year but fade thereafter. Fins 
transparent. 

Sexual dimorphism. Morphometric differences are not immedi- 
ately apparent between the sexes; however, males and females may be 
distinguished on the basis of genital papillae (Fig. 3). The papilla of 
the male is slender, curved and tapers to a point posteriorly. The 
female's papilla is greater in diameter than that of the male and 
uniform in thickness, terminating in a round opening with a bilobed 
flap bilaterally; it is also directed posteriorly but it is not as long as 
that of the male. Immature gobies exhibit no sexual difference in the 
shape of the papilla (see also Egami 1960). 

There are significant differences in both shape and size of male and 
female genitalia among Trimma eviotop.s. T. taylori and T. caesiura. 
The genitalia of T. taylori and T. caesiura are very similar in shape 
and structure but differ in relative length of the papilla of the male. 



1979 GOBIID FISH OF THE GENUS TRIMMA 7 

The relative lengths of male papillae (in percent of standard length) 
are: T. taylori6%, T. caesima 2)A%. and T. eviotops 2.2%. The papilla 
of T. eviotops is not curved as is that of T. taylori (Fig. 3), but extends 
straight posteriorly and is comparatively thinner and not tapering to 
a point, ending bluntly. A pigment line runs ventrally along the length 
of the male papilla of T. eviotops. 

Arai (1964) illustrated the genital papillae of 25 goby species. Four 
are particularly relevant here: Eviota abax (Jordan and Snyder), 
Quisquilius eugenius (Jordan and Evermann). Zonogobius semi- 
doliatus (Cuvier and Valenciennes) and Fusigobius neophytus 
(Giinther) (Fig. 4 modified from Arai. 1964). The genitalia in both 
sexes of T. taylori and T. caesiura appear most like that of Fusigobius 
neophytes. The genital papilla of T. eviotops is more similar to those 
of Zonogobius semidoliatus. The genitalia of Quisquilius and 
Zonogobius are superficially more similar to each other than either is 
to those of Fusigobius. Such comparisons may become important 
when the phylogeny of these taxa is revised. 



1^:3= 



XH"^' ^.^--^ ' EVIOTA ABAX 



QUISQUILIUS EUGENIUS 








ZONOGOBIUS SEMIDOLIATUS 



FUSIGOBIUS NEOPHYTES 
Figure 4. Genital papillae of some related Gobiids. Modified from Arai (1964). 



BREVIORA 



No. 456 




Figure 5. Triinma tayloii. detail of scales on flank, S.E.M. photo by E. Seling. 

Scale Morphology. The number of ctneii in a single row along the 
posterior margin of the scale may differentiate species of Tritnnia. 
The pattern and structure of scales of T. taylori (Fig. 5). and of T. 
caesiura (Fig. 6), is characteristic for each species. Trimma taylori 
possess fewer and larger scale ctenii (mean 10.9, range 8- 14 per scale, 
N = 1 2) than does T. caesiura (mean 3 1 .6, range 28 36, per scale, N = 
5). Comparison to other species was not possible due to the 
unavailability of specimens. Consequently, the taxonomic value of 
such scale morphology is uncertain. 

Zoogeography. Triinma taylori is known only from the island of 
Oahu, Hawaiian archipelago. All other Triinma species occur in the 
Indopacific oceanic region or in the Indian Ocean. None are 
recognized from the Atlantic Ocean or within the temperate zones. 
All Trimma spp. with known habits are reef dwellers. 



Entymology. Trimma taylori is named in honor of Dr. Leighton R. 
Taylor, Jr. on the occasion of his appointment as director of the 
Waikiki Aquarium. A tradition of the Waikiki Aquarium has been 



1979 



GOBIID FISH OF THE GENUS TRIMMA 



that each of its directors have a uniquely Hawaiian fish as a 
namesake. 

DISCUSSION 

The nominal species of Trimma 

The original description of the genus Trimma was not adequate 
and there has consequently been confusion in the generic assignment 
of the species. The genus Trimma was defined on the basis of T. 
caesiura by Jordan and Scale (1906). It was diagnosed as "near Eviota 
but the region before the dorsal is fully scaled. In texture the species is 
firmer and less translucent" (Jordan and Scale, 1906, p. 361). This 
description does not adequately define the genus with regard to some 
of the species that have since been included in it. Additional 
morphometric data of T. caesiura are listed in Table 1 with 
comparisons to T. taylori. 

The species included in the genus Trimma which appear closely 
related to T. caesiura based on the original generic characters and 
other characters described below are Trimma naudei Smith (1956) 
and T. tevegae Cohen and Davis (1969). Other species placed in 




Figure 6. Trimma caesiura, details of scales on flank. S.E.M. photo by E. Seling. 



10 BREVIORA No. 456 

Trimma present an enigma. Trimma eviotops Schultz (1943) is 
presently considered a valid species but its placement in Trimma will 
soon be reconsidered (S. Karnella. pers. comm.). It clearly differs 
from T. caesiura in the lack of scales along the nape. Trimma eviotops 
does not fit comfortably in other closely related genera. For example, 
it differs from species oi Eviota by lacking head pores, although this 
may not be taxonomically significant. Smith (1956) remarked that T. 
eviotops may well become the type of a new genus. Consequently, it is 
included here only provisionally. However, there is no difficulty in 
assigning Trimma taylori to the genus Trimma. 

I follow Hoese's (pers. comm.) unpublished classificatron for 
species of Trimma. The species (as originally designated) are Eviota 
macrophthalma Tomiyama (1936), Eviota grammistes Tomiyama 
(1936), Zonogobius flammeus Smith (1959) and Zonogobius coral- 
linus Smith (1959). These all differ from Trimma caesiura in lacking 
predorsal scales along the nape but resemble Trimma eviotops in that 
character. Until related species and genera are better known the 
decision to include these species in Trimma involves the fewest 
complications. However, if T eviotops is removed from Trimma, 
then the generic placement of these other species should also be 
carefully reconsidered since some are more similar to T eviotops 
than to T. caesiura, the species on which the genus is based. Meristic 
and color traits of the nominal species of Trimma are contrasted in 
Table 2. 

There exists numerous problems with the classification of the 
genera and species of gobies. However, most of the conflicts cannot 
be solved until both more species are described and the characters 
defining the genera are delineated. The genus Tritnma is no exception 
but since assignment of Trimma taylori into this genus offers no 
conflicts, discussion of other issues is postponed until more data are 
available. 

Ecology. Trimma taylori appears to be an exclusively deep reef 
species that inhabits the recesses of caves and crevices. Its habits are 
somewhat like those described for T. tevegae (Cohen and Davis, 
1969). During the daytime, swarms of 50 to 150 individuals hover 
near the ceiling, in about the midsection of a cave or pocket. The 
shallowest location at which adults have been observed is at a depth 
of 30 meters. A few juveniles have been seen somewhat shallower. 



1979 GOBIID FISH OF THE GENUS TRIMMA 11 

The cave habitat in which all specimens were collected was typically a 
small crevice (one to two meters deep) along a vertical wall ot the deep 
reef. Oahu, Hawaii. Perhaps it is significant that these shallow 
pockets did not house holocentrids or priacanthids. Caves containing 
the latter were inspected for Trimma taylori but none were found. 
Sympatric cave inhabitants which were conspicuous included a 
prawn. Stetiopus (an undescribed species) and the fishes Pterois 
sphex (Scorpaenidae). Apogon macidiferus (Apogonidae), Gymno- 
thorax spp. (Muraenidae). an undescribed species of Quisquilius 
(Gobiidae). Scorpaenid sp., and Pseudanthias sp. (Serranidae- 
Anthiinae). Of these fishes, only Pterois sphex, Gymnothorax spp. 
and Apogon niaciiliferus seem likely potential predators of T. taylori. 
Trimma taylori feeds on harpacticoid copepods. Eight specimens 
were examined for gut contents and three of these were empty. 

Key to the Nominal Species of Trimma 

1 . a. Predorsal scales on nape 2 

b. No scales on nape 5 

2. a. Dorsal fin with 8 rays 3 

b. Dorsal fin with more than 8 rays 4 

3. a. Gill raker on first arch 3+I + I6 T. caesiura 

b. Gill raker on first arch 3+I + I3 or 14 T. naudei 

4. a. Dorsal fin VI-IIO. Pectoral 14 T. taylori 

b. Dorsal fin VI-I9. Pectoral 13 T. levegae 

5. a. Pectoral fin with fewer than 19 rays 6 

b. Pectoral fin with 19 rays T. coralinus 

6. a. Pectoral fin with 1 7 rays 7 

b. Pectoral fin with fewer than 1 7 rays 8 

7. a. Body with eight dark bars or saddles T. eviotops 

b. Body without bars or saddles T.flammeus 

8. a. Pectoral with 15 rays, 1 1 scales transverse series T. grammistes 

b. Pectoral with 16 rays. 9 scales transverse series T. macrophthalma 

Supplementary key to Hawaiian gobiid fishes 

1 . a. pelvic fins separate (no frenum) 2 

b. pelvic fins united to form a cup See Gosline and Brock I960, p. 266 

2. a. each scale with a dark crescent shaped mark, no spines on the preopercle 

3 

b. no markings on scales, head compressed. 3 to 5 spines on the preopercle . . 
Asterropteryx semipumtalus 

3. a. no predorsal scales on nape Eviota epiphanes 

b. predorsal scales on nape present Trimma taylori 



12 



BREVIORA 



No. 456 



Nominal 
Species 



TABLE 2 
Comparison of the Nominal species of Trimma 



Author 







Pec- 


Pel- 


Longi- 


Dorsal 


Anal 


toral 


vic 


tudinal 


fin 


fin 


fin 


fin 


scales 



T. caesiura^ 



Jordan and VI-L8 
Scale 1906 



15 L5 23-25 



T. coralinus 



(Smith 1959) VI-LIO L9 19 



30 



T. eviolops 



Schultz 
1943 



VII.9-I0 L8-9 17 L5 26-27 



T.flammeus (Smith 1959) VI-1,9-10 1,8-10 17 



26-36 



T. grammistes- (Tomiyama VI-1.10 

1936) 



T. macruph- 
ihalina- 

T. naiidei 



(Tomiyama VL1,9 
1936) 

Smith 1956 VI-L8 



1.9 
L9 



15 1.5 



16 1.5 



16 



30 

23 
24-25 



T. tavlori 



n.sp. 



VI-LIO LIO 14 L5 22-23 



T. teve^ae 



Cohen and V1-L9 L9 

Davis 1969 



13 L5 



28 



'1 examined 21 specimens for additional counts. 
-Pectoral fin count from drawing with description. 



1979 



GOBIID FISH OF THE GENUS TRIMMA 



13 



TABLE 2 
Comparison of the Nominal species of Trimma 



Scale Gillrakers 

Trans- on first Verte- 

verse arch brae Locality 



Live Coloration 



6-7 



10 



8-9 



8-9 



3-1-16 



3+1+10 



3+1+11-12 



25 Samoa 



Seychelles 



3+1 + 1 1 -12 25 Phoenix & 

Samoan Islands 



Australia & 
S. Africa- 

Japan 



Plain bright red with grey spots 
on back of tail and grey spots on 
many scales of back and head. 
Fins all bright red. 

Body red with two silvery areas 
on back below first, and three 
below second dorsal fin. Orange 
spots on head and peduncle. 
Rays red with pink membrane. 

Pale with eight vertical bars, 
saddles on head and body. 

Light orange with darker spots; 
edge of soft dorsal and caudal, 
dusky. 

Longitudinal dark bands. 



Japan 



(not described) 



3+1+13-14 



Seychelles 



3+1 + 12-14 25 Hawaii 



25 New Britain 



Body and fins brilliant orange- 
scarlet, irregularly mottled with 
iridescent light areas; dorsal fin 
with two light pink bars between 
the orange. 

All straw yellow with orange-red 
and dark pigment spots on head. 

(Live body color not described) 
Dark reddish spot at end of 
caudal peduncle. 



14 BREVIORA No. 456 

NOTE: Asterropteryx semipunctatus and Eviola epiphanes were placed in the 
Eleotridae by Gosline and Brock (1960). However, the only eleotrid in Hawaii is 
Eleotris samivicensis, which is restricted to freshwater. It can be easily distinguished 
from the gobies. E. sandvicensis possesses about 75 scales in longitudinal series 
whereas gobies generally have less than 30 longitudinal scales. In addition, eleotrids 
are primarily freshwater and have six branchiostegal rays; gobies are mostly marine 
and have five branchiostegal rays. There are 21 other gobies presently known in the 
Hawaiian Islands. 

ACKNOWLEDGEMENTS 

I thank Douglas Hoese ( AMS) who first examined my specimens 
and confirmed them as a new species. D. Hoese also generously 
supplied valuable information which made my task considerably 
more manageable. Ernest Lachner (USNM) and Susan Karnella 
(USNM) were especially helpful by explaining to me many of the 
problems associated with goby systematics and loaning specimens. I 
thank Karel Liem (MCZ), John Randall (BPBM) and especially 
William Fink (MCZ) for their comments. Ed Baughman. John Earle 
and Janie Culp helped to collect the specimens. Ed Seling (MCZ) 
prepared the scanning electron microphotographs with support from 
NSFgrantBMS 7412494. Karsten Hartel (MCZ) prepared Figure 3. 
Kathie Cunningham and Terry Dash kindly typed the manuscript. 
Support for field work was provided by the Department of Zoology, 
University of Hawaii while I was an undergraduate there. Additional 
support and travel were provided by the Biology Department, 
Harvard University. 

LITERATURE CITED 

Arm, R. 1964. Sex characters of Japanese gobioid fishes (I). Bull. Nat. Sci. Mus. 

Tokyo, 9: 295 306. 
CoHFN, D. M. AND W. p. Davis. 1969. Vertical orientation in a new gobioid fish 

from New Britain. Pac. Sci., 23: 317-324. 
Egami, N. 1960. Comparative morphology of the sex characters in several species 

of Japanese gobies, with reference to the effects of sex steroids on the characters. 

J. Fac. Sci. Univ. Tokyo, 9:67-100. 
Gosline, W. A. and V. E. Brock. I960. Handbook of Hawaiian fishes. Honolulu: 

University of Hawaii Press, IX + 372 pp. 
Jordan. D. S. and A. Seale. 1906. The fishes of Samoa. Bull. Bur. Fish., 25: 

173-^55. 
Randall, J. E. 1976. The endemic shore fishes of the Hawaiian Islands, Lord 

Howe Island and Easter Island. Colloque Commerson 1973. O.R.S.T.O.M. 

Travaux et Documents., No. 47: 49-73. 



1979 GOBIID FISH OF THE GENUS TRIMMA 15 



ScHL'LTZ, L. P. 1943. Fishes of the Phoenix and Samoan islands collected in 1939 

during the expedition of the U.S. S. "Bushnell". Bull. U.S. Nat. Mus.. 180: 1-316. 
Smith, J. L. B. 1956. The fishes of Aldabra. Part 6. Annals and Magazine of 

Natural History, Ser. 12., Vol. 9. pp. 817-829. 
Smith, J. L. B. 1959. Gobioid fishes of the families Gobiidae. Periophthalmidae, 

Trypauchenidae, Taenioididae. and Kraemeriidae of the Western Indian Ocean. 

Rhodes Univ. Ichthy. Bull.. 13: 185-225. 
ToMiYA.MA, I. 1936. Gobiidae of Japan. Jap. J. of ZooL, 7; 37-112. 



MAR 1 8 ttb 

HARVARD 

B R E V I 0' R A 

Muiseium of Comparative Zoology 

us ISSN 0006-9698 

Cambridge, Mass. January 31, 1980 Number 457 

TWO NEW SPECIES OF ELEUTHERODACTYLUS 

(AMPHIBIA: LEPTODACTYLIDAE) 

FROM THE LOWLANDS AND 

LOWER CLOUD FORESTS OF 

WESTERN ECUADOR 

John D. Lynch' and Kenneth Miyata- 

Abstract: Eleutherudactylus muricatus sp. nov., an ally of the large, flare- 
snouted frogs of the rubicundus assembly, is named from lowland and lower cloud 
forest localities in Provincia Pichincha, Ecuador. The new species is smaller than its 
sympatric allies E. creminguis and E. laiidiscus. E'.eutherodactylus tenehriunis sp. 
nov., is also found in primary lowland and lower cloud forests in west-central Ecua- 
dor. It is allied to a species found in the high cloud forests in western Ecuador but 
differs in color pattern and lacks a calcar and ulnar tubercles. 

Introduction 

Approximately 15 species of Eleutherodactylus inhabit the 
Pacific lowlands of Ecuador (Lynch, in press) and perhaps another 
40 species are found in the cloud forests of the Pacific versant in the 
Ecuadorian Andes. Most of these frogs are small organisms (less 
than 35 mm SVL) but two members of the rubicundus assembly 
{Eleutherodactylus crenunguis Lynch and E. latidiscus {^ou\tngtr)\ 
are much larger species having long, slender limbs and digits and 
large emarginate (or notched) digital pads. 

Our field work in western Ecuador over the past several years has 
revealed that crenunguis is a frog of the lower cloud forests 
(800-1500 m) and is not an altitudinal replacement for latidiscus 
(20 1500 m). In the course of this field work two additional species 
having notched or indented digital pads were found in sympatry 



'John D. Lynch, School of Life Sciences, The University of Nebraska, Lincoln, 
Nebraska 68588 

-Kenneth Miyata, Museum of Comparative Zoology, Harvard University, 
Cambridge, Massachusetts 02138 



2 BREVIORA No. 457 

with both of the larger species. In spite of sharing the notched pads, 
only one of these species seems allied to the ruhicundus assembly as 
defined by Lynch (1979). The other species appears to be most 
closely allied to a species found in the high cloud forests of western 
Ecuador (Lynch and Trueb, in press). 

The format of the descriptions follows Lynch's long-used style. 
Abbreviations employed below include the following: SVL — 
snout-vent length; HW — head width; lOD — interorbital distance; 
E-N — eye to nostril distance. 

Eleutherodactylus nniricatus sp. nov. 

Holotype: MCZ 94469, an adult male collected at the Rio Fai- 
sanes where it is crossed by Ecuador Highway 28 (the road from La 
Palma to Quito via Chiriboga), 14.4 km from the junction with 
Highway 30 (the Aloag to Santo Domingo de los Colorados road) 
at La Palma. Provincia Pichincha, Ecuador. 1380 m, on 12 Novem- 
ber 1977 by Ken Miyata. 

Paratypes. MCZ 92091, 92095, 92100-01, 94848, 97528-31, 
USNM 211 172-74, topotypes; MCZ 97592, Centinela, 14.1 km SE 
Patricia Pilar by road, Provincia Pichincha, 570 m; MCZ 90337, 
94456, 94460, Centro Cientifico Rio Palenque, 47 km S Santo 
Domingo de los Colorados, Provincia Pichincha, 220 m. 

Diagnosis. 1 ) skin of dorsum smooth with numerous conical 
tubercles, that of venter feebly areolate; no dorsolateral folds; no 
anal sheath; 2) tympanum moderately distinct, its length Va eye 
length; 3) snout subacuminate in dorsal view, rounded in lateral 
profile; canthus rostralis moderately distinct; 4) upper eyelid much 
wider than lOD, bearing many conical warts: no cranial crests; 5) 
vomerine odontophores large, triangular in outline, narrowly separ- 
ated; 6) males with vocal slits, subgular vocal sac; no nuptial pads; 
7) first finger shorter than second; all digits bearing broad discs, 
pads on fingers H-IV, those of HI and IV notched; 8) fingers bear- 
ing lateral keels; 9) 1-2 small ulnar tubercles; 10) one large conical 
tubercle on heel; low tubercles along outer edge of tarsus; short 
inner tarsal fold; 11) two metatarsal tubercles, inner elongate, 10 
times size of round, subconical outer; supernumerary plantar tuber- 
cles at base of toes; 12) toes bearing lateral fringes, not webbed; toe 



1979 



TWO NEW SPECIES OF ELEUTHERODACTYLUS 




X E 
- E 






^ 5: > 

2 -- E 
S^ E 



C £ -' 

-? E £ 

r-' O^ ro 






^ I^ 



^ ID -^ 
r ^ 'S 



c uj 



> 



4 BREVIORA No. 457 

pads notched, smaller than those of outer fingers; 13) dorsum brown 
with black spots; ventral surfaces brown with cream flecks; slightly 
darker brown chevrons on throat; posterior surfaces of thighs 
brown; 14) adults from type-locality moderate-sized, males 31.8- 
40.7 (x= 36.0, n = 4) mm SVL, one female 46.3 mm SVL; two 
gravid females from the lowlands (Centinela and Rio Palenque) are 
only 33.8-36.0 mm SVL. 

Eleutherodactylus muricatus is most similar to E. crenunguis but 
differs in coloration (no orange patch on the breast) and in having 
fewer, larger tubercles on the dorsum (Fig. 1 ). It is also smaller than 
E. crenunguis (Lynch 1976). 

Description. Head as wide as or wider than body, wider than 
long: HW 37.1-39.9 (x= 38.4, n= 7) per cent SVL; snout subacum- 
inate in dorsal view, rounded in lateral profile; nostrils weakly pro- 
tuberant, directed dorsolaterally; canthus rostralis relatively sharp 
(swollen), straight or weakly convex; loreal region concave, sloping 
gradually to lips; lips fiared; E-N 80.7 100.0 per cent (x = 89.4, n = 
7) eye length; upper eyelid 100.0-132.3 per cent (x = 1 19.7, n = 7) 
lOD, bearing many pungent tubercles; no cranial crests; supratym- 
panic fold moderately 'distinct, obscuring upper edge of tympanum; 
tympanum not prominent, round, separated from eye by distance 
equal twice tympanum length; tympanum length 22.0-27.9 (x = 
25. 1 , n = 6) per cent eye length except in MCZ 94456 (small female 
from Rio Palenque has ratio of 37.5 per cent); postrictal tubercles 
present, not prominent; choanae round, not concealed by palatal 
shelf of maxillary arch; vomerine odontophores median and poste- 
rior to choanae, large, triangular in outline, separated on midline by 
distance less than 1/3 an odontophore width, each larger than a 
choana, bearing 5 12 teeth in a transverse row; tongue longer than 
wide, its posterior edge notched; posterior 1 / 3 not adherent to floor 
of mouth; males with vocal slits posterolateral to tongue, median 
subgular vocal sac. 

Skin of dorsum smooth but bearing many warts, most numerous 
on lower back (where skin is tuberculate); many elongate, subcorn- 
eal warts on flanks; no dorsolateral folds; no anal sheath; skin 
posterior and posterolateral to anus areolate but also bearing coni- 
cal warts; skin of throat areolate, that on other ventral surfaces 
smooth with very feeble areolations; discoidal folds well anteriad to 
groin; one or two small ulnar tubercles; palmar tubercle bifid, larger 



1979 TWO NEW SPECIES OF ELEUTHERODACTYLUS 5 

than oval thenar tubercle; several prominent supernumerary palmar 
tubercles; subarticular tubercles round, pungent; fingers bear lateral 
keels; all digits with pads, that of thumb scarcely wider than digit 
below pad, on II twice as wide, on III-IV three times as wide (wider 
than tympanum); pads of fingers III-IV notched apically; discs on 
all pads broader than long; fingers long, first slightly shorter than 
second; thurjib of male lacking nuptial pad or swelling. 

Numerous tubercles on heel, one large conical tubercle (not a 
calcar); low tubercles along outer edge of tarsus; short fold at base 
of inner metatarsal tubercle which is three times as long as wide, 10 
times size of round, subconical outer metatarsal tubercle; supernu- 
merary plantar tubercles at bases of each toe; subarticular tubercles 
longer than wide, pungent; toes bearing distinct lateral fringes, 
broad discs, expanded pads; pads notched apically, smaller than 
those of outer fingers; heels of flexed hind limbs broadly overlap; 
shank 54.7 62.2 per cent (x = 58.4, n = 7) SVL. 

Brown above with black spots enclosing tubercles; bars evident 
on thighs but remainder of pattern (limbs, labial bars, canthal- 
supratympanic stripe, dorsal chevrons, etc.) only suggested by black 
spots; venter brown with cream flecks and small spots; inverted 
brown chevrons on chin and throat; undersides of limbs and groin 
brown with cream flecks; anterior and posterior surfaces of thighs 
brown with cream flecks. In small individuals, ground color gray 
and with evident shank bars (narrow and oblique), labial bars, and 
canthal-supratympanic stripe. 

In life, E. muricatus from the type locality are pale to dark brown 
with black rings around large tubercles; flanks paler brown with 
yellowish wash; venter dark purplish-brown with yellow-brown 
mottling; iris gold flecked with black. Lowland specimens lack the 
black rings around the tubercles; the venter is a muddy yellow 
mottled with dark brown; the lower flanks, groin, and underside of 
the limbs have a purplish-brown wash; the iris is copper. The above 
color descriptions apply to daytime patterns; at night all individuals 
are much paler in dorsal coloration, ranging from pale buff to olive 
brown, but the various markings remain intact. 

Measurements of holotype (in mm). SVL 32.3; shank 20.1; HW 
12.5; head length 1 1 .6; upper eyelid 2.9; lOD 2.9; tympanum length 
I.O; eye length 4.3; E-N 3.9. 



6 BREVIORA No. 457 

Etymology. The specific epiphet is derived from the Latin, 
meaning spiny, in reference to the pungent tubercles on the dorsum 
and upper eyeHd. 

Variation. The two adult females from low elevations (MCZ 
94456 from Rio Palenque. 220 m. and MCZ 97592 from Centinela, 
570 m) are much smaller than the only adult female from the type- 
locality. They are also peculiar (Fig. 1) in having prominent verte- 
bral stripes which are yellow-cream to buffy orange in life. All of the 
material from the lower elevations has venters more pale than the 
topotypic material; in life the ventral surfaces of the topotypic popu- 
lation are predominantly brown with some yellowish mottling while 
those of the low elevation populations are predominantly yellowish 
with brown mottling. The lowland populations also have more 
prominent tubercles on the eyelids and dorsum. All of these differ- 
ences are rather minor and probably represent slight geographical 
or altitudinal variation. We are taking a conservative approach in 
assigning all of these specimens to the same taxon; further work 
may well prove we have confused two species. 

Remarks. Adults of E. cremmguis and E. latidiscus are seldom 
encountered. No adult females of E. crenunguis have been found, 
although juvenile females to 41.3 mm have been examined, and 
adult males range in size from 35.0 to 49.2 mm SVL. Very few adults 
of E. latidiscus are available even though juveniles are often quite 
common. Three males with vocal slits are 43.9-50.0 mm SVL and 
four females having convoluted oviducts are 49.7-64.5 mm SVL. 
Both of these frogs are considerably larger than the sympatric E. 
muricatus. 

Natural History. Eleutherodactylus muricatus from the type- 
locality have been collected on streamside vegetation within 1.5 m of 
the water surface. Several specimens were taken on logs just above 
water level, but the majority were perched on large leaves adjacent 
to the stream. For a description of the type-locality see Miyata 
(in press). Individuals are found in low density on most nights at this 
locality. 

The lowland specimens have all been taken from primary forest 
or, in the case of the Centinela specimen, from very recently eut 
primary forest. The Rio Palenque specimens were taken from low 
vegetation in forest with a dense canopy on rainless nights. 



1979 



TWO NEW SPECIES OF ELEUTHERODACTYLUS 



Eleutherudactylus nmricatus appears to have a spotty distribu- 
tion, somewhat analogous to that seen in other anurans in this 
region (Lynch 1977). E. muricatus has not been collected at the Rio 
Orito. a locality very close to the type locality, despite several visits 
by field parties from the University of Kansas. More field work is 
needed to confirm the spottiness of the distribution. 

Eleutherodactyliis tenehrionis 

Holotype: MCZ 90326, an adult male collected at the Hotel 
Tinalandia, 16 km E Santo Domingo de los Colorados by road, 
Provincia Pichincha, Ecuador, 800 m, on 6 August 1975 by Ken 
Miyata. 






Figure 2. Outlines of heads of western Ecuador Eleutherodactylus. (A) E. 
cremmguis. MCZ 92099; (B) E. muricatus. MCZ 92095; (C) E. latidiscus. cotype, 
BM 98.4.28.109; (D) E. tenehrionis. MCZ 92081. Scale for A-C is between figs. 
A and C; that for D is to its right. Scales equal 5 mm. 



8 BREVIORA No. 457 

Paratypes: MCZ 88890, 90325, 90327-29, 92079 81, 94712, top- 
otypes; KU 179224-27, Santo Domingo de los Colorados, Provincia 
Pichincha, 580 m; KU 179228 30, 2 km E. 1 km S Santo Domingo 
de los Colorados, Provincia Pichincha, 600 m; MCZ 97596 97, 
USNM 21 1 176, Centinela, 14.1 km SE Patricia Pilar by road, Pro- 
vincia Pichincha, 570-600 m; KU 146171, 165874-77, MCZ 94864 
65, 94867, 98164 66, USNM 211175, Centro Cientifico Rio 
Palenque, 47 km S Santo Domingo de los Colorados by road, 
Provincia Pichincha, 170 220 m. 

Diagnosis. 1 ) Skin of dorsum smooth with occasional tubercles, 
that of venter areolate; no dorsolateral folds; no anal sheath; 2) 
tympanum distinct, round, its length 1/4-2/5 eye length; 3) snout 
round in dorsal view, truncate in lateral profile; canthus rostralis 
distinct; 4) interorbital space narrower than upper eyelid; low cra- 
nial crests in females; small tubercle on upper eyelid; 5) vomerine 
odontophores elevated, triangular in outline, narrowly separated; 6) 
males with vocal slits; males lack nuptial pads on thumb; 7) first 
finger shorter than second; all digits bearing broad discs on 
expanded pads, pads of fingers III IV largest, emarginate, those of 
I-II smaller, rounded apically; 8) fingers lack lateral fringes; 9) no 
ulnar tubercles; 10) small tubercles on heel, none on knee or tarsus; 
11) two metatarsal tubercles, inner oval, 4 times size of flat outer; 
low supernumerary tubercles at bases of toes II-IV; 12) toes lack 
lateral fringes; discs broader than long on weakly emarginate pads, 
pads smaller than those on fingers; 13) brown above with little 
indication of pattern (Fig. 1); venter cream with extensive brown 
reticulation; undersides of limbs brown with cream flecks; anterior 
and posterior surfaces of thighs brown with small cream Jlecks; in 
life, iris blue; 14) adults moderate-sized, males 20.8-26.8 (x = 23.8 
± 0.9, n= 16) mmSVL, females 30.6 36.9 (x = 33.8 ± 1.6, n= 7) 
mm SVL. 

Eleutherodactylus tenebrionis has no close relatives known to us 
except for an undescribed species from the upper cloud forests 
(2000-2700 m) in Provincias Imbabura and Pichincha, Ecuadpr 
(Lynch and Trueb, in press) which differs from E. tenebrionis in 
having prominent conical tubercles on the forearm, tarsus, and 
upper eyelid and a calcar on the heel. 

Description. Head as wide as or wider than body, wider than 



1979 TWO NEW SPECIES OF ELEUTHERODACTYLUS 9 

long; HW 37.0 41.4 per cent (x= 39.3, n= 23) SVL; snout round in 
dorsal view, truncate in lateral profile; E-N in males 72.0 84.8 per 
cent(x= 77.9, n= 16) eye length, in females 75.9-87.0 (x= 82.9, n 
= 7); nostrils protuberant, directed laterally; canthus rostralis dis- 
tinct, weakly concave; loreal region concave, sloping abruptly in 
males and gradually in females to lips; lips not flared (Fig. 2); upper 
eyelid 110.3-165.7 per cent (x = 125.8, n = 23) lOD, bearing 1-2 
small conicaT tubercles; adult females have low cranial crests (not 
evident in smaller frogs); lower 3/4 of tympanum distinct, upper 
edge hidden by diffuse supratympanic fold; tympanum separated 
from eye by almost its length; tympanum length in males 25.0 35.0 
per cent (.x= 29.4, n = 16) eye length, in females 29.6-38.5 (x= 32.9. 
n = 7); small postrictal tubercles present; skin on rest of head 
smooth; choanae longer than wide, not concealed by palatal shelf of 
maxillary arch; vomerine odontophores median and posterior to 
choanae, pungent, triangular in outline, bearing 5-7 teeth in a trans- 
verse row, separated on midline by a space equal to 1/3 1/2 of 
odontophore width; odontophores nearly as large as a choana; 
tongue longer than wide, its posterior border notched, posterior 1/5 
not adherent to floor of mouth; male with vocal slits. 

Skin of dorsum essentially smooth but some small, low. flat warts 
on lower back (in a few examples these warts are pungent) and a few 
tiny warts scattered on back; no dorsolateral folds; flanks become 
areolate, areolation continuing onto venter; throat smooth; discoi- 
dal folds present; no anal sheath; no ulnar tubercles; palmar tuber- 
cles bifid (outer lobe the smaller), as large as or smaller than oval 
thenar tubercle; if present, supernumerary palmar tubercles flat, 
indistinct; subarticular tubercles relatively low, round; fingers lack- 
ing lateral fringe or keel; all fingers bearing broader than long discs; 
pad smallest on I, intermediate on II, largest on III and IV; pad of 
III as large as tympanum; pads on III and IV feebly emarginate; II 
longer than I; males lack nuptial pads. 

No tubercles on knee or tarsus; 1-2 small tubercles on heel; inner 
metatarsal tubercle twice as long as wide, outer I /4 (or less) size of 
inner, flat; supernumerary plantar tubercles low, at bases of toes 
II IV; subarticular tubercles longer than wide, not conical; toes lack 
lateral fringes, bearing broad discs on expanded, feebly emarginate 
pads (toe pads smaller than those on outer fingers); heels of flexed 
hind legs overlap; shank of males 52.8 62.5 (x = 57.9, n = 16) per 
cent SVL, of females 52.2-59.7 (x = 56.1, n = 7). 



10 BREVIORA No. 457 

Brown above with diffuse dark brown or black scapular "W", 
supratympanic stripe, interorbital bar, and indefinite sacral chev- 
ron; canthal stripe, labial bars, anal triangle brown, not distinct; 
limb bars brown, nearly transverse on shanks, about as wide as 
interspaces; flanks pale brown (cream invasion of brown); venter 
cream with dense fine brown reticulation; some cream flecks on 
limbs, fewest on underside of shank; throat and breast heavily 
stippled with brown (relative to venter); anterior and posterior sur- 
faces of thighs (and groin) brown with small cream flecks. 

In life, E. tenebrionis is burnt umber with black and dull golden 
flecks above; the venter is gray with gray-brown mottling. The iris is 
gray-blue. At night they tend to be a much paler yellowish-tan 
dorsally with prominent black spots in the shoulder region. 

Measurements of Holotype('\n mm). SVL 26.4; shank 15.0; HW 
10.8; head length 10.4; upper eyelid 3.0; lOD 2.6; tympanum length 
1.4; eye length 4.4; E-N 3.3. 

Etymology. The specific epiphet is derived from the Latin, 
meaning a lover of darkness, in reference to the restriction of E. 
tenebrionis to primary forest. 

Natural History. Eleutherodactylus tenebrionis is restricted to 
primary forest, where it is most frequently found at night on low 
vegetation along stream courses. The lowland rain forests in western 
Ecuador which this species inhabits are characterized by full cano- 
pies and relatively sparse understories. Epiphytes are especially 
prominent in these wet forests, and some of the lowland forests 
appear superficially like cloud forest. Much of this region is covered 
with clouds during the dry season which encourages this lush 
growth even though annual rainfall is only moderately heavy (Dod- 
son and Gentry 1978). 

The preference of E. tenebrionis for streamside vegetation 
appears to be real and not a sampling bias. One of us (KM) has 
spent considerable time working study plots in primary forest at Rio 
Palenque located on top of a plateau and lacking any streams and 
has never encountered E. tenebrionis there. Along the small creeks 
elsewhere in the Rio Palenque forest they are encountered regularly, 
if not commonly. The specimens from Centinela likewise all came 
from along a small forest stream rather than from the primary 
ridgetop forest or the recently cleared hillside forest. At Tinalandia 



1979 TWO NEW SPECIES OF ELEUTHERODACTYLUS 1 1 

occasional individuals were found perched on low vegetation within 
2 m of the ground along a hillside about 30 m above stream level, 
but this area at 800 m elevation is more mesic than the lower 
localities. 

Discussion 

The flared lips and long snouts of £. crenunguis, E. latidiscus. E. 
muricatus, atid E. ruhicundus are in marked contrast to the condi- 
tion seen in E. lenehrionis (Fig. 2). All of these species share the 
narrow 10 D, a relatively uncommon trait among Eleutherodactylus 
(Fig. 2). Low cranial crests are present in most of these species 
(except E. muricatus) but are generally evident only in adult 
females. These traits are also exhibited by E. cruentus (Peters), a 
species sometimes confused with E. latidiscus which may be allied to 
E. crenunguis and E. latidiscus (Lynch 1976). As noted by Lynch 
(in press) only E. crenunguis, E. latidiscus. and E. ruhicundus of the 
ruhicundus assembly have emarginate digital pads. We can now add 
E. muricatus to this list but we are not convinced that these four 
species are more closely related to one another than any is to the 
other two known species of the assembly. The weakly emarginate 
digital pads of E. tenehrionis are not convincing evidence of its 
relationship with the ruhicundus assembly; its snout physiognomy 
suggests that it is allied elsewhere. 

Acknowledgements 

Lynch's fieldwork was supported by the University of Nebraska 
Research Council and Miyata's by the Barbour Fund of the 
Museum of Comparative Zoology, the Anderson and Richmond 
Funds of the Department of Biology, Harvard University, the 
Museu de Zoologia of the Universidad de Sao Paulo, and Earth- 
watch and the Center for Field Research of Belmont, Massachu- 
setts. Facilities were provided and specimens were loaned by 
William E. Duellman, Museum of Natural History, University of 
Kansas (KU), Ernest E. Williams, Museum of Comparative Zool- 
ogy, and Roy W. McDiarmid, National Museum of Natural History 
(USNM). We are grateful to Fernando Ortiz and Eugenia del Pino 
of the Universidad Catolica del Ecuador in Quito for logistic sup- 
port in the field. For permission to collect specimens we would like 
to thank Calaway Dodson and Alcides Marmol (Centro Cientifico 
Rio Palenque) and Alfredo and Tina Garzon (Tinalandia). 



12 BREVIORA No. 457 

I riERATURE CITED 

DoDSON, C. H. AND A. H. Gemrv. 1978. Flora of the Rio Palenque Science 

Center. Selbyana, 4(1-6): xxx + 628 p. 
Lynch. J. D. 1976. New species of frogs (Leptodactylidae: Eleutherodaciylus) 

from the Pacific versant of Ecuador. Occas. Pap. Mus. Nat. Hist. Univ. 

Kansas. 55:1 33. 
1977. A new frog ( Leptodactylidae: Eleiiiherodactvlus) from the Pacific 

lowlands of Ecuador. Copeia, 1977:282 284. 
In press. A new species of Eleutherodactylus from northern Ecuador 

(Amphibia: Leptodactylidae). Proc. Biol. Soc. Washington. 

In press. Taxonomic and distributional notes on poorly-known frogs 



(Eleutherodaciylus: Leptodactylidae) from the chocoan lowlands of South 

America. Herpetologica. 
Lynch, J. D. and L. Trueb. In press. A new species of Eleutherodaciylus from 

the cloud forests of western Ecuador (Leptodactylidae) Copeia. 
MiYATA, K. In press. A new species of Atelopus (Anura: Bufonidae) from the 

cloud forests of northwestern Ecuador. Breviora. 



2 



R E V I O 



IS ISS\ OOOd 469S 



LfORARY 

MAR 1 8 1985 

f^VEJagftTY 



iiseiim of C ()]ii])arative Zoology 



Cambridge. Mass. Januarv 31. 1980 



Number 458 



A NKV\ SPECIES OF ATELOPUS 

(AM RA:Bl FONIDAE) 

FROM THE CLOl D FORESTS 

OF NORTHWESTERN ECUADOR 

Keweih Miyata' 

Ahsir\( I: Atel(iiHi\ (omivi is described trom the Rio Faisanes in Pichincha 
l'ri)\ince. Ecuador. It appears to ha\e an extremeU restricted distribution and may 
be HI danger ot extinction as its liabitat is altered b\ man. The new species is most 
closei_\ to Aiclopus iiuiuUivnMs^ 

INTRODUCTION 

The western sUipc ot the Andes in northern Ecuador has a rich 
anuran fauna, much ot which has been described only within the 
past decade. Until recentK access to this area has been difficult due 
to its precipitous terrain and dense blanket of montane forest. Many 
species appear to have restricted distributions and it is not surpris- 
ing that they were missed b\ earl\ collectors. Within this forested 
zone, which extends up to at least 3000 m elevation, there are several 
distinct asemblages of frogs. Ma.ximum species diversity apparently 
occurs between 1000 and 1600 m. Many of these species are very 
restricted in distribution; few are known from outside Ecuador. My 
field work in this region revealed the presence of an apparently new 
species of frog of the genus Aie/opu.s that seems to exhibit an excep- 
tionally restricted distribution even in comparison with other ele- 
ments of this tauna. Despite extensive collecting efforts in the cloud 
forests of western Ecuador by myself and field parties from the 
Museum of Natural Historv of the University of Kansas, this new 
species remains know n from onlv two localities within a few kilome- 



'Kenneth Miyata. Museum of Comparative Zoology. Harvard University 
Cambridge. Massachusetts 02138 



BREVIORA 



No. 458 



ters of each other. Gi\en the rather precarious status of these habi- 
tats as human dexelopment encroaches, 1 take the opportunity here 
to describe this species before the populations become e.xtinct. 




Figure 1. Dorsal view ot hololype of Awlnpus coynei (MCZ 91444). 



1979 



A NEW SPECIES OF ATELOPUS 




Figure 2. Ventral vieu of holotype of Atelopus cornel (MCZ 91444). 

Alelopus coynei sp. nov. 

Holoivpe: MCZ 91444. an adult male, one of a series collected 
on the banks of the Rio Faisanes where it crosses Ecuador Highway 
28 (the road from La Palma to Quito via Chiriboga). 14.4 km from 
the junction with Highway 30 (the Aloag to Santo Domingo de los 



4 BREVIORA No. 458 

Colorados road) at La Palma, Pichincha Province. Ecuador. 1380 
m, on 1 1 July 1976 by Godfrey Guynn. Kay Marker. Steven Kaal, 
Ken Miyata, David Paul, and Harrison Weed. 

Paratvpes. Topotypes: MCZ 9144^-91449, 96775-96756. col- 
lected with the holotype; MCZ 91450, collected on 7 August 1976 by 
Jerry Coyne and Ken Miyata; MCZ 9541 1, collected on 8 January 
1978 by Lauren CarduUo. Andrea Dion. Ken Miyata. Hugh Tor- 
bert, and Lisa Schwadron; MCZ 95676. collected on 30 April 1978 
by Paul Greenfield and Ken Miyata; MCZ 96754. collected on 12 
November 1977 by Ken Miyata; USNM 211171. collected on 17 
February 1979 by Roy McDiarmid. From 4 km E Dos Rios. Pichin- 
cha Province, Ecuador. 1140 m: KU 164744. collected on 2 April 
1975 by William Duellman. 

Diai^nosis. A small Atelopus (males to 23 mm, females to 32 
mm) distinguished from all other known species by the following 
combination of characters: 1.) Hind limbs relatively long, the heels 
overlapping slightly when held parallel to femora at right angles to 
the body and reaching or just falling short of the orbit when 
adpressed. 2.) First finger almost entirely buried in a thick, rather 
fleshy webbing. 3.) Ventral pattern consisting of a sparse network of 
fine dark reticulations on a light opaque background. 

Description. Head narrower than body, somewhat longer than 
wide. Snout projecting past tip of lower jaw, rounded from above. 
Projecting snout forming fairly sharp right angle above and in front 
of nostril in lateral profile. Nostrils opening laterally about 2/3 of 
way from anterior margin of orbit to tip of snout, directly above or 
slightly behind tip of lower jaw. From above, canthi diverging 
slightly from behind nostrils to a point just anterior of orbits where 
they diverge outward more abruptly. Canthus rostralis rounded 
with slight depression in loreal region. Interorbital space wider than 
upper eyelid. Tympanum hidden. Skin on head generally smooth 
with some very sparse and fine granulation. 

Dorsum finely shagreened, dorsolateral folds absent. Venter and 
sides with numerous plate-like folds, smallest and most distinct on 
throat and neck, becoming larger and less distinct on sides and 
towards cloaca. Pair of narrow low ridges on dorsum in parotoid 
region. 

Forearm slightly thicker than upper arm. Forefeet fleshy with 
thick webbing. First finger almost buried in fleshy webbing with 
only the tip free. Other fingers basally webbed with lateral fringes 



1979 A NEW SPECIES OF ATELOPUS 5 

extending past distal subarticular tubercles. Subarticular tubercles 
indistinct; palmar tubercle prominent, oval in shape. Males with 
cornified pad on inner margin of first finger. 

Heels overlap slightly when tibiofibulae held parallel to femora at 
right angles to body. Adpressed heels reach to. or fall just short of, 
posterior margin of orbit. Tarsal fold absent. Hind feet fleshy with 
extensive thick webbing to tips of all toes except the fourth which is 
free distally." Subarticular tubercles indistinct; outer metatarsal tu- 
bercle small and rounded. 

Measurements. Mean standard length for adult males is 22.6 
mm, the single known adult female is 32.1 mm. The holotype is 22.4 
mm. Table I summarizes the measurements and ratios used by 
Peters (1973) in his review of Ecuadorian Ate/opus. 

Color in life. The dorsum of males varies from green with dark 
brown reticulations to dark brown with green spots and blotches. In 
all specimens the dorsal green becomes turquoise blue laterally. The 
single adult female (MCZ 96754) was bright green with sparse dark 
brown dorsal reticulations and no trace of turquoise blue on the 
sides. The ventral surfaces of the males were opaque white, occa- 
sionally with a yellowish wash, with a sparse network of dark brown 
or black reticulations (Fig. 2). The female was a bright opaque 
yellow ventrally with dark brown reticulations and a reddish-orange 
wash on the palms and soles. This same reddish-orange wash on the 
palms and soles was seen on one male as well (MCZ 9451 1 ). but it 
was not as prominent as on the female. The iris varies from golden 
yellow to orange-copper. 

Color in preservative. The parts of the dorsum which were green 
in life are pale lavender. The dark brown parts of the dorsal pattern 
remain brown, but are somewhat paler and have a reddish wash. 
The venter remains white in the males, but all traces of the yellow 
wash are lost. The female retains some of the yellow ventral color 
after 20 months in preservative. The ventral reticulations range from 
pale to medium brown. 

Etymology. The specific epiphet is a patronymi for Dr. Jerry 
Coyne, whose timely financial assistance helped stave off the wolves 
on several occasions and allowed me to complete the description. 

Natural History 

The Rio Faisanes is a small mountain stream flowing through a 
narrow forested canyon where it crosses Ecuador Highway 28, 14.4 



6 BREVIORA No. 458 

km from the village of La Palma on the road to Quito. The elevation 
of the stream where it crosses the road is given as 1380 m on a 
topographical map of the area prepared by the Institute Geografico 
Militar of Ecuador m 1969 (Alluriquin quadrat, CT-NIII-A3, 3893- 
111). This falls into the "bosque muy humedo Pre Montano" eco- 
logical zone (Institute Geografico Militar 1977), and the forest is 
characterized by a relatively low canopy height with an extremely 
heavy epiphyte growth. The Rio Faisanes is rarely more than 5 m in 
width and few places are more than 0.5 m deep. The bottom consists 
of pea-sized gravel and the bed sometimes flows over or against 
large rocks. The water normally runs clear and in most places the 
canopy completely overhangs the water (see below). The other 
known locality where coynei has been collected (4 km E Dos Rios, 
I 140 m) is along the Rio Orito and is located about 1.3 km NSW of 
the type locality. According to the topographical map, the elevation 
is 1280 m. The two small rivers flow into the Rio El Transito, which 
then flows into the Rio Pilaton a few kilometers to the SW. 

All of the specimens taken on 1 1 July 1976 were collected along 
the banks of the Rio Faisanes. Individuals were captured in the late 
afternoon while they were active on the rocky banks of the river 
during a light rain. At night additional specimens were collected 
sleeping on streamside vegetation, usually within 1 m of the ground 
and frequently on leaves overhanging the water. A total of 4 1 speci- 
mens was collected that day in appro.ximately 18 person-hours of 
collecting along a 250 m stretch of the river. Most of these speci- 
mens were used for biochcTiical analysis of possible skin toxins by 
Harrison Weed and were not available for this description. 

Subsequent visits to the Rio Faisanes in July and August of 1976, 
May and November of 1977, January and April of 1978, and Febru- 
ary of 1979 resulted in the collection of very few additional speci- 
mens. All of these later visits were made at night and hence no more 
active individuals were found. All of these specimens were sleeping 
on the tops of leaves within 1 m of the ground. These specimens 
were not taken along the Rio Faisanes proper, but were found on 
the banks of a small tributary creek which crosses the road about 50 
m SW of the Rio Faisanes bridge. 

Since the first visit to the type locality in July of 1976 the Rio 
Faisanes seems to have become more silted as a result of small-scale 
logging operations and the canopy has been opened to a considera- 
ble degree in places. On the first visit the river was running clear 
despite the fact that it was high from the rains. On later visits the 



Males (\=1) 


Fenwic (S=\) 


22.6 mm 


32.1 mm 


20.3 


29.2 


10.6 


15.5 


6.6 


8.7 


7.8 


10.5 


87.7 


91.0 


47.1 


48.3 


34.4 


32.7 


84.5 


82.9 


137.1 


147.6 


52.5 


53.1 



1979 A NEW SPECIES OF ATELOPUS 

Table i. Measurements and ratios of Aielnpus covnei (as in Peters 1973). 



Standard distance 
Knee-knee 
Tibofibula 
Head width 
Head length 

Knee-knee standard distance 
Tibiofibula standard distance 
Head length standard distance 
Head width head length 
Tibiofibula head length 
Tibiofibula knee-knee 

water was often cloudy and turbid even though the level was low 
and there had been no recent rains. The scarcity of frogs after the 
first visit may be due partly to the increase in human activity along 
the stream which may have rendered it unsuitable for larval devel- 
opment. An increase in suspended silt in the water due to this 
human encroachment may cause the scraping of the algae off the 
surface of the rocks before the tadpoles can get to it. 

Only a single female coynei has been found out of a total of 47 
specimens (including the ones used for biochemical work). This is 
not necessarily indicative of a skewed sex ratio. All of the collections 
have been made along stream banks and McDiarmid (197!) has 
suggested that male Atelopus may spend considerably more time 
along the creeks than females. The large number of males taken on 
the first visit to the type locality is indicative of a breeding concen- 
tration and it is likely that the females had not yet arrived. Since 
then, individuals have been found in low concentrations and the 
bulk of the population may have dispersed into the forest. The 
apparent absence of coynei on recent visits may also be due in part 
to an absence of breeding concentrations. 

Other anurans collected syntopically and synchronously with 
active Atelopus coynei include Atelopus longirostris, Bufo chancha- 
nensis, Eleutherodactylus achat inus. E. necerus, and Colostethus sp. 

Comparisons and Discussion 

The relatively long hind limbs and the extensively webbed thumbs 
distingush A. coynei from all but a few species o^ Atelopus. Of these 
species, the hind limbs oiA. elegans, A. longibranchius, A.palmatus, 



8 BREVIORA No. 458 

and A. rui^'u/osu.s are very long and the adpressed heel reaches 
beyond the anterior margin of the orbit, rather than just reaching or 
falling short of the posterior margin. In A. crucii^er from Venezuela 
and A. flavescens from French Guiana the adpressed heels reach to 
near the posterior margin of the orbit, but they lack webbing 
between the second and third fingers and are larger than ,-1. covnei 
(male A. cnuif^er to 29 mm, male A. flavescens to 32 mm). 

Aielopus covnei most closely resembles A. mincioensls. The most 
striking difference between these two species is ventral pattern, with 
A. mindoensis lacking the reticulations which are characteristic of 
A. cuynei. There are a number of other differences as well (Table 2) 
and there is little doubt that the taxa are distinct. 

The description of a new species oi' A fe/o/yus based on external 
morphology from two neighboring populations may seem unwise 
given the known degree of variation in such characters in some 
members of the genus (Peters 1973, Savage 1972). The close similar- 
ity of A. covnei to A. mindoensis may make it seem particularly 
suspect. Although A. covnei and A. mindoensis are almost certainly 
closely related, the differences outlined in Table 2 are consistent 
enough to warrant their recognition. 

Although color pattern can be quite variable both within and 
between populations of some species oi Ate/opus, the ventral patt- 
ern of A. covnei is unique and allows the immediate recognition of 

Table 2. Aielopus coviivi and mindoensis compared 
A TELOPUS CO YNEI A TIA.DPI S \IIMH)i:SSIS 

Venter white to \ellou with brown Venter reddish brown with \ellow spots. 

reticulations. 

■ ,.,,, ,, Ventral skin smooth or with small tolds 

Ventral skin with numerous small 

1114 11 .1 "n throat, 

scale-like tolds anteru)rl\. 

., . I I , ,■ Ventral skin translucent to transparent. 

Ventral skin completely opaque. ' 

r . ■ ui .u ' 1. 1 ■ I-SJi^ clearU \isible through skin in 

hggs not visible through skin in one ^*^ ^ 

■ , ,• I , gravid temales. 

gravid female known. "^ 

V, . , I ,, , , Small tubercles and enamelled pustules 

iNo tubercles or enamelled pustules ()n ' 

, , , on back and sides 

back or sides. 

t: . J 1 . i_ , Snout more pointed Irom above and 

Snout more rc)unded trom abo\e and 

, , , , coming to more rounded angle trom side 

coming to sharp angle Irom side. ^ ^ 

V. , . ., . , , Adult males with loose skin on throat. 

^o CMdence ol throat pt)uch in adult 

males. Smaller si/e; adult males average 

Larger size; adult males average 22.6 lX.7mm(N .^7), adult temales 24.9 mm 

mm. adult lemale .''2.1 mm. (N 22). 



1979 A NEW SPECIES OF ATELOPUS 9 

this species. 1 have examined the holdings oi Aielopus in the MCZ 
collection and made a literature survey of all of the 40 presently 
recognized species (those listed by McDiarmid 1971 and subse- 
quently described species; descriptions consulted in lieu of speci- 
mens include Bokermann 1962. Boulenger 1902. 1903, Cochran and 
Coin 1970, Donoso-Barros 1969, McDiarmid 1973, Noble 1921, 
Peters, 1973, Ruiz-Carranza and Hernandez-Camacho 1978, Sav- 
age 1972) and the only species that exhibit a reticulate ventral patt- 
ern are A. chlriquiensis, A. loni^lrosiris, A. pachydermus, A. 
pulvher, A. tricolor, and A. varius. In each of these species the 
reticulate pattern, when present (some species are variable in this 
character), is very bold and consists of thick lines mixed with 
blotches and spots. In contrast, the ventral reticulations of ,4. coynei 
are thin and sparsely distributed. Although the degree of this ventral 
marbling in A. coynei is variable, it is immediately recognizable in 
all of the specimens I have examined. 

Although the 40 known species of A telopus make it a relatively 
large assemblage of frogs, all of which are Neotropical, the species 
are distributed with remarkably little geographical overlap. Exten- 
sive sympatry between species of A telopus is unusual, a situation 
which is in striking contrast to that seen in other Neotropical anuran 
genera of similar diversity. The rather specialized breeding require- 
ments of Atelopus may somehow inhibit overlap. In areas where two 
species of Atelopus occur in sympatry they are usually of quite dif- 
ferent habitus. To date, A. coynei has been collected with the larger 
and more gracile A. longirostris but has not been found with the 
similar. 4. mindoensis. Although .4. mindoensis is known from eleva- 
tions between 20 m and 2100 m. the lower records are from the 
drainage of the Rio Cachabi in Esmeraldas Province. On the Rio 
Toachi drainage in Pichincha Province A. mindoensis appears to be 
restricted to elevations above 1 500 m and does not come into contact 
with either A. coynei or A. longirostris. Since A. mindoensis and A. 
longirostris do come into contact in the Rio Guayllabamaba drain- 
age just to the north of the Toachi. it appears that A. coynei may be 
limited to the relictual populations described here. 

ACKNOWl EDGEMENTS 

The field work resulting in the discovery of Atelopus coynei was 
funded by the Richmond and Anderson Funds of the Department 



10 BREVIORA No. 458 

of Biology. Harvard University, the Barbour Fund of the Museum 
of Comparative Zoology, The Museu de Zoologia. Universidade de 
Sao Paulo, Brasil, and by Earthwatch and the Center for Field 
Research of Belmont. Massachusetts. For assistance and compan- 
ionship in the field 1 would like to thank J. Bell. L. Burnham. L. 
Cardullo. L. Chanler. J. Coyne. A. Dion. P. Greenfield. S. Gross. 
G. and R. Guynn. M. Harker. S. Kaal. D. Low. R. McDiarmid. W. 
Morrow. D. Paul. E. Schupp. L. Schwadron. and G. Vigle. I would 
also like to thank W. E. Duellman (KU). R. I. Crombie and W. R. 
Heyer (USNM) for the loan of material. E. E. Williams read and 
commented upon the manuscript. 

I I I F R \1 1 Kl ( 1 lEl) 

BoKiRM'WN. W. C. A. 1962. LIna nueva espccie de Awlopus del nordeste de 

Brasil (Amphibia, Salientia, Brachycephalidae). Neotropica. 8(26):42 44. 
Boi I \ Mil K. G. A. 1902. Descriptions of new batrachians and reptiles from the 

Andes of Peru and Bolivia. Ann. Mag. Nat. Hist., ser. 7, 10(59);394 402. 
1902. Descriptions of new batrachians in the British Museum. Ann. 

Mag. Nat. Hist., ser. 7, 12(71 ):552 557. 
Cochran, D. M. wd C. .1. GoiN. 1970. Frogs of Colombia. U.S. Nat. Mus. 

Bull.. 288:1-655. 
D()N()S()-B.\RR()s, R. 1969. Una nue\a especie de Atelopodidae de Bolivia 

(Anura). Physis, 2H(77):327 3.^0. 
Instituto Geograhco Mil mar. 1977. Atlas Geografico de la Republica del 

Ecuador. Quito, Ecuador. 82 pp. 
McDiarmi[5, R. W. 1971. Comparative morphology and e\olution of frogs of 

the Neotropical genera Aielopus. Dendrophrvniscus. Melanophryniscus, and 

Oreophrvnella. Los Angeles County Mus. Nat. Hist., Science Bull. 12:1-66. 
1973. .\ new species of Aielopus (Anura, Bufonidae) from northeastern 

South America. Los Angeles County Mus. Nat. Hist., Contr. Sci., 240:1-12. 
NoBi E, G. K. 1921. Five new species of Salientia from South America. Amer. 

Mus. Nat. Hist. Novitates, 29:1 7. 
Peters, J. A. 1973. The frog genus Aielopus in Ecuador (Anura: Bufonidae). 

Smiths. Contr. Zool.. 145:1 49. 
Ritz-CARRANZ \. P. M. \M) J. L H ER N \\ DEZ-C A M AC HO. 1978. Una nueva 

especie colombiana de Aielopus (Amphibia: Bufonidae). Caldasia, 12(57): 

181 197. 
SavaciI . J. M. 1972. The harlequin frogs, genus Aielopus. of Costa Rica and 

western Panama. Herpetologica. 28(2):77 94. 



fur 



Mb'3. CCMP. ZOOL 
LIBRARY 



B R E V I O WX 

L-MiVERfilTY 

Miiseiim of Comparative Zoology 



us ISSN 0006 9698 



Cambridge, Mass. January 31, 1980 Number 459 

A NEW SPECIES OF DENDROBATES 

(ANURA: DENDROBATIDAE) 

FROM THE LOWLAND RAIN FORESTS 

OF WESTERN ECUADOR 

Gregory O. Vigle' and Kenneth Miyata' 

Abstract: Dendrohates erythromos sp. nov. is described from the lowlands of 
western Ecuador. The new species is tentatively assigned to the genus Dendrohates 
on morphological grounds and appears to be most closely allied to the Colombian D. 
m^eri. This species may have specific microhabitat requirements which result in a 
spotty distribution. 

INTRODUCTION 

In January of 1978 several specimens of an apparently unde- 
scribed species of dendrobatid frog were collected at the Centro 
Cientifico Rio Palenque in the Pacific lowlands of Ecuador. Prepa- 
ration of an account of the herpetofauna of the region requires that 
a name be assigned to this population, and here we describe the 
species on the basis of material in the Museum of Comparative 
Zoology (MCZ) and the National Museum of Natural History 
(USNM). 

The generic partitioning of dendrobatid frogs is in a state of flux 
and the characters which have traditionally been used to define the 
genera are being supplanted by new ones. Following the criteria of 
Savage (1968) and Silverstone (1976) this new species would be 
asigned to Phyllobates. However, this genus has recently been 
defined as a monophyletic group (in the cladistic sense) on the basis 
of a biochemical synapomorphy by Myers et al. (1978) and consists 
of only four of the species recognized by Silverstone plus one newly 



'Gregory O. Vigle, Biology Department, Eckerd College, St. Petersburgh, Florida 
33733. 

^Kenneth Miyata. Museum of Comparative Zoology, ' Harvard University, 
Cambridge, Massachusetts 02138. 



BREVIORA 



No. 459 



described species. All of the remaining species of Phyllohates recog- 
nized by Silverstone (the femoralis, picius, and irivitiatus groups) 
were reassigned to Dendrohates. The new species most closely 
resembles those in Silverstone's pictus group on the basis of size, 
coloration, and the absence of toe webbing and we tentatively assign 
it to the genus Dendrohates. 

Dendrohates erythromos sp. nav. 

Holotype: MCZ 96384, an adult female collected at the Centro 
Cientifico Rio Palenque, 47 km S of Santo Domingo de los Colora- 
dos, Provincia Pichincha, Ecuador, 170 m, on 17 January 1978 by 
Gregory O. Vigle. 

Paratypes. All are topotypes: MCZ 96381-83, 96385, 94896 (lot 
of 3 tadpoles from back of 96385), USNM 211169-70. 

Diagnosis. A dendrobatid frog with the following combination 
of characters: 1.) Teeth present. 2.) Omosternum present. 3). First 




Figure I. Paratype of DeiiJrohaltw erythromos (MCZ 96381). The slightly paler 
area on the dorsal surface of the forearm is the bright orange flash mark. 



1979 A NEW SPECIES OF DENDROBATES 3 

finger longer than second. 4.) Toe webbing absent. 5.) Muscle tissue 
flecked with black pigment. 6.) In life, the dorsum is dark brown 
and the venter is black with pale blue marbling. 7.) A bright reddish- 
orange flash mark on the posterodorsal surface of the upper arm in 
life. 8.) Size medium, to 23.5 mm snout-vent length (SVL). 

The above combination of morphological characters, combined 
with the lack of any dorsal or dorsolateral striping and the bright 
orange flasK marks distinguish Dendrohates erythromos from all 
other known dendrobatid frogs. 

Description. Snout-vent length to 23.4 mm; four adult females 
21 .5-23.4 mm; two adult males 20.4-22.6 mm; one immature female 
18.6 mm. 

Skin smooth to very finely pitted dorsally, slightly rugose laterally 
and on posterior surface of thighs; smooth on venter and limbs. 
Head as wide as body or slightly wider, widest at jaw articulations. 
Eyes prominent, diameter about one and one-half times the length 
of the snout; width of upper eyelid less than interorbital width. 
Snout short, truncate in dorsal and lateral aspect. Canthus rostralis 
rounded; loreal region flat to slightly concave. Tympanum round, 
about one-half diameter of orbit; posterodorsal portion concealed. 

Relative lengths of appressed fingers 3>4>2>1, each having a 
distinct disc expanded about 1 .5 times the width of the distal end of 
the adjacent phalanx. First finger longer than second if not ap- 
pressed. A large outer metacarpal tubercle and a smaller, less promi- 
nent inner metacarpal tubercle at the base of the palm. One slightly 
prominent subarticular tubercle on fingers one and two and two 
slightly less prominent tubercles on fingers three and four; all are 
low with rounded surfaces. Relative lengths of appressed toes 
4>5>3>2>l. Toes have distinct discs approximately 1.5 times the 
width of the distal end of the adjacent phalanx. A small elongated 
outer metatarsal tubercle and a smaller, round inner metatarsal 
tubercle at the base of the foot. Indistinct subarticular tubercles on 
toes; one on toes one and two, two on toes three and five, and three 
on toe four. A slightly prominent curved tarsal fold on the inner side 
with a weakly developed tubercle at the proximal end. Palms and 
soles flattened on bottom and slightly fleshy. Digits lacking lateral 
fringes (or with very narrow ones on toes), webbing, and supernu- 
merarv tubercles. 



4 BREVIORA No. 459 

Palatine bones absent. Maxillary and premaxillary teeth present. 
Omosternum present. Muscle tissue flecked with black pigment. 
Paired vocal slits in adult males; single subgular vocal sac. 

Measurements of holotype. The undissected holotype is the 
largest individual in the type series. The following measurements are 
to the nearest 0.1 mm as measured on dial calipers. Snout to vent 
23.4; tibia from heel to fold of skin on knee 9.6; greatest width of 
body 8.4; greatest width of head 8.5; head length from tip of snout 
to angle of jaws 7.6; length of snout from anterior edge of orbit to 
tip of snout 2.4; diameter of orbit 3.6; diameter of tympanum 1.9; 
length from proximal edge of large palmar tubercle to tip of third 
finger 5.7; width of disc of third finger 0.6; width of distal end of 
adjacent phalanx 0.4. 

Color in Life. Dorsum dark brown, appearing almost black in 
some specimens. The sides are black or very dark gray. Bright 
reddish-orange flash marks on the posterodorsal surfaces of the 
upper arms, extending from the axilla to the elbow and sometimes 
slightly beyond to the forearm. Anterior surface of arms and hands 
dark brown. The dorsal surfaces of the hind limbs are banded dark 
brown and black. The throat and ventral surface of the body are 
black with pale blue marbling which extends slightly up the sides. 
The iris is brown. 

Color in Preservative. The dorsal and lateral surfaces are black 
to very dark brown. The banding on the dorsal surface of the hind 
limbs is obscure. The flash marks on the upper arm are white. The 
ventral surface is black with dull gray marbling. 

Tadpoles. The following description is based on the paratypic 
lot of tadpoles (MCZ 94896) taken from the back of an adult male 
(MCZ 96385) on 18 January 1978. Only two of the three tadpoles 
are in good condition and these are the ones used for the descrip- 
tion. 

The tadpoles are in stage 25 (Gosner 1963) and measure 9.04 9.60 
mm total length (measurements taken with ocular micrometer). The 
head and body measure 3.20-3.68 mm and constitute 35-38% of the 
total length. The body is somewhat flattened with the width 
about 1.25 times the depth of the body. The eyes and nostrils are 
directed dorsolaterally. The spiracle is sinistral and the anus is 
dextral. 



1979 A NEW SPECIES OF DENDROBATES 5 

The tail has a relatively low fin and is as deep as the body or 
slightly deeper posteriorly; the dorsal fin extends forward slightly 
past the level of the anus. 

There is one row of pigmented teeth on the anterior labium. The 
beak is keratinized, the upper jaw slightly convex anteriorly and the 
lower jaw in a strong "V" shape. The oral disc is laterally indented. 
There is a single row of small, blunt papillae which occur continu- 
ously along'the lateral and posterior edges of the oral disc and are 
absent from the anterior edge. 

The ground color is pale buff, heavily speckled with brown on the 
dorsal surface of the body. The venter is lightly pigmented ante- 
riorly and heavily speckled posteriorly with brown. The brown 
speckling on the tail is restricted to the dorsal aspect and the fins are 
unmarked. 

Etymology. The specific epiphet is derived from the Greek ery- 
thros. meaning red, and omos. meaning upper arm, in reference to 
the distinctive reddish-orange flash marks on the posterodorsal sur- 
face of the upper arm. 

Natural History. Dendrobates erythromos is known only from 
the Centro Cientifico Rio Palenque. The forest there is considered 
to be tropical wet forest in the sense of Holdrige (1967) although 
available clmatic data do not seem to match this physiognomy 
(Dodson and Gentry 1978). The frogs are apparently restricted to a 
single creek system within this patch of forest. All of the specimens 
have come from along the banks of Lodo and Sherd Creeks above 
trail 3a (see Dodson and Gentry 1978:ix for map of station). The 
forest along these creeks has a somewhat broken canopy with signif- 
icant amounts of bamboo and other second-growth vegetation 
mixed in with the tall trees. The relief is rather steep and much of the 
openness of the canopy appears to be due to tree falls, although 
there has been some cutting in this area. The creeks flow over rocky 
or sandy beds and parts of them become dry during the end of the 
dry season. Dendrobates erythromos has been taken along these 
stream beds in January, when water levels were still low prior to the 
onset of the rainy season. Several specimens were collected along a 
dry portion of the bed of Lodo Creek although the majority were 
taken near flowing water. The frogs were never observed in the 
water but came from areas near the stream with good deposits of 
leaf litter and fairly heavy understory vegetation. They seemed to 



6 BREVIORA No. 459 

prefer those areas with relatively closed canopies, avoiding the 
numerous light gaps along the streams. 

All of the frogs of this species were taken during the day as they 
were active on the surface of the litter and ground. They were quite 
wary and difficult to approach and when pursued they sought shel- 
ter under the litter and in dense thickets of understory vegetation. 
Calls tentatively associated with this species were soft repetitive 
chirps. 

Three other species of dendrobatids are known from the Centro 
Cientifico Rio Palenque. They are Dendrobates espinosai, D. histri- 
onicus, and Colostethus infraguttatus. Only the much smaller D. 
espinosai has been found syntopically with d. erythromos, although 
D. histrionicus is common in the adjacent forest away from the 
creeks. 

Discussion 

Dendrohates erythromos most closely resembles D. ingeri, a spe- 
cies known only from a single locality in Amazonian Colombia. The 
two species are very similar morphologically, both sharing maxil- 
lary and premaxillary teeth and a light spot in the axilla and upper 
arm and both lacking lateral stripes. The dorsum of D. ingeri is very 
granular and it has a light calf spot and is easily distinguished from 
D. erythromos. The two species are probably closely allied; Silver- 
stone (1976) placed D. ingeri closest to D. pictus within hx'i pictus 
group. Although D. pictus does have lateral stripes and some speci- 
mens lack premaxillary and maxillary teeth, this arrangement does 
seem to have validity. The tadpoles of D. erythromos resemble those 
of D. pictus rather closely; those of D. ingeri are unknown. A fourth 
species, D. parvidus, was also associated with D. pictus by Silver- 
stone, but it seems to be somewhat more distantly allied with the 
other three species. Silverstone believed that his/?/c7w5 group might 
not be a natural group and recognized three different assemblages 
within it. The three species he associated with D. pictus. along with 
D. erythromos, do seem to constitute a group of similar organisms, 
but further information regarding calls and skin toxins will be 
necessary before the relationships can be worked out. 

Acknowledgements 

We thank the following for financial support of our fieldwork: 
Museu de Zoologia, Universidade de Sao Paulo, Brasil; the Barbour 



1979 A NEW SPECIES OF DENDROBATES 7 

Fund of the Museum of Comparative Zoology; and Earthwatch and 
the Center for Field Research of Belmont, Massachusetts. For help 
in the field we thank Bruce Jayne, Roy McDiarmid, Eugene 
Schupp, and participants in the Earthwatch programs; for permi- 
sion to collect we thank Calaway Dodson and Alcides Marmol. We 
are grateful to Joseph Kronbichler, John Lynch. Charles Myers, 
Jose Rosado, Susan Rhodin, and Ernest Williams for assistance of 
varied sorts' 



LITERATURE CITED 

Dodson. C. H. and A. H. Gentry. 1978. Flora of the Rio Palenque Science 
Center. Selbyana. 4( 1 -6):I -628+.\x.\. 

GosNER, K. L. 1963. A simplified table for staging anuran embryos and larvae 
with notes on identification. Herpetologica, 16:183 190. 

HoLDRiDGE, L. R. 1967. Life Zone Ecology. Tropical Science Center. San Jose, 
Costa Rica. 206 pp. 

M'lERS. C. W., J. W. Dai V. and B. Mai kin. 1978. A dangerously toxic new 
frog (Phyllohates) used by Embera Indians of western Colombia, with dis- 
cussion of blowgun fabrication and dart poisoning. Bull. Amer. Mus. Nat. 
Hist., 161:307 366. 

Savage, J. M. 1968. The dendrobatid frogs of Central America. Copeia. 1968: 
745-776. 

SiLVERSTONE, P. A. 1976. A revision of the poison-arrow frogs of the genus 
Phyllohates Bibron in Sagra (Family Dendrobatidae). Nat. Hist. Mus. Los 
Angeles Co.. Sci. Bull., 27:1-53. 



LIHRARY 

MAR 1 8 1985 



B R E V I d'"F°A 

Museum of Comparative Zoology 

us ISSN 0006 9698 

Cambridge, Mass. June 30, 1980 Number 460 



ON THE EVOLUTION OF THE JAW ADDUCTOR 
MUSCULATURE IN PRIMITIVE GNATHOSTOME FISHES 

George V. Lauder, Jr.' 



Abstract. The primitive condition of the adductor mandibulae musculature in 
gnathostomes is a large unsubdivided, fan-shaped muscle mass originating from the 
palatoquadrate and inserting on the lateral aspect of the mandible. Unspecialized 
suborbital fibers extend posteriorly from the palatoquadrate to insert laterally on the 
lower jaw, and an intramandibular adductor division is absent. The Actinopterygii, 
Actinistia, Dipnoi, and Choanata primitively possess three distinct components of 
the adductor mandibulae: a suborbital division, a medial division, and a posterolat- 
eral division, as well as medial intramandibular adductor fibers. The suborbital 
division of the adductor mandibulae has been lost in teleosts and, independently, in 
coelacanths and living lungfishes. Devonian lungfishes and early choa nates possessed 
both the suborbital and intramandibular adductor muscle components. 

INTRODUCTION 

Towards the end of the nineteenth and in the first half of the 
twentieth century, a large number of investigators were concerned 
with the homologies of the jaw musculature between the different 
vertebrate classes (AlUs, 1897, 1917, 1923; Edgeworth, 1935; Kes- 
teven, 1942, 1943, 1944; Lightoller, 1939; Luther, 1913; Souche, 
1932; Vetter, 1874). The musculature of the mandibular arch was 
the subject of special attention and several attempts were made to 
precisely link each branchial arch muscle with its serial homologue 
on the mandibular arch [see Lightoller (1939) for an example of this 
procedure carried to an extreme]. In recent years, little work has 
been done on the evolution of the musculature in primitive gnathos- 
tomes, despite the discovery of significant new fossil material that 
allows more accurate reconstruction of the musculature in extinct 
taxa. 



'The Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138 



2 BREVIORA No. 460 

Lauder (1980) has proposed that the primitive condition of the 
jaw musculature in the Actinopterygii (ray-finned fishes) is a tripar- 
tite division into distinct components: an anterior or suborbital 
division, a medial division, and a posterolateral division. In this 
paper, the consideration of adductor muscle components, including 
the intramandibular division, is extended to other primitive gna- 
thostome taxa to elucidate the evolution and homologies of the 
gnathostome jaw adductor musculature. 



COMPARATIVE ANATOMY 

Elasmohranchiomorphi 

The adductor musculature in the primitive living selachian Chla- 
mydoselachus anguineus has been described by Allis (1923) and 
Smith (1937). Based on their descriptions and on the comparative 
anatomy of the adductor mandibulae in other living sharks, the 
adductor musculature in the primitive Devonian selachian Clado- 
dus has been reconstructed (Fig. lA). 

In living selachians, the adductor musculature is divided into two 
main components. The adductor mandibulae proper [= quadrato- 
mandibularis of Kesteven (1942) and Lightoller (1939)] is a single 
unsubdivided muscle mass having a fan-shaped origin on the palato- 
quadrate. The fibers converge to a tendinous inscription before fan- 
ning out ventrally to insert on the lower jaw. The adductor insertion 
is on the lateral aspect of the mandible and no intramandibularis 
division is present. 

A suborbitalis muscle [= pterygoideus of Kesteven (1942), adduc- 
tor (i of Vetter (1874), levator labii superioris of Allis (1923), preor- 
bitalis of Luther (1913)] originates tendinously from the 
"ectethmoidal process" (Allis, 1923) of the cranium anteriorly and 
inserts onto the upper labial cartilage and the skin of the lip. In 
Cladodus this muscle is represented as distinct suborbital fibers of 
the adductor mandibulae originating from the palatoquadrate (Fig. 
lA: SO). Some fibers may also have originated from the nasal 
capsule, as in living sharks, and inserted onto the labial cartilages, 
when present. 



1979 



EVOLUTION OF JAW MUSCULATURE 



A 



SO— 





MD 



Figure 1. A. Reconstruction of the adductor mandibuiae musculature in a 
primitive shark (Cladodus). Osteologicai features after Gross (1938). B. Reconstruc- 
tion of adductor mandibuiae musculature in an acanthodian (Acanthodes). 
Osteologicai features after Jarvik (1977) and Miles (1973). Abbreviations: AM, 
adductor mandibuiae; MD, mandible; PQ, palatoquadrate; SO, suborbital fibers of 
the adductor mandibuiae (separated into a distinct muscle in Cladodus). 



4 BREVIORA No. 460 

In many living sharks, but not in Chlamydoselachus. a small 
superficial muscle overlies part of the adductor mandibulae. This 
muscle, the adductor mandibulae superficialis of Edgeworth (1935), 
adductor y of Smith (1937), Vetter (1874), and Allis (1923), origi- 
nates from the skin overlying the adductor mandibulae and inserts 
onto the posterior aspect of the palatoquadrate and mandible. 
There is currently no evidence to indicate whether the lack of this 
muscle in Chlamydoselachus is a specialization or if the primitive 
elasmobranchiomorph condition (absence of this muscle) has simply 
been retained. This muscle represents a small superficial component 
of the main adductor muscle, and I have reconstructed Cladodus 
(Fig. 1) with an unsubdivided adductor. 



Acanthodii 

The structure of the adductor mandibulae in acanthodians 
appears to have been very similar to that hypothesized to be primi- 
tive for elasmobranchiomorphs ( Fig. 1 ). The large adductor mandib- 
ulae had a fan-shaped origin from the palatoquadrate and a lateral 
insertion on the mandible. An intramandibular division is hypothe- 
sized to have been absent since the "adductor fossa" is located on 
the lateral aspect of the mandible and the medial surface is convex 
in shape (Jarvik, 1977; Miles, 1973). 

The adductor mandibulae is proposed to have extended ante- 
riorly ventral to the orbit (Fig. IB: SO); the suborbital fibers origi- 
nating from the palatoquadrate and extending posteriorly to join 
the other adductor fibers at a tendinous inscription before inserting 
on the mandible. 



Actinopterygii 

In ray-finned fishes, the adductor mandibulae complex may be 
divided into three separate divisions each inserting on the lower jaw, 
and an intramandibular adductor component (Fig. 2; Lauder, 
1980). Suborbital adductor fibers are present in Amia, Lepisosteus, 
and Polyodon. In Amia, these suborbital fibers have been called the 
levator maxillae superioris by Allis (1897) while the homologous 
adductor component in Lepisosteus is the palatomandibularis minor 
and major. Polyodon possesses a large separate anterior adductor 



1979 EVOLUTION OF JAW MUSCULATURE 5 

division originating on the palatoquadrate (Danforth, 1913). The 
suborbitalis adductor component has been lost in all teleosts (Fig. 2: 
character 6; also see Winterbottom, 1974). 

In palaeoniscoid actinopterygians, the suborbital fibers were 
enclosed in the anterior part of the maxillary — palatoquadrate 
chamber (Schaeffer and Rosen, 1961) and extended posteroven- 
trally to insert in the mandibular fossa. 

The medial adductor division represents one of two components 
of the primitive unsubdivided postorbital adductor fibers. Homolo- 
gies of the medial division in actinopterygians are given in Lauder 
(1980). The medial adductor fibers in Polypterus are called the "pte- 
rygoideus" and "temporalis" muscles by Allis(1922). The "pterygoi- 
deus" of Allis (1922) is not homologous to the "pterygoideus" of 
Kesteven (1942), which represents the suborbital division. 

A distinct posterolateral adductor component is also hypothes- 
ized to be primitive for actinopterygians (Lauder, 1980). This mus- 
cle represents the superficial lateral adductor fibers which run 
dorsoventrally from the palatoquadrate to insert in the adductor 
fossa. 

An intramandibular adductor division is present in most teleosts 
and in Amia, Polypterus, Polyodon, and palaeoniscoids (as inferred 
by the presence of an adductor fossa). The presence of intramandib- 
ular fibers is thus proposed to be a primitive actinopterygian charac- 
ter (Fig. 2: character 4). 

Actinistia 

Millot and Anthony (1958) describe three adductor muscles in 
Latimeria. A superficial adductor arises posteroinferiorly from the 
quadrate and inserts on the mandible in the mandibular fossa. This 
muscle appears to represent the posterolateral adductor division. 

The two remaining adductors arise from the ethmosphenoid por- 
tion of the cranium posterior to the orbit. Millot and Anthony 
(1958) note that these muscles appear to correspond to the "ptery- 
goideus" and "temporalis" adductors of Polypterus, and thus are 
both comparable to the medial adductor complex in primitive acti- 
nopterygians. No anterior (suborbital) division is present in Latime- 
ria; all jaw adductors extend dorsoventrally behind the orbit. 

A large intramandibular adductor occurs in Latimeria (Millot 
and Anthony, 1958: Fig. 19). 



BREVIORA 



No. 460 



GNATHOSTOMATA 




Figure 2. The distribution of various character states of the adductor mandibulae 
musculature in gnathostomes superimposed on one hypothesis of gnathostome 
phylogeny. For other characters supporting this phylogeny see Liem and Lauder (in 
press). Character 1: fan-shaped adductor mandibulae with unspecialized suborbital 
and postorbital fibers and a lateral insertion on the mandible; 2: suborbitalis muscle 
differentiated from suborbital fibers; 3: medial insertion of the adductor mandibulae 
on the mandible; 4: intramandibular adductor division present; 5: postorbital 
adductor component divided into medial and posterolateral divisions; 6: loss of 
suborbital adductor component; 7: specialization of suborbital fibers into the levator 
maxillae superioris 3 and 4 muscles (see AUis, 1897); 8: loss of the suborbital 
adductor component; 9: loss of suborbital component in living forms. 



Dipnoi 

While extant dipnoans lack both intramandibular and suborbital 
adductor divisions, Devonian lungfishes appear to have possessed 
both. A space between the infraorbital bones laterally and the pala- 
toquadrate medially may have housed the anterior adductor (subor- 
bital) division, and an adductor fossa for the intramandibular 
adductor is found on the medial surface of the lower jaw (see Miles, 
1977). 



1979 EVOLUTION OF JAW MUSCULATURE 7 

The main adductor mass in living lungfishes is divisible into pos- 
terolateral and medial components. The adductor mandibulae pos- 
terior (Edgeworth, 1935; = retractor anguli oris of Luther, 1913), 
corresponds to the posterolateral adductor division, and lies lateral 
to the ramus maxillaris and mandibularis V. The adductor mandib- 
ulae anterior (Edgeworth, 1935; = temporalis of Luther, 1913) is 
located anteromedially to the posterior adductor. In adult lung- 
fishes the adductor mandibulae anterior is a large muscle with a 
broad postorbital origin from the skull. In larval lungfishes (Edge- 
worth, 1935), the anterior adductor is nearly equal in mass to the 
posterior adductor and lies medial to it, and lateral to the trabeculae. 

Choanata 

Olson (1961) and Thomson (1967) both provide partial recon- 
structions of the adductor mandibulae complex in primitive cho- 
anates. A suborbital component of the adductor mandibulae was 
located anteriorly and presumably had its origin from the palato- 
quadrate. This division was housed in a chamber formed between 
the palatoquadrate medially and the dermal skull bones laterally. 
Postorbitally, the lateral wall of the suborbital adductor chamber 
was formed by the postorbital and jugal bones; while anteriorly the 
prefrontal, jugal, and maxilla enclosed the suborbital adductor. The 
posterolateral and medial adductor divisions both appear to have 
been present (Thomson, 1967) and thus the configuration of the 
adductor mandibulae in primitive choanates closely resembles that 
of palaeoniscoid actinopterygians. An intramandibular adductor di- 
vision is inferred to have been present by the occurrence of an 
adductor fossa in the lower jaw (see Romer, 1966). 

DISCUSSION 

In figure 2, the various configurations of the adductor mandibu- 
lae complex are superimposed on one hypothesis of gnathostome 
phylogeny. Other characters supporting this hypothesis may be 
found in Liem and Lauder (in press). The primitive configuration of 
the adductor mandibulae in gnathostomes is hypothesized to be a 
single unsubdivided fan-shaped muscle mass with unspecialized sub- 
orbital fibers originating anteriorly from the palatoquadrate (Fig. 2: 
character 1). Elasmobranchiomorphs retain the fan-shaped adduc- 
tor configuration but possess specialized suborbital fibers (Fig. 2: 



8 BREVIORA No. 460 

character 2); these fibers originate, in most cases, from the nasal 
capsule. A fan-shaped adductor muscle is also hypothesized for 
acanthodians (Fig. IB), and the main postorbital adductor mass is 
reconstructed as being unsubdivided. 

One notable aspect of the adductor mandibulae in elasmo- 
branchiomorphs and acanthodians is its lateral insertion on the 
lower jaw. This condition contrasts sharply with the medial 
adductor insertion in actinopterygians, actinists, dipnoans, and 
choanates (Fig. 2: character 3). The medial insertion correlates with 
the presence of an intramandibularis adductor division (Fig. 2: 
character 4) and the occurrence of an adductor fossa on the medial 
surface of the lower jaw, in contrast to the lateral fossa of elasmo- 
branchiomorphs and acanthodians. 

In the Actinopterygii, Actinistia, Dipnoi, and Choanata, an 
hypothesis of a tripartite division of the adductor mandibulae into 
suborbital, medial, and posterolateral components best fits the 
embryological data and morphological relationships of the mus- 
culature in the adult (Fig. 2: character 5). Although among living 
taxa, Polypterus lacks the suborbital adductor fibers and Lepiso- 
steus the intramandibular division, other primitive actinopterygians 
(including palaeoniscoids) possess or show evidence of three 
extramandibular adductor components. 

Teleost fishes have lost the suborbital adductor division (Fig. 2: 
character 6) but halecomorphs (Amia) retain specialized derivatives 
of the suborbital musculature, the levator maxillae superioris 3 and 
4 of Allis (1897) (Fig. 2: character 7). The cylindrically-shaped 
suborbital muscles of Amia are not morphologically homologous to 
the similarly shaped and located suborbitalis muscle of elas- 
mobranchiomorphs. Both muscles, however, are derived from the 
same geographical area of the adductor anlagen, and are thus 
homologous as suborbital fibers of the adductor mandibulae. 

Coelacanths retain both the medial and posterolateral adductor 
components but have lost all suborbital fibers (Fig. 2: character 8). 

The primitive condition for dipnoans and choanates is the 
presence of well-developed suborbital, medial, and posterolateral 
adductor divisions as well as an intramandibular section. Devonian 
dipnoans and early choanates appear to possess a very primitive 
configuration of the jaw adductor musculature, similar in all 
essential features to that of palaeoniscoids. Living lungfishes have 
lost all suborbital adductor fibers (Fig. 2: character 9). 



1979 EVOLUTION OF JAW MUSCULATURE 9 

Based on this analysis, the following classification of jaw adductor 
muscle states is proposed. The primitive gnathostome condition 
consisted of a jaw adductor composed of unspecialized suborbital 
and postorbital components. The presence of both suborbital and 
postorbital fibers defines the "fan-shaped" condition. Suborbital 
fibers occur in elasmobranchiomorphs as a distinct suborbitalis 
muscle while the postorbital component is divided into two 
divisions in actinopterygians, dipnoans, and choanates. The 
mandibular component represents the third adductor component. It 
lies lateral to the mandible in elasmobranchiomorphs and 
acanthodians, and medial to it in all other groups (Fig. 2). 

ACKNOWLEDGEMENTS 

During this study I was supported by an NIH Predoctoral 
Fellowship in musculoskeletal biology (5T32 GM077117) and by a 
Junior Fellowship in the Society of Fellows, Harvard University. 

I thank Drs. W. Fink, K. Liem, R. Winterbottom, E. O. Wiley, 
and H.-P. Schultze for comments on the manuscript and Dr. R. S. 
Miles for his communication on lungfish anatomy. 

LlTERATL'RE ClTED 

Allis. E. p. 1897. The cranial muscles and cranial and first spinal nerves in 

Amia calva. J. Morph. 72.-487-772. 
1917. Homologies of the muscles related to the visceral arches of the 

gnathostome fishes. Quart. J. Microsc. Sci. 62.'303-406. 
1922. The cranial anatomy of Polypterus. with special reference to 

Polvpierus hichir. J. Anat. 5(5.180-294. 
1923. The cranial anatomy of Chlamydoselachus anguineus. Acta. Zool. 



4.123-221. 
Danforth. C. H. 1913. The myology of Polyodon. J. Morph. .V.- 107- 146. 
Edgeworih, F. H. 1935. The Cranial Muscles of vertebrates. Cambridge Univ. 

Press, London. 493 pp. 
Gross, W. 1938. Das Kopfskelett von Cladodus wildungensis Jaekel. 2. Teil: 

Der Kieferbogen. Senckenbergiana 20.- 123-145. 
Jarvik, E. 1977. The systematic position of acanthodian fishes, pp 199-225. 

//; Andrews, S. M., Miles, R. S., and Walker, A. D. (Eds.), Problems in 

Vertebrate Evolution, Academic Press, London. 
Kesteven. H. L. 1942. The evolution of the skull and the cephalic muscles. 

Part 1. The fishes. Mem. Aust. Mus. 5.1-62. 
1943. The evolution of the skull and the cephalic muscles Part L The 

fishes (continued). Mem. Aust. Mus. (S.-63-I32. 
1944. The evolution of the skull and the cephalic muscles. Part. 11. 



The Amphibia. Mem. Aust. Mus. 5.133-236. 



10 BREVIORA No. 460 

Lauder, G. V. 1980 Evolution of the feeding mechanism in primitive actino- 

pterygian fishes: a functional anatomical analysis of Polypterus, Lepisosteus, 

and Amia. J. Morph. 163. 
LiEM, K. F. AND G. V. Lauder, in press. The evolution and interrelationships 

of the actinopterygian fishes. In Fish Neurobiology and Behavior, R. Davis 

and R. G. Northcutt (Eds). University of Michigan Press. 
LiGHTOLLER, G. H. S. 1939. Probable homologues. A study of the comparative 

anatomy of the mandibular and hyoid arches and their musculature. Trans. R. 

Soc. Lond. 24.- 349-445. 
Luther, A. 1913. Uber die von N. trigeminus versorgte Muskulatur der 

Ganoiden und Dipneusten. Acta. Soc. Scient. Fenn. 41:1-12. 
Miles, R. S. 1973. Relationships of acanthodians. pp. 63 104. In Interrelation- 
ships of Fishes, Greenwood, P. H., Miles, R. S., and Patterson, C. (Eds.), 

Academic Press, London. 
1977. Dipnoan (lungfish) skulls and the relationships of the group: a 

study based on new specimens from the Devonian of Australia. Zool. J. Linn. 

Soc. 6/.T-328. 
MiLLOT, J. AND J. Anthony. 1958. Anatomic de Laiimeria chalumnae L 

Squelette, muscles, et formations de soutiens. Centre National de la Recherche 

Scientifique, Paris. 
Olson, E. C. 1961. Jaw mechanisms: rhipidistians, amphibians, reptiles. Am. 

Zool. /.•205-215. 
Romer, a. S. 1966. Vertebrate Paleontology. University of Chicago Press, 

Chicago, 468 p. 
ScHAEFFER, B. AND D. E. RosEN. 1961. Major adaptive levels in the evolution 

of the actinopterygian feeding mechanism. Am. Zool. /.•187-204. 
Smith, B.G. 1937. The anniomy oi ihe W\\\ed shark Chlamydoselachus anguineus 

Garman. Bashford Dean Memorial Volume VI pp 333-505. American Museum 

of Natural History, New York. 
SoucHE, G. 1932. Morphologic comparative des muscles elevatei "s de la 

mandibule chez les poissons. Mem. Soc. Scient. Phy. Nat. de Bordeau 7e 

Serie J.T 292. 
Thomson, K. S. 1967. Mechanisms of intracranial kinetics in fossil rhipidistian 

fishes (Crossopterygii) and their relatives. J. Linn. Soc. (Zool.) </6.-223-253. 
Vetter, B. 1874. Untersuchungen zur vergleichenden Anatomic der Kiemen- 

und Kiefermuskulatur der Fische. L Selachien. Jena, Zeitschr. Bd. (9.-405-458. 
WiNTERBOTTOM, R. 1974. A descriptive synonymy of the striated muscles of the 

Teleostei. Proc. Acad. Nat. Sci. Phil. 725.-225-317. 



B R E ¥-lt-#HR A 



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Cambridge, Mass. June 30, 1980 Number 461 

GEOGRAPHIC VARIATION IN 

A NOUS BREVIROSTRIS 

(SAURIA: IGUANIDAE) IN HISPANIOLA 

Douglas L. Arnold^ 

Abstract. The nominal Hispaniolan anole species, Anolis hreviroslris Bocourt, 
is considered to be comprised of four distinct parapatric sibling species: A. breviros- 
tris, A. caudalis. A. websteri. and A. marron. These species and the subspecies of .-1. 
brevirostris are described. 

INTRODUCTION 

Anolis brevirostris Bocourt is a rather small and stocky anoline 
lizard that occurs within an irregular range in xeric habitats on 
Hispaniola and associated islets. In 1870 Bocourt named Anolis 
brevirostris from Haiti. The name fell into disuse and obscurity, 
with specimens oi A. brevirostris generally being included with A. 
dominicensis Reinhardt and Lutken. Cochran (1941) described and 
named two satelUte island subspecies of /4. dominicensis: wetmorei 
from Isla Beata, Dominican Republic; and caudalis He de la 
Gonave, Haiti. Barbour (1937) first combined A. dominicensis with 
A. distichus Cope. This combination was followed by Mertens 
(1939) who resurrected brevirostris as a subspecies oi A. distichus 
from the vicinity of Barahona in southwestern Dominican Republic. 
Ernest E. Williams, at the Museum of Comparative Zoology at 
Harvard University, recognized that some of the forms in Haiti and 
the Dominican Republic associated with A. distichus actually per- 
tain to another similar species, A. brevirostris (the differential char- 
acters were reported in Schwartz, 1968). Named forms correctly 
associated with A. brevirostris are caudalis Cochran and wetmorei 
Cochran. Webster and Burns (1973) used electrophoretic observa- 



' University of Kentucky. Albert E. Chandler Medical Center, P. O. Bo,\ 269, Lexing- 
ton. Kentucky 40536, U.S.A. 



2 BREVIORA No. 461 

tions on several proteins to assign mainland Haitian populations of 
""A. hrevirostris" to three unnamed sibling species. Webster (1978a) 
later expanded his electrophoretic studies to include most of the 
remaining distribution of A. hrevirostris. His accidental death left 
behind two near-final-form manuscripts on A. hrevirostris which 
lack the tables, figures, and numerical data he would have prepared 
to give his statements full documentation. These manuscripts subse- 
quently became available in the Third Anolis Newsletter (Williams, 
editor, 1977). Thus, as understood by Webster, the hrevirostris com- 
plex consisted of three unnamed sibling species referred to as A, B, 
and C, and two named satellite island subspecies. Williams (1976) 
referred the mainland Haitian species B to caudalis Cochran and 
regarded species C as nominate hrevirostris. 

ACKNOWLEDGMENTS 

This paper is the result of: 1) Collections made by Albert 
Schwartz and parties in Hispaniola, partly under National Science 
Foundation grants G-7977 and B-023603 and 2) information 
gathered by Albert Schwartz, Ernest E. Williams, and T. Preston 
Webster under National Science Foundation grant B-019801X and 
previous grants. Webster suggested that Schwartz undertake the 
formal description of the sibling species presented in Webster and 
Burns (1973) and a discussion of the ecology and distribution of the 
entire A. hrevirostris complex. In the fall of 1974 I began investiga- 
tion of the A. hrevirostris complex under Schwartz's direction. I am 
most grateful to him for his guidance, time, patience, and faith 
throughout the course of this project. 

Most of the specimens I have examined are in the Albert 
Schwartz Field Series (ASFS). A more limited amount of material 
has been borrowed from the American Museum of Natural History 
(AMNH), Museum of Comparative Zoology (MCZ), Museum 
National d'Histoire Naturelle (MNHP), Florida State Museum 
(UF/FSM), and the National Museum of Natural History (USNM). 
I am grateful to the following curators and their assistants for the 
loan of this material: Albert Schwartz, George W. Foley, Ernest E. 
Williams, Jean Guibe, Walter A. Auffenberg, W. Ronald Heyer, 
and Ronald J. Crombie. Burt L. Monroe provided me with work 
space at the University of Louisville and was instrumental in facili- 
tating loans. 



1979 GEOGRAPHIC VARIATION IN /lA'O/,/5 3 

METHODS 

I have not considered it worthwhile to study all the available 
collected specimens of the hrevirosiris complex since many of the 
older specimens are long-preserved and now much faded and since 
coloration and pattern play such a major role in differentiation of 
the various species and subspecies. I have attempted to examine all 
the material which might be assignable to new taxa proposed herein 
and have examined all specimens designated as holotypes, syntypes, 
and paratypes. 

The hreyirostris complex needs fine sampling to be understood 
(Webster, in lift., 1972); therefore I have restricted the type-locality 
oi A. brevirostris. Lists of associated specimens, not from the type- 
locality itself, include localities and museum numbers which I have 
assigned to certain taxa on the basis of my examination. A list of 
referred specimens in Appendix I includes localities and museum 
numbers which I have assigned to taxa on the basis of provenance; 
specimens so listed have not been examined by myself. The proba- 
bility is high that almost all lizards listed as referred specimens are 
correctly designated specifically and subspecifically, since they are 
delimited by fresh material that I have examined. Exceptional cases 
or uncertain allocations are noted in the text. 

Various scale counts and relationships were used by Oliver ( 1948) 
to describe Bahamian populations of A. distichus. Schwartz (1968) 
applied the same counts to non-Bahamian populations. Hoping that 
an application of these counts to A. brevirostris complex members 
might reveal differences other than coloration and pattern between 
various sibling species and their subspecies, I have followed the 
techniques of Oliver and of Schwartz and applied them to the mate- 
rial I have examined. The following data were recorded on each 
specimen: 

1) Snout-vent length, in millimeters. 

2) Number of scales across the snout at the level of the second 
canthal scale, reckoned from the anterior border of the orbit. 

3) Number of horizontal rows of loreal scales. 

4) Minimum number of scales between the supraorbital semi- 
circles. 

5) Number of rows of scales between the interparietal scale and 
the supraorbital semicircles on each side [this datum written as a 
fraction] (e.g., 1/1 = one row of scales on each side). 



4 BREVIORA No. 461 

6) Number of scales bordering the postfrontal laterally (see 
Oliver, 1948:16, for drawings showing these scales in A. disiichus). 
In a small number of specimens oi A. brevirostris, the postfrontals 
may extend so far laterally as to make contact with one or two of the 
canthals. In such instances, I have not included the canthal as a scale 
in contact with the postfrontal, since the condition is anomalous. 

7) Number of median (usually azygous) scales posterior to the 
posterior-most paramedian pair of snout scales, usually restricted to 
the midline from the anterior border of the postfrontals posteriorly 
to the anterior border of supraorbital semicircle contact, thus 
excluding the highly variable preoccipital region. (This count differs 
from those of Oliver and Schwartz but is made comparable by 
subtracting 1 from their counts — the nearly-always present preoc- 
cipital of A. disiichus.) 

8) Number of supraorbital semicircle scales in contact with the 
interparietal scale. This count is partly correlated with (5), the 
number of rows of scales between the semicircles and the interparie- 
tals; for instance, if the latter count is 1 / 1 , the number of supraorbi- 
tal scales in contact with the interparietal will of necessity be 0/0. 
However, if the count of (5) is 0/0 (i.e., there are no scales between 
the semicircles and the interparietal) then (8) may have a fairly wide 
fluctuation. 

9) Number of postmental scales. 

10) Condition of the preoccipital region categorized as: preoccip- 
ital present ( + ); series of small scales bordering interparietal ante- 
riorly which may be incomplete (ss); preoccipital present but 
separated from interparietal by a series of small scales (ps); multiple 
small scales in preoccipital region (ms); preoccipital present, but 
tiny (t); and preoccipital absent ( — ), supraorbital semicircles in con- 
tact with anterior border of interparietal. 

11) Condition of nuchal patch, categorized as large, medium, 
reduced, or absent; dark or faint; with or without white border. A 
medium-sized nuchal patch is reckoned to approximate the size of 
the external ear opening. 

The above counts and notations were taken on 770 specimens 
from Haiti and the Dominican Republic. Test counts taken on the 
number of subdigital lamellae on phalanges II and III of the fourth 
toe of the hind foot and the number of scales in tail verticils were 
discontinued as no significant variation was observed. Complete 



1979 GEOGRAPHIC VARIATION IN /^yVOL/S 5 

scale counts for each specific and subspecific samples are given in 
Table 1, where frequencies and modes are also shown. The reader is 
referred to this table for details of scutellar variation. 

A few scale counts as modes, not as absolute counts, have proven 
to be useful in defining the species and subspecies. The degree of 
overlap between various counts for different samples is generally 
extensive, so that it is impossible to identify the species and subspe- 
cies of a particular specimen solely on the basis of any set of counts. 
Therefore reliance must be placed on such features as dewlap pat- 
tern and coloration, nuchal patch pattern, and coloration and pat- 
tern of the head and body in general. These additional data were 
taken from series having field notes on coloration. (As mentioned 
above, museum specimens as such have very limited utility.) The 
colors of many dewlaps were noted in the field with reference to 
Maerz and Paul (1950). 

THE ANOLIS BREVIROSTRIS COMPLEX 

As noted in the Introduction, the brevirostris complex is presently 
understood to consist of three sibling species. The material on which 
the name A. brevirostris was based had its provenance merely as 
"Haiti". I have examined the three syntypes loaned by the Museum 
National d'Histoire Naturelle in Paris and find that two are A. 
distichus. The remaining one is A. brevirostris and has similarities 
to those lizards which are currently called "Species C". With the 
recognition of new species within the brevirostris' complex, it is 
appropriate to restrict the type-locality of /I. brevirostris in order to 
clarify my concept of the species; I hereby designate the vicinity of 
Fond Parisien, Departement de I'Ouest, Haiti, as the type-locality of 
A. brevirostris. It is not unlikely that the original specimens, col- 
lected by M. Braconnier, actually came from the vicinity of Port-au- 
Prince, for the Haitian capital has long been a prominent Caribbean 
seaport. No brevirostris complex anoles have been collected recently 
from Port-au-Prince, yet sibling species A, B, and C have all been 
collected within 20 km airline of that city. Therefore I have chosen 
to designate the type-locality oi A. brevirostris in the known distri- 
bution of Species C. Henceforth in this paper, A. brevirostris will 
refer only to Species C and ""A. brevirostris'' will refer to the com- 
plex in general. 



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1979 GEOGRAPHIC VARIATIION IN /lA'CL/S 9 

''A NO LIS BRE\ IROSTRIS" NATURAL HISTORY 

Anoles of the brevirosiris complex and A. distichus are usually 
allotopic, but broadly sympatric. However, in certain localities 
where the generally xeric habitat of "/I. hrevirostris'" intergrades 
with the generally mesic habitat of /I. distichus, the two are precisely 
syntopic. In these situations hybridization is known to occur, to 
varying degrees, producing in the analyzed cases sterile offspring 
(Webster, 1978b). 

"/I. hrevirostris'^ is cryptic. A generalized, composite head and 
dorsal pattern for the entire complex consists of: barred limbs and 
tail, two longitudinal dorsolateral stripes, an interocular bar, inter- 
parietal U, interocular V, and a series of five to seven dorsomedian 
chevrons pointed posteriorly. The first chevron is an internuchal 
patch. The upper longitudinal stripe extends from the angle of the 
jaw posteriorly to the level of the scapulae. Localized emphasis of 
this stripe forms the nuchal patch and scapular stripe. The lower 
longitudinal stripe extends from beneath the jaw across the shoulder 
to the hindlimbs. The pigmentation for the median dorsal stripe 
with chevrons is darker than the region below the upper dorsolateral 
stripe. All the markings are in general, faint with poor contrast, and 
interrupted outlines. 

The largest specimens of "/I. hrevirostris" are from inland north- 
ern Haiti. In the Vallee de I'Artibonite in the vicinity of Dessalines 
and Pont Sonde, males reach a snout-vent length of 51 mm and 
females 47 mm. In general, females of all populations reach a maxi- 
mum size about 4 to 8 mm less than do males. The smallest speci- 
mens are from Isla Beata off the coast of the Peninsula de 
Barahona. where males reach a snout-vent length of only 43 mm 
and females 38 mm. 

"/4. hrevirostris"" has an extensive but narrow distribution on 
Hispaniola. It occupies situations varying from mesic moderate ele- 
vations (Sierra Martin Garcia. 915 m) to extreme xeric regions (Cul 
de Sac-Valle de Neiba plain, at or below sea level). Typically an 
inhabitant of xeric habitats. '\4. hrevirostris"" prefers the larger trees 
of xeric woods and flourishes in the proximity of springs and in the 
conspicuously taller Acacia around villages. It is present through 
much of the low (often dense) Acacia scrub between the more favor- 
able habitats. 



10 BREVIORA No. 461 

In the mesic regions along the east coast of the Peninsula de 
Barahona, ""A. hrevirosiris'" is frequently observed on fenceposts in 
variegated sun and shade situations as well as on trees. In addition, 
"/I. hreyirostris"" has been collected on vines, shrubs, trees, and on a 
cliff face 90 m above the ocean. In the extreme xeric region of the 
Cul de Sac-Valle de Neiba plain it is more frequently associated with 
Cocos, being found diurnally on the ground under palm logs, stacks 
of palm fronds, in coconut piles, and in Cocos trash. It is possible 
that in this region "A. brevirostris" resorts to such situations for 
nocturnal retreats. Generally, when it is found asleep it is one to 
three m above the ground on vines, shrubs, or particularly Acacia. 

SYSTEMATIC ACCOUNT 

Anolis brevirostris brevirostris Bocourt 

Anolis brevirostris Bocourt, 1870, Nouv. Arch. Mus. Hist., Paris 
6:11. 

Lectotype. MNHP 2467B, an immature female collected by M. 
Braconnier. 

Type-locality. "Haiti"; restricted here to within 5 km of Fond 
Parisien, Dept. de I'Ouest, Haiti. 

Associated specimens. Haiti, Dept. de I'Ouest, Leogane (MCZ 
13781); Fond Michelle, 549 m (ASFS V37159); 7.4 km W. Thoma- 
zeau (ASFS V8156); Terre Rouge (ASFS V24266-94); La Source, 
Fond Parisien on edge of Etang Saumatre (ASFS V8 1 29-31); 
Source Fond Parisien, 1.6 km NE Fond Parisien (ASFS V36921- 
58); 3.2 km NW Fond Parisien (ASFS V36980-81); Dominican 
Republic, Independencia Province, Tierra Nueva (ASFS V42367- 
88); 10.4 km NE Jimani (ASFS X9506); 5 km N Jimani (ASFS 
V35476-77); Boca de Cachon (ASFS V4379, ASFS V397 19-41); Las 
Baitoas(ASFS V143I0-22); 6 km NW Duverge (ASFS V17142-46); 
5 km NNW Duverge (ASFS X9937); just E Duverge (ASFS 
V23274); Los Saladillos (ASFS V41744-45). 

Definition. A sibling species of the /I. brevirostris complex char- 
acterized by: moderate size (males to 48 mm, females to 45 mm 
snout-vent length); dorsum dark gray to medium gray and brown; 
dewlap pale, monochromatic, shades of orange, peach, yellow, gray, 
olive, and brown; nuchal patch variable, from reduced but dark, 
with no white border to medium and dark, with a white border; 
modally 1/1 scales between the supraorbital semicircles and the 



1979 GEOGRAPHIC VARIATION IN /1/VOZ./5' II 

interparietal; modally 0/0 supraorbitals in contact with the interpa- 
rietal; modally a single row of small scales bordering interparietal 
anteriorly; modally 3/3 scales in contact laterally with the postfron- 
tals (Species C of Webster and Burns, 1973). 

Distribution. Hispaniola: in the Dominican Republic from Inde- 
pendencia Province (Las Salinas), the Valle de Neiba, west into the 
Haitian Cul de Sac Plain, Dept. de I'Ouest (Daspinasse) and an 
isolated population on the Cap de Leogane, Dept. de I'Ouest 
(Leogane). 

Description of lectotype. The lectotype has the following meas- 
urements and scale counts: snout-vent length 34 mm, tail 55 mm, 
four scales across snout, four loreal rows, semicircles in contact, I / 1 
scales between supraorbital semicircles and the interparietal, 0/0 
supraorbitals in contact with the interparietal, a single row of small 
scales bordering interparietal anteriorly, 2/2 scales in lateral contact 
with postfrontals, one median azygous head scale, six postmentals. 
Syntypes MHNP 2467 and 2467 A are both A. distichus. 

Variation. Scale counts for the series of 146 specimens of /I. b. 
brevirostris are shown in Table 1. 

The dorsum and markings of A. b. brevirostris vary clinally. In 
the east (Duverge), the dorsum is medium gray-brown. The interoc- 
ular bar is present. The interparietal U and interocular V are often 
present. Dorsomedian chevrons are occasionally found. Both dorso- 
lateral stripes are present in reduced form. The upper remains as a 
scapular stripe and a nuchal patch is medium and dark, with a white 
border. The lower dorsolateral stripe is irregularly interrupted. The 
limbs and tail are barred. The more western the population, the 
more vague are the head and body markings, and the darker and 
more uniform gray the dorsum. At the western extreme the interpar- 
ietal U and the dorsomedian chevrons are absent. The tail and limbs 
are vaguely barred. The upper dorsolateral stripe is reduced, often 
to a scapular spot and a reduced, dark, borderless nuchal patch. 

Three geographical regions of dewlap variation in A. b. breviros- 
tris are discernible. At the western extreme (Daspinasse to Thoma- 
zeau) dewlaps are pale, monochromatic in hues of gray, brown, 
yellow, yellow-gray, cream, peach, and rarely white. In the southern 
Cul de Sac Plain and Valle de Neiba the dewlaps are predominately 
pale orange and peach. Dark and bright orange dewlaps have been 
noted in the vicinity of Fond Parisien and Duverge. Bicolored dew- 
laps, pale orange with a thin pale yellow margin, are found within 



12 BREVIORA No. 461 

this region from Las Salinas to 10 km west of Duverge. A third 
region, the northern Valle de Neiba between Etang Saumatre and 
Lac Enriquillo, has predominately pale yellow and yellow-brown 
dewlaps. Hues noted for A. h. hrevirostris are: PI. 916, PI. 9K5, PI. 
14C1, and PI. 14H5. 

The isolated population on the Cap de Leogane is represented by 
a single specimen (MCZ 13781) collected by G. M. Allen, August 
1919. The dorsum is a medium to dark brown with a dark colored 
and medium sized nuchal patch with a white border. The limbs and 
tail are barred. The scapular stripe is present, but other markings 
are obscured by the dark dorsum. Scale counts for the Leogane A. 
b. hrevirostris are: six scales across the snout, four loreal rows, 
supraorbital semicircles in contact, 1 / 1 scales between semicircles 
and interparietal, 0/0 supraorbitals in contact with interparietal, 
4/3 scales in lateral contact with postfrontals, four median head 
scales, four postmentals, and preoccipital present but separated 
from interparietal by a row of small scales. 

Anolis hrevirostris deserticola, new subspecies 

Holotype. MCZ 132391, an adult male. 

Type-locality. 2.1 km S San Jose de Ocoa, Peravia Province, 
Dominican Republic one of a series taken 19 November 1971, col- 
lected by natives. Original number ASFS V34034. 

Paratypes (all from Peravia Province. Dominican Republic). 
ASFS V34031-33, ASFS V34035-67, same data as holotype; ASFS 
V724-25, 1.8 km S San Jose de Ocoa, 24 August 1963, A. Schwartz; 
ASFS V15615-25, San Jose de Ocoa, 3 August, 1968, native 
collectors. 

Associated specimens. Dominican Republic, La Estrelleta Prov.. 
2.5 km NW El Llano, 365 m (ASFS V3 15 10-12); San Juan Prov., 
6.8 km NW Juan Herrera, 490 m (ASFS V31401); 1.7 km NW 
Sabaneta, 670 m (ASFS V3I394); 10.2 km S Las Matas de Farfan, 
550 m (ASFS V31327, ASFS V31468); 7.8 km NW Vallejuelo, 732 
m (ASFS V3I312); I km W Sabana Alta (MCZ 125547-48); SE 
Sabana Alta (MCZ 125554-55); 3 km W Guanito (MCZ 128236-47); 
15 km SE San Juan (ASFS V412, ASFS V470-86); Azua Prov., 
Padre las Casas (MCZ 58474-76); Sierra Martin Garcia, Mt. Busii, 
610 854 m (ASFS V31214-24); Sierra Martin Garcia, above Barre- 
ras, ca. 915 m (ASFS V2 11 74-202); Barreras, 10 km SW Puerto 
Viejo (ASFS V21 110-16); Peravia Prov., 3 5 km S San Jose de 



1979 GEOGRAPHIC VARIATION IN /l,VOZ./5 13 

Ocoa (ASFS V2 1396-97); 14.2 km S San Jose de Ocoa, 396 m 
(ASFS V715); 16.2 km S San Jose de Ocoa, 396 m (ASFS V712); 10 
km S Los Ranchitos (ASFS V15627); 19 km NW Bani (ASFS 
VI 5632); Limonal (ASFS V 1492 1-28); Barahona Prow, west side 
Punta Martin Garcia (ASFS VI 11-16); 3.2 km NE Fondo Negro 
(ASFS X9688); 4 km SE Canoa (ASFS V40785); 15 km ESE Canoa 
(ASFS V40796); 0.8 km NE Cachon, 122 m (ASFS V30749); 7.5 km 
E Cabral (ASFS X9604-05); ca. 3 km E Cabral (ASFS V20540-41); 
1 1.8 km S Cabral, 701 m (ASFS X9849-53); 12.3 km S Cabral. 640 
m (ASFS X9834-37). 

Definition. A subspecies of A. hrevirosiris characterized by: mod- 
erate size (males to 48 mm, females to 45 mm snout-vent length); 
dorsum medium gray-brown to medium brown; venter white; dew- 
lap bicolor, red-orange basal spot, margin cream or pale yellow to 
pale, or monochromatic, peach, salmon, or yellow; nuchal patch 
large and dark, with white border; modally 0/0 scales between the 
supraorbital semicircles and interparietal; modally supraorbitals 
in contact with the interparietal; modally one or more rows of small 
scales bordering interparietal anteriorly; modally 2 2 scales in con- 
tact laterally with the postfrontals. 

Distribution. Dominican Republic Valle de San Juan (El Llano, 
La Estrelleta Prov.), east onto the southern slopes of the Sierra de 
Ocoa and to the Llanos de Azua (Limonal, Peravia Prov.) and 
south onto the northern slopes of the Sierra de Baoruco (south of 
Cabral, Barahona Prov.). 

Description ofholotype. The holotype has thfe following measure- 
ments and scale counts: snout-vent length 45mm, tail (broken) 
48mm, four scales across snout, four loreal rows, semicircles in 
contact, 0/0 scales between supraorbital semicircles and interparie- 
tal, 1/1 supraorbitals in contact with interparietal, preoccipital 
absent, 2 2 scales in lateral contact with postfrontals, one median 
azygous head scale, seven postmentals. 

Variation. Scale counts for the series of 179 A. h. deserticola are 
shown in Table 1. 

The dorsum of A. b. deserticola is medium gray-brown to 
medium brown. The head markings are distinct and generally 
include the interocular bar, parietal U, and interocular V. Dorsome- 
dian chevrons are absent, except for the internuchal chevron which 
is rare. The lower dorsolateral stripe is absent. The upper is present 
and extends from the scapula to the hindlimbs in an interrupted 



14 BREVIORA No. 461 

pattern. The nuchal patch is large and dark, with a white border. 
The Hmbs and tail are barred. The venter is white to pale yellow with 
pale yellow on the underside of the tail. The dewlap is uniformly 
pale yellow, peach, or salmon to bicolor peach or red-orange with a 
pale yellow or cream margin; hues noted for the dewlap are: PI. 
10D8, PI. 18F1, PI. 17EI, with centers of PI. 5B12 and Pi. 6B11. 

The dorsum is a darker gray and the markings most distinct in 
specimens from the Sierra Martin Garcia and from the southern 
slopes of the Sierra de Ocoa. The browner dorsum and least distinct 
markings characterize specimens from the eastern Valle de San Juan 
(San Juan). 

Seven specimens collected west of San Juan are noteworthy. The 
samples are small (one to three each) and were collected in ravines 
peripheral to the Valle de San Juan. The preoccipital is present in 
71% of the specimens— significantly different from all other A. h. 
deserticola. The nuchal patch is reduced to absent. 

Scale counts for specimens from the moderate elevations between 
Lago de Rincon and the Bahia de Neiba and along the northern 
slopes of the Sierra de Baoruco are intermediate between those oi A. 
b. deserticola to the north and A. h wetmorei to the south. The scale 
counts and dewlap more closely favor A. b. deserticola and have 
thus been included with them. 



Anolis brevirostris wetmorei Cochran 

Anolis dominicensis wetmorei Cochran, 1931, Proc. Biol. Soc. 
Washington 44:89. 

Anolis brevirostris wetmorei: Schwartz, 1968, Bull. Mus. Comp. 
Zool. 137(2): 257. 

Holotype. USNM 83881, an adult male, 13 May 1931, collected 
by A. Wetmore and F. C. Lincoln. 

Type-locality. Isla Beata, Dominican Republic. 

Associated specimens. Dominican Republic, Barahona Prow, 
Barahona (ASPS X9530-60, ASPS VI40I7-38, ASPS X9456-58); 5 
km S Barahona (ASPS V20545-48); 6 km SE Barahona (ASPS 
V40787-88); 6.6 km SW Barahona, 137 m (ASPS V30240-43, 
V30418-I9); 15 km SW Barahona (ASPS V4078I-83); 5.3 km NE 
La Cienaga (ASPS X9379-8 1 ); 6.4 km SW La Cienaga, 46 m ( ASPS 
V39858); 9 km SW La Cienaga, 91 m (ASPS V42710); 2 km NW 
Paraiso (ASPS VI 7-24); 1 km NE Paraiso, Rio Nizaito (ASPS 



1979 GEOGRAPHIC VARIATIION IN /4yvOL/5 15 

V40762-71); 1 km W Paraiso (ASFS V40772-80); 7.7 km W Paraiso, 
152 m (ASFS V30928-31); 17.9 km NE Caleton (ASFS V30790- 
802); Los Patos (ASFS V16926); 6 km N Enriquillo, 427 m (ASFS 
V42190-209); 2.2 km NE Enriquillo (ASFS V14078-80); Pedernales 
Prov., 21 km SW Enriquillo (ASFS X9416); 8 km NE Oviedo 
(viejo) (ASFS X9964-65); 7 km N Oviedo (viejo) (ASFS V23121); 5 
km NE Oviedo (viejo) (ASFS V 14073-75); 3 km NW Oviedo 
(nuevo) (ASFS V40678-87, ASFS V40471-82); 7 km N, thence 20 
km SE Cabo Rojo, 183 m (ASFS V29769-70); 7 km N, thence 17.6 
km SE Cabo Rojo, 152 m (ASFS V30088-89); 2 km E turn to Cabo 
Rojo (MCZ 128257); behind Cabo Rojo police station (MCZ 
143407-08); 21 km NE Cabo Rojo, 396 m (ASFS V16782-85); 21.2 
km N Cabo Rojo on Alcoa mine road, 440 m (MCZ 143403-04); 
L'Eglise, near La Mercedes (ASFS V21510); 10 km N Pedernales, 
244 m (ASFS V301 15-17); 4 km SE Pedernales (ASFS V16729-30); 
south center of Pedernales (ASFS V29846); Pedernales (ASFS 
V2505-06, ASFS V2919); Isla Beat a (ASFS V2670-71, UF/FSM 
36969-70, MCZ 37520, MCZ 58695-702, AMNH 41414, AMNH 
41416-18, AMNH 44854-58); just E Punta Beata (ASFS V17220); 
Haiti. Dept. de l' Quest, near Sahrou (MCZ 49274-79); 4 km NNE 
Marigot, 61 m (ASFS V37372-77). 

Definition. A subspecies oi A. brevirostris characterized by: large 
size (males to 50 mm, females to 45 mm snout-vent length); dorsum 
pale grayish tan to gray to tan to brown to mottled greenish tan; 
venter white to yellow; dewlap color very variable from locality to 
locality (see below); nuchal patch reduced and gray to brown with 
no white border to large and dark with white border; modally 1/1 
scales between supraorbital semicircles and interparietal; modally 
0/0 supraorbitals in contact with interparietal; modally a single row 
of small scales bordering the interparietal anteriorly; bimodally 2/2 
or 3/3 scales in contact laterally with postfrontals. 

Description ofholotype. The holotype has the following measure- 
ments and scale counts: snout-vent length 43 mm, tail (broken) 62 
mm, five scales across snout, four loreal rows, semicircles in contact, 
1/1 scales between supraorbital semicircles and interparietal, 0/0 
supraorbitals in contact with interparietal, a single row of small 
scales bordering interparietal anteriorly, 3/3 scales in lateral contact 
with postfrontals, three median head scales, five postmentals. 

Variation. Scale counts for the series of 232 A. h. wetmorei are 
shown in Table 1. 



16 BREVIORA No. 461 

Disirihuiion. Hispaniola: in Haiti from the Dept. de TOuest 
(Marigot), the southern coast of the Tiburon Peninsula east across 
the southern portion of the Dominican Peninsula de Barahona 
(Enriquillo), Isla Beata, and north along the eastern coast of the 
Peninsula de Barahona to the city of Barahona. 

A. h. wetmorei varies geographically. On Isla Beata, whence wet- 
tnorei was first described, the dorsum is nearly uniform tan or light 
gray. In general the head and body markings are faint to absent. The 
interocular bar is faint and the interocular V is reduced to a short 
postocular stripe. The dorsomedian chevrons and dorsolateral 
stripes are absent. The nuchal patch is present and varies from large 
and dark to reduced and gray, but always with a white border. The 
limbs and tail are faintly barred. The venter is whitish and the 
dewlap is pale yellow. 

On mainland Hispaniola wetmorei dewlaps vary from pale yellow 
to bicolored yellow with an orange central spot to deep orange or 
red-orange. However, regionally the variation is less extreme, favor- 
ing either the deep orange, red-orange, and bicolored orange with a 
thin pale margin (Enriquillo to la Cienaga and Pedernales to Caii- 
ada la Cerca) or the pale yellow to yellow with orange blush basally 
characteristic of the remainder of the extensive wetmorei distribu- 
tion. The full range of variation has been observed at the northeast- 
ern distribution extreme, Barahona, and at the regional interfaces at 
Pedernales, La Cienaga, and south of Enriquillo. 

The eastern orange-dewlap wetmorei have a dorsum that is pale 
greenish tan to tan. The markings of the head and body are cryptic 
to the point that the dorsum has a woodgrain pattern. The interocu- 
lar bar is regularly present. The interparietal U is wide and dark. 
Mediandorsal chevrons are occasionally present, although most 
often they are reduced to dorsomedian spots or are absent. The 
upper and lower dorsolateral stripes are interrupted to absent. The 
legs and tail are lightly barred. The nuchal patch is reduced and 
faint, often gray, tan, or reddish brown, and with no white border. 
Hues noted for the dewlap in this series are: PI. 1017. PI. lOGII, 
PI. 3F12, PI. 4B11, and Pi. 1 1J8. 

The western orange-dewlap wetmorei are similar to the eastern, 
but there are no reports of any green in the dorsal color. The mark- 
ings have greater contrast. The scapular stripe is long and there are 
remnants of the lower dorsolateral stripe. PI. lOCIO is the only hue 
noted for the dewlap. 



1979 GEOGRAPHIC VARIATION IN /lA'C>Z./5 17 

The yellow-dewlap wetmorei are in general more conspicuously 
marked, except in the vicinity of Oviedo where the dorsum is a 
darker and grayer version of wetmorei on Isla Beata. The nuchal 
patch for these populations is highly variable, from large and dark 
with a white border to reduced and faint with no white border. 
Additional hues noted for the dewlaps are: PI. I OF 1 , P 1 . 11 F3, P 1 . 
IIEI, with PI. IIBIO and PI. 1107 centrally. 

Anolis caudalis Cochran 

Anolis dowiniceusis caudalis Cochran, 1932, Proc. Biol. Soc. 
Washington 45:185. 

Anolis brevirostris caudalis: Schwartz, 1968, Bull. Mus. Comp. 
Zool. 137(2):257. 

Holotype. USNM 76801, an immature male, March 1929, 
collected by A. J. Poole and W. M. Perrygo. 

Type-locality. Nan Cafe, He de la Gonave, Haiti. 

Paraivpes. Haiti, lie de la Gonave. USNM 76799-800. same data 
as holotype; USNM 77080-82. Pointe a Raquettes, "1927", W. J. 
Eyerdam; USNM 80388-89, 8.3-16.7 km inland Anse-a-Galets, 23 
March 1930, L. H. Parish and W. M. Perrygo; He de la Petite 
Gonave, USNM 80390, 23 March 1930, L. H. Parish and W. M. 
Perrygo. 

Associated specimens. Haiti, Dept. de l' Quest, Trou Forban 
(ASFS X1939, ASFS V8204-14); 3.5 km E Trou Forban (ASFS 
X4001-03); 0.3 km NE Carries, Ouanga Bay Hotel (ASFS V36876- 
82); 4 km N Arcahaie (MCZ 125008-18); 3 km SE Arcahaie (ASFS 
V43778-81); 6 km N Duvalierville (MCZ 125022-37); 1.4 km SE 
Duvalierville (ASFS V36872-74); 6 km SE Duvalierville (ASFS 
V9834-36); He a Cabrit, 6 km airline SE Duvalierville (ASFS 
V9853-56, ASFS V44806-25); Source Matelas (MCZ 125019-21); 
Dept. du Sud, Presqu'ile de Baraderes, vicinity of Grand Boucan 
(ASFS V26342-48); 4.2km W Jeremie (USNM 59255-56); He de la 
Gonave, Source Picmi, above Picmi (ASFS V266I0-15, ASFS 
V26652); Anse-a-Galets (ASFS V22413-22); Etroits (ASFS X2410- 
15). 

Definition. A sibling species of the /i. brevirostris complex char- 
acterized by: moderate size (males to 48 mm. females to 44 mm 
snout-vent length); dorsum medium brown to light gray-tan to light 
gray; venter pale yellow; dewlap highly variable from uniformly 



18 BREVIORA No. 461 

white, yellow or yellow-gray to bicolor yellow with red or orange 
centers to bicolor gray-green with rusty centers; nuchal patch 
medium, dark with white border; modally 11 scales between the 
supraorbital semicircles and the interparietal; modally 0/0 supraor- 
bitals in contact with the interparietal; modally preoccipital present; 
modally 2/2 scales in contact laterally with the postfrontals (Species 
B of Webster and Burns, 1973). 

Distribution. Haiti: Dept. de I'Ouest, from Trou Forban south 
along the coast of the Canal de St. Marc to the Bale de Port-au- 
Prince (Source Matelas); He de la Gonave; He de la Petite Gonave; 
and two isolated populations on the northern shore of the Tiburon 
Peninsula (Dept. du Sud: Presqu'ile de Baraderes and Jeremie). 

Description ofholotype. The holotype has the following measure- 
ments and scale counts: snout-vent length 42 mm, tail broken, six 
scales across snout, four loreal rows, semicircles in contact, 0/0 
scales between supraorbital semicircles and interparietal, 1 / 1 supra- 
orbitals in contact with interparietal, preoccipital region with five 
small scales with an incomplete row of small scales bordering inter- 
parietal anteriorly, 2/2 scales in lateral contact with postfrontals, 
one median azygous head scale, five postmentals. 

Variation. Scale counts for the series of 13 1 .4. caiidalis are shown 
in Table 1. 

The dorsum of A. caudalis is medium brown to light gray; the 
lightest specimens examined are from Anse-a-Galets, He de la 
Gonave. The head markings are generally absent. Dorsomedian 
chevrons are absent. The nuchal patch is large and dark with a white 
border. The scapular stripe is short but distinct. The remainder of 
the dorsolateral stripes are reduced, interrupted, and vague to 
absent. The limbs and tail are weakly barred. The venter is pale 
yellow. The dewlap color is highly variable from uniformly white, 
yellow, yellow-gray, or gray-green, to bicolored white or yellow with 
variable orange centrally to bicolored yellow-green or gray-green 
with yellow, mustard, rusty, or darker gray-green centers. Hues 
noted for the dewlap color are: PI. 12E1, PI. lOCl, and PI. 6K10 
centrally with PI. 1011 and PI. 17G1 peripherally. 

At the northern mainland extreme oi A. caudalis (Trou Forban) 
the dewlap is uniformly white or pale yellow to bicolored with a 
small orange basal spot. There is a clinal increase in the size and 
intensity of the orange spot the farther south along the coast (Web- 
ster and Burns, 1973). However, the extent of the orange, even at the 



1979 GEOGRAPHIC VARIATION IN /lA'OZ./5 19 

southern mainland extreme (Source Matelas), is not so great as that 
of /4. wehsteri. Two exceptions to this general cline are recorded. At 
Trou Forban. one series (ASFS V8204-14) had dewlaps gray to 
greenish gray with mustard or darker gray-green centers. On the lie 
a Cabrit in the Baie de Port-au-Prince the dewlaps are gray-green. 

On He de la Gonave no such cline exists, and the dewlap color is 
variable and includes all dewlap colors described above for A. 
caudalis. 

The two disjunct populations of A. caudalis are of particular 
interest. Jeremie caudalis is represented by only two specimens 
(USNM 59255-56) collected by Henderson and Bartsch, 10 April 
1917. The dorsum is a faded gray-tan, with a dorsomedian stripe 
and short scapular stripe. The nuchal patch is large and dark with a 
white border. The limbs and tail r le barred. The head has a faint 
interocular bar, short ocular stripe, and a broad and dark interpa- 
rietal U. Scale counts for these two caudalis are: six scales across 
snout, four loreal rows, supraorbital semicircles in contact, 0/0 and 
1/1 scales between semicircles and interparietal, 0/0 and 1/1 
supraorbitals in contact with interparietal, 2/2 scales in lateral con- 
tact with postfrontals, no median head scales, and preoccipital 
present. 

The other disjunct population. Grand Boucan caudalis, is repre- 
sented by seven specimens (ASFS V26342-48) collected by Richard 
Thomas and natives, 4 August 1971. Of the three known disjunct 
populations of "A. brevirostris" on the Tiburon Peninsula, only 
Grand Boucan caudalis have dewlap color notes. The dewlaps are 
colored "pale (greenish) with faint basal dull yellow smudge". The 
dorsum is medium to dark brown with a scapular stripe. The lower 
dorsolateral stripe is discernible. The interparietal U and interocular 
bar and V are present. The limbs and tail are barred. The nuchal 
patch is large and dark with a white border. Grand Boucan caudalis 
differ from mainland and Gonave caudalis in two modal scale 
counts: five scales across the snout and three median head scales. 
The preoccipital, although present, is fragmented into multiple 
scales in three of seven specimens. 

Anolis marron, new species 

Holotype. MCZ 124732, an adult male. 

Type-locality. Jacmel, Departement de I'Ouest, one of a series 
taken 15 June 1970, collected by T. P. Webster. 



20 BREVIORA No. 461 

Paratypes. MCZ 124730-31, MCZ 124733-36, same data as 
holotype. 

Associated specimens. Haiti, Dept. de I'Ouest, Marigot (MCZ 
124723-29); 3.2 km W Marigot (ASFS V9771-78); 4.8 km E Cayes 
Jacmel (ASFS V971 1-16); 8 km W Jacmel (ASFS V9696-97); Terre 
Noire, 19.2 km SW Jacmel (ASFS V37509). 

Definition. A sibling species of the /I. brevirostris complex char- 
acterized by: moderate size (males to 50 mm, females to 42 mm 
snout-vent length); dorsum pale gray and tan to medium brown; 
venter pale yellow to off-white; dewlap uniformly olive-gray to bico- 
lor tan with reddish center; nuchal patch large and dark with faint 
white border; modally 0/0 scales between supraorbital semicircles 
and the interparietal; modally 0/0 supraorbitals in contact with the 
interparietal; modally one or more rows of small scales bordering 
interparietal anteriorly; modally 2/2 scales in contact laterally with 
postfrontals. 

Distribution. Haiti; the southern coast of the Tiburon Peninsula, 
from Marigot west to Terre Noire. 

Description of holotype. The holotype has the following measure- 
ments and scale counts: snout-vent length 47 mm, tail 62 mm, four 
scales across snout, four loreal rows, semicircles in contact, 0/0 
scales between supraorbital semicircles and the interparietal, 2/1 
supraorbitals in contact with the interparietal, a series of small 
scales bordering interparietal anteriorly, 2/2 scales in lateral contact 
with postfrontals, one median azygous head scale, five postmentals. 

Variation. Scale counts for the series of 31 A. marron are shown 
in table 1. 

The dorsum of A. marron is pale gray and brown to medium 
brown. A thin interocular bar is regularly present, but other head 
markings are vague or absent. The dorsomedian chevrons are 
reduced to small dorsal spots or are absent in all but the darkest 
specimens. The thin, dark scapular stripe broadens and becomes less 
distinct posteriorly. The lower dorsolateral stripe is interrupted and 
vague. The nuchal patch is large and dark with a white border. The 
limbs and tail are barred. The venter is pale yellow to off-white. The 
dewlap is either uniformly dark olive-gray or bicolored tan with a 
reddish center; hues noted for the dewlap are: PI. 15H5, PI. 6B10, 
and PI. 6C10. 



1979 GEOGRAPHIC VARIATION IN /^A'OL/S 21 

Anolis websteri, new species 

Holotype. MCZ 132390, an adult male. 

Type-locality. 1 A km NW Dessalines, Departement de TArtibo- 
nite, Haiti, one of a series taken 8 July 1974, collected by natives. 
Original number ASFS V39323. 

Paratypes. ASFS V39322, ASFS V39324-37, same data as holo- 
type. 

Associated specimens. Haiti, D6pt. de Nord Quest. Mole St. 
Nicholas (MCZ 63143-53); Dept. de TArtibonite, 6.7 km NW 
Gonaives .(ASFS V39295-302); 1 km NE Pont Sonde (ASFS 
V39342-50); Bains de Amani-y (ASFS V36900-03, ASFS V43742); 
2.6 km NW Montrouis (ASFS V36888-89). 

Definition. A sibling species of the A. hrevirostris complex char- 
acterized by: large size (males to 51 mm, females to 47 mm snout- 
vent length); dorsum medium to dark gray and brown; venter bright 
yellow to yellow-orange; dewlap orange with a thin pale yellow 
edge; nuchal patch large and dark with no white border; modally 
1 / 1 scales between the supraorbital semicircles and the interparietal; 
modally 0/0 supraorbitals in contact with interparietal; modally a 
single row of small scales bordering interparietal anteriorly; modally 
2/2 scales in contact laterally with postfrontals. (Species A of Webs- 
ter and Burns, 1973). 

Distribution. Haiti; from the Dept. de Nord Quest (Mole St. 
Nicholas) and northwestern Dept. de I'Artibonite (Marche aux 
Poteux; between Gonaives and Ennery), south into the Vallee de 
I'Artibonite (Dessaliens and Pont Sonde) and along the coast of the 
Golfe de la Gonave to Pointe de Montrouis (Montrouis). 

Description of holotype. The holotype has the following measure- 
ments and scale counts: snout-vent length 47 mm, tail 70 mm, five 
scales across snout, four loreal rows, semicircles in contact, 1 / 1 
scales between supraorbital semicircles and interparietal, 0/0 supra- 
orbitals in contact with interparietal, a single row of small scales 
bordering interparietal anteriorly, 3 3 scales in lateral contact with 
postfrontals, one median azygous head scale, four postmentals. 

Variation. Scale counts for the series of 51 A. websteri are shown 
in Table 1. 

The dorsum of A. websteri is moderate to dark gray or brown. 
Head and body markings, if present, are generally dark and diffuse. 



22 BREVIORA No. 461 

often disrupted. The nuchal patch is large and dark, with no white 
border. The scapular stripe is dark and short. The interocular bar is 
present in the lighter colored specimens, as is sometimes the case 
with the interparietal U and interocular V. Dorsal chevrons are 
always absent. The dorsolateral stripes are disrupted and diffuse. 
The limbs and tail are vaguely barred. The interparietal is yellow, 
but appears as a copper flake in preserved specimens. The venters 
are bright yellow to yellow-orange. The dewlap is uniformly orange 
or orange with a narrow yellow margin; hues noted for the dewlap 
are: PI. 9L7, PI. 9L8, PI. 9L10, PI. 10L8, and PI. IIKIO. 

A series of eight A. websteri from Pont Sonde is notable in the 
variation in dewlap color and modal scale counts. The dewlaps vary 
from orange (PI. 10L8) to mustard (PI. 12L7) to pale yellow (Pi. 
1 1L4). The venters are not so bright as other wehsteri. Modal scale 
counts differences for the series are: six scales across the snout; five 
loreal rows; and 3/3 scales in contact laterally with postfrontals. 

Well outside its normal distribution, A. wehsteri was found in 
abundance 3.5 km NW Bon Repos, Dept. de I'Ouest, in young 
native mahogany {Swietenia mahagoni Jacq.). Electrophoretic evi- 
dence indicates that it is slightly more similar to northern A. weh- 
steri than to southern specimens of that species. Presumably, A. 
wehsteri in this location was a recent introduction by man to an 
artificial habitat. Recent road improvement has led to the total 
destruction of the localized habitat (Webster, 1978a). No specimens 
were preserved. 

DISCUSSION 

INTERACTION OF THE 'ANOLIS BREVIROSTRIS"— 
A NOUS DISTICH US COMPLEXES 

Species of the "A. brevirostris" — A. distichus group often present 
contrasting dewlap colors in areas of contact or proximity. Such 
differences suggest that dewlap color may be important for species 
identification and reproductive isolation (Webster and Burns, 1973). 
Both body size and color appear to be important in Anolis species 
identification, especially in simple (e.g., two species) faunas (Will- 
iams and Rand, 1977). 

The A. hrevirostris complex is closely associated with four sub- 
species of A. distichus: dominicensis, ravitergum, favillarum, and 
suppar. In areas of sympatry "A. hrevirostris" and distichus gener- 



1979 GEOGRAPHIC VARIATION IN /lA'OZ./5 23 

ally vary in body size, body color, and dewlap color. In brief, the 
associated subspecies have the following characteristics: dominicen- 
sis, males to 58 mm, females to 48 mm snout-vent length, dorsum 
bright marbled green to rich chocolate brown, dewlap usually pale 
yellow; ravitergunu males to 56 mm, females to 45 mm snout-vent 
length, dorsum ashy gray to dark tan to pale greenish, dewlap pale 
yellow to bicolored with an orange basal spot: favillarum, males to 
54 mm, females to 47 mm snout-vent length, dorsum dark green, 
dewlap vivid orange centrally with narrow pale yellow edge; suppai\ 
males to 54 mm, females to 44 mm snout-vent length, dorsum pale 
green, dewlap pale yellow to yellow-green, at times with a dull 
yellow-orange basal smudge. For more detailed descriptions, see 
Schwartz (1968) and Webster (1978b). In general, A. distichus is 
larger than "A. brevirostris" and the green and brown unmarked 
dorsa of distichus contrast with the gray and marked dorsa of 
"brevirostris". 

A. caudalis is the only species within the brevirostris complex 
which is not extensively sympatric with. 4. d. dominicensis. Quite to 
the contrary, both mainland and Gonave caudalis appear to exclude 
dominicensis. except for two specimens, one from Trou Forban and 
another from nearby Ste. Philomene. At the disjunct populations on 
the Tiburon Peninsula, Grand Boucan caudalis also appear to 
exclude dominicensis and the Jeremie population is sympatric with 
another subspecies, A. d. suppar. No color notes exist for the Jere- 
mie caudalis, nor can any be assumed. A. caudalis here is the smaller 
lizard and has two prominent markings. The nuchal patch is large 
and faint with a white border and the interparietal U is broad and 
dark. 

A. d. dominicensis is broadly sympatric with A. websteri and A. 
marron throughout the latters' respective ranges. Locally domini- 
censis is precisely syntopic with websteri. A. websteri has the largest 
snout-vent length for the brevirostris complex and at this region of 
syntopy is larger than dominicensis. The reduced, borderless nuchal 
patch and generally vague and diffuse markings of gray-brown web- 
steri probably indicate an increased reliance on dewlap color for 
species recognition and reproductive isolation. The orange dewlap 
of websteri sharply contrast with the typical pale yellow of domini- 
censis and white to gray-yellow with greenish basally oi A. caudalis. 
The isolation is incomplete, for Webster (1978b) reports a low inci- 



24 BREVIORA No. 461 

dence of hybridization at Montrouis with duininicensis; and (in Htt.) 
some hybridization and introgression (two of five proteins, only one 
way, only a short distance) with caudalis at Trou Forban. 

Though populations oi A. marron are never far from those of /4. 
d. dominicensis, no areas of syntopy are known. The dark olive-gray 
or bicolored tan with reddish center dewlap contrast with the pale 
yellow of dominicensis. The dark scapular stripe and large, dark, 
white bordered nuchal patch oi marron are more prominent dorsal 
markings than those oi A. wehsteri and may provide additional cues 
for species recognition and reproductive isolation. A. marron and 
A. h. wetmorei come in contact near Marigot. The bicolored pale 
yellow with orange basal spot of .^. h. wetmorei contrasts with the 
marron dewlap. 

In the eaistern Cul de Sac Plain (Thomazeau; Manneville), A. h. 
hrevirostris is sympatric (but apparently not syntopic) with A. d. 
dominicensis. Of the two species in the area, A. b. hrevirostris is 
distinctly the inhabitant of the xeric scrub and A. d. dominicensis 
the inhabitant of more mesic situations (oases and cultivated areas). 
A. b. brevirostris is the widespread lizard in open areas, whereas /I. 
d. dominicensis is restricted to certain less rigorous habitats and is in 
effect surrounded by A. b. brevirostris. Here, the dewlap color of ^. 
d. dominicensis is atypically a deep orange (PI. 4CI 1, PI. 4GI0, and 
PI. 4G1 1), at times with a faintly brown cast (Schwartz, 1968). This 
contrasts strongly with the dewlap of A. b. brevirostris, which 
locally is dully pigmented with no conspicuous pattern in white to 
pale shades of yellow, gray-yellow, and gray-brown. 

There is a clinal decrease in the amount of orange in the dewlap of 
A. d. dominicensis away from A. hrevirostris, until at the southern 
edge of the Cul de Sac and Valle de Neiba plain, pale yellow is again 
the typical color oi A. d. dominicensis dewlaps. It is important to 
note that in this area (Fond Parisien to Duverge) A. h. brevirostris 
dewlaps are predominantly pale orange, dull orange, to bright 
orange. 

Another subspecies, A. d. ravitergum, is sympatric with A. b. 
hrevirostris at the latter's eastern distribution. Webster (1978b) 
reports hybridization locally at Balneario la Zurza, 5 km WNW 
Duverge. Here the dewlaps oi A. d. ravitergum range from uni- 
formly light yellow-orange to strongly bicolored with an orange 
basal spot. Dewlaps oi A. b. brevirostris are all monochromatic 



1979 GEOGRAPHIC VARIATION IN /4/VOZ./5 25 

orange in varying degrees of intensity. Dewlaps do not appear con- 
sistently to differentiate these populations despite the strong dewlap 
color contrast. 

A. h. wetmorei, like /I. caudalis, appears to exclude .4. distichus 
but is locally sympatric with two subspecies of /I. distichus at three 
localities. A. d. dominicensis and A. h. wetmorei are sympatric at 
Belle Anse (=Saltrou), Dept. de POuest, and Las Mercedes, Peder- 
nales Province. Here, wetmorei is bicolored with an orange basal 
spot and a yellow or cream outer margin. A. d. dominicensis dewlap 
is typically pale yellow but occasional specimens in the Sierra de 
Baoruco iji Pedernales Province have the dewlap very pale yellow to 
practically white. 

A. d. favillarum and A. h. wetmorei are sympatric at Hermann's 
finca, near Paraiso, Barahona Province. No color notes exist for the 
single specimen of wetmorei taken from this locality. (These are, 
however, old records and may not imply real sympatry.) 

At all three of these widely separated localities, wetmorei is gener- 
ally the lizard of the lower elevations with distichus occupying the 
moderate to higher elevations (300^ 1 250 m) of the Massif de la Selle 
and Sierra de Baoruco. 

A. h. deserticola is sympatric with one of three subspecies oi A. 
distichus throughout most of its range. A. d. dominicensis and A. b. 
deserticola are widely sympatric in the Valle de San Juan from 7 km 
NW Vallejuelo and 15 km SE San Juan westward to the Dominico- 
Haitian border. The dewlaps of both are pale yellow except at Valle- 
juelo where deserticola is bicolored with a pale orange blush. A. d. 
ravitergum is sympatric throughout most of the remaining range of 
deserticola and in local situations is even syntopic (0.5 and 2.5 km E 
Cachon). Webster (1978b) notes the similarity in dewlaps. Both are 
bicolored, with a pale margin surrounding a darker or brighter spot 
of variable size. Corresponding to this high degree of similarity is a 
higher incidence of hybridization at these localities. 

A. h. deserticola extends onto the northern slopes of the Sierra de 
Baoruco, where in the intermediate altitudes and transitional vege- 
tation it is sympatric and locally syntopic with. 4. d. favillarum. The 
dewlaps are here identical; both are dark orange, but the nuchal 
patch and gray dorsum of deserticola contrast strongly with the 
green dorsum of favillarum. The incidence of hybridization here has 
not been investigated. 



26 BREVIORA No. 461 

The sample examined in this study represents the known distribu- 
tion of the "A. brevirostris" complex. Three of the six populations 
recognized by scale counts correspond to the informal siblings A, B, 
and C of Webster and Burns (1973). A. brevirostris is restricted to 
Species C and the name A. caudalis is assigned to Species B as 
suggested by Williams (1976). The distribution of /I. b. wetmorei of 
Isla Beata is expanded to include the Peninsula de Barahona and 
part of the southern coast of the Tiburon Peninsula. Species A {A. 
websieri), a new sibling species (A. marro?i), and a new subspecies 
(A. b. deserticola) are described. Scale counts were taken as supple- 
mentary evidence of geographic variation along with that in dewlap 
color within the brevirostris complex, and to provide a basis for the 
formal description of parapatric siblings. 

A. caudalis and each of the three subspecies of A. brevirostris 
overlap in dewlap color variation. Dewlaps bicolored yellow with 
orange centers and/ or uniformly pale yellow are found to varying 
degrees in the distribution of each species. A. caudalis is, by modali- 
ties of scale counts, the most readily distinguishable of these species 
and of the entire brevirostris complex. Modally, has six scales 
across the snout (all others four), preoccipital present (all others a 
single row of small scales bordering the interparietal anteriorly), and 
two median scales (all others modally one or bimodally one and 
two). 

Webster (1978a) noted the presence of shared alleles of A. cauda- 
lis with A. marron and A. b. deserticola. The scale counts of the 
latter two populations differ from the former not only by the unique 
modalities of caudalis noted above, but further by the modes shared 
by marron and deserticola: scales between interparietal and supra- 
orbital semicircles 0/0 (all others 1/1); postmentals, bimodes five 
and six, with the highest mean values deserticola 5.8 and marron 5.6 
(others, mode four or five, or trimode four, five, and six; means 4.7 
to 5.3). Despite the wide geographic separation (ca. 120 km airline 
across the Sierra de Baoruco and Massif de la Selle at the closest 
points), these two populations have essentially the same scale 
counts. The modalities for the scale counts taken are the same, the 
percentages of the modalities are within three to 17% of each other, 
and the distribution of the non-modal counts is similar. Further, 
these are two of the three populations that are extensively sympatric 
with A. distichus. A. marron differs from A. b. wetmorei and the 



1979 GEOGRAPHIC VARIATION IN /1/VOZ./5 27 

entire hrevirostris complex by its unique dewlap color (dark olive- 
gray to tan with reddish center). 

A. h. hrevirostris differs from other populations in the "A. hrevi- 
rostris" complex by the unique modes of 3/3 scales in contact with 
postfrontals laterally (all others modally 2/2 or bimodally 2 2 and 
3/3) and four postmentals, with 4.7 the lowest mean value (others 
modally five, bimodally five and six or trimodally four, five, and six; 
means 4.7 to 5.8). 

A. b. wetmorei differs from other populations in iht hrevirostris 
complex by the unique bimode of 2 2 and 3 3 scales in contact with 
postfrontals laterally. A. h. wetmorei differs from the other subspe- 
cies oi A. hrevirostris in having modally five postmentals {hreviros- 
tris modally four and deserticola bimodally five and six), but share 
this characteristic with A. caudalis. Variation in scale counts and 
dorsum occur within A. h. wetmorei. At Enriquillo there is a dra- 
matic change in vegetation and geography from the mesic slopes of 
the Sierra do Baoruco in the north to the arid plain that forms the 
southern tip of the Peninsula de Barahona. This change appears to 
effectively inhibit genetic exchange between /I. h. ner/?zore'/ popula- 
tions demonstrated by Webster's observation of different frequen- 
cies for variants of three proteins and the modal shift in loreal rows 
and the dorsum coloration differences for the two regions docu- 
mented by the present study. On the eastern coast of the Peninsula 
de Barahona (Barahona to Enriquillo) wetmorei modally has five 
loreal rows and a greenish tan dorsum in association with the 
orange, red-orange, and bicolored orange with a thin yellow margin 
dewlap color. To the south and west (Oviedo to Marigot) wetmorei 
modally has four loreal rows and no greenish dorsa have been 
recorded. 

A. wehsteri differs from other populations in the hrevirostris 
complex by the unique trimode of four, five, and six postmentals 
(others modally four or five, or bimodally five and six). A. wehsteri 
and each of the three subspecies of A. hrevirostris overlap in dewlap 
color variation. Dewlaps uniformly orange and bicolored orange 
with a thin yellow margin are found to varying degrees in the distri- 
bution of each species. The subspecies of A. hrevirostris are further 
distinguishable from A. wehsteri by their unique modal scale counts 
noted above. 



28 BREVIORA No. 461 

The distribution presented in this paper follows closely that of 
Webster and Burns (1973), but varies from that of Webster ( 1978a). 
Webster noted electrophoretic similarities between A. caudalis and 
the Jacmel-Marigot populations and between A. caudalis and the 
San Juan-San Jose de Ocoa populations. He implied that the 
Jacmel-Marigot populations be referred to caudalis. Here the 
southwestern Tiburon populations are designated as a new species, 
A. marron. For the hrevirostris complex, marron varies greatly 
from caudalis in modal scale counts, dewlap color, and in relation- 
ships to A. distichus. Modal scale count variations are: scales across 
snout (four versus five), loreal rows (five versus four), scales betwen 
supraorbital semicircles and interparietal (0/0 versus 1/1), median 
head scales (bimodes one and two versus mode two), postmentals 
(bimodes five and six versus mode five), and preoccipital condition 
(modally a series of small scales bordering interparietal anteriorly 
versus preoccipital modally present). The dark olive-gray to tan 
with reddish centered dewlap of marron distinguishes it not only 
from caudalis but is unique in the whole hrevirostris complex. A. 
caudalis excludes A. distichus dominicensis where A. marron and 
A. distichus are sympatric. Webster also noted shared alleles of 
marron with A. h. wetmorei at Bell Anse. 

The northern Dominican populations having electrophoretic sim- 
ilarities to A. caudalis are designated as a new subspecies, A. b. 
desert icola. A. h. desert icola and A. marron have the same scale 
count modes, and thus the same substantial difference from cauda- 
lis. Webster (1978b) noted variation in hybridization oi A. distichus 
with A. h. hrevirostris from that with A. h. desert icola. However, 
neither this variation nor the electrophoretic similarities discour- 
aged Webster from associating deserticola with hrevirostris. 

A. marron would then appear to be an extension of "/I. hreviros- 
tris" from the Peninsula de Barahona, a pattern found in other 
reptilian species (Ameiva leheri, Ameiva lineolata privigna, Ameiva 
taeniura vulcanalis). 

APPENDIX 1 

1) Anolis hrevirostris hrevirostris: Haiti. Dipt, de l-Ouest, Source 
Trou Caiman (MCZ 124737-49); Thomazeau (MCZ 13769-70, 
MCZ 59388-89, MCZ 124705); between Thomazeau and Manne- 
ville (MCZ 59381); Manneville (MCZ 59382-87, MCZ 63154-58, 



1979 GEOGRAPHIC VARIATION IN /1.VO^/5 29 

MCZ 121777-78, MCZ 123125-55) Eaux Gaillees (MCZ 59943-48, 
MCZ 63159-71, MCZ 118790-92, MCZ 121096-101); Ganthier 
(MCZ 124706-22); Dominican Republic, Independencia Prow, Las 
Lajas (AMNH 41456, AMNH 41458-59, AMNH 41461); La Descu- 
bierta (AMNH 41433, AMNH 41437-38); Las Baitoas (AMNH 
50149-50, AMNH 50199, AMNH 50210-16); 8.8 km W Duverge 
(MCZ 125541); 2 km SW Duverge (UF/FSM 42738). 

2) Anolis brevirostris deserticola: Dominican Republic, Azua 
Prow, 4 km SW Sabana Yegua, 420 m(UF FSM 42739); Barahona 
Prov., Cabral (MCZ 58511-25, MCZ 58526-36); 5 km S Cabral 
(MCZ 5850'9-10); La Cueva (MCZ 58425, MCZ 58429, MCZ 58432, 
MCZ 58436); lower part of road to Polo (MCZ 125551-53, MCZ 
128219-25); 7.5 km N Canoa (MCZ 125549-50); Sierra Martin 
Garcia, 1540 m (UF FSM 36971); Peravia Prow, 2 km N San Jose 
de Ocoa (MCZ 107079-80); 1 km N San Jose de Ocoa (MCZ 
128255-56); 1 km S San Jose de Ocoa (MCZ 126012-44); 2 km S San 
Jose de Ocoa (MCZ 128248-54); Rio Limon, 8 km S San Jose de 
Ocoa (MCZ 107081-82); 5 km N Bani, 110 m (UF FSM 36972). 

3) Anolis brevirostris wetmorei: Dominican Republic. Barahona 
Prow. Barahona (AMNH 41307-15, AMNH 41317, AMNH 41322, 
AMNH 51430, AMNH 51446, AMNH 51533, MCZ 58694, MCZ 
93106, UF FSM 42733-35, USNM Field No RIC 040037-43); avia- 
tion field, Barahona (AMNH 51508-22, AMNH 51525, AMNH 
51527); Hotel Guarocuya, Barahona (MCZ 107083-86); 1.7 km S 
Barahona (MCZ 9311); SW Barahona (AMNH 51535-40); Paraiso 
(AMNH 51630-31); Hermann's finca, near Paraiso (AMNH 
516145); Los Patos (MCZ 58559-60); Pedernales Prow. Oviedo 
(viejo) (MCZ 58537-38); 2 km E turn to Cabo Rojo (MCZ 128215- 
18, MCZ 128258-59); 1 km SW Las Mercedes, 380 m (UF FSM 
42736); 2 km E Las Mercedes, 250 m (UF, FSM 42737); Haiti. Dept. 
de I'Ouest Lan Banane, near SaUrou (MCZ 68685-90). 

4) Anolis caudalis: Haiti. Dept. de I'Ouest. 6.3 km N Duvalierville 
(MCZ 123072-99); Ste. Philomene (USNM 123351-56); He de la 
Gonave. no other locality (MCZ 13783-89); Anse-a-Galets (MCZ 
29046-50, MCZ 37507-16, MCZ 85299-309); Nan Cafe (MCZ 
85225-45, UF FSM 12286-1-9, UF, FSM 12287-1-9, UF FSM 
12288, UF FSM 12290); Pointe a Raquettes (MCZ 25509-18, MCZ 
80613-38, UF/FSM 12291-1-2); vicinity of Pointe a Raquettes 
(MCZ 80581-86); Ti Roche, 9.5 km Pointe a Raquettes (MCZ 
80587-612); Nan Palmiste, 4 km from Pointe a Raquettes (MCZ 



30 



BREVIORA 



No. 461 




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1979 GEOGRAPHIC VARIATION IN /(.VC>Z./5 31 

80639-58); Nan Saline (MCZ 85169-74, MCZ 85175-81); Tete 
Source (MCZ 85182-205); Platon Yeye (MCZ 85206-24); Lan 
Coupe (MCZ 85281-98); He de la Petite Gonave (AMNH 49745, 
AMNH 49747-48). 

5) Anolis wehsteri: Haiti, Dept. de I'Artibonite, Marche aux 
Poteaux (MCZ 125117-25); Passe Reine, between GonaTves and 
Ennery (MCZ 63141-42); Pont de 1-Estere (MCZ 63135-40); bridge 
over Riviere de TArtibonite, St. Marc road (MCZ 59355-80); 
Lafond (MCZ 118783-89, MCZ 123101-24, MCZ 125110-16); St. 
Marc (AMNH 42593, AMNH 49696, AMNH 49699-700, AMNH 
49702-04, 'AMNH 49708-09, AMNH 49711-12, AMNH 77566, 
USNM 59219); Montrouis (MCZ 125126-69). 

Literature Cited 

Barbour, T. 1937. Third list of Antillean reptiles and amphibians. Bull. Mus. 
Comp. Zool., 82(2):77-166. 

Cochran, D. M. 1941. The herpetology of Hispaniola. Bull. U.S. Nat. Mus.. 177: 
vii + 398 pp. 

Maerz, a., and M. R. Paul. 1950. A dictionary of color. New York. McGraw- 
Hill, Book Co., N.Y.. vii + pp. 1-23 and 137-208, 56 pis. 

Mertens, R. 1939. Herpetologische Ergebnisse einer Reise nach der Insel His- 
paniola, Westindien. Abh. senckenberg. naturf. Ges. (449): 1-84. 

Oliver, J. A. 1948. The anoline lizards of Bimini, Bahamas. Amer. Mus. 
Novitates (1383): 1-36. 

Schwartz, A. 1968. Geographic variation in Anolis distichiis Cope (Lacertilia, 
Iguanidae) in the Bahama Islands and Hispaniola. Bull. Mus. Comp. Zool., 137 
(2):255-309. 

Webster, T. P. 1978a. Geographic variation in "Anolis hrevirosiris": evidence 
from proteins and a consideration of dewlap color. Third Anolis Newsletter: 
153-164. 

1978b. Hybridization of Hispaniolan lizards in the Anolis disiichus 

species group. Third Anolis Newsletter: 166 170. 

and J. Burns. 1973. Dewlap color variation and electrophoretically 



detected sibling species in a Haitian lizard Anolis hrevirosiris. Evolution. 27: 

368-377. 
Williams, E. E. 1976. West Indian anoles: a ta.xonomic and evolutionary 

summary. I. Introduction and species list. Breviora, 440: 1-21. 
and a. S. Rand. 1977. Species recognition, dewlap function and faunal 

size. Amer. Zool., 17: 261-270. 



-iOC- 



ML'S. COMP. ZOOL 
LIRRARY 



B R E V*I'0 R A 

MiiseiiinQ of ConiparatiTe Zoology 

us ISSN 0006 9698 

Cambridge, Mass. June 30, 1980 Number 462 



ELEVTHERODA CTYLVS EREMITVS, 

A NEW TRANS-ANDEAN SPECIES OF 

THE LACRIMOSUS ASSEMBLY FROM ECUADOR 

(AMPHIBIA: LEPTODACTYLIDAE) 

John D. Lynch' 

Abstract: Eleutherodactylus eremiius is named from cloud forests along the Rios 
Mindo, Pilaton. and Saloya, in Provincia Pichincha. Ecuador, between 1540 and 
2100 m. The new frog is most closely related to £. bromeliaceus from the Amazonian 
versant of the Andes in southern Ecuador. 

Lynch and Duellman (1980) included the following four species 
(all cis-Andean) in the lacrimosiis assembly of the imistrigatus 
group o{ Eleutherodactylus — E. bromeliaceus Lynch, E. lacrimosus 
(Jimenez de la Espada), E. mendax Duellman, and E. petersi Lynch 
and Duellman. Eleutherodactylus lacrimosus is distributed below 
1000m from eastern Ecuador east to the mouth of the Amazon river 
(Lynch and Schwartz, 1972; Lynch, 1979), whereas the other three 
species occur on the eastern slopes of the Andes — E. petersi, from 
the headwaters of the Rio Magdalena (Colombia) south to the val- 
ley of the Rio Pastaza (Ecuador); E. bromeliaceus. south of the 
Pastaza Trench to the Cordillera de Zamora in southern Ecuador; 
and E. mendax. from central Peru (Huanuco) south into adjacent 
Bolivia. 

In 1970 while collecting on the Pacific slopes in Andean Ecuador, 
I found a small, green frog in a bromeliad. Subsequent study of the 
collections of the late James A. Peters provided two additional 
examples from nearby localities. Additional specimens were 
obtained by William E. Duellman, Kenneth Miyata, and myself in 
the course of field work between 1975 and 1978. Sufficient material 
is now available to provide a description of the first trans-Andean 
representative of the lacrimosus assembly. 



'School of Uife Sciences, The University of Nebraska, Lincoln, Nebraska 68588 



2 BREVIORA No. 462 

Eleutherodactylus eremiius sp. nov. 

Holotype: MCZ 92103, a gravid female, taken on the La Palma- 
Chiriboga road (hwy 28), 25.7 km above (NE) La Palma, Prov. 
Pichincha, Ecuador, 1820 m, on 12 July 1976 by Kenneth Miyata. 

Paratypes (all from Prov. Pichincha, Ecuador). Topotype, MCZ 
92104; 2 km W Campamento Silante (La Palma-Aloag road), 
2100m. KU 140878; 5 km ESE Chiriboga, Quebrada Zapadores, 
2010m, KU 179085-86, 180248; km 58, ± 8 km W Chiriboga, 
USNM 211209; 3.5 km NE Mindo, 1540m, KU 165884; 6.2 km E 
Tandapi, 1750m, MCZ 92105; Tandayapa, USNM 211208. 

Diagnosis. 1 ) skin of dorsum finely areolate, that of venter coarse- 
ly areolate; no dorsolateral folds; no anal sheath; 2) tympanum 
partially visible, round, its length 2/5-1/2 eye length; 3) snout sub- 
acuminate in dorsal view, rounded or weakly protruding in lateral 
profile (papilla at tip); canthus rostralis moderately distinct; 4) 
upper eyelid bearing a conical tubercle, narrower than lOD; no 
cranial crests; 5) vomerine odontophores oval in outline, elevated; 6) 
males with vocal slits, vocal sac, and non-spinous nuptial pads; 7) 
first finger shorter than second; all fingers with broad discs, pads on 
II-IV expanded; numerous supernumerary palmar tubercles; 8) fin- 
gers bear lateral fringes; 9) 2-3 small, subconical ulnar tubercles; 10) 
small tubercles on knee, heel, outer edges of tarsus; 1 1 ) two metatar- 
sal tubercles, inner oval, 4 times size pungent outer; numerous 
supernumerary plantar tubercles; 12) toes bear lateral fringes, broad 
discs, dilated pads; pads smaller than those of fingers; 13) cream 
with faint brown stippling; vague canthal-supratympanic and flank 
stripe; colorless areas in groin and on posterior surfaces of thighs; 
14) adults small, males 17.2-20.0 (x=18.9, N=6) mm, one aduh 
female 27.6 mm SVL. 

Eleutherodactylus eremitus most closely resembles E. mendax but 
differs in having areolate skin on the dorsum and more prominent 
vomerine odontophores (odontophores absent or indistinct in E. 
mendax). Both have larger tympana than do E. hromeliaceus, E. 
lacrimosus, and E. peter si. 

Description. Head broader than body (except in gravid female), 
wider than long; H W 35. 1-38.4 (x=36.4, N=8) per cent SVL; snout 
subacuminate in dorsal view with conical papilla at tip, in profile 
rounded or truncate but with papilla, sometimes protruding; snout 
moderately long, E-N in males 81.5-88.5 (x=85.4, N=4) per cent 



1979 ELELTHERODACTVLUS EREMITUS SP. NOV. 3 

eye length, in females 86.4-100.0 (x=95.6, N=4) per cent; nostrils 
weakly protuberant, directed dorsolaterally; canthus rostralis mod- 
erately distinct, straight; loreal region weakly concave, sloping grad- 
ually to lips, lips not flared; upper eyelid width 70.4-93.4 (x=81.5, 
N=8) per cent lOD, bearing one conical tubercle in center (Fig. 
la-b) and several low warts (areolations) posterolaterally; no cran- 
ial crests; tympanum moderately distinct, annulus visible through 
thin skin, directed dorsolaterally, round, its length 42.3-50.0 (x= 
47.2, N=8) per cent eye length, separated from eye by distance equal 
to tympanum length; supratympanic fold indistinct amidst areola- 
tions; large, subconical, postrictal tubercles posterolateral to 
tympanum. 

Choanae round, large, not concealed by palatal shelf of maxillary 
arch; vomerine odontophores elevated, median and posterior to 
choanae, oval in outline, separated on midline by distance equal to 
1-1/2 odontophore widths, each bearing transverse row of 4-5 
teeth; tongue longer than wide, bearing a shallow notch along poste- 
rior border or not, posterior 2/5 not adherent to floor of mouth; 
males with vocal slits posterolateral to tongue; vocal sac median, 
subgular, external. 

Skin of dorsum finely areolate (most obvious paravertebrally and 
on upper flanks), venter coarsely areolate; throat areolate in 
females, no dorsolateral folds or anal sheath; larger warts posterior 
and posterolateral to vent; discoidal fold well anteriad to groin. 

Three to four small, subconical ulnar tubercles; palmar tubercle 
bifid, much larger than oval thenar tubercle; palm areolate, but 1-2 
more pungent supernumerary tubercles evident at base of each fin- 
ger; subarticular tubercles pungent, round (or slightly broader than 
long); KU 179086 has a bifid basal subarticular tubercle on finger 
III; fingers bear prominent lateral fringes; similar fringe on outer 
edge of IV continuing along most of palm (Fig. 1); fingers bear 
broad discs, pads (except on thumb) moderately large, apically 
rounded; first finger shorter than second; thumbs of males swollen, 
bearing white, non-spinous nuptial pads. 

Small (but prominent), subconical tubercles on knee, heel, outer 
edge of tarsus; less distinct tubercle on inner edge of tarsus; inner 
metatarsal tubercle twice as long as wide, relatively flat; outer sub- 
conical, 1/4 size of inner; numerous supernumerary plantar tuber- 
cles (some more prominent than others, in rows); subarticular 
tubercles round to slightly longer than wide; toes bear prominent 



BREVIORA 



No. 462 




1979 ELEUTHERODACTYLUS FREMITUS SP NOV 5 

lateral fringes, no webbing; toe pads smaller than those of fingers, 
apically rounded, bearing broad discs; heels of flexed hind limbs 
overlap; shank 47.1-55.0 (x=51.9, N=8) per cent SVL. 

Most individuals are cream with diffuse brown stippling above 
and less below; pigmentless areas in groin and on posterior surfaces 
of thighs (bordered by more dense brown stippling); no labial or 
limb bars; indefinite canthal-supratympanic stripes, continuing as a 
diffuse band onto anterior flanks. KU 140878 and 179086 have dark 
brown dorsolateral stripes continuous with canthal-supratympanic 
stripe (Fig.^2). KU 179085 is golden brown above with a brown 
blotch in the center of the back and prominent canthal-supra- 
tympanic stripes. 

In life, E. eremitus is green above with a brown to reddish brown 
head and white below (throat stippled with brown). The groin and 
posterior surfaces of the thighs are pale yellow. The iris is bright 
copper with brown flecks. 

Measurements ofholotype in mm. SVL 27.6; shank 13.0; HW 9.7; 
head length 8.5; upper eyelid width 2.8; lOD 3.0; tympanum length 
1.5; eye length 3.1; E-N 3.1. 

Etymology. Latin, meaning lonely or solitary, in allusion to its 
relatives which are all cis-Andean. 

Natural history. KU 140878 was found during the day in a large 
terrestrial bromeliad beside hwy 30, before the habitat was severely 
altered by road construction. Specimens from the Quebrada Zapa- 
dores were collected at night as they sat on broad leaves in the forest 
beside the streams. Eleutherodactylus eremitus is either quite uncom- 
mon or several collectors have failed to discover its preferred micro- 




HRF 




B 









Figure 2. Eleutherodactylus eremitus. (A) KU 179085, male. 19,8 SVL; (B) KU 
179086, juvenile female, 23.0 mm SVL. 



6 BREVIORA No. 462 

habitat in the cloud forests (on only one occasion has more than one 
specimen been found [Miyata secured two on a single night in the 
type-locality]). 

Remarks. Frogs of the lacrimosus assembly have been discovered 
slowly. Eleutherodaciylus lacrimosus was named in 1875 but not 
reported again until 1952 (Lutz and Kloss, 1952). Duellman (1978), 
Lynch (1979), and Lynch and Duellman (1980) each named species 
found on the Amazonian slopes of the Andes from Colombia to 
Bolivia, but the first specimens of these were not obtained until the 
late 1950's or early 1960's. In eastern Ecuador, these small frogs are 
especially common in arboreal bromeliads. In July 1977 at Mera 
(Prov. Pastaza), at least a dozen examples of E. lacrimosus and E. 
petersi were found in one bromeliad in remaining patches of forest 
along the Rio Pastaza. The bromeliad microhabitat is occupied by a 
variety of frogs on the Pacific versant. At the base of the Andes, E. 
suhsigillatus (Boulenger) normally is found only in arboreal brome- 
liads. In high cloud forests, E. celator Lynch, E. phoxocephalus 
Lynch, and E. thymelopsoides Lynch are found almost exclusively 
in bromeliads. I suspect E. eremitus is actually more abundant on 
the Pacific versant between 1500 and 2100 m than is suggested by 
the ten specimens now available. Too infrequently I have failed to 
exploit the bromeliad microhabitat betwen 800 and 2400 m, in part 
because frogs were abundant at night and I collected infrequently by 
day. Inspection of only three bromeliads in July 1977 at Tandapi 
(Prov. Pichincha) yielded one example each of E. crucifer (Bou- 
lenger) and E. parvillus Lynch. 

Eleutherodaciylus bromeliaceus seems to be the closest known 
relative of E. eremitus, with E. mendax being less closely related. 
Eleutherodaciylus lacrimosus and E. petersi are considered more 
closely allied to one another than either is to the other three species. 

ACKNOWLEDGMENTS 

Specimens were loaned by: William E. Duellman, Museum of 
Natural History, University of Kansas (KU); W. Ronald Heyer, 
National Museum of Natural History (USNM); and Ernest E. Willi- 
ams, Museum of Comparative Zoology (MCZ). Field work in Ecua- 
dor was supported by funds from the American Philosophical 
Society (Penrose Fund) and the University of Nebraska Research 
Council. David Cannatella and Marsha Lynch aided me in field 
work. 



1979 ELEUTHERODACTYLUS EREM/TUS SP^NOV. 7 

LITERATURE CITED 

DuELLMAN, W. E. 1978. Three new species ot /:7('(/?/;t'r(;>(/arM7i« from Amazonian 

Peru (Amphibia: Anura: Leptodactylidae). Herpetologica 34: 264-270. 
LuTZ, B. & Kloss, G. R. 1952. Anfibios anuros do alto Solimoes e Rio Negro 

apontamentos sobre algumas formas e suas vicariantes. Mem. Inst. Oswaldo 

Cruz 50: 626-78. 
Lynch, J. D. 1979. Leptodact\'lid frogs of the genus Eleuiherodactylus from the 

Andes of southern Ecuador. Misc. Publ. Mus. Nat. Hist. Univ. Kansas (66): 1- 

62. 
Lynch, J. D. & Dieli.m.^n, W. E. 1980. The Eleuiherodactylus of the Amazonian 

slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Misc. Publ. Mus. 

Nat. Hist. Univ. Kansas, in press. 
Lynch, J. D. & Sch\v.\rtz, A. 1972. Ta.xonomic disposition of some 19th 

century leptodactylid frog names. J. Herpetol. 5: 103-114. 



f^U^. comp zctqi 



B R E V rtJ R- A 

MAR 18 1985 

Miiseum of Conipan^feV^^QZoology 



us ISSN 0006 9698 



^[:^|VKH«,Ty 



Cambridge, Mass. August 15, 1980 Number 463 

JAW MUSCULATURE OF THE WEST INDIAN SNAKE 

ALSOPHIS CANTHERIGERUS BROOKSI 

(COLUBRIDAE, REPTILIA). 

Kenneth V. Kardong^ 
ABSTRACT 

The West Indian colubrid snake Alsophis cantherigerus brooksi is presently 
assigned to the subfamily Xenodontinae (sensu Maglio, 1970). It is a member of an 
assemblage of related snakes from continental stocks that have spread to and diversi- 
fied within this Caribbean archipelago. The head myology proves to be representa- 
tive of other .xenodontine snakes of the West Indies and thus is a convenient reference 
for comparison to other subspecies and may be useful in later comparisons to other 
species" assemblages of the region. Its jaw musculature conforms basically to that of 
other generalized colubrid snakes. Special attention is given to the internal fascicular 
bundles within each muscle. 

The adductor mandibulae externus superficialis passes posteroventrally from its 
origin on the skull, inserting mostly by a broad aponeurosis except, for a rostral slip 
that forms its anterior edge and inserts directly on the mandible. The adductor 
mandibulae externus profundus consists of anterior and posterior wedges of muscle 
delineated by a vertical suture between them. Adductor mandibulae externus media- 
lis is separable into superficial and deep slips. One significant difference in general 
muscle anatomy from some other colubrids concerns the protractor quadrati. This 
muscle inserts on the retroarticular process of the mandible which differs from the 
genus Opheodrys in which it is reported to insert on the quadrate. 

Lateral jaw musculature proves to be very similar within the subspecies of Alsophis 
cantherigerus. The only detectable difference occurs in the deep division of the 
depressor mandibulae. In most subspecies (adspersus. brooksi. cantherigerus, cayma- 
nus, schwartzi), this deep division is a single slip of the depressor arising from a single 
site. However, in fuscicauda. pepei, and rutiyi it splits lengthwise so as to arise from 
two sites of origin. One instance of individual asymmetry was identified in adspersus 
wherein the insertion of the superficialis differs on left and right sides. 

The conformity of lateral jaw musculature among subspecies related to brooksi 
emphasizes the uniform nature of this group. Other authors also found this unifor- 
mity in features of the integument, skull osteology, and hemipenial characteristics. 

1 Department of Zoology 
Washington State University 
Pullman, Washington 99164 



BREVIORA No. 463 



INTRODUCTION 



Examination of the West Indian colubrid snakes of the subfamily 
Xenodontinae by Maglio (1970) served to emphasize the special 
island biogeographic features of these species. Compared to conti- 
nental species. West Indian insular members of this subfamily have 
well delineated distributions and their probable origin, namely from 
mainland genera, seems better established. Routes of initial coloni- 
zation by and diversification within the different species assem- 
blages through this Caribbean archipelago were proposed by Maglio 
as well. With a view to testing this and related proposals on the 
phylogeny of this group, I examined the jaw musculature of 
members from the various assemblages, following any changes 
through the various species on the different islands. This work and 
its implications will be the subject of later reports. The present paper 
is restricted to the species Alsophis cantherigerus and, in particular, 
the subspecies A. c. brooksi. 

This subspecies is found only on the Swan Islands, located almost 
200 kilometers north of Honduras. It deserves special attention for 
several reasons. It is part of the assemblage of closely related popu- 
lations that spread to and diversified over the islands. Its jaw mus- 
culature is representative of the condition in other West Indian 
xenodontine species. Its jaw anatomy can thus serve as a central 
point of reference in later analysis of these island snake species. It is 
also a rare subspecies even in museum collections. 

MATERIALS AND METHODS 

All descriptions of the head anatomy of Alosphis cantherigerus 
brooksi are based on a single alcohol preserved female specimen 
from the Museum of Comparative Zoology (MCZ) 1 1979. The spec- 
imen, taken on Swan Islands and donated to the collection in 1916, 
measured 85.5 cm snout-vent length. The tail, missing its severed 
tip, measured 23.4 cm. The cranial musculature was exposed 
initially by cutting through the integument along the lips and reflect- 
ing the skin backwards. This revealed that the specimen had sus- 
tained a deep cut in the neck musculature that also severed the 
vertebral column through the fourth cervical vertebra and conse- 
quently limited the description of structures in the immediate vicin- 
ity. This study centered upon the jaw musculature and therefore 
often required the disruption of nerves to first confirm muscle at- 



1979 Jaw muscul&iUTe of Alsophis cantherigerus brooksi 3 

tachments with certainty. As a result, some of the peripheral distri- 
butions of the nerves were lost before they could be traced with 
confidence. The blood vessels proved impractical to follow for the 
same reasons. Consequently, only those pathways and distributions 
of nerves and blood vessels that could later be confirmed are 
included in the descriptions. No bilateral asymmetry in jaw muscu- 
lature was detected. Muscle and ligament terminology follows Haas 
(1973), Kardong (1973, 1974), and Pregill (1977); interpretation of 
cranial nerves relies upon Auen and Langebartel (1977); and 
cephalic gland nomenclature is based on Taub (1966). One depar- 
ture in nomenclature concerns the muscle running between the neck 
and dorsal end of the quadrate, which is often referred to as M. 
retractor quadrati especially in descriptions of venomous snakes 
(e.g. DuUemeijer, 1956). Others employ this same term for a quite 
different muscle (e.g. Kochva, 1962). To avoid confusion herein, the 
term M. cervicoquadratus is used instead. 

The other subspecies oi Alsophis cantherigerus examined include: 
adspersus (MCZ 68727), cantherigerus (MCZ 13288), caymanus 
(MCZ AA'^m^fuscicauda (MCZ 44875), pepe/ (MCZ 13289), ruttyi 
(MCZ 44877), and schwartzi (MCZ 56430). 

GENERAL DESCRIPTION 

Cephalic Glands 

The nasal glands, like other lateral cephalic glands, are bilaterally 
paired. Each rests in an anterior depression on the anteromedial 
face of the prefrontal and is bounded medially by a dorsal process of 
the respective septomaxilla and by the nasal capsule. The Harderian 
gland (Figs. 2, 3a, 4b) is found ventral to and extends behind the 
eye. Beneath the eye the gland lies within the orbit and is com- 
pressed between eyeball and the ligamentum orbitale interioris (sep- 
tum interorbitale of DuUemeijer, 1956) which forms the ventral 
floor of the orbit. Passing posteriorly out of the orbit it enlarges, 
occupying the space lateral to the levator pterygoidei muscle and 
medial to the postorbital bone and the adductor mandibulae exter- 
nus superficialis muscle. The infralabial glands (Fig. 4c) extend pos- 
teriorly along the lateral aspect of the dentary and compound bones 
to approximately the level of the anterior insertion of the adductor 
mandibulae externus superficialis muscle. The supralabial glands 



4 BREVIORA No. 463 

extend posteriorly slightly farther. However, they lie along the 
upper lips and pass lateral to the superficialis muscles and maxillae. 
Pressed into the side of each supralabial gland and residing behind 
the eye is the large Duvernoy's gland (Fig. 4c), which in the pre- 
served state appears lighter in color and more distinctly lobed. The 
small, unpaired premaxillary gland lies beneath the upper lip on the 
superficial face of the premaxillary bone. 

In the chin, the sublingual gland (Fig. 6a, b) is represented by 
three parts, a single medial and paired lateral glands. The medial 
sublingual gland is stationed along the ventroanterior end of the 
tongue and anchored to its connective tissue sheath. The lateral 
sublingual glands insert into the oral epithelium and serve as attach- 
ment sites for the pars glandularis of the transversus branchialis and 
the protractor larygeus muscles. 

Ligaments 

A number of small, unnamed ligaments link the cranial elements 
together, but only the prominent or most commonly cited in the 
literature are discussed here. Two primary ligaments serve to check 
the motion of the supratemporal relative to the braincase. One is the 
parieto-supratemporal ligament that runs from the dorsal edge of 
the supratemporal to the dorsal surface of the exoccipital. It also 
serves as the site of origin for parts of the adductor mandibulae 
externus medialis and depressor mandibulae muscles. The other is 
the prooto-supratemporal ligament that attaches to the ventral edge 
of the supratemporal and to the adjacent region of the braincase. 
Rotation of the prefrontal is controlled in part by the fronto- 
prefrontal ligament (Fig. 2) from the ventromedial corner of the 
prefrontal to the adjacent parasphenoid. 

Several prominent ligaments are associated with the palato- 
maxillary arch (pterygoid, ectopterygoid, maxilla, and palatine). 
Within the arch, the intramaxillary ligament (Fig. 2) reaches from 
the medial ectopterygoid process of the maxilla forward to attach 
jointly on the palatine process of the maxilla and maxillary process 
of the palatine. The short, fan-shaped quadrato-pterygoid ligament 
runs from the narrow posterior end of the pterygoid to a wider 
medial attachment site along the mandible that includes the retroar- 
ticular process and extends below the articular notch. The strong, 
cord-like maxillo-postorbital ligament (Fig. 4) reaches from the 



1979 Jaw musculature of Alsophis cantherigerus brooksi 5 

downward directed tip of the postorbital to the lateral side of the 
maxilla just anterior to its articulation with the ectopterygoid. The 
quadrate-maxillary ligament (Fig. 4) begins along the laterodorsal 
edge of the retroarticular process, passes forward above the lateral 
condyle of the quadrate, and divides. The smaller division attaches 
to the nearby dermis beneath the most posterior supralabial scale, 
whereas the larger division continues forward to attach to the pos- 
terolateral region of the maxilla. 

Cranial Nerves 
Figs. 2, 3a-d, 4a-c 

Trigeminal Nerve (V). The trigeminal is one of three major nerves 
supplying the jaw musculature. Trigeminal branches that supply the 
jaw musculature exit from the cranium through two openings in the 
prootic bone, the maxillary and mandibular foramina. Five 
branches could be found passing through the maxillary foramen. 
The largest, the maxillary division (V^), passes lateral to the pseudo- 
temporalis then bends forward continuing in an anterior path dorsal 
to the origin of the pterygoideus. A second nerve divides lateral to 
the pseudotemporalis with one branch passing laterally to glandular 
tissue along the upper Hp and the second swinging dorsally, superfi- 
cial to the facial vein, to enter the ventral surface of the adductor 
mandibulae externus superficialis. Two other nerves also course 
dorsally after exiting from the foramen. Both pass deep to the facial 
vein, but one enters the medial face of the adductor mandibulae 
externus medialis (deep division) and the other enters the medial 
face of the adductor mandibulae externus superficialis. The fifth 
nerve leaving this foramen passes laterally. 

Through the mandibular foramen exit six branches of the trigemi- 
nal nerve. The largest branch is the mandibular division (Vg) that 
passes posteriorly and then turns ventrally following a route that 
carries it between the adductor mandibulae externus profundus and 
the adductor mandibulae posterior. Eventually it enters a foramen 
in the mandible located at the anterior end of the mandibular fossa. 
A second branch of the trigeminal courses posteriorly, then swings 
dorsally, bifurcates, and enters the medial face of the adductor man- 
dibulae externus profundus and medioposterior face of the adduc- 
tor mandibulae externus medialis (superficial division). A third 
nerve passes posteriorly deep to the mandibular division. It sends 



6 BREVIORA No. 463 

one branch between medial and lateral divisions of the adductor 
mandibulae posterior, another between the pars minimus and the 
lateral division of adductor mandibulae posterior, and several 
branches into the medial surface of the adductor mandibulae exter- 
nus profundus. The remaining three nerves of the trigeminal that 
exit through the mandibular foramen pass ventrally — one disap- 
pears between the pterygoideus and the levator pterygoidei, one 
enters the lateral face of the pseudotemporalis, and the last runs 
laterally into the glandular tissue along the upper lip. 

Facial Nerve (VII). Two divisions of the facial nerve exit through 
the mandibular foramen. The larger is the hyomandibular nerve 
that passes posteriorly over the columella, under the quadrate, and 
enters the depressor mandibulae. A slender communication arising 
from the base of the hyomandibular travels posteriorly and merges 
with craniocervical trunk. The second division of the facial nerve is 
the palatine nerve. It courses ventrally along the underside of the 
braincase, is joined by the cranial sympathetic nerve, and then 
enters the posterior Vidian canal in the ventral aspect of the basi- 
sphenoid bone. 

Craniocervical Trunk. The glossopharyngeal (IX), vagus (X), and 
hypoglossal (XII) cranial nerves combine to form a single large 
nerve or plexus, the craniocervical trunk (Auen and Lagebartel, 
1977). In addition, a slender communication runs between the base 
of the hyomandibular and this trunk. Finally, a spinal nerve (pre- 
sumably the ventral ramus of the first spinal nerve) emerges from 
between the rectus capitus anterior pars dorsalis and longissimus 
dorsi (ventral head) cervical muscles to join with the craniocervical 
trunk. The trunk sweeps around the side of the body at about the 
angle of the jaws and supplies various members of the hypobran- 
chial musculature. 

Lateral Jaw Musculature 

M. cervicomandibularis (cm) 
Figs. 1, 4c 

The epimysium of the spinalis-semispinalis complex near the dor- 
sal midline serves as the surface of origin. This origin extends along 
a line even with neural spines 3 to 8 in the nape of the neck and 



1979 J SLV/ musculsituTe of A Isophis cant herigerus brooksi 3 

tachments with certainty. As a result, some of the peripheral distri- 
butions of the nerves were lost before they could be traced with 
confidence. The blood vessels proved impractical to follow for the 
same reasons. Consequently, only those pathways and distributions 
of nerves and blood vessels that could later be confirmed are 
included in the descriptions. No bilateral asymmetry in jaw muscu- 
lature was detected. Muscle and ligament terminology follows Haas 
(1973), Kardong (1973, 1974), and Pregill (1977); interpretation of 
cranial nerves relies upon Auen and Langebartel (1977); and 
cephalic gland nomenclature is based on Taub (1966). One depar- 
ture in nomenclature concerns the muscle running between the neck 
and dorsal end of the quadrate, which is often referred to as M. 
retractor quadrati especially in descriptions of venomous snakes 
(e.g. Dullemeijer, 1956). Others employ this same term for a quite 
different muscle (e.g. Kochva, 1962). To avoid confusion herein, the 
term M. cervicoquadratus is used instead. 

The other subspecies oi Alsophis cantherigerus examined include: 
adspersus (MCZ 68727), cantherigerus (MCZ 13288), caymanus 
(MCZ 44^m, fuscicauda (MCZ 44^15), pepei (MCZ 13289), ruttyi 
(MCZ 44877), and schwartzi (MCZ 56430). 

GENERAL DESCRIPTION 

Cephalic Glands 

The nasal glands, like other lateral cephalic glands, are bilaterally 
paired. Each rests in an anterior depression on the anteromedial 
face of the prefrontal and is bounded medially by a dorsal process of 
the respective septomaxilla and by the nasal capsule. The Harderian 
gland (Figs. 2, 3a, 4b) is found ventral to and extends behind the 
eye. Beneath the eye the gland lies within the orbit and is com- 
pressed between eyeball and the ligamentum orbitale interioris (sep- 
tum interorbitale of Dullemeijer, 1956) which forms the ventral 
floor of the orbit. Passing posteriorly out of the orbit it enlarges, 
occupying the space lateral to the levator pterygoidei muscle and 
medial to the postorbital bone and the adductor mandibulae exter- 
nus superficialis muscle. The infralabial glands (Fig. 4c) extend pos- 
teriorly along the lateral aspect of the dentary and compound bones 
to approximately the level of the anterior insertion of the adductor 
mandibulae externus superficialis muscle. The supralabial glands 



4 BREVIORA No. 463 

extend posteriorly slightly farther. However, they lie along the 
upper lips and pass lateral to the superficialis muscles and maxillae. 
Pressed into the side of each supralabial gland and residing behind 
the eye is the large Duvernoy's gland (Fig. 4c), which in the pre- 
served state appears lighter in color and more distinctly lobed. The 
small, unpaired premaxillary gland lies beneath the upper lip on the 
superficial face of the premaxillary bone. 

In the chin, the sublingual gland (Fig. 6a, b) is represented by 
three parts, a single medial and paired lateral glands. The medial 
sublingual gland is stationed along the ventroanterior end of the 
tongue and anchored to its connective tissue sheath. The lateral 
sublingual glands insert into the oral epithelium and serve as attach- 
ment sites for the pars glandularis of the transversus branchialis and 
the protractor larygeus muscles. 

Ligaments 

A number of small, unnamed ligaments hnk the cranial elements 
together, but only the prominent or most commonly cited in the 
literature are discussed here. Two primary ligaments serve to check 
the motion of the supratemporal relative to the braincase. One is the 
parieto-supratemporal ligament that runs from the dorsal edge of 
the supratemporal to the dorsal surface of the exoccipital. It also 
serves as the site of origin for parts of the adductor mandibulae 
externus medialis and depressor mandibulae muscles. The other is 
the prooto-supratemporal ligament that attaches to the ventral edge 
of the supratemporal and to the adjacent region of the braincase. 
Rotation of the prefrontal is controlled in part by the fronto- 
prefrontal ligament (Fig. 2) from the ventromedial corner of the 
prefrontal to the adjacent parasphenoid. 

Several prominent ligaments are associated with the palato- 
maxillary arch (pterygoid, ectopterygoid, maxilla, and palatine). 
Within the arch, the intramaxillary ligament (Fig. 2) reaches from 
the medial ectopterygoid process of the maxilla forward to attach 
jointly on the palatine process of the maxilla and maxillary process 
of the palatine. The short, fan-shaped quadrato-pterygoid ligament 
runs from the narrow posterior end of the pterygoid to a wider 
medial attachment site along the mandible that includes the retroar- 
ticular process and extends below the articular notch. The strong, 
cord-like maxillo-postorbital ligament (Fig. 4) reaches from the 



1979 Jaw musculature of Alsophis cantherigerus brooksi 5 

downward directed tip of the postorbital to the lateral side of the 
maxilla just anterior to its articulation with the ectopterygoid. The 
quadrato-maxillary ligament (Fig. 4) begins along the laterodorsal 
edge of the retroarticular process, passes forward above the lateral 
condyle of the quadrate, and divides. The smaller division attaches 
to the nearby dermis beneath the most posterior supralabial scale, 
whereas the larger division continues forward to attach to the pos- 
terolateral region of the maxilla. 

Cranial Nerves 
Figs. 2, 3a-d, 4a-c 

Trigeminal Nerve (V). The trigeminal is one of three major nerves 
supplying the jaw musculature. Trigeminal branches that supply the 
jaw musculature exit from the cranium through two openings in the 
prootic bone, the maxillary and mandibular foramina. Five 
branches could be found passing through the maxillary foramen. 
The largest, the maxillary division (V2), passes lateral to the pseudo- 
temporalis then bends forward continuing in an anterior path dorsal 
to the origin of the pterygoideus. A second nerve divides lateral to 
the pseudotemporalis with one branch passing laterally to glandular 
tissue along the upper Up and the second swinging dorsally, superfi- 
cial to the facial vein, to enter the ventral surface of the adductor 
mandibulae externus superficialis. Two other nerves also course 
dorsally after exiting from the foramen. Both pass deep to the facial 
vein, but one enters the medial face of the adductor mandibulae 
externus mediaUs (deep division) and the other enters the medial 
face of the adductor mandibulae externus superficialis. The fifth 
nerve leaving this foramen passes laterally. 

Through the mandibular foramen exit six branches of the trigemi- 
nal nerve. The largest branch is the mandibular division (V3) that 
passes posteriorly and then turns ventrally following a route that 
carries it between the adductor mandibulae externus profundus and 
the adductor mandibulae posterior. Eventually it enters a foramen 
in the mandible located at the anterior end of the mandibular fossa. 
A second branch of the trigeminal courses posteriorly, then swings 
dorsally, bifurcates, and enters the medial face of the adductor man- 
dibulae externus profundus and medioposterior face of the adduc- 
tor mandibulae externus mediaUs (superficial division). A third 
nerve passes posteriorly deep to the mandibular division. It sends 



6 BREVIORA No. 463 

one branch between medial and lateral divisions of the adductor 
mandibulae posterior, another between the pars minimus and the 
lateral division of adductor mandibulae posterior, and several 
branches into the medial surface of the adductor mandibulae exter- 
nus profundus. The remaining three nerves of the trigeminal that 
exit through the mandibular foramen pass ventrally — one disap- 
pears between the pterygoideus and the levator pterygoidei, one 
enters the lateral face of the pseudotemporalis, and the last runs 
laterally into the glandular tissue along the upper lip. 

Facial Nerve (VII). Two divisions of the facial nerve exit through 
the mandibular foramen. The larger is the hyomandibular nerve 
that passes posteriorly over the columella, under the quadrate, and 
enters the depressor mandibulae. A slender communication arising 
from the base of the hyomandibular travels posteriorly and merges 
with craniocervical trunk. The second division of the facial nerve is 
the palatine nerve. It courses ventrally along the underside of the 
braincase, is joined by the cranial sympathetic nerve, and then 
enters the posterior Vidian canal in the ventral aspect of the basi- 
sphenoid bone. 

Craniocervical Trunk. The glossopharyngeal (IX), vagus (X), and 
hypoglossal (XII) cranial nerves combine to form a single large 
nerve or plexus, the craniocervical trunk (Auen and Lagebartel, 
1977). In addition, a slender communication runs between the base 
of the hyomandibular and this trunk. Finally, a spinal nerve (pre- 
sumably the ventral ramus of the first spinal nerve) emerges from 
between the rectus capitus anterior pars dorsalis and longissimus 
dorsi (ventral head) cervical muscles to join with the craniocervical 
trunk. The trunk sweeps around the side of the body at about the 
angle of the jaws and supplies various members of the hypobran- 
chial musculature. 

Lateral Jaw Musculature 

M. cervicomandibularis (cm) 
Figs. 1, 4c 

The epimysium of the spinalis-semispinalis complex near the dor- 
sal midline serves as the surface of origin. This origin extends along 
a line even with neural spines 3 to 8 in the nape of the neck and 



1979 Jaw musculature oi Alsophis cantherigerus hrooksi 7 

partially overlaps the anterior origin of the neuromandibularis. I his 
broad muscle travels ventrally, passing over the deep ccrvico- 
quadratus, and narrows abruptly to insert on the lateral epicondyle 
of the quadrate and on the adjacent part of quadrato-maxillarv 
(dermal) ligament, which passes posteriorly to its own attachment 
on the dorsolateral retroarticular process. 

M. depressor mandibulae (dm) 

(M. occipito-quadrato-mandibularis) 

Figs. 1, 4 a-c 

There are two parts of this muscle, superficial and deep, divided 
by the cervicoquadratus that passes between these two parts en 
route to its own insertion. Branches of cranial nerve VII also pass 
through the belly of the deep division, then along the medial side of 
the superficial division. 

The superficial division has an origin that runs from the parieto- 
supratemporal ligament next to the adductor mandibulae externus 
medialis, across the supratemporal to the anterodorsal corner of the 
quadrate, and extends down along the surface epimysium of adduc- 
tor mandibulae externus profundus near its own origin. It inserts 
directly on the lateral rim of the mandibular retroarticular process, 
overlapping the attachment site of the quadrato-maxillary ligament. 

The deep division arises from the posterior tip of the supratem- 
poral and the posterior, dorsomedial edge of the quadrate. It is a 
parallel muscle that inserts on the dorsomedial surface of the retro- 
articular process. 



M. cervicoquadratus (cq) 
Figs. I, 4 a-c 

This is a long, tapering muscle that takes origin in the dermis of 
the lateral integument posterior to the belly of the neuromandibula- 
ris over the intercostal muscles. It gradually narrows as it courses 
forward, slipping under the more superficial cervicomandibularis, 
becoming a cord-like tendon that passes between the superficial and 
the deep divisions of the depressor mandibulae to insert laterally on 
the proximal end of the quadrate just below its posterior corner. 



BREVIORA No. 463 



Figure I. Lateral view of the skull and ventral view of the quadrate and lower jaw 
showing sites of muscle attachments. 

Figure 2. Ventral view of head. The bones of the left palato-maxillary arch have 

been removed to expose deep structures. On the right, the M. pterygoideus (pg) has 

been cut and partially reflected and the mandible rotated outward to better expose 
some of the underlying structures. 

Abbreviations: 

am M. adductor mandibulae externus medialis 

ap M. adductor mandibulae externus profundus 

apo M. adductor mandibulae posterior 

as M. adductor mandibulae externus superficialis 

br transverse ridge on basisphenoid 

c columella 

cm M. cervicomandibularis 

cp choanal process of palatine 

cq M. cervicoquadratus 

ct craniocervical nerve trunk (IX, X, XII) 

dm M. depressor mandibulae 

gg M. genioglossus 

gt M. geniotrachealis 

Hd Harderian gland 

ima M. intermandibularis anterior 

ipa M. intermandibularis posterior, pars anterior 

ipp M. intermandibular posterior, pars posterior 

Ifp ligamentum fronto-prefrontale 

lim ligamentum intramaxillare 

Imp ligmentum maxillo-postorbitale 

loi ligamentum orbitale inferioris 

Ip M. levator pterygoidei 

Iqm ligamentum quadrato-maxillare 

nm M. neuromandibularis 

pg M. pterygoideus 

pga M. pterygoideus accessorius 

pp M. protractor pterygoidei 

pq M. protractor quadrati 

pst M. pseudotemporalis 

red M. rectus capitus anterior, pars dorsalis 

rev M. rectus capitus anterior, pars ventralis 

rp M. retractor pterygoidei 

rv M. retractor vomeris 

V2 Trigeminal nerve, maxillary division 

V3 Trigeminal nerve, mandibular division 



1979 Jaw musculsLtme of Alsophis cantherigerus brooksi 9 





10 BREVIORA No. 463 

M. adductor mandibulae externus superficialis (as) 
Figs, 1, 3 c-d, 4 a-c 

A temporal crest, prominent posteriorly, becomes lower as it 
passes forward along the dorsolateral side of the skull. Anteriorly it 
curves outward to terminate on the postorbital process of the parie- 
tal. The superficialis originates directly from the anterior half of this 
temporal crest and from the base of the postorbital bone. It passes 
posteriorly and downward, partially overlying the adductor mandi- 

Figure 3. Lateral view of head showing progressively more structures (A-D) in 
place. 

Abbreviations: 

am M, adductor mandibulae externus medialis, deep (d) and superficial (s) 

divisions 

ap M. adductor mandibulae externus profundus 

apo M. adductor mandibulae posterior, lateral division (1), medial division (m), 

pars minimus (mi) 

as M. adductor mandibulae externus superficiaHs 

c columella 

ct craniocervical nerve trunk (IX, X, XII) 

ec ectopterygoid 

f frontal 

Hd Harderian gland 

if infralabial nerve foramen (posterior) 

ilc M. iliocostalis 

Id M. longissimus dorsi 

Ip M. levator pterygoidei 

mf mental foramen 

mx maxilla 

n nasal 

pf prefrontal 

pg M. pterygoideus 

pga M. pterygoideus accessorius 

po postorbital 

pp M. protractor pterygoidei 

pq M. protractor quadrati 

pst M. pseudotemporalis 

q quadrate 

rev M. rectus capitus anterior, pars ventralis 

ssp M. spinalis-semispinalis 

St supratemporal 

to temporal crest 

V2 Trigeminal nerve, maxillary division 

V3 Trigeminal nerve, mandibular division 



1979 Jaw musculature oi Alsophis cantherigerus brooksi 11 





J-TTPP^ 





12 



BREVIORA 



No. 463 






1979 Jaw musculature of Alsophis canlherigerus brooksi 13 

bulae externus medialis. Its anterior fibers arch around the corner of 
the mouth without forming any attachments to the buccal mem- 
brane and pass to the mandible, where they directly insert on the 
compound bone just posterior to the end of the dentary tooth row 
and lateral to the anterior insertion of the adductor mandibulae 
externus profundus. The remaining fibers of this flat muscle termi- 
nate superficial to the profundus in a broad aponeurosis that 
spreads across the surface epimysium of the profundus and can be 
followed to an insertion extending in a narrow line from the lateral 
epicondyle of the quadrate along the mandible, ventral to the inser- 
tion of the profundus, and forward to a point not quite reaching the 
directly inserting anterior fibers described above. 

M. adductor mandibulae externus profundus (ap) 
Figs. 1. 3 (^d. 4 a-c 

This is a large, triangular block of muscle divisible into two mus- 
cle wedges that separate cleanly along a vertical cleft between them. 
The anterior wedge takes origin from the lateral, anterodorsal 
corner of the quadrate. The origin of the posterior wedge also begins 
on this corner of the quadrate beneath the attachment of the ante- 
rior wedge. However, the origin of the posterior wedge extends 

Figure 4. Lateral view of skull showing progressively (A-C) more structures in 
place. 

Abbreviations: 

am M. adductor mandibulae externus medialis 

ap M. adductor mandibulae externus profundus 

as M. adductor mandibulae externus superficialis 

cm M. cervicomandibularis 

cq M. cervicoquadratus 

dm M. depressor mandibulae, deep division (d), superficial division (s) 

Dv Duvernoy's gland 

Hg Harderian gland 

ig infralabial gland 

ipp M. intermandibularis posterior, pars posterior 

Imp ligmentum maxillo-postorbitale 

Ip M. levator pterygoidei 

Iqm ligamentum quadrato-maxillare 

nm M. neuromandibularis 

pg M. pterygoideus 

sg supralabial gland 



14 BREVIORA No. 463 

distally along the anterolateral edge of the quadrate reaching the 
lateral epicondyle. Both parts form a large, thick muscle that inserts 
directly on the lateral side of the compound bone in a low depres- 
sion from below the quadrato-mandibular articulation forward to 
the posterior infralabial nerve foramen. 

M. adductor mandibulae externus medialis (am) 
Figs. 1, 3 c^d, 4 a-c 

This muscle takes origin from the parieto-supratemporal ligament 
and the posterior half of the temporal crest, filling a depression and 
area adjacent to the crest. As the muscle passes ventrally its fibers 
gather into two divisions — superficial and deep. Dorsally a few fib- 
ers intermingle between both but as they pass downward they 
become more discrete. The deep division takes origin along the 
posterior temporal crest only. This division descends as a wide, thin 
sheet that inserts directly to the compound bone medial to the 
adductor mandibulae posterior. This insertion then extends forward 
and upward to the dorsal edge of the compound bone reaching a 
point posterior to the end of the dentary tooth row. The superficial 
division contains most of the fibers of this muscle and originates 
from both the posterior temporal crest and the parieto-supra- 
temporal ligament. It narrows and gives rise to a short tendon that 
inserts on the dorsal, posterior end of the dentary. 

M. adductor mandibulae posterior (apo) 
Figs. 1, 3 c^d 

This muscle, composed of parallel fibers, is triangular in shape 
and concealed beneath the adductor mandibulae externus profun- 
dus. It passes to both sides of the prearticular crest and so is divided 
naturally into two major parts — lateral and medial. In addition, a 
distinctive broad, flat muscle sheet forms the most lateral division, 
here referred to as the pars minimus. 

The pars minimus takes origin at the base of the proximal end of 
the quadrate and then extends along its entire anterior, lateral edge 
to a point above the lateral condyle. It inserts along the dorsal edge 
of the surangular crest forward to a point just posterior to the 
adjacent mandibular nerve. The lateral division of the muscle takes 
origin along the anteromedial length of the quadrate. Its parallel 
fibers pass ventrally and forward to insert directly on the floor of the 
mandibular fossa and lateral side of the prearticular crest. 



1979 JsLW musculature of Alsophis cantherigerus brooksi 15 

The medial division of this muscle takes origin along the medial 
side of the quadrate but more posterior in position than the lateral 
division. A few central fibers arise from a small, short, common 
tendon, but most are parallel and all collectively form a sheet that 
passes in an anteroventral direction eventually inserting directly on 
the dorsal edge of the prearticular crest and along a low depression 
on its medial face. The anterior part of this insertion passes lateral 
to the posterior insertion of the deep division of adductor mandibu- 
lae externus medialis. 

M. pseudotemporalis (pst) 
Figs. 1, 3 b-d 

This muscle arises beneath the anterior part of the adductor man- 
dibulae externus medialis and the posterior part of the adductor 
mandibulae externus superficialis along the mid-temporal crest. It is 
a ribbon-like muscle of parallel fibers that courses ventrally, passing 
deep to the maxillary division of the trigeminal (V) nerve, the facial 
carotid artery, and jugular vein. It attaches directly to the com- 
pound bone medial and slightly anterior to the front of the deep 
division of the adductor mandibulae externus medialis. 

M. pterygoideus (pg) 
Figs. 1, 2, 3 c^d, 4 a-b, 5 a-b 

The pterygoideus takes origin from the lateral maxillary process 
of the ectopterygoid, primarily by a prominent tendon. The tendon 
remains superficial and passes ventrally reaching the middle of the 
muscle. Most muscle fibers arise along the length of this prominent 
tendon although a few directly arise from a small area on the ecto- 
pterygoid immediately medial to the origin of this tendon. The fib- 
ers sweep backward forming, on approach to the insertion, the 
swollen belly of the muscle that projects below the compound bone. 
The fibers curve upward to insert on the ventral surface of the 
mandibular retroarticular process. Specifically, this insertion 
includes the ventroposterior surface of the retroarticular process 
and extends a short distance forward along the ventrolateral edge of 
the process to a point even with the articular notch. 

There are two further noteworthy anatomical features of this 
muscle. First, along its anterolateral surface the epimysium is drawn 
up into a loose fascial connection that joins it with the anterior edge 



16 BREVIORA No. 463 

of the nearby adductor mandibulae externus medialis. Second, fib- 
ers composing the anterodorsal part of the muscle form a separate 
fascicle. Though anatomically an integral part of the pterygoideus, 
these fibers insert, via a wide aponeurosis, on the ventrolateral edge 
of the retroarticular process. Most fibers of this slip arise directly 
from the prominent ventral tendon near its anterior end, but a few 
arise directly from the lateral maxillary process of the ectopterygoid 
medial to this tendon. 

M. pterygoideus accessorius (pga) 
Figs. I, 2, 3b 

This muscle arises directly from the ventral surface of the ptery- 
goid and base of the ectopterygoid. Specifically, this origin includes 
the ventral surface of the lateral projection of the pterygoid, its 
lateroventral groove, and lateroposterior base of the ectopterygoid. 

The muscle passes backward to insert on the inner side of the 
retroarticular process along its medial curvature, parallel with but 
medial to the insertion of the pterygoideus. 

M. protractor quadrati (pq) 
Figs. 2. 3 a-b 

This muscle arises from a tendon, shared with its contralateral 
partner, that originates from the midventral basioccipital. It is a flat 
muscle, passing horizontally in a posterolateral direction over the 
dorsal surface of the protractor pterygoidei. A few fibers insert on 
the medial epicondyle of the quadrate, but most insert directly to the 
dorsomedial side of the mandibular retroarticular process, dorsal to 
the origin of the pterygoideus accessorius. 

M. protractor pterygoidei (pp) 
Figs. 1, 2, 3b 

This muscle takes origin from the basisphenoid, specifically from 
a mid-transverse ridge, and forward parasagittally along the basi- 
sphenoid to a point just past the anterior Vidian foramen. It passes 
posteriorly, becoming fusiform in shape, and inserts across the pos- 
terodorsal end of the pterygoid with an especially firm attachment 
to the caudal tip of the bone. 



1979 Jaw musculature of Alsophis cantherigerus brooksi 17 

M. levator pterygoidei (Ip) 
Figs. K 2, 3 a-d. 4a 

A low depression in the posterior, ventral face of the postorbital 
process of the parietal serves as the surface from which this muscle 
takes origin. It passes ventrally, widening along an anteroposterior 
axis to form a long insertion on the pterygoid. This insertion begins 
posteriorly on the dorsolateral side of the pterygoid, runs forward to 
the ectopterygoid-pterygoid articulation, across the base of the 
ectopterygoid, and ends on the adjacent outer side of the pterygoid 
short of the pterygo-palatine articulation. The insertion thus lies 
along the shaft of this bone occupying the dorsolateral fossa. 

M. retractor pterygoidei (rp) 
Fig. 2 

This muscle arises directly from the anterior slope of the trans- 
verse ridge on the basisphenoid, adjacent to the origin of the pro- 
tractor pterygoidei. It passes forward to insert directly on the 
anterodorsal surface of the pterygoid and posterodorsal surface of 
the palatine, and by a broad aponeurosis to the posterior edge of the 
choanal process of the palatine. 

M. retractor vomeris (rv) 
Fig. 2 

This muscle arises from the sphenoid, lateral to the anterior end 
of the origin of the protractor pterygoidei. It passes forward as a 
spindle-like muscle bounded laterally by the retractor pterygoidei. 
Anteriorly, its fibers converge into a slender cord-like tendon that 
accounts for half the total muscle length. This tendon inserts on 
posterodorsal corner of the vomer. 

Throat Musculature 

M. neuromandibularis (nm) 
Figs. 1, 4c, 5 a-b. 6b 

Beginning in the neck, this broad, flat muscle sweeps downward 
and forward around the side of the body, passing under the cervico- 
quadratus and over the axial musculature, to insert at several sites 



18 



BREVIORA 



No. 463 





1979 Jaw musculature of Alsophis cantherigerus brooksi 19 

along the mandible via aponeuroses. It has been variously treated by 
other authors, some dividing it into several parts (Langebartel, 
1968; Kardong, 1973; Haas, 1973) while others include it as part of a 
composite muscle (Albright and Nelson, 1959; Cundall, 1974). For 
descriptive purposes, it is here treated as a single unit. It originates 
from the epimysium of the spinalis-semispinalis muscles adjacent to 
neural spines 6-11. The anterior part of this origin lies under the 
posterior part of the cervicomandibularis. As it passes into the 
throat, it is crossed by two inscriptions. A separate muscle, the 
costomandibularis arises by three separate slips from the ventrolat- 
eral tips and costal cartilages of ribs 2-4. From here the costomandi- 
bularis passes forward to join and most fibers terminate on the 
medial face of neuromandibularis along the posterior inscription. 
Farther anteriorly, the neuromandibularis is joined by a few addi- 
tional fibers originating from the lingual process and anterior end of 
the ceratobranchial of the hyoid; these fibers contribute to the 
medial border of the muscle. 

Insertion on the mandible is by a broad aponeurosis, but along 
three specific sites. One site of insertion, via an aponeurosis, is along 
the ventral edge of the angular and posteroventral edge of the sple- 
nial. A second insertion, also via an aponeurosis, begins along the 
posterolateral edge of the dentary where it articulates with the com- 

Figure 5. Ventral view of throat musculature. At bottom (B). a section of each 
intermandibularis posterior, pars posterior (ipp) has been removed. The left neuro- 
mandibularis (nm) has been cut and reflected to e.xpose the underlying structures, 
including a prominent blood vessel (bv). 

Abbreviations: 

cc M. constrictor colli 

cos M. costomandibularis 

ht M. hyotrachealis 

hy hyoid 

iaa M. intermandibularis anterior, pars anterior 

iag M. intermandibularis anterior, pars glandularis 

iap M. intermandibularis anterior, pars posterior 

ig infralabial gland 

is M. interscutali 

ipa M. intermandibularis posterior, pars anterior 

ipp M. intermandibularis posterior, pars posterior 

nm M. neuromandibularis 

tb M. transversus branchialis 

th M. transversus hyoideus 



20 BREVIORA No. 463 

pound bone. This insertion extends forward along the lateral face of 
the dentary, passing below the anterior mental foramen and termi- 
nates behind the intermandibularis anterior. Thirdly, the epimysium 
along the superior face of the neuromandibularis forms an aponeu- 
rosis that passes dorsal to the origin of intermandibularis posterior, 
pars anterior to insert adjacent to it on the mandible. This forms a 
sleeve through which intermandibularis posterior, pars anterior 
passes from its origin forward toward its insertion (5b, 6b). 

M. intermandibularis posterior (ipa, ipp) 
Figs. 1, 4c, 5 a-b, 6b 

The intermandibularis posterior, pars posterior takes origin 
beneath the insertions of the adductores mandibulae profundus and 
superficialis along the ventral surface of the compound bone. It is a 
strap-like muscle that passes forward superficial to the neuroman- 
dibularis to join its contralateral partner in inserting in the dermis at 
the central midline, just posterior to the origin of transversus 
branchialis. 

The origin of intermandibularis posterior, pars anterior is also 
along the ventral edge of the mandible, but anterior to the insertion 
of the pars posterior. Forming into a strap-like muscle, it passes 
through a sleeve formed by the insertion of the neuromandibularis 
and then runs medial to the mandibular ramus. It inserts in the 
posteroventral epimysium of the transversus branchialis. 

M. intermandibularis anterior (iaa, iap, iag) 
Figs, 1, 5 a-b, 6 a-b 

The muscle originates from the anteroventral tip of the dentary as 
the bone narrows and bends inward. This is lateral to the origins of 
the genioglossus and geniotrachealis. Though not anatomically 
separate, two regions of this muscle can be recognized — an anterior 
region (pars anterior), whose fibers pass inward to insert on the 
interramal pad, and a posterior region (pars posterior), that also 
passes inward to insert in dermis with its contralateral partner on a 
midventral raphe. 

Anterior fibers of the intermandibularis anterior, pars glandularis 
(=constrictor of Langebartel, 1968) are fan-shaped and attach to the 
lateral edge of the interramal pad. The narrow posterior part of the 
muscle wraps around the ventral aspect of the lateral subhngual 
gland and attaches along the posteroventral surface of this gland. 



1979 Jaw musculature of Alsophis cantherigerus brooksi 21 

M. transversus branchialis (tb) 
Figs. 5 a-b, 6 a-b 

This muscle originates from two heads, glandular and mucosal 
(pars glandularis and pars mucosalis respectively, of Albright and 
Nelson, 1959). The glandular head arises on the posterior end of the 
lateral sublingual gland. It passes as a ribbon-like band, arching 
posteriorly around the more medial genioglossus and geniotrachea- 
lis, then ventromedially to form the anterior part of the muscle. It is 
joined by fibers of the mucosal head arising from the dermis of the 
buccal epithelium midway between mandible and trachea to form 
the middle and posterior parts of the muscle. It inserts in the dermis 
of the chin integument at the ventral midline, between insertions of 
intermandibulares anterior and posterior. 

M. genioglossus (gg) 
Figs. I, 6 a-b 

Posteriorly this paired muscle is entwined around the tongue. 
Anteriorly, it originates from two heads. The medial head arises 
from the posterior tip of the interramal pad, then passes caudally 
around the lateral face of the medial sublingual gland. 

The lateral head receives a few fibers from the dorsal surface of 
the intermandibularis anterior, but most arise from the anterior tip 
of the dentary along its inward inflection ventral to the geniotra- 
chealis and dorsal to the intermandibularis anterior. It slants inward 
as a narrow cord-like muscle to join with the medial head near the 
middle of the medial sublingual gland. Just posterior to this gland, 
the muscle passes to the tongue sheath along which it runs caudally. 
About half way along the tongue, right and left genioglossus mus- 
cles change sides. The right muscle passes across the ventral surface 
to the left side and, at the same point, the left muscle passes across 
the dorsal surface to the right side. They stay in these reversed 
positions until reaching their posterior insertions on the hyoglossus, 
at the point of termination of the tongue sheath. 

M. geniotrachealis (gt) 
Figs. 1, 5b, 6 a-b 

This muscle inserts along the side of the trachea beginning just 
posterior to the intrinsic laryngeal musculature (Kardong, 1972) 
spanning 9 cartilaginous rings. It passes forward as a ribbon-hke 



22 



BREVIORA 



No. 463 




1979 Javj musculature of Alsophis cantherigerus brooksi 23 

muscle to its origin along the medial curve of the anterior dentary 
dorsal to and expanding across a wider area of origin than the 
genioglossus. 

M. hyotrachealis (ht) 
Figs. 5b, 6 a-b 

This slender, ribbon-like muscle inserts along the side of the tra- 
chea dorsal to the anterior origin of geniotrachealis. This insertion 
begins immediately posterior to the intrinsic largyngeal muscles (A/. 
dilatator laryngis) and spans three cartilaginous rings. The muscle 

Figure 6. Ventral view of deep throat musculature. At bottom (B), the left 
neuromandibularis (nm) and right transversus branchialis (tbm) are reflected. The 
middorsal surface of the neuromandibularis develops a short attachment to the 
nearby mandible that forms a sheath-like (sh) channel for the origin of the 
intermandibularis posterior, pars anterior (ipa). The label for the hyoid (hy) here lies 
on the lingual process. The intermandibularis anterior (iaa and iap) has been 
sectioned along its midventral insertion and pulled slightly to one side. At top (A), 
most of the muscles and infralabial gland have been removed to reveal the course of 
the transversus branchialis (tbm and tbg), intermandibularis anterior, pars glandu- 
laris (iag), and geniotrachialis (gt). The anterior part of the tongue (tg) has been 
removed to expose the trachea (tr) and a few muscle attachments to it. 

Abbreviations: 

gg M. genioglossus 

gt M. geniotrachealis 

gt M. hyotrachealis 

hy hyoid 

iaa M. intermandibularis anterior, pars anterior 

iag M. intermandibularis anterior, pars glandularis 

iap M. intermandibularis anterior, pars posterior 

ig infralabial gland 

im intrinsic musculature of larynx 

ipa M. intermandibularis posterior, pars anterior 

ir inter-ramal pad 

nm M. neuromandibularis 

pi M. protractor laryngeus 

sh sleeve formed by neuromandibularis about origin of intermandibularis 

posterior, pars anterior 

sll sublingual gland, lateral 

slm sublingual gland, medial 

tbg M. transversus branchialis, glandular head 

tbm M. transversus branchialis, mucosal head 

tg tongue 

tr trachea 



24 BREVIORA No. 463 

passes posteriorly along the floor of the buccal cavity coursing next 
to the trachea and reaches its origin on about the middle of the 
hyoid cornu, which lies, at this point, on the dorsal face of the 
neuroma ndibularis. 



M. protractor laryngeus (pi) 
Fig. 6 a-b 

This short, flat muscle takes origin from the posterior tip of the 
interramal pad and adjacent lateral head of genioglossus. It passes 
medially, running in the dermis of the buccal integument. Its ante- 
rior fibers meet their contralateral partners at the midline posterior 
to the orifice of the tongue sheath. The posterior fibers insert on the 
anterior tip of the glottis. 



DISCUSSION 

Jaw musculature of Alsophis cantherigerus brooksi conforms 
closely to the descriptions of several other colubrids (Albright and 
Nelson, 1959; Varkey, 1973; Haas, 1973; Cundall, 1974). The inter- 
nal divisions of the adductor mandibulae externus medialis, adduc- 
tor mandibulae externus profundus, adductor mandibulae posterior, 
and depressor mandibulae that were noted are often not included in 
these earlier papers. This probably does not indicate species differ- 
ence, but rather differences in investigator emphasis in selecting 
subtle internal detail of muscle to include in the descriptions. There 
does, however, seem to be an actual anatomical difference in pro- 
tractor quadrati. As suggested by the name, this muscle inserts on 
the quadrate in Opheodrys (Cundall, 1974), but here in A. c. 
brooksi, Elaphe (Albright and Nelson, 1 959), and Nerodia (Varkey, 
1973) it inserts on the retroarticular process of the mandible. As in 
most other colubrids, a definitive M. levator anguli oris is absent, 
but a weakly developed M. constrictor colli is present. 

In regard to lateral jaw musculature, the subspecies oi Alsophis 
cantherigerus are a uniform morphological group (sensu Bock, 
1963). As in A. c. brooksi, the adductor mandibulae externus super- 
ficialis arises from the anterior temporal crest and inserts as a broad 
aponeurosis on the lateral side of the mandible. A shp of fibers 



1979 Ja-w musculature of Alsophis cantherigerus brooksi 25 

persists at the anterior edge of the muscle, continues around the 
corner of the mouth, and inserts directly to the mandible. The only 
exception is found in A. c. adspersus where this muscle inserts not 
on the mandible but to the epimysium of the adductor mandibulae 
externus profundus. However, this condition occurs only on the 
right side, the left being like other subspecies, and thus likely repre- 
sents an individual, rather than a subspecific difference. 

In all subspecies, the triangular adductor mandibulae externus 
profundus consists of anterior and posterior wedges of muscle 
separated by a vertical cleft that is often tendinous and forms a site 
of origin for^some fibers. Similarly in all, the adductor mandibulae 
externus medialis is separable into superficial and deep divisions, 
each with different sites of insertion. The adductor mandibulae pos- 
terior always possesses three parts — two major parts, lateral and 
medial, that pass to the respective sides of the prearticular crest and 
a third part, the pars minimus. In all, the protractor quadrati arises 
from a tendon shared with its partner and inserts along the medial 
retroarticular process. The general attachments of the depressor 
mandibulae are similar in all subspecies, and the muscle is divisible 
into superficial and deep portions divided by the insertion of the 
cervicoquadratus. However, the deeper part tends to split length- 
wise mfuscicauda, pepei, and natyi into two parallel divisions, one 
arising from the posterior corner of the quadrate, the other from 
here and from the posterior end of the supratemporal. Both, how- 
ever join to insert directly on the retroarticular process. 

In none of the subspecies examined did the lateral jaw muscula- 
ture differ fundamentally as to specific bones involved in the attach- 
ment sites. The observation that the only differences are in the 
nature of internal muscle anatomy again emphasizes the uniform 
nature of these island subspecies. 



ACKNOWLEDGMENTS 

I am grateful to Dr. Ernest E. Williams (Museum of Comparative 
Zoology) for the loan of the specimens of Alsophis examined in this 
paper. My very special thanks go to Drs. David L. Cundall and 
Alan Savitsky for critically reading and offering thoughtful sugges- 
tions upon the manuscript. 



26 BREVIORA No. 463 

LITERATURE CITED 

Albright, R. G. and E. M. Nelson. 1959. Cranial kinetics of the generalized 

colubrid snake Elaphe ohsoleta quadrivittata. I. Descriptive morphology. II. 

Functional morphology. J. Morph., 105 (2): 193-239; 241-291. 
AiEN, E. L. .\ND D. A. Langebartel. 1977. The cranial nerves of the colubrid 

snakes Elaphe and Thamnophis. J. Morph., 154(2): 205-222. 
Bock. W. J. 1963. Evolution and phylogeny in morphologically uniform groups. 

Amer. Nat., 97: 265-285. 
Ci'NDALL, D. L. 1974. The cranial osteology and myology of the green snakes, 

genus Opheodrys. Unpublished Ph.D. dissertation. University of Arkansas, 

Fayetteville & Little Rock. University microfilm 74-28, 078. 
DtLLEMiJER, P. 1956. The functional morphology of the head of the common 

viper, I'iper herns (L.). Archives Neerlandaises de Zoologie, 11: 387-497. 
Haas, G. 1973. Muscles of the jaws and associated structures in the rhyncho- 

cephalia and squamata. In C. Gans and T. S. Parsons (Eds.), Biology of the 

Repiilia. 4: 285-490. 
Kardong, K. V. 1972. Morphology of the respiratory system and its musculature 

in different snake genera. (Part I) Crolalus viridis oreganus and Elaphe ohsoleta 

quadrivittata. (Part II) Charina hottae. Gegenbaurs morph. Jahrb., 117(1972): 

285-302; 364-376. 
1973. Lateral jaw and throat musculature of the cottonmouth snake 

Agkistrodon piscivorus. Gegenbaurs morph. Jahrb., 119(3):31fr-335. 
1974. Kinesis of the jaw apparatus during the strike in the cottonmouth 



snake, Agkistrodon piscivorus. Forma et functio, 7: 327-354. 
KocHVA, E. T. 1962. On the lateral jaw musculature of the Solenoglypha with 

remarks on some other snakes. J. Morph., 110:277-284. 
Langebartel, A. 1968. The hyoid and its associated muscles in snakes. Illinois 

Biological Monographs, 38:1-156. 
Maglio, V. J. 1970. West Indian .xenodontine colubrid snakes: their probable 

origin, phylogeny, and zoogeography. Bull Mus. Comp. Zool., 141(1): 1-53. 
Pregill, G. K. 1977. Axial myology of the racer Coluher constrictor with 

emphasis on the neck region. Trans. San Diego Soc. Nat. Hist., 18(11): 185- 

206. 
Taub, a. M. 1966. Ophidian cephalic glands. J. Morph., 118: 529-542. 
Varkey, a. 1973. Comparative cranial myology of North American natricine 

snakes. Unpublished Ph.D. dissertation, Louisiana State University and 

Agricultural and Mechanical College, Baton Rouge. University microfilms, 

73-27, 877. 



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