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HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 






B R E V I R A 



11 



seuin of Comparative Zoology 



HARVARD UNIVERSITY 



Numbers 464-487 
1981-1986 



MC2 
LIBRARY 

NOV 1 9 1986 

HARVARD 
UN 



CAMBRIDGE, MASSACHUSETTS 02138 U.S.A. 
1986 



BREVIORA 

Museum of Comparative Zoology 

CONTENTS 

Numbers 464-487 

1981 

No. 464. The Origin of the Crocodiloid Tarsi and the Inter- 
relationships of Thecodontian Archosaurs. By Donald 
Brinkman. 23 pp. December 30. 

No. 465. The Structure and Relationships of the Dromasaurs 
(Reptilia: Therapsida) By Donald Brinkman. 34 pp. 
December 30. 

1982 

No. 466. Systematics of the Mexicana Species Group of the 
Colubrid Genus Lampropeltis, with an Hypothesis 
Mimicry. By William R. Garstka. 35 pp. June 30. 

No. 467. Three New Species of the Anolis Punctatus Complex 
from Amazonian and Inter-Andean Colombia, with 
Comments on the Eastern Members of the Punctatus 
Species Group. By Ernest E. Williams. 38 pp. June 
30. 

No. 468. A New Forest Skink from Ponape. By A. Ross Kiester. 
10 pp. June 30. 

No. 469. Catalog of the Primary Types of Bombyliidae (Diptera) 
in the Entomological Collections of the Museum of 
Comparative Zoology, with Designations of Lecto- 
types. By Neal L. Evenhuis. 23 pp. June 30. 

No. 470. Systematic Implications of Innervation Patterns in Tele- 
ost Myotomes. By Quentin Bone and R. Dana Ono. 
23 pp. June 30. 



No. 471. Arthur Loveridge — A Life in Retrospect. By Ernest E. 
Williams. 12 pp. June 30. 

No. 472. Fishes of the Suborder Labroidei (Pisces: Perciformes): 
Phylogeny, Ecology, and Evolutionary Significance. 
By Leslie S. Kaufman and Karel F. Liem. 19 pp. June 
30. 

1983 

No. 473. The Interrelationships of Pelycosaurs. By Donald 
Brinkman and David E. Eberth. 35 pp. April 29. 

No. 474. Ruthiromia Elcobriensis, A New Pelycosaur from El 
Cobre Canyon, New Mexico. By David A. Eberth 
and Donald Brinkman. 26 pp. April 29. 

No. 475. New or Problematic Anolis from Colombia. I. Anolis 
Calimae, New Species, from the Cloud Forest of 
Western Colombia. By Stephen Ayala, Dennis Har- 
ris, and Ernest E. Williams. 1 1 pp. April 29. 

No. 476. Townsend's Unmapped North Atlantic Right Whales 
(Eubalaena Glacialis). By William E. Schevill and 
Karen E. Moore. April 29. 

1984 

No. 477. New or Problematic Anolis from Colombia. II. Anolis 
Propinquus, Another New Species from the Cloud 
Forest of Western Colombia. By Ernest E. Williams. 
7 pp. September 7. 

No. 478. New or Problematic Anolis from Colombia. III. Two 
New Semiaquatic Anoles from Antioquia and Choco, 
Colombia. By Ernest E. Williams. 22 pp. September 7. 

No. 479. Agonistic and Courtship Displays of Male Anolis Sagrei. 
By Michelle P. Scott. 22 pp. September 7. 



1985 

No. 480. Three New Lizards of the Genus Emoia (Scincidae) from 
Southern New Guinea. By Walter C. Brown and Fred 
Parker. 12 pp. June 21. 

No. 481. A New Anolis of the Lionotus Group from Northwest- 
ern Ecuador and Southwestern Colombia (Suaria: 
Iguanidae) By Kenneth Miyata. June 21. 

No. 482. New or Problematic Anolis from Colombia. IV. Anolis 
Antioquiae, New Species of the Anolis Eulaemus 
Subgroup from Western Colombia. By Ernest E. 
Williams. 9 pp. June 21. 

No. 483. Notes on Pristidactylus (Sqamata: Iguanidae). By 
Richard Etheridge and Ernest E. Williams. 18 pp. 
June 21. 

No. 484. Male Aggressive Behavior in a Pair of Sympatric Sibling 
Species. By Jonathan B. Losos. 30 pp. June 21. 

1986 

No. 485. The Anatomy and Relationships of Stereophallodon 
and Baldwinonus (Reptilia, Pelycosauria). By Donald 
Brinkman and David A. Eberth. 36 pp. August 30. 

No. 486. Thelodus "Macintoshi" Stetson 1928, The Largest 
Known Thelodont (Agnatha: Thelodonti). By Susan 
Turner. 20 pp. August 30. 

No. 487. The Identity of Larval Parasudis (Teleostei, Chloroph- 
thalmidae), with Notes on the Relationships of Auli- 
poform Fishes. By Karsten E. Hartel and Melanie L. 
J. Stiassny. 24 pp. August 30. 



BREVIORA 

Museum of Comparative Zoology 

Index of Authors 

Numbers 464-487 

1981-1986 

Ayala, Stephen 475 

Bone, Quentin 470 

Brinkman, Donald 464, 465, 473, 474, 485 

Brown, Walter C , 480 

Eberth, David E 473, 474, 485 

Etheridge, Richard 483 

Evenhuis, Neal L 469 

Garstka, William R 466 

Hartel, Karsten E 487 

Harris, Dennis 475 

Kaufman, Leslie S 472 

Losos, Jonathon B 484 

Liem, Karel F 472 

Miyata, Kenneth 481 

Moore, Karen E 476 



Ono, R. Dana 470 

Parker, Fred 480 

Scott, Michele P 479 

Shevill, William E 476 

Stiassny, Melanie L. J 487 

Turner, Susan 486 

Williams, Ernest E 467, 471, 475, 477, 478, 482, 483 






B R W;X°1 ORA 

Museum of Comparative Zoology 



I S ISSN 0006 %9K 



Cambridge, Mass. 30 December 1981 Number 464 

THE ORIGIN OF THE CROCODILOID TARSI 

AND THE INTERRELATIONSHIPS OF 

THECODONTIAN ARCHOSAURS 

Donald Brinkman 1 

Abstract. The tarsus of the proterosuchian Chasmatosaurus represents the 
primitive archosaur tarsus. This kind of tarsus is also present in rhynchosaurids, 
trilophosaurids, prolacertids, and Protorosaurus, and suggests that these reptiles are 
members of a single radiation. Two distinct kinds of crocodiloid tarsi are present in 
thecodonts, a crocodile-normal tarsus and a crocodile-reversed tarsus. The crocodile- 
reversed tarsus could have originated from the crocodile-normal tarsus, but the 
reverse relationship is not plausible. Gracilisuchus, the only "ornithosuchid" with a 
crocodile-normal tarsus, shows features of the skull that are not consistent with its 
placement in the Ornithosuchidae. Euparkeria. on the basis of both cranial and 
postcranial characters, is a plausible ornithosuchid ancestor but could not be 
ancestral to a pseudosuchian with a crocodile-normal tarsus. The tarsus of 
Erythosuchus neither contradicts nor supports a relationship between Erythrosuchus 
and rauisuchids. 

INTRODUCTION 

In recent years, it has been recognized that a number of 
structurally distinct kinds of tarsi are present in archosaurs, and an 
understanding of the evolution of this structural complex is 
necessary for an understanding of the interrelationships of the 
group. In the tarsus of crocodiles and typical pseudosuchians, the 
ankle joint passes between the astragalus and calcaneum, the 
astragalus being locked to the tibia and the calcaneum integrated 
with the pes. In dinosaurs, the ankle joint passes distal to the 
astragalus and calcaneum. Krebs (1963, 1973) argued that this 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachsuetts 
02138. 



2 BREVIORA No. 464 

difference precludes derivation of dinosaurs from any known 
pseudosuchian. Recently, two additional kinds of tarsi have been 
recognized in thecodonts. Proterosuchian thecodonts of the family 
Proterosuchidae have a distinct tarsus that is in many ways 
primitive (Cruickshank, 1972; Carroll, 1976). Bonaparte (1971) 
recognized that two distinct kinds of crocodilelike tarsi are present 
in pseudosuchians, one like that of crocodiles in which the 
astragalus has a peg that fits in a socket on the calcaneum, and one 
seen in advanced ornithosuchids in which the calcaneum has a 
process that fits in a socket on the astragalus. Chatterjee (1978) 
termed these the crocodile-normal and crocodile-reversed tarsus 
respectively. Also, two pseudoschians with mesotarsal ankle joints 
have been described (Romer, 1971). One of these, Lagerpeton, has a 
fully developed mesotarsal joint. The second, Lagosuchus, retains a 
posteriorly directed calcaneal tuber and a complex articulation 
between the astragalus and calcaneum (Bonaparte, 1975a). 

The evolution of these tarsal patterns was recently discussed by 
Cruickshank (1979). Cruickshank showed that the proterosuchian 
tarsus is an excellent structural ancestor to the crocodile-normal 
tarsus and argued that the two kinds of crocodile tarsi can be used 
to separate pseudosuchians into two groups. Based on this, 
Cruickshank suggested that Gracilisuchus, which has a crocodile- 
normal tarsus, be removed from the Ornithosuchidae, all other 
members of which have a crocodile-reversed tarsus. However, the 
origin of the crocodile-reversed tarsus remains unknown. If the 
crocodile-normal tarsus was ancestral to the crocodile-reversed 
tarsus, then a crocodile-normal tarsus could have been present in 
primitive ornithosuchids, and the presence of a crocodile-normal 
tarsus would not bar Gracilisuchus from the Ornithosuchidae. 
Thus, in order to use the structure of the tarsus as a basis for 
interpreting the interrelationships of archosaurs, it is necessary to 
obtain a more precise understanding of both the origin of the 
crocodile-reversed tarsus and the phylogenetic position of Gracili- 
suchus. 

THE CROCODILE-NORMAL TARSUS 

In extant crocodiles, five elements are present in the tarsus: the 
astragalus, the calcaneum, and the second to fourth distal tarsals. 



!981 



THE ORIGIN OF THE CROCODILOID TARSI 



The astragalus (Fig. 1C-D) supports the tibia and contacts the fibula 
by articular surfaces that almost completely cover the proximal 
surface of the bone. Anteriorly, the astragalus has a strongly covex 
surface that articulates with the proximal end of the first two 
metatarsals and the medial surface of the second and third distal 
tarsals. Above this, the anterior face of the astragalus is formed by a 
concave area covered by finished bone. Laterally, a distinctive 
articular surface for the calcaneum is present. This is divisible into 
two separate areas. The ventral area has the shape of a portion of a 




a, bond-shaped 
art surf 



med flange 



5 mm 



Figure I. The right astragalus and calcaneum of Caiman sclerops. A) calcaneum 
in proximal view; B) calcaneum in medial view; C) astragalus in dorsal view; D) 
astragalus in medial view. 

Abbreviations: band-shaped art surf, band-shaped articular surface; cal tub, 
calcaneal tuber; cone-shaped art surf, cone-shaped articular surface; fib art surf, 
fibular articular surface; med flange, medial flange; met art surf, metatarsal 
articular surface; tib art surf, tibial articular surface; wheel-shaped art surt. 
wheel-shaped articular surface; ast art surf, astragalar articular surface. 



4 BREVIORA No. 464 

cone, its apex forming the tip of the laterally directed peg. Dorsal to 
this, a notch in the lateral edge of the astragalus leads to a band- 
shaped articular surfce. These two areas meet along a ridge that 
terminates on the tip of the lateral peg. 

The calcaneum (Fig. 1 A-B) has three characteristic areas: a dorsal 
area that has the form of a portion of a wheel, a medially directed 
flange that underlies the astragalus, and a posteriorly directed tuber. 
The medial half of the wheel-shaped articular surface fits in the 
notch on the lateral edge of the astragalus and is overlapped by the 
astragalus. The lateral half supports the fibula. The fibular and 
astragalar surfaces are differentiated by a slight change in the 
curvature of the articular surface. The medially directed flange 
articulates behind the cone-shaped articular surface of the astra- 
galus. The calcaneal tuber extends across the full width of the bone. 
The distal end of the tuber is expanded and has a vertical groove in 
which lie tendons of the long pedal flexors. Anterodistally, the 
calcaneum as a flat articular surface that abuts the fourth distal 
tarsal. 

A crocodile-normal tarsus is present in the Rauisuchidae (Krebs, 
1965, 1973; Sill, 1974), the Aetosauridae (Sawin, 1947; Walker, 
1961; Bonaparte, 1971; Sill, 1974), and Gracilisuchus Bonaparte, 
1975b). A crocodile-normal calcaneum from the uppermost Lower 
or lowermost Middle Triassic was figured by Young (1964, Fig. 60) 
and attributed to Wangisuchus. 

The astragalus in these pseudosuchians, where known, differs 
from that of crocodiles in having more extensive development of 
finished bone on its anterior face and in the proportions of the 
articular surfaces, the metatarsal articular surface being narrower 
mediolaterally in most genera, as in Gracilisuchus (Fig. 4B). The 
proportions of the calcaneum also show some variation, the 
calcaneal tuber of aetosaurs being considerably broader than in 
crocodiles and other pseudosuchians (Sawin, 1947). Despite this 
variation, the structure of the joint between the astragalus and 
calcaneum is like that of crocodiles. 

THE TARSUS OF CHASMATOSAVRUS 

The tarsus of the early proterosuchid Chasmatosaurus ] (Fig. 2) is 
primitive in the presence of a separate astragalus and centrale, and 



198 



THE ORIGIN OF THE CROCODILOID TARSI 



the retention of a foramen between the astragalus and calcaneum. 
However, comparison with an eosuchian tarsus, such as that of a 
tangasaurid (Fig. 3), demonstrates that a number of derived features 
are present. The articular surface between the astragalus and 
calcaneum in eosuchians is flat and forms a straight line when the 
tarsus is seen in dorsal view. In Chasmatosaurus, the portion of the 
articular surface proximal to the perforating foramen is inclined 
relative to the distal portion, with the astragalus overlying the 
calcaneum. The articular surface between the astragalus and 
calcaneum is a complex concave-convex joint. The portion of the 
articular surface distal to the perforating foramen is a ball and 
socket joint, with the socket on the calcaneum. Proximal to the 
perforating foramen, a concave-convex joint is also present, but the 
concavity is on the astragalus. The proximal edge of the calcaneum 
is not preserved in Chasmatosaurus vanhoepeni, but in the 






Figure 2. The tarsus of Chasmatosaurus vanhoepeni. Left calcaneum, astragalus, 
and centrale in A) ventral, B) dorsal, and C) proximal views. Drawing based on cast 
of NM C3016, Nasionale Museum, Bloemfontein. 

Abbreviations: ast, astragalus; cal, calcaneum; cen, centrale. 



■Following Charig and Sues ( 1976), Proterosuchus is considered a nomen dubium, 
and the remaining proterosuchid material from South Africa is referred to the genus 
Chasmatosaurus. 



BREVIORA 



No. 464 




Figure 3. The left tarsus of Hovasaurus, a tangasaurid eosuchian, in dorsal view. 
Drawn from cast of MNHN 1925-5-61, National Museum of Natural History, Paris. 

Abbreviations: ast, astragalus; cal, calcaneum; cen, centrale. 



1981 THE ORIGIN OF THE CROCODILOID TARSI 

calcaneum of Chasmatosaurus yuani illustrated by Young (1936, 
Fig. 9D) the convexity of the calcaneal surface is seen to continue 
laterally to form a dorsoventrally convex fibular articular surface, 
as it does in the early rhynchosaur Noteosuchus (Fig. 6A). 

The calcaneum has been modified from the primitive platelike 
condition by the development of a laterally extending tuber. The 
distal edge is thin, and a thickened medial buttress extends 
transversely across the bone to the expanded cartilage-covered 
lateral edge. Judging from Young's illustration of the calcaneum of 
Chasmatosaurus yuani, the proximal edge was covered by unfin- 
ished bone, as it is in the early rhynchosaur Noteosuchus (Fig. 6 A). 

The centrale has shifted its position so that it is now located 
laterally, rather than distally, to the astragalus. It may aid in 
support of the tibia. 

Cruickshank (1972), based on the similarity of the tarsus of 
Chasmatosaurus and early rhynchosaurs and the presence in both of 
a downturned premaxilla, suggested that Chasmatosaurus was a 
carnivorous rhynchocephalian. If the proterosuchian tarsus was 
restricted to these animals, it would suggest that Chasmatosaurus is 
phylogenetically a rhynchosaur and thus would bring into question 
the validity of using the proterosuchian tarsus as the primitive 
archosaur pattern. However, a number of additional groups of 
diapsid reptiles have a tarsus that, in so far as comparison is 
possible, is like that of Chasmatosaurus. The tarsus of Prolacerta 
(Gow, 1975, Fig. 33) shows all the advanced features seen in 
Chasmatosaurus and early rhynchosaurs. As would be expected, the 
aquatic members of the Prolacertiformes show a decrease in the 
ossification of the tarsus. Despite this, mature specimens of 
Tanystropheus (Wild, 1973; Fig. 75) and Macrocnemus (Peyer, 
1937; PI. 56, Fig. 2) show the diagnostic features of a laterally 
directed dorsoventrally compressed tuber on the calcaneum, and a 
complex concave-convex articulation between the astragalus and 
calcaneum. A proterosuchian tarsus is also seen in Protorosaurus 
from the Permian of Europe (von Meyer, 1856, PI. 9), and 
Trilophosaurus from the Triassic of Texas (Fig. 4). 

At first glance, this assemblage of reptiles seems to be an 
unnatural one, bringing together animals with markedly different 
adaptations and skull configurations. However, a more detailed 
consideration shows that this assemblage is not as artificial as first 



BREVIORA 



No. 464 




Figure 4. The right astragalus and calcaneum of Trilophosaurus. Calcaneum in 
A) ventral, B) dorsal, C) proximal, and D) distal views; astragalus in E) ventral, F), 
medial, and G) dorsal views. Calcaneum: specimen TMM 31025-258; astragalus: 
specimen TMM 31025-259, Texas Memorial Museum. 



appears. Prolacerta and Protorosaurus have long been recognized 
as being closely related (Camp, 1945; Watson, 1958). Romer (1956) 
denied the presence of such a relationship, choosing to place 
Protorosaurus in the Euryapsida because of the presumed presence 
of a solid cheek. However, as noted by Chatterjee (1980), much 
uncertainty about the structure of the skull of Protorosaurus exists. 
Details of the skull and postcranial skeleton that are known for 
certain are similar to Prolacerta, and Chatterjee unites these genera 
in the suborder Prolacertiformes. Gow (1975) concluded on the 
basis of evidence from the skull of Prolacerta that the Prolacerti- 
formes are related to archosaurs. 

The remaining groups in this assemblage, rhynchosaurids and 
trilophosaurids, have highly specialized skulls that could be derived 
from any primitive diapsid, so the skull neither supports nor negates 
a relationship between these groups and the archosaur-protoro- 



1981 THE ORIGIN OF THE CROCODILOID TARSI 9 

saurid-prolacertid group. Postcranially, both trilophosaurids and 
rhynchosaurids are less specialized, and a similarity in the structure 
of their postcranial skeletons and the postcranial skeleton of some 
members of the archosaur-protorosaurid-prolacertid group has 
been noted (Gregory, 1945; Carroll, 1976), although it is not certain 
whether these represent derived features or primitive features 
widespread in diapsid reptiles. 

Thus, there seems to be no need to hypothesize a multiple origin 
of the proterosuchian tarsus. If rhynchosaurs, prolacertiformes, 
prolacertids, trilophosaurids, and archosaurs are a natural group, 
the proterosuchian tarsus could have originated only once. One 
implication of this is that the proterosuchian tarsus is the primitive 
archosaur tarsus and is the ultimate structural ancestor of the 
various kinds of tarsi seen in advanced archosaurs. In some cases, 
an intermediate structural complex may have been present, but, as 
shown by Cruickshank (1979), the proterosuchian tarsus was 
probably the direct structural antecedent of the crocodile-normal 
tarsus. It is useful to identify the structural changes that would have 
occurred during this transition before considering the structure and 
origin of the crocodile-reversed tarsus. 

ORIGIN OF THE CROCODILE-NORMAL TARSUS 

As recognized by Cruickshank (1979), the astragalus-centrale unit 
of the proterosuchian tarsus is directly comparable to the astragalus 
of the crocodile-normal tarsus (Fig. 5). The area distal to the 
perforating foramen is homologous to the cone-shaped articular 
surface of the crocodile-normal tarsus, but differs in being smaller 
and less strongly curved. The area proximal to the perforating 
foramen is homologous to the dorsal half of the notch on the lateral 
edge of the crocodile-normal astragalus, the main difference being 
that in the proterosuchian tarsus this surface is separated from the 
distal surface by finished bone. 

The calcaneum of the crocodile-normal and proterosuchian tarsi 
(Fig. 6) differs in the orientation of the calcaneal tuber; in the 
proterosuchian tarsus, this is directed laterally, while in the 
crocodile-normal tarsus, this is directed more posteriorly. If the 
calcaneal tuber of the proterosuchian tarsus were oriented so that it 
extended posteriorly, the articular surface for the fibula and 
astragalus would be oriented along the long axis of the bone rather 



10 



BREVIORA 



No. 464 





1 cm 



Figure 5. The left astragalus of the proterosuchian and crocodile-normal tarsus 
in anterior view. A) astragalus and centrale of Noteosuchus; B) astragalus of 
Gracilisuchus. Not drawn to scale. Noteosuchus based on cast of Albany Museum 
3591. Grasilisuchus based on cast of specimen in collection of Paleontologia 
Vertebrados de la Fundacion M. Lillo. 



cal tub 




fib art surf 



ast art surf 



ast art surf 




ib < 
art surf 



Figure 6. The left calcaneum of the proterosuchian and crocodile-normal tarsus. 
A) calcaneum of Noteosuchus: B) calcaneum of Gracilisuchus. 

Abbreviations: ast art surf, astragalar articular surface; cal tub, calcaneal 
tuber; fib art surf, fibular articular surface. 



1981 THE ORIGIN OF THE CROCODILOID TARSI 1 1 

than across it. A simple enlargement of the articular surface, 
together with the extension of the proximal portion of the astragalar 
articular surface onto the medial edge of the perforating foramen, 
would form the wheel-shaped articular surface of the crocodile- 
normal calcaneum. An enlargement of the ventral half of the 
articular surface for the astragalus would form the medial flange of 
the crocodile-normal calcaneum. 

Thus structurally, the proterosuchian tarsus is an excellent 
ancestor of the crocodile-normal tarsus. The mechanical changes 
involved in this transition were probably minor, since, as noted by 
Thulborn (1980), the joint between the astragalus and calcaneum 
was probably movable in the proterosuchian tarsus. 

THE CROCODILE-REVERSED TARSUS 

The crocodile-reversed tarsus is best known in the ornithosuchid 
Riojasuchus (Bonaparte, 1971, 1975b). Dorsally, the calcaneum 
(Fig. 7B) has an articular surface that is convex both mediolaterally 
and proximodistally. This surface supports the fibula along its 
lateral edge and the astragalus along its medial edge. These are 
exactly the relationships of the wheel-shaped articular surface of the 
crocodile-normal tarsus (Fig. 8A-B). The major difference is that 
the medial edge of this area is hypertrophied in Riojasuchus to form 
a medially directed process. This process is functionally equivalent 
to the ventral flange of the crocodile-normal calcaneum in that it 
underlies the astragalus (Fig. 8C-D). Consequently, it is not 
surprising that the ventral flange or an homologous area is not 
present in the crocodile-reversed calcaneum. The absence of the 
ventral flange is associated with a reduction in the width of the 
calcaneal tuber; in the crocodile-reversed tarsus, the calcaneal tuber 
does not extend the full width of the bone. In addition, the tuber of 
the crocodile-reversed tarsus is distinctive in that its distal end 
curves medially and is without a groove for the tendon of the 
gastrocnemial muscles. 

The differences in structure of the astragalus of the crocodile- 
normal and crocodile-reversed tarsus correspond to the differences 
in structure of the calcaneum: the hypertrophy of the medial edge of 
the wheel-shaped articular surface is associated with the elongation 
of the overlying portion of the astragalus, and the loss of the ventral 
flange is associated with the loss of the cone-shaped articular 
surface. 



12 



BREVIORA 



No. 464 



tib art surf. 




met art surf 



Figure 7. The left astragalus and calcaneum of the crocodile-reversed tarsus. A) 
Astragalus and B) calcaneum of Riojasuchus. Drawn from cast of PVL 3827, 
Paleontologia Vertebrados de la Fundacion M. Lillo. 

Abbreviations: ast art surf, astragalar articular surface; cal tub, calcaneal 
tuber; fib art surf, fibular articular surface; met art surf, metatarsal articular 
surface; tib art surf, tibial articular surface. 



From this comparison, it can be seen that the crocodile-normal 
tarsus is a plausible ancestor of the crocodile-reversed tarsus. The 
major changes involved in such a transition would be the medial 
elongation of the wheel-shaped articular surface of the calcaneum 
and the loss of the ventral flange. Derivation of the crocodile- 
reversed tarsus directly from the primitive archosaur tarsus is also 
possible. However, derivation of the crocodile-normal tarsus from 
the crocodile-reversed tarsus can be discounted as being improbable 
since it would involve the redevelopment of the ventral flange, a 
structure that is present in the crocodile-normal tarsus and the 
primitive archosaur tarsus but absent in the crocodile-reversed 
tarsus. 

Given this relationship of the crocodile-normal and crocodile- 
reversed tarsi, the structure of the tarsus cannot be used to exclude 
Gracilisuchus from the Ornithosuchidae. Rather, the systematic 
position of Gracilisuchus has implications for the evolution of the 
tarsus. If Gracilisuchus is a true ornithosuchid, the crocodile- 
normal tarsus must have given rise to the crocodile-reversed tarsus, 
and the structure of the tarsus has little real phylogenetic 
significance. If however, Gracilisuchus is not an ornithosuchid, the 
crocodile-reversed tarsus may have originated independently from 



1981 



THE ORIGIN OF THE CROCODILOID TARSI 



13 







Figure 8. The left calcaneum of the crocodile-normal and crocodile-reversed 
tarsi. A) crocodile-normal calcaneum of Gracilisuchus in proximal view; B) 
crocodile-reversed calcaneum of Riojasuchus in proximal view; C) section through 
an articulated crocodile-normal astragalus and calcaneum; D) section through an 
articulated crocodile-reversed astragalus and calcaneum. Not drawn to scale. 



the proterosuchian tarsus, and the presence of the crocodile- 
reversed tarsus can be used as the defining feature of some 
taxonomic group. Thus it is necessary to reconsider the relation- 
ships of Gracilisuchus. 



THE RELATIONSHIPS OF GRACILISUCHUS 

The skull of Gracilisuchus was reconstructed by Romer ( 1972) on 
the basis of MCZ 4117, a complete, three dimensional skull. One of 



14 BREVIORA No. 464 

the unusual features seen in this skull is a small lower temporal 
opening. However, other material, particularly MCZ 4116 and 
MCZ 41 18, show that MCZ 41 17 has been slightly crushed and the 
quadratojugal and squamosal have been displaced. In MCZ 4116, 
the preorbital bar is slender and the antorbital opening is larger than 
in MCZ 41 17 (Fig. 9A). In MCZ 41 18, the postorbital bar is tall and 
slender, the quadratojugal and squamosal are separated from the 
postorbital, and the lower temporal opening is large and rectangular 
(Fig 9B). Based on these skulls, the arrangement of the temporal 
region and the height of the face in the reconstruction of the skull of 
Gracilisuchus is modified (Fig. 9B). 

Gracilisuchus differs from the advanced ornithosuchids as 
defined by Bonaparte (1975b) (Fig. 10) in the following features: 

1) The antorbital fenestra is rectangular in Gracilisuchus and is 
triangular in the advanced ornithosuchids. 

2) The ventral border of the orbit is round in Gracilisuchus, and a 
distinct antorbital ramus of the jugal is not present. In the 
advanced ornithosuchids, a distinct preorbital ramus of the 
jugal is present; this is close to the postorbital ramus at its base, 
so the ventral margin of the orbit is pointed. 

3) The quadratojugal of Gracilisuchus is a tall, slender element 
that extends nearly the full height of the lower temporal 
opening. In the advanced ornithosuchids, the quadratojugal is 
broader and is limted to the ventral half of the lower temporal 
opening. 

4) The upper tooth row is complete, and all the teeth of the lower 
jaw fit inside the upper teeth row in Gracilisuchus. In advanced 
ornithosuchids, a gap is present between the anterior tooth of 
the maxilla and the posterior tooth of the premaxilla, with the 
anterior one or two dentary teeth passing lateral to the maxilla 
in this gap. 

5) In Gracilisuchus, the lower temporal fenestra is rectangular, 
and no anterior inflection of the quadratojugal and squamosal 
is present. In the advanced ornithosuchids, a large anterior 
inflection of the quadratojugal and squamosal results in the 
presence of an L-shaped lower temporal fenestra. 

6) In Gracilisuchus, the squamosal has a peculiar, posteriorly 
concave flange on its dorsal end. No such flange is present in 
the advanced ornithosuchids. 



1981 



THE ORIGIN OF THE CROCODILOID TARSI 



15 






Figure 9. The skull of Gracilisuchus. A) specimen drawing of MCZ 4116; B) 
specimen drawing of MCZ 4118; C) reconstruction of skull. 



16 BREVIORA No. 464 

7) In Gracilisuchus, the posterior end of the dentary extends 
dorsal to the mandibular fenestra. In the advanced ornitho- 
suchids, the posterior end of the dentary is forked, with one 
branch extending dorsal and one branch extending ventral to 
the lateral mandibular fenestra. 

8) In Gracilisuchus, the splenial forms the ventral margin of the 
jaw along the posterior half of the dentary. In the advanced 
ornithosuchids, the splenial is restricted to the inner surface of 
the jaw. 

9) In Gracilisuchus, the cervical vertebrae are not keeled. In the 
advanced ornithosuchids, the cervical vertebrae, where known 
(Riojasuchus and Ornithosuchus), are keeled. 

10) In Gracilisuchus, there are two pairs of dermal scutes per 
vertebra in the cervical region of the vertebral column. In the 
advanced ornithosuchids, where known {Ornithosuchus and 
Riojasuchus), there is one pair of scutes per vertebra. 
This list of differences between Gracilisuchus and the advanced 
ornithosuchids shows that the advanced ornithosuchids are more 
similar to each other than to Gracilisuchus. But these features by 
themselves do not demonstrate that Gracilisuchus is not an 
ornithosuchid; Gracilisuchus occurs much earlier in time than the 
advanced ornithosuchids, so the differences may represent succes- 
sive grades of evolution in a single radiation. In order to be used 
phylogenetically, it is necessary to determine which of the character- 
states represent derived features. Euparkeria (Fig. 10D), the oldest 
known pseudosuchian, had traditionally been considered to be the 
structural ancestor of later pseudosuchians and thus can be used as 
the outgroup in determining which character-states are primitive or 
advanced. 

Features that are shared by Gracilisuchus and Euparkeria, and 
thus can be considered primitive, are the presence of a complete 
upper tooth row with all the dentary teeth fitting inside the upper 
tooth row, the shape of the lower temporal fenestra, and the shape 
of the antorbital fenestra (features 1, 4, and 5 in the above list). In all 
other features, Euparkeria is like the advanced ornithosuchids and 
unlike Gracilisuchus: the jugal has a well-developed antorbital 
process, and the base of this is near the postorbital process; the 
squamosal is without the peculiar posteriorly concave flange seen in 
Gracilisuchus; the posterior end of the dentary is forked, with a 
branch above and a branch below the lateral mandibular fenestra; 



1981 



THE ORIGIN OF THE CROCODIEOID TARSI 



17 




Figure 10. The skulls of the advanced ornithosuchids and Euparkeria. A) 
lenaiicosuchus; B) Ornithosuchus; C) Riojasuchus; D) Euparkeria. From Bona- 
parte, 1975. 



the splenial is restricted to the internal surface of the lower jaw; the 
cervical vertebrae are keeled; and one pair of dermal scutes is 
present per vertebral segment. For these features, the character-state 
present in the advanced ornithosuchids must be considered primi- 
tive, and the Gracilisuchus condition derived. 

Thus, if an ornithosuchid, Gracilisuchus is derived from the 
ornithosuchid pattern in a way different from the advanced 
ornithosuchids. Alternatively, Gracilisuchus may be a member of a 
radiation distinct from that of ornithosuchids. This latter possibility 
is suggested by the presence of some of the derived features of 
Gracilisuchus in Sphenosuchus, Pseudohesperosuchus, and Lewi- 
suchus, pseudosuchians that are thought to be unrelated to 
ornithosuchids (Romer, 1972). These features include the tall. 



18 BREVIORA No. 464 

slender quadratojugal, the posterior flange on the squamosal 
(known in Pseudohesperosuchus and Sphenosuchus), and the 
presence of unkeeled cervical vertebrae (known in Lewisuchus). 

THE TARSUS OF EUPARKERIA 

The similarity between ornithosuchids and Euparkeria raises the 
possibility that Euparkeria is closely related to the Ornithosuchidae, 
rather than being a generalized primitive pseudosuchian. Such a 
relationship is supported by the structure of the tarsus. The 
astragalus and calcaneum of Euparkeria were illustrated by 
Bonaparte (1975) and Cruickshank (1979). As noted by these 
authors, these elements are directly comparable to those of the 
crocodile-reversed tarsus (Fig. 11). The dorsal portion of the 
articular surface of the calcaneum is divisible into two areas, a 
narrow area along the lateral edge of the bone that would have 
articulated with the fibula, and a medial area that would have 
articulated with the astragalus. The astragalar area is in the shape of 
a portion of a cone with its apex directed medially. No area 
comparable to the ventral flange of the crocodile-normal tarsus is 
present. The calcaneal tuber is directed posteriorly and is narrow. 

Thus the calcaneum of Euparkeria is similar to the crocodile- 
reversed tarsus in all the features by which the crocodile-normal and 
crocodile-reversed tarsi differ. Of special importance is the absence 
of an area homologous to the ventral flange of the astragalus of the 
crocodile-normal tarsus. As was argued above, the absence of this 
area prevents derivation of a crocodile-normal tarsus from a 
crocodile-reversed tarsus without the redevelopment of a lost 
structure. On this basis, Euparkeria can be considered to be more 
closely related to the ornithosuchids than to any pseudosuchian 
with a crocodile-normal tarsus. 

THE TARSUS OF ERYTHROSUCHVS 

An additional problem to be considered is the relationship of 
Erythrosuchus. Sill (1974) showed that a striking similarity exists 
between the skull of Erythrosuchus and rauisuchids, so that a 
relationship between these two groups is probable. Cruickshank 
(1978, 1979), however, felt that the tarsus of Erythrosuchus was 
most similar to that of Euparkeria, and placed Erythrosuchus with 
Euparkeria and the ornithosuchids in his Ornithosuchoidea. 



1981 THE ORIGIN OF THE CROCODILOID TARSI 



19 



cal tub 



fib art 
surf b 




fib art surf 




Figure 1 I. The right astragalus and calcaneum of Euparkeria. A) astragalus; B) 
calcaneum; and C) astragalus and calcaneum in articulation. A and B from 
Bonaparte (1975), C based on drawings of South African Museum specimen SAM 
6049 made by J. Bonaparte. 

Abbreviations: ast art surf, astragala/ articular surface; cal tub, calcaneal 
tuber; fib art surf, fibular articular surface; tib art surf, tibial articular surface. 

A general similarity between the tarsus of Erythrosuchus and 
Euparkeria is present, but this can be attributed to the poor 
ossification of the bones. More fundamental is the similarity 
between the tarsus of Erythrosuchus and Chasmatosaurus. A 
comparison of the calcaneum of these two animals (Fig. 12) shows 
that the calcaneal tuber extends laterally, has thin proximal and 
distal edges, and has a thickened medial buttress that extends across 
the tuber to an expanded cartilage-covered lateral edge. The poor 
ossification of the astragalus of Erythrosuchus obscured the 
structure of the articulation between the astragalus and calcaneum. 



20 



BREVIORA 



No. 464 







Figure 12. The left calcaneum of Erythrosuchus in A) ventral, and B) distal 
views; and Chasmatosaurus in C) ventral, and D) distal views. Erythrosuchus 
drawing based on cast of Bernard Price Institute F. 2069 M 405, Chasmatosaurus 
drawing based on cast of NM C 3016, Nasionale Museum, Bloemfontein. 



although Cruickshank (1978) identifies a perforating foramen, a 
primitive feature that is also present in Chasmatosaurus. 

Thus the tarsus of Erythrosuchus is best considered a poorly 
ossified proterosuchian tarsus. A reduction of ossification is 
commonly seen in aquatic animals, and probably reflects an 
adaptation for an aquatic habitat, rather than the development of a 
mechanically distinct structural complex. A fully terrestrial erythro- 
suchid would, therefore, be expected to have a tarsus like that of 
Chasmatosaurus. Since this kind of tarsus is the structural ancestor 
of the crocodile-normal tarsus, the structure of the erythrosuchid 
tarsus neither supports nor negates a relationship between erythro- 
suchids and rauisuchids. If a relationship between erythrosuchids 
and rauisuchids is accepted on the basis of similarities in the skull, it 
is necessary to assume that the crocodile-normal tarsus originated 
after the origin of this radiation of thecodonts, and therefore the 
tarsus cannot be used as the defining feature of the group. 



1981 



THE ORIGIN OF THE CROCODIl OID TARSI 



21 



SUMMARY 

A summary of the relationships suggested above may be made in 
the form of a phylogenetic diagram (Fig. 13). According to this 
phylogeny. many of the higher taxa of diapsid reptiles are artificial 
assemblages, and major modifications of reptilian classification are 
necessary to accurately reflect reptilian relationships. It would, 
however, be useful to test these relationships through examination 
of other structural complexes before making the necessary changes. 



Ornithosuchidae 



Stagonolepdidae 
Rauisuchidae Gracilisuchus 



Lacertilia 




Tnlophosaundae 



Sauropterygia 



Figure 13. The interrelationships of thecodonts. 



ACKNOWLEDGMENTS 

I would like to thank Dr. R. L. Carroll, at whose suggestion this 
project was undertaken. For reading the manuscript and making 
many useful suggestions, 1 thank Hans Dieter Sues, Museum of 
Comparative Zoology, Harvard University; Michael Parrish, Uni- 
versity of Chicago; Dr. Sankar Chatterjee, Texas Tech University; 
and Robert Long and Dr. S. Welles, University of California, 



22 BREVIORA No. 464 

Berkeley. I also thank Lillian Maloney, who proofread the 
manuscript. Figure 9 was drawn by Linda Krause. Publication costs 
of this study were covered in part by a grant from the Wetmore 
Colles Fund. 



LITERATURE CITED 

Bonaparte, J. F. 1971. Los tetrapodos del sector superior de la formation Los 
Colorados, La Rioja, Argentina. Opera Lilloana, 22: 1-183. 

1975a. Nuevos materials de Lagosuchus talampayensis Romer (Theco- 

dontia-Pseudosuchia) y su significade en el origen de los Saurischia. Acta Geol. 
Lilloana, 13: 5-87. 

1975b. The family Ornithosuchidae (Archosauria: Thecodontia). Col- 



loques int. Cent. Natn. Rech. Scient. No. 218, pp. 485-506. 
Camp, C. L. 1945. Prolacerta and the protorosaurian reptiles. Am. J. Sci., 243: 

17-32, 84-101. 
Carroll, R. L. 1976. Noteosuchus, the oldest known rhynchosaur. Ann. S. Afr. 

Mus., 72: 37-57. 
Charig, A. J., and H. -D. Sues. 1976. Proterosuchia, pp. 1 1-39. In A. J. Charig, 

B. Krebs, H. -D. Sues, and F. Westphal, Handbuch der Palaoherpetologie, 13, 

Stuttgart and Portland, Gustav Fischer, 137 pp. 
Chatterjee, S. 1978. A primitive parasuchid (phytosaur) reptile from the Upper- 

Triassic Maleri Formation of India. Paleontology, 21: 83-127. 
1980. Malerisaurus, a new eosuchian reptile from the late Triassic of 

India. Phil. Trans. Roy. Soc. Lond. (B). 291: 163-200. 
Cruickshank, A. R. I. 1972. The proterosuchian thecodonts, 89-119. In K. A. 

Joysey, and T. S. Kemp (eds.). Studies in Vertebrate Evolution: Essays Pre- 
sented to Dr. F. R. Parrington F. R. S. Edinburgh, Oliver and Boyd, 284 pp. 
1978. The pes of Erythrosuchus africanus Broom. J. Linnean Soc. 

(Zool.), 62: 161-178. 
1979. The ankle joint in some early archosaurs. S. Afr. J. Sci., 75: 



168-178. 
Gow, C. E. 1975. The morphology and relationships of Youngina capensis 

Broom and Prolacerta broomi Parrington. Paleontol. Afr., 18: 89-131. 
Gregory, J. T. 1945. Osteology and relationships of Trilophosaurus. U. of Texas 

Publ. No. 4401, pp. 273-359. 
Krebs, B. 1963. Bau und Function des Tarsus eines Pseudosuchiers aus der Trias 

des Monte San Giorgio (Kanton Tessin, Schweiz.). Palaontol. Z., 37: 88-95. 
1965. Tictinosuchus ferox nov. gen., nov. sp. Ein neuer Pseudosuchier 

aus der Trias des Monte San Giogio. Schweiz. Palaontol. Abh., 81: 1-140. 
1973. Der Tarsus von Rauisuchus (Pseudosuchia, Mittel Trias). Mitt. 



Bayer. Staatsamml. Palaontol. Hist. Geol., 13: 95 101. 
Peyer, B. 1937. Die Triasfauna der Tessiner Kalkalpen, XII Macrocnemus 
bassanii Nopsca. Schweiz. Palaontol. Abh., 59: 1-140. 



1981 THE ORIGIN OF THE CROCODILOID TARSI 23 

Romer, A. S. 1956. The Osteology of the Reptiles. Chicago, University of Chica- 
go Press, 772 pp. 

1971. The Chanares (Argentina) Triassic reptile Fauna X. Two new but 

incompletely known long-limbed pseudosuchians. Mus. Comp. Zool., Breviora 
No. 378. pp. 1-10. 

1972. The Chanares (Argentina Triassic Reptile Fauna, XIII. An early 



ornithosuchid pseudosuchian, Gracilisuchus slipaniaicorum, gen. et sp. nov. 

Mus. Comp. Zool., Breviora No. 389, pp. 1-24. 
Sawin, H. J. 1947. The pseudosuchian reptile Typothorax meadei. J. Paleontol., 

21: 201-238. 
Sill, W. D. 1974. The anatomy of Saurosuchus galilei and the relationships of the 

rauisuchid thecodonts. Bull. Mus. Comp. Zool., 146: 317-362. 
Thllborn, R. A. 1980. The ankle joints of archosaurs. Alcheringa. 4: 241-261. 
von Meyer. H. 1856. Zur Fauna der Vorwelt, Saurier aus dem Kupferschiefer 

der Zechstein-Formation. Frankfut-am-main. 
Walker, A. D. 1961. Triassic reptiles from the Elgin area: Stagonolepis, 

Dasygnathus and their allies. Phil. Trans. Roy. Soc. Lond. (B), 244: 103-204. 
1970. A revision of the Jurassic reptile Hallopus victor (Marsh), with 

remarks on the classification of crocodiles. Phil. Trans. Roy. Soc. Lond. (B), 

257: 323-372. 
Watson, D. M. S. 1958. On Millerosaurus and the early history of the sauropsid 

reptiles. Phil. Trans. Roy. Soc. Lond. (B), 240: 355-400. 
Wild, R. 1973. Die Triasfauna der Tessiner Kalkalpen XXIII. Tanystropheus 

longobardicus (Bassani) (Neue Ergebnisse). Schweiz. Palaontol. Abh., 95: 
, 1-162. 
Young, C. C. 1936. On a new Chasmatosaurus from Sinckiang. Bull. Geol. Soc. 

China, 15: 291-320. 
1964. The pseudosuchians in China. Palaeontol. Sinica, New Ser. C, 19: 

1-205. 



MUS. COMP. ZOOL 

LfHRARY 



MAR 1 8 1985 

BRE fc±?0 R A 

Museum of Comparative Zoology 

US ISSN 0006 9698 

Cambridge, Mass. 30 December 1981 Number 465 



THE STRUCTURE AND RELATIONSHIPS 
OF THE DROMASAURS (REPTILIA: THERAPSIDA) 

Donald Brinkman 1 

Abstract. The dromasaurs are primitive anomodonts that are advanced over the 
venjukoviamorph grade of evolution in the reduction of the postorbital and 
zygomatic branches of the jugal, the reduction of the septomaxilla, and the 
enlargement of the area on the dentary for the insertion of the external mandibular 
muscle. Derived features that distinguish dromasaurs from dicynodonts are the lack 
of canines, the presence of a tall, slender postorbital, and the rodlike lower temporal 
bar. The three known dromasaurs form a structural series showing changes in the 
proportions of the face, a loss of premaxillary teeth, changes in the curvature of the 
humerus, and a loss of the ectepicondylar foramen. 



INTRODUCTION 

Frequently, the earliest representatives of higher taxa include 
groups that are at a primitive grade of evolution but are, in some 
features, highly specialized. Within the Therapsida, one such group 
is the Dromasauria. The first dromasaur to be recognized, Gale- 
chirus, was initially considered to be more primitive than any other 
therapsid known at that time (Broom, 1907). Later, two additional 
genera, Galepus and Galeops, were recognized, with Galeops being 
considered sufficiently distinct to be placed in its own family, the 
Galeopsidae (Broom, 1910, 1912). The better understanding of the 
dromasaur skull provided by these genera showed that they were, in 
many respects, derived from the primitive therapsid condition, and 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. 



2 BREVIORA No. 465 

Broom (1914: 12) suggested that they were "an aberrant group of 
small primitive therapsids." 

While there has been little question of the reality of the group, its 
relationships have been a matter of much uncertainty. Romer and 
Price (1940) noted that many of the features that separated droma- 
saurs from other therapsids were primitive for pelycosaurs of the 
suborder Edaphosauria, implying that a close relationship existed 
between the two groups. Later, in a review of therapsid classifica- 
tion, Romer and Watson (1956) suggested that a relationship 
between dromasaurs and dicynodonts was likely, and placed the 
dromasauria within the Suborder Anomodontia as an infraorder 
equal in rank to the Dicynodontia. The possibility that both of these 
relationships are correct was raised by Olson ( 1962), who suggested 
that the therapsids had a multiple origin, with the anomodonts 
originating from within the Edaphosauria. 

Uncertainty about the relationships of the dromasaurs is a result 
of an incomplete understanding of the anatomy of the genera 
comprising the group. Although much of the skeleton is preserved, 
the original material consists of natural casts, so that Broom was 
able to determine little more than the outlines of the bones. 
Recognizing these problems, it was decided to reexamine the 
specimens studied by Broom using the methods developed for the 
study of specimens preserved as natural casts by Baird (1951, 1955) 
and Carroll (1976), in which latex peels of the specimens provide a 
positive image of the bones. Because it is the most completely 
preserved of the dromasaurs, Galeops will be described first. 

G A LEO PS 
Materials and Methods: Galeops is represented by a single 
species, Galeops whaitsi, and by a single specimen, AMNH 5536, 
that comes from the Tapinocephalus zone of South Africa (Kitch- 
ing, 1977). It consists of the impressions of the skull and the front 
half of the postcranial skeleton in a fine-grained sandstone. Both 
part and counterpart are present. Latex peels of the specimen 
provided a positive image of the skeleton, and in some cases, it was 
possible to cast the two halves of the individual elements and join 
them together to give three-dimensional replicas of the original 
bone. 



1981 THE DROMASAURS 3 

Skull Roof: The skull of Galeops (Figs. 1-5) is small for a 
therapsid, its length from snout to quadrate being 55 mm. The 
orbits and external narial openings are large and face laterally. The 
face and postorbital regions are short. Sclerotic rings are present. 
The ventral margin of the postorbital region is deeply emarginated, 
reducing the zygomatic bar to a slender rod. The posterior edge of 
the upper temporal opening is folded backwards, providing a large 
area for the temporal muscles. No such outfolding of the squamosal 
is present ventral to the zygomatic bar. The occiput slopes forward 
and grades into the skull roof. A large pineal opening is present at 
the level of the frontal-parietal suture ( Fig. 4). The teeth are reduced 
in size and are restricted to the maxilla — presumably a horny beak 
would have been present anteriorly. 

Much of the lateral surface of the face is formed by the maxilla. In 
lateral view, this bone is triangular with its apex directed dorsally. It 
is sculptured by numerous small pits and grooves, with these being 
most strongly developed anteriorly. The relationship of the maxilla 
to the surrounding bones of the face is similar to that of dicyno- 
donts: it articulates with the jugal, lacrimal, and prefrontal poste- 
riorly: the nasal dorsally; and the premaxilla midway below the 
external narial opening. The contacts of the maxilla with the 
prefrontal and nasal are strongly interdigitating sutures. The lacri- 
mal appears to overlay the maxilla, with the suture line being 
straight in external view. The details of the remaining contacts are 
uncertain. 

Ventrally, the maxilla has a well-developed palatal flange that 
borders the internal narial opening (Fig. 5). Four sockets are present 
in the left and five in the right maxilla. These increase in size 
posteriorly from the first to third, with the fourth being about equal 
in size to the second in the case of the right maxilla. A distinct 
canine is not present. 

A small portion of the lateral surface of the premaxilla is present 
ventral to the external narial opening. This has a roughened surface 
like that of the maxilla. Ventrally, the paired premaxillae have large 
palatal flanges that form a dicynodontlike secondary palate ( Fig. 5). 
The ventral surfaces of these flanges are roughened by numerous 
posteriorly projecting spicules of bone. A distinct suture separates 
the two premaxillae from each other. 



BREVIORA 



No. 465 




1981 THE DROMASAURS 5 

A small septomaxilla spans the premaxilla-maxilla suture (Fig. 
1). It has no exposure on the lateral surface of the face. The 
septomaxilla foramen is a small opening located between the 
septomaxilla and maxilla. 

The nasal has a large exposure on the lateral surface of the face 
and forms the dorsal half of the posterior margin of the external 
narial opening. The anterior three quarters of the bone has a 
roughened pitted texture like that of the maxilla. Posterior to this, 
the lateral surface of the nasal is smooth. 

The lacrimal, in lateral view, appears to be a small subcircular 
bone fitting between the maxilla, jugal, and prefrontal. The orbital 
edge of the bone has an internally directed flange, although this is 
incompletely preserved (Fig. 2). 

The jugal is reduced to a slender rodlike bone forming the ventral 
margin of the orbit and articulating with the postfrontal and 
squamosal posteriorly and with the maxilla and lacrimal anteriorly. 
No contribution to the formation of the postorbital or zygomatic 
bar is made by the jugal. Anteriorly, a well-defined groove is present 
on the lateral surface of the jugal. This may have been associated 
with the horny beak. 

The prefrontal is a triangular bone forming the anterodorsal edge 
of the orbit. It contacts the lacrimal ventrally, the maxilla and nasal 
anteriorly, and the frontal dorsally. All of these sutures are squa- 
mous interdigitating contacts. The prefrontal is restricted to the 
lateral surface of the face, the transition from the lateral to dorsal 
surface occurring at the area of the prefrontal-frontal contact. The 
lateral surface of the prefrontal is smooth. 

The frontals are seen in ventral view. Each frontal contacts the 
nasal and prefrontal anteriorly, the postfrontal and parietal poste- 
riorly, and its fellow at the midline. No preparietal bone is present. 



Figure 1. Galeops whaitsi. Drawing based on latex peel taken from AMNH- 
5536. 

Abbreviations: an, angular; ar. articular; C, clavicle; Co, coracoid; d, dentary; f, 
frontal; H, humerus; i, intermedium; IC, interclavicle; j, jugal; I, lacrymal; m, 
maxilla; n, nasal; p, parietal; pa, prearticular; PCo, procoracoid; pf, postfrontal; pm. 
premaxilla; po, postorbital; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; 
qj, quadratojugal; r, radiale; R, radius; S, scapula; sa, surangular; sm, septomaxilla; 
sq, squamosal; u, ulnare; U, ulna. 



BREVIORA 



No. 465 




1981 THE DROMASAURS 7 

so the frontals form the anterior border of the pineal foramen. 
Laterally, they contribute to the margin of the orbit. Ventrally, 
strong ridges form the border of the nasal tracts. 

The postfrontal is a small, triangular bone wedged between the 
frontal and postorbital. It has a smooth concave lateral surface. 

The postorbital is a tall, narrow bone separating the orbit from 
the temporal opening. It is supported by the jugal and squamosal 
ventrally and articulates with the parietal and postfrontal dorsally. 
It does not extend posteriorly along the parietal to meet the 
squamosal as is the case in dicynodonts and primitive therapsids 
generally. 

The squamosal is a complex bone that is, in its general structure, 
like that of dicynodonts. As in dicynodonts, the main body of the 
bone forms the lateral portion of the occipital plate. The dorsal 
portion of this area is folded posteriorly so that the posterior rim of 
the temporal fenestra overhangs the occiput. No such outfolding is 
seen ventral to the zygomatic bar. A ventrally directed ramus 
supports the quadrate and quadratojugal and forms the lateral 
portion of the occipital plate. Laterally, an anteriorly directed 
zygomatic ramus extends to the jugal. This is a narrow rodlike bone, 
rather than being flat and beamlike as in dicynodonts. The zygoma- 
tic ramus of the squamosal overlaps the jugal, extending well 
anterior to the posterior edge of the orbit. 

The parietals are seen in ventral view (Fig. 4). They have a sutural 
contact with the frontals and postorbitals anteriorly and with the 
postparietal posteriorly. Large tabulars are probably present as 
well, but sutures cannot be clearly identified. Ventrally, each 
parietal has a crescent-shaped ridge that served as the area of 
attachment of the epipterygoid and braincase to the skull roof. The 



Figure 2. Galeups whaitsi. Drawing based on latex peel taken from counterpart 
slab of AMNH 5536. Skull and lower jaw bones are shown from medial surfaces. 

Abbreviations: aa, atlas arch; ac, atlas centrum; an. angular; ar, articular; ax, axis; 
C, clavicle; Co. coracoid; d, dentary; f, frontal; H. humerus; ic. intercentrum; IC, 
interclavicle; j. jugal; 1, lacrymal; m. maxilla; n. nasal; p. parietal; PCo, procoracoid; 
pf, postfrontal; pm, premaxilla; prf, prefrontal; pro, prootic; q, quadrate; R, radius; 
S. scapula; sp. splenial; sq, squamosal; st pit. stapedial pit; U. ulna. 



BREVIORA 



No. 465 




Figure 3. Galeops whaitsi. A) reconstruction of skull and lower jaw; B) 
reconstruction of lower jaw, medial view; C) reconstruction of articular, dorsoposte- 
rior view. 

Abbreviations: see Figure 2. 



parietals extend lateral to this ridge as a ventrally facing shelf that 
would have provided an area for muscle attachment. This shelf 
forms a portion of the lateral edge of the temporal fenestra. 
Anteriorly, a cup-shaped depression is present just anterior and 
lateral to the area of attachment of the braincase. 

The quadratojugal is a small splintlike bone resting on the 
posterolateral corner of the quadrate ventrally and fitting in a 
groove on the anterior face of the squamosal dorsally. An oval 
quadratojugal foramen lies between the quadrate and quadrato- 
jugal. 

The quadrate is a large bone with a dorsally directed process that 
fits in a groove in the squamosal, an anteriorly directed flange that 



1981 



THE DROMASAURS 



meets the pterygoid, and a ventral expansion that forms the 
articular surface for the lower jaw. The articular surface is divided 
into two condyles separated by a deep groove. The lateral condyle is 
located slightly posterior to the inner condyle. A pit is present on the 
internal surface of the quadrate that would have received the stapes, 
although the stapes itself is not preserved. 

Braincase: The braincase of Galeops is represented by the internal 
surface of the dorsal half of the occiput (Fig. 4), the lateral surface 
of the prootic (Fig. 1), and the ventral surface of the basisphenoid 
(Fig. 5). No sutures can be seen in the occiput. A small posttemporal 
fenestra is present, surrounded by the squamosal laterally and the 
occipital plate medially. A groove on the lateral surface of the 
prootic leads to the temporal fenestra. Two large paired internal 
carotid foramina are present in the basisphenoid just medial and 



par f 




Figure 4. Galeops whaitsi. Internal view of posterior surface of skull drawn 
from latex peel of AMNH 5536. 



Abbreviations: oc, occiput; p, parietal; par f, parietal foramen; pp. postparietal; 
pro, prootic; pt, pterygoid; ptf, posttemporal fenestra; q, quadrate; qj, quadratojugal; 
sq, squamosal; t, tabular. 



10 BREVIORA No. 465 

posterior to the basipterygoid articulation. Grooves in the basi- 
sphenoid lead to these foramina from the posterior edge of the bone. 
Posterior to the basipterygoid articulation, the lateral edge of the 
basisphenoid extends ventrally as a large semicircular flange that 
forms the ventral edge of the fenestra ovalis. This is located well 
above the level of the stapedial pit on the quadrate, so the stapes 
would have sloped ventrally, rather than being nearly horizontal as 
in dicynodonts. 

Palate: The palate of Galeops (Fig. 5) is primitive in its general 
structure; a full complement of palatal bones is present, the. 
transverse flange of the pterygoid is present, and the interptery- 
goidal vacuity is small. An advanced feature is the enlargement of 
the palatal flanges of the premaxillae to form a secondary palate. 
The vomers, as in dicynodonts, form vertical plates, although in 
Galeops they are separate ventrally. The palatines have the relation- 
ships seen in the primitive anomodont Otsheria: each palatine 
contacts the maxilla and ectopterygoid laterally, the vomers and 
opposite palatine medially, and the pterygoid posteriorly. The 
palatines are arched to roof the posterior portion of the internal 
narial openings. A sharp ridge borders this vaulted area laterally, 
suggesting that a soft palate was present. The ectopterygoid is a 
small rectangular bone extending from the transverse flange of the 
pterygoid to the maxilla. The bone has been displaced slightly, so it 
is impossible to determine if a lateral palatal foramen was present as 
in dicynodonts (Cluver, 1975). The pterygoids are partially sepa- 
rated by a small interpterygoid vacuity similar in size and propor- 
tions to that of Otsheria. The two pterygoids contact one another 
anterior to this vacuity but do not meet posteriorly, as is the case in 
dicynodonts, including Eodicynodon, the earliest known dicyno- 
dont (Barry, 1974). The basipterygoid articulation is unfused. The 
lateral edge of the transverse flange of the pterygoid is expanded 
and covered by unfinished bone. 

Lower Jaw: The lower jaw of Galeops (Figs. 1-3) is short and 
deep with a vertical unfused symphysis and a straight ventral edge. 
A fenestra is present on the lateral surface of the jaw between the 
dentary, surangular, and angular. A distinct coronoid process is not 
present, although the posterior edge of the dentary extends above 



1981 



THF DROMASAIRS 



I I 





Figure 5. The palate of Galeops whaitsi. A) as preserved in AMN'H 5536; B) 
reconstruction. 

Abbreviations: bsp. basisphenoid; ect, ectopterygoid: ic, foramen for internal 
carotid artery; m, maxilla; pal, palatine; pm, premaxilla; pt, pterygoid; ptf, transverse 
flange of pterygoid; v, vomer. 

the level of the postdentary bones. A small ventrally directed 
retroarticular process is present. 

The dentary forms the anterior half of the lower jaw. Except for a 
small posterior area, the lateral surface of the dentary is covered by 
numerous small pits and grooves. Posteriorly, the surface is smooth 
and is depressed below the anterior portion of the bone. This 
arrangement, which is similar to that of dicynodonts, suggests that 
the muscles extended onto the posterior portion of the dentary and 
that a horny beak was present anteriorly. In contrast to most 
dicynodonts, no longitudinal groove is present on the upper edge of 
the dentary lateral to the tooth row. 



12 BREVIORA No. 465 

Internally, only the upper half of the dentary is exposed, the 
ventral half being covered by the splenial. The anterior end of the 
dentary is without teeth; posterior to this, eight sockets are present 
for teeth along the lateral margin of the bone. The anterior sockets 
are slightly larger than the posterior sockets, indicating that tooth 
size increased anteriorly. There is no indication that a distinct 
canine was present. All of the sockets are vertical. The tooth row of 
the dentary is longer than the tooth row of the maxilla, suggesting 
that some anteroposterior movement of the jaw occurred during 
closing. 

The splenial is restricted to the internal surface of the lower jaw. 
Posteriorly, the splenial is wedged between the angular and prear- 
ticular. It is incompletely preserved anteriorly, although grooves in 
the dentary show that it extended to the symphysis. 

The angular forms most of the lateral surface of the posterior half 
of the jaw. Its most prominent feature is the reflected lamina, which 
separates from the angular at the posterior edge of the lateral 
mandibular fenestra and extends posteriorly and ventrally as a thin 
sheath of bone. 

The prearticular is a broad crescent-shaped bone that extends 
from the articular to the dentary. It forms the ventral edge of the 
lower jaw posteriorly, but its lateral exposure is reduced anteriorly 
so that at the posterior edge of the reflected lamina it is restricted to 
the internal surface of the jaw. The prearticular most probably 
formed the margin of the adductor fossa, but since the dorsal 
portion of the bone is not preserved, the full extent of the fossa is 
unknown. 

The surangular extends from the dentary to the articular, forming 
the dorsal margin of the postdentary region of the jaw. It is 
extensively overlapped by the angular so that, in lateral view, it is 
restricted to the dorsal margin of the jaw, but in medial view, it 
extends halfway down the internal surface of the jaw. 

The articular is supported by the prearticular, angular, and 
surangular. It forms both the articular surface for the quadrate and 
the retroarticular process. The articular surface faces strongly 
posteriorly. It is differentiated into two grooves separated by a 
rounded ridge. The lateral groove is elongate anteroposteriorly, 
strongly concave mediolaterally and slightly so anteroposteriorly. A 
small medially directed shelf forms the medial groove. 

The internal surface of the postdentary region is partially covered 



1981 



THE DROMASAURS 



13 



by the transverse flange of the pterygoid, so the presence of a 
coronoid is uncertain. 

Vertebrae: The first fourteen vertebrae of Galeops are present 
( Figs. 6, 7), although many of these are incompletely preserved. Cox 
(1959) differentiated the cervical from the trunk vertebrae of the 
dicynodont Kingoria on the basis of the size of the parapophysis 
and diapophysis and the thickness of the associated ribs, the cervical 
vertebrae having more poorly developed parapophyses and more 
slender ribs. In Galeops, a well-developed parapophysis is first seen 
in the seventh vertebra. No parapophysis can be seen on the 
corresponding area of the sixth vertebra, although a small one may 
have been present ventrally. Thus, the first six vertebrae can be 
considered to be cervicals. 

The atlas-axis complex is well-preserved (Fig. 6). It is primitive in 
its general construction: the atlas centrum and axis are separate, and 
judging from the articular surfaces, separate atlas and axis inter- 
centra and proatlases would have been present. The atlas arches 



pro at 
art surf < ? a 




Figure 6. The atlas-axis complex of Galeops whaitsi. A) lateral view; B) anterior 
view of atlas arch and centrum. Drawing based on latex peels from AMNH 5536. 

Abbreviations: aa. atlas arch; aa art surf, articular surface for atlas arch; ac. atlas 
centrum; ax, axis; intc, intercentrum; intc art surf, articular surface for intercentrum; 
prezyg, pre/ygopophysis; pro at art surf, articular surface for proatlas. 



14 BREVIORA No. 465 

meet above the neural canal. Transverse processes are present on 
these elements sloping posteriorly and ventrally. The atlas centrum 
does not extend to the ventral edge of the vertebral column. It has a 
trefoil-shaped articular surface anteriorly. The neural spine of the 
axis is not preserved. The prezygapophysis is convex and faces 
dorsolaterally. A well-developed transverse process is present on the 
axis extending posteriorly and ventrally from the base of the neural 
arch. 

Both the cervical and dorsal vertebrae are deeply amphicoelous. 
A sharp ventral keel is present on the centra. The neural arches are 
not fused to the centrum, although the sutural attachment is 
intimate. The base of the neural arch is located anteriorly on the 
centrum. A large intercentrum is present between the sixth and 
seventh vertebrae, and presumably intercentra would have been 
present between the more anterior vertebrae. A small intercentrum 
is located between the seventh and eighth vertebrae. None is present 
posterior to this. 

A number of structural changes can be seen posteriorly along the 
vertebral column: 

1) The transverse process moves dorsally. The transverse process 
of the axis is located at the dorsal margin of the centrum. On the 
fourth vertebra, it is located more dorsally on the lateral surface of 
the neural arch. The position of the process of the following three 
vertebrae is similar. A further dorsal migration is seen between the 
seventh and ninth vertebrae, with the transverse process reaching 
the level of the zygapophysis. 

2) The inclination of the transverse process changes from ventral 
to dorsal. On the axis, the transverse process slopes ventrally and 
posteriorly. On the fourth vertebra, the process is nearly horizontal 
and is directed laterally. On the eighth vertebra, a distinct dorsal 
inclination is seen. This is further accentuated on the more posterior 
vertebrae. 

3) The parapophyses move dorsally. A distinct parapophysis is 
first seen on the seventh vertebra, where it is located well down on 
the anterior edge of the centrum. The intercentrum and posterior 
edge of the sixth vertebra also contribute to its formation. The 
parapophysis of the eighth vertebra is located slightly higher, and 
the articular surface does not extend onto the seventh vertebra. The 
parapophysis of the eleventh vertebra is located about midway 



1981 THE DROMASAURS 15 



J U U a vfv^v., v jv 



tp v_, — * — J \ .—^V 




Figure 7. The vertebral column posterior to the axis of Galeops whaitsi in A) 
dorsal and B) lateral views. Based on latex peels taken from AMNH 5536. 

Abbreviations: ic, intercentrum; pp. parapophysis; tp, transverse process. 



between the dorsal and ventral edges of the centrum. On the 
fourteenth vertebra, the parapophysis is located on the dorsal 
margin of the centrum close to the transverse process, although the 
capitular and tubercular articular surfaces of the rib head remain 
distinct. There is no indication that the diapophysis migrates 
ventrally from the transverse process, as in dicynodonts (Cox, 
1959). 

4) The zygapophyses move closer to the midline. On the sixth 
vertebra, the first vertebra on which the zygapophyses are clearly 
preserved, the zygapophyses are located at the edge of the centrum 
and are inclined about 45° to the horizontal. The seventh to 
eleventh vertebrae show a medial movement of the zygapophyses 
and an increase in the angle that they make with the horizontal from 
about 45° to about 60°. The medial movement of the postzygapo- 
physes results in a coalescence of the processes supporting the 
articular surfaces, resulting in a single posteriorly directed process. 

Ribs: Of the cervical ribs of Galeops, only those associated with 
the fifth and sixth vertebrae are preserved (Figs. 1, 2). Although 
both of these are incomplete, they are distinctly less robust than the 
more posterior ribs. 

The first nine thoracic ribs are at least partially preserved. All of 
these are double-headed. The tuberculum terminates the shaft of the 
rib, and the capitular articular surface is formed by a process that 
extends ventrally at an angle of about 35° to the shaft. The thoracic 
ribs are only slightly curved. In the right rib associated with the 



16 BREVIORA No. 465 

twelfth vertebra, the shaft of the rib extends nearly straight from the 
tubercular surface for about a quarter of the length of the rib. Thus 
Galeops probably had a tall laterally compressed body. 

Pectoral Girdle: The pectoral girdle of Galeops is formed by the 
paired clavicles, coracoids, procoracoids, and scapulae and by a 
single median interclavicle (Figs. 1, 2). There is no evidence that 
either a cleithrum or an ossified sternum was present. 

The scapula has a tall, slender blade and an expanded platelike 
base. A prominent tubercle for the scapular head of the triceps is 
present on the posterior edge of the bone just proximal to the 
glenoid. A distinct acromion process is not present, nor is a scapular 
spine developed as is the case in dicynodonts (Boonstra, 1966). 

The coracoid and procoracoid are subequal in size. The glenoid is 
formed by the scapula and coracoid. The primitive screw-shaped 
structure of the glenoid has been lost; the glenoid of Galeops is short 
and faces posterolateral^. The coracoid extends posterior to the 
glenoid. The coracoid foramen is located within the procoracoid. 

The clavicle has a triangular base and a rodlike stem. The 
interclavicle is a long paddle-shaped element. Its head is not 
preserved. Distinct grooves for a sternum are not present in the 
stem. 

Humerus: The humerus of Galeops is a slender bone with 
moderately expanded proximal and distal ends in a distinct shaft 
(Fig. 8). The proximal and distal ends are distinctly, but not 
strongly, twisted on one another; when viewed proximally, the angle 
between the long axes of the two ends is about 40°. The proximal 
end of the humerus curves strongly dorsally; when seen in antero- 
dorsal view (Fig. 8B), the dorsal third of the humerus makes an 
angle of about 120° with the distal third of the humerus. The 
entepicondyle is well developed, and an entepicondylar foramen is 
present. The ectepicondyle is poorly developed, and no ectepicon- 
dylar foramen is present. A groove passing proximodistally along 
the lateral edge of the bone presumably held the nerves and vessels 
that usually pass through the ectepicondylar foramen. The ulnar 
articular surface is distinct and well developed. It faces primarily 
ventrally, although it extends onto the distal end of the humerus. 
The radial condyle, although distinct, is not a strongly rounded 
capitulum like that seen in gorgonopsians and dicynodonts (Boon- 
stra, 1965, 1966). 



1981 



THE DROMASAURS 



17 







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18 BREVIORA No. 465 

Radius: The radius of Galeops is a slender bone about 60 percent 
of the length of the humerus. The proximal articular surface is oval 
in outline and is slightly concave. The distal articular surface is also 
oval. 

Ulna: The ulna of Galeops is slightly longer than the radius (Figs. 
I, 2). The olecranon process is prominent, extending well past the 
humeral articular surface. The humeral articular surface is a well- 
defined notch. Lateral to this, a distinct articular surface for the 
radius is present. 

Carpus: Only the dorsal surface of the proximal carpal bones of 
Galeops is seen (Fig. 1). The radiale is short and has a small area of 
finished bone on its dorsal surface. The intermedium and ulnare are 
elongate. They have a large area of contact with one another 
proximal to the perforating foramen. A portion of the lateral 
centrale is present. This supports the intermedium and forms the 
distolateral corner of the perforating foramen. 

GALE PUS 

Materials and Methods: Galepus is represented by a single 
species, Galepus jouberti, and by a single specimen, AMNH 5541, 
that comes from the Cistecephalus zone of the Beaufort series of 
South Africa. Much of the skull and skeleton is represented by 
impressions in a coarse sandstone from which latex peels were 
made. Unfortunately, the counterpart block was not preserved, so it 
was not possible to cast individual elements in three dimensions. 

Skull: The skull of Galepus is represented by a cast of the internal 
surface of the roofing bones and of the palatal flanges of the 
premaxillae (Figs. 9, 12). It is similar to that of Galeops in its 
general structure: the orbits and external narial openings are large; 
the face and postorbital region of the skull are short; a large oval 
pineal opening is present; the ventral margin of the postorbital 
region of the skull is deeply emarginated; the occiput slopes 
forwards; the orbit is bounded posteriorly by a tall narrow postor- 
bital; a secondary palate formed by the palatal flanges of the 
premaxillae is present; and the internal surface of the parietal is 
marked by a crescentic ridge and a cup-shaped depression antero- 



198 



THE DROMASAURS 



19 




Figure 9. Galepus jouberti. Drawing based on latex peel taken from AMNH 
5541. Skull and lower jaw bones viewed from medial surface. 



Abbreviations: C, clavicle; cal, calcaneum; cen, centrale; Co, coracoid; ect for. 
ectepicondylar foramen; F, femur; H, humerus; IC, interclavicle; IL, ilium; IS, 
ischium; mc 1, first metacarpal; mt 5, fifth metatarsal; PCo, procoracoid; PU. pubis; 
r, radiale; R, radius; U, ulna. 



20 BREVIORA No. 465 

lateral to this ridge. Unfortunately the sutures referred to by Broom 
(1914) have been obliterated, so the exact relationships of the 
individual skull elements are uncertain. 

The left lower jaw is visible in internal view. It is more slender 
than the jaw of Galeops, and its outline differs: the ventral margin of 
the jaw, rather than being straight as in Galeops, is slightly concave 
in profile, and the dorsal margin of the postdentary region extends 
nearly straight from the dentary to the articular, rather than being 
gently convex as in Galeops. A splenial is not preserved, but grooves 
in the dentary indicate that it extended to the symphysis. The 
sutures between the postdentary bones cannot be identified. 

Six small peglike teeth are preserved in the dentary and an empty 
aveolus is present anterior to these. A small edentulous region is 
present anterior to the tooth row. Unlike Galeops where the sockets, 
and presumably the teeth they contained, are vertical, the teeth of 
Galepus slope anteriorly. 



Postcranial Skeleton: A series of eighteen dorsal vertebrae of 
Galepus is present. These are deeply amphicoelous and show a 
sutural attachment between the neural arch and centrum. The 
structure of the neural arch is seen clearly only in the fifth vertebra 
from the anterior end of the preserved series. It is like the neural 
arch of the corresponding vertebra in Galeops: the zygapophyses are 
strongly tilted and are near the midline, the postzygapophyses are 
coalesced so the articular surfaces appear to be formed by a single 
posteriorly directed process, and the transverse processes are large 
dorsolateral^ directed structures located at the level of the zyga- 
pophyses. 

The caudal region is represented by two disconnected series of 
vertebrae, an anterior series of nine vertebrae and a posterior series 
of eight vertebrae. They are separated by a space that would have 
been occupied by six vertebrae and are separated from the sacral 
region by a space that would have been occupied by about eight 
vertebrae. Thus the tail would have included at least 31 vertebrae. 
These decrease in both height and length posteriorly. The zyga- 
pophyses slope less steeply than in the dorsal series. A prominent 
transverse process is located at the base of the neural arch on the 
first vertebra in the series. No transverse process is seen on any of 
the more posterior vertebrae. Haemal spines and arches are present 



1981 THE DROMASAURS 21 

between all the vertebrae in the proximal series and between the first 
two vertebrae in the posterior series. 

Eighteen ribs are at least partially preserved on the right and six 
on the left side. In all the ribs, the curvature is greatest near the 
head, with the distal half of the rib being nearly straight and at an 
angle of 30° to the proximal portion. The heads of the first three 
ribs on the left side and the last two ribs on the right side are visible. 
The anterior ribs are double-headed. The posterior ribs are single- 
headed. 

The pectoral girdle is seen in internal view. A large interclavicle is 
present with a paddle-shaped stem and a diamond-shaped head. No 
ossified sternum is present. The clavicle has a long narrow riblike 
stem and a triangular head. The coracoids, procoracoids, and 
scapulae are too poorly preserved for their structure to be fully 
determined. 

The humerus of Galepus is like that of Galeops in its proportions 
and general features. In both, the entepicondyle is well developed, 
the ectepicondyle is reduced, the proximal and distal ends are 
distinctly twisted on one another, the ulnar surface faces strongly 
ventral but also extends onto the distal surface of the bone, and the 
pectoral crest is restricted to the proximal third of the bone. The 
humerus of Galepus differs from that of Galeops in having an 
ectepicondylar foramen and in the lack of an upturning of the 
proximal end, the dorsal edge being nearly straight in anterodorsal 
view. 

Only the medial surface of the proximal end of the ulna is 
preserved. As in Galeops an olecranon process is present, and the 
medial surface of the proximal end of the ulna is concave. The right 
radius is seen in lateral aspect. It is like the radius of Galeops in its 
proportions and in having a slight S-shaped curve. 

The hand overlies the foot, although much of the hand is missing, 
exposing the metatarsals and phalanges of the foot. 

Only three elements of the carpus are present, these being the 
radiale, lateral centrale and first distal carpal. The radiale is short 
and without finished bone. The first distal carpal is subcircular and 
has a large area of finished bone on its ventral surface. 

Of the metacarpus, only the first metacarpal is present. This is a 
short phalangelike element. A large tubercle is present on the lateral 
edge of the bone near its proximal end. The toes are represented by 
three long, clawlike terminal phalanges. 



22 breviora No. 465 

The pelvis is seen in internal view. It is primitive in having a large 
pubis and ischium and in having well-developed anterior and 
posterior extensions of the iliac blade. An advanced feature is the 
enlargement of the obturator foramen. 

The dorsal surface of the proximal end of the left femur is visible. 
The proximal end of the femur is tilted dorsally about 35° from the 
shaft of the bone. 

The tarsus is represented by an incompletely preserved right 
calcaneum, centrale, and the first four distal tarsals, all visible in 
ventral view. The calcaneum is a tall, platelike element with a notch 
for the perforating foramen on its medial edge. The centrale is a 
cubical cartilage-covered element. The first three distal tarsals are 
subequal in size. The fourth is larger, although it is less than twice 
the size of the third in its linear dimensions. 

All five metatarsals are present, visible in ventral view. They 
increase in size from the first to the fourth with the fifth being about 
equal to the third in length. The phalangeal formula of the pes is 
2, 3,?,?, 3. The terminal phalanges, where known, are elongate, 
clawlike elements. 



GALECHIRUS 

Materials and Methods: Galechirus is represented by a single 
species, Galechirus scholtzi, and by two specimens: SAM 1068, 
which shows the front half of the skull and the lower jaw in lateral 
view and much of the postcranial skeleton (Fig. 10); and AMNH 
5516 (now housed in the South African Museum), which shows a 
pelvic and pectoral limb and fragmentary ribs and vertebrae (Fig. 
1 1). Both of these specimens come from the Cistecephalus zone of 
the Beaufort series of South Africa (Kitching, 1977). They are 
preserved as natural casts. Molds taken from these specimens 
provided a positive image that served as the basis for the descrip- 
tions and drawings. 

Skull: The skull of Galechirus (Fig. 10) is like that of Galeops in 
having large orbits and external narial openings, a short temporal 
region, and a deep emargination of the cheek posterior to the orbit. 
In contrast to the condition in Galeops, Galechirus retains premax- 
illary teeth and has a more elongate face. 



1981 



THE DROMASAURS 



23 




Figure 10. Galechirus scholtzi. Drawing based on latex peel taken from SAM 
1068. 



Abbreviations: a. astragalus; ac, atlas centrum; ax. axis; ax ic, axis intercentrum; 
C. clavicle; cal. calcaneum; cer rib, cervicle rib; dt 1, first distal tarsal; ect for, 
ectepicondylar foramen; F, femur; FIB, fibula; H, humerus, i, intermedium; IS 
ischium; j, jugal; m cen. medial centrale; pi, pisiform; pm, premaxilla; prf, prefrontal; 
PU, pubis; T. tibia; u. ulnare; U, ulna. 



24 BREVIORA No. 465 

The arrangement of the bones of the face in Galechirus is similar 
to that of Galeops. The premaxilla-maxilla contact is located 
midway below the external narial opening. A separate bone, 
presumably the septomaxilla, is present internal to the premaxilla- 
maxilla contact. The maxilla meets the nasals midway along the 
posterior edge of the external narial opening. The maxilla-nasal 
suture slopes dorsoposteriorly from this point. Unfortunately, the 
sutures between the maxilla and more posterior bones are obscure, 
so the relationship of these bones is uncertain. Six small peglike 
teeth are present in the maxilla, and an empty socket is present 
posterior to these. The anterior three teeth are subequal in size and 
are slightly larger than the posterior three teeth. 

The premaxillae are separate. Each premaxilla has a well- 
developed dorsal ramus that extends between the nasals and a 
ventral ramus that meets the maxilla midway below the external 
narial opening. The right premaxilla, which is seen in internal view, 
contains two large, procumbent, chisel-shaped teeth. Three teeth are 
present in the preserved portion of the left premaxilla. The posterior 
two are approximately equal to the maxillary teeth in size and are 
vertically oriented. The more anterior tooth is slightly larger and 
slopes anteriorly. The missing portion of the premaxilla probably 
held two large procumbent teeth similar to those seen in the right 
premaxilla. 

The prefrontal, which has been displaced slightly, is a triangular 
element forming the anterodorsal margin of the orbit. Its lateral 
surface is smooth and curves onto the dorsal surface of the skull. 

The suborbital portion of the jugal is a narrow rodlike bone. 
Posteriorly, a groove is present on its lateral surface, probably for 
an overlying zygomatic branch of the squamosal. 

Lower Jaw: The lower jaw of Galechirus is long and slender. Its 
ventral edge curves upwards, and the symphysis slopes forward. The 
dentary forms the anterior two-thirds of the jaw. Its lateral surface is 
covered by numerous small pits and grooves anteriorly. Posteriorly, 
the surface is smooth and is recessed below the more anterior 
portion of the dentary. Presumably the adductor muscles inserted 
on this area of the dentary. Only one tooth is preserved in the 
dentary. This is a small peglike element located well anteriorly. 
There does not appear to be room anterior to this for large 



1981 THE DROMASAURS 25 

procumbent teeth similar to those seen in the premaxilla, nor is 
there any evidence that such teeth were present. 

The dorsal edge of the postdentary region of the jaw curves evenly 
from the posterior edge of the dentary to the articular. A fenestra is 
present between the dentary and the postdentary bones. The 
reflected lamina of the angular separates from the angular at the 
posterior edge of the fenestra. A small ventrally directed retroarticu- 
lar process is present beneath the jaw joint. The sutures between the 
individual bones of the postdentary region of the jaw are not visible. 

Postcranial Skeleton: The vertebral column of Galechirus is 
partially preserved in SAM 1068. Anteriorly, four cervical vertebrae 
are present between the posterior end of the jaw and the pectoral 
girdle. The first vertebra in this series is the atlas, the centrum of 
which is seen in lateral view. As in Galeops, it does not reach the 
ventral surface of the vertebral column. A large axis intercentrum is 
present as a distinct element between the atlas centrum and the axis. 
The following vertebrae are short and broad, their length being 
about equal to their width across the posterior end of the centrum. 
The axis has a posteroventrally directed transverse process located 
well down on the neural arch. The centrum does not have a sharp 
ventral keel, although a low ridge is present. The following two 
vertebrae are seen in ventral view. A distinct ventral keel is present, 
and intercentra are present between each of these vertebrae. 

A series of three dorsal vertebrae, probably the tenth to twelfth, 
lies between the shoulder girdle and the hand. These are seen in 
ventral view. The keel has been reduced and intercentra are absent. 
Distinct articular surfaces for the heads of the ribs are not visible. 

Faint traces of eight presacral vertebrae are present between the 
wrist and the knee. Little of their structure can be determined. 

The caudal vertebrae are represented by two series: an anterior 
series of twelve vertebrae and a more posterior series of three 
vertebrae. A space equal in length to four centra is present between 
the anterior series and the pelvis, and the two sections of the tail are 
separated by a space equal in length to about ten centra. Thus, the 
tail would have had in excess of 29 vertebrae. 

The caudal vertebrae decrease both in height and length poste- 
riorly. Haemal arches are present between each of the first ten 
vertebrae, and transverse processes are present on each of the 



26 



BREVIORA 



No. 465 



vertebrae in the anterior series. The more posterior vertebrae have 
neither transverse processes nor haemal arches. 

Three ribs are preserved in SAM 1068. These are associated with 
the three vertebrae seen between the shoulder girdle and the hand. 
The more anterior of these is clearly double-headed. The separation 
of the two articular surfaces is less distinct posteriorly. 

Small, needle-like gastralia are present posteriorly, but these are 
disturbed so their natural arrangement is unknown. This is the only 
evidence of gastralia in therapsids (Romer, 1956). 




Figure 11. Galechirus scholtzi. Drawing based on cast of AMNH 5516 (now 
housed in the South African Museum). 



Abbreviations: a, astragalus; cal, calcaneum; cen, centrale; ect for, ectepicondylar 
foramen; ent for, entepicondylar foramen; F, femur; FIB, fibula; IL, ilium; IS, 
ischium; 1 cen, lateral centrale; PU, pubis; r, radiale; R, radius; S, scapula; T, tibia; u, 
ulnare; U, ulna. 



1981 THE DROMASAURS 27 

The pectoral girdle is seen in ventral view in SAM 1068 and in 
lateral view in AMNH 5516. The scapulae have tall, slender blades 
and expanded platelike bases. A distinct rugosity on the posterior 
edge of the scapular blade just proximal to the glenoid marks the 
site of the origin of the scapular head of the triceps. An acromion 
process is not present. 

The coracoid and procoracoid are both large, with the coracoid 
extending well posterior to the glenoid. The procoracoid foramen is 
located within the procoracoid just proximal to the glenoid. The 
interclavicle has a diamond-shaped head and a broad paddle-shaped 
stem. The head of the interclavicle is recessed ventrally to receive the 
medial end of the clavicle. 

The left humerus is seen in ventral view in both specimens. The 
proportions of this element are similar to the proportions of the 
humerus of Galeops: it is a slender bone with a well-defined shaft 
and with little expansion of the proximal and distal ends. The 
pectoral crest is restricted to the proximal third of the bone. In 
contrast to the condition in Galeops, both an ectepicondylar and 
ectepicondylar foramen are present. 

The radius is a slender bone, slightly shorter than the humerus. It 
has a slight S-shaped curvature. The ulna is almost identical to that 
of Galeops in its proportions and in the development of the articular 
surfaces, although the olecranon process is less well developed. This 
difference may be a result of differences in the ontogenetic stage of 
development of the specimens. 

A complete carpus is present in SAM 1068 and most of the carpus 
is present in AMNH 5516. All the elements primitively present in 
the synapsid carpus are present, although these are all shortened 
proximodistally so the length of the carpus is less than that of the 
metacarpus. A distinctive feature is the expansion of the ulnare to 
form a flat platelike bone. 

The metacarpals increase in length from the first to the fourth, 
with the fifth being about equal to the second in length. The 
phalangeal formula is 2,3,3,3,3. The terminal phalanges are elon- 
gate, laterally compressed, clawlike elements. 

The pelvis is seen in ventral view in SAM 1068 and in lateral view 
in AMNH 5516. The pubis and ischium are large, platelike ele- 
ments. The pubis is damaged, so the size of the obturator foramen is 
uncertain. The ilium is not completely known, although the pre- 



28 BREVIORA No. 465 

served portion shows that it had a tall, slender blade that extends 
posterior to the acetabulum. 

The femur is seen in postaxial view in both specimens. It is equal 
in length to the humerus. The head of the femur is incompletely 
preserved, although there is no indication of a dorsal tilting of the 
proximal end such as is seen in Galepus. Also, no downturning of 
the distal end of the femur is seen. 

The tibia and fibula are elongate, slender bones approximately 
equal in length to the femur and substantially longer than the radius 
and ulna. Both bones are slightly curved with the concave edges of 
the bones facing one another. The tibia is more robust than the 
fibula, the diameter of the tibia at its narrowest point being about 
twice the minimum diameter of the fibula. 

A nearly complete tarsus is seen in dorsal view in AMNH 5516, 
and an astragalus is seen in dorsal view in SAM 1068. The 
calcaneum is a large platelike bone. The astragalus is L-shaped. A 
perforating foramen passes between the astragalus and calcaneum. 
Distal to the perforating foramen, the articular surface on the 
astragalus for the calcaneum is strongly convex and extends onto 
the dorsal surface of the bone. Proximal to the perforating foramen, 
the articular surface on the astragalus for the calcaneum is concave. 
The tibial articular surface is restricted to the medial edge of the 
astragalus. The centrale is a small, cartilage-covered bone located 
between the astragalus and first four distal tarsals. The distal tarsal 
row consists of four elements, the fifth distal tarsal having been lost. 
The first three are subequal in size, the fourth is about three times 
the size of the third in its linear dimensions. 

The metatarsals increase in length from the first to the fourth, 
with the fifth being about equal to the third in length. The 
phalangeal formula is 2,3,3,3,3. The terminal phalanges are elon- 
gate, laterally compressed clawlike elements. 

DISCUSSION 

In considering the general evolution of therapsids, Hopson(1969) 
recognized three basic groups: the Dinocephalia, characterized by 
an interlocking of the incisors and the presence of a jaw joint that 
permits no anteroposterior movement of the jaw; the Theriodontia, 
characterized by the development of a movable quadrate permitting 
some anteroposterior movement of the lower jaw; and the Anomo- 



1981 THE DROMASAURS 29 

dontia, characterized by the ventral emargination of the cheek 
posterior to the orbit and a jaw joint that permits considerable 
anteroposterior movement of the lower jaw on the fixed quadrate. 
The emargination of the cheek in the dromasaurs immediately 
indicates that they belong within the Anomodontia. This placement 
is supported by features of the skull of dromasaurs that are similar 
to other anomodonts, including: 1) the formation of a secondary 
palate by palatal flanges of the premaxillae and 2) the presence of a 
lateral mandibular fenestra. 

The most primitive anomodonts known are the venjukovia- 
morphs Otsheria and Venjukovia from the Middle Permian of 
Russia. These show the diagnostic feature of an emarginated cheek, 
but, relative to dicynodonts, retain a primitive jaw joint, a primitive 
palate, and primitive features jn the temporal region (Watson, 1948; 
Olson, 1962; Barghusen, 1976). In many respects, the dromasaurs 
are at the same grade of evolution as are the venjukoviamorphs. 
This is demonstrated by the following features, which are shared by 
dromasaurs and the venjukoviamorphs and which are different 
from the more derived condition seen in dicynodonts: 

1) No outfolding of the squamosal is present ventral to the 
zygomatic rarhus of the squamosal. In dicynodonts, such an 
outfolding is present. 

2) The palate is more primitive in showing a small interpterygoid 
vacuity, transverse flanges on the pterygoids, an unfused 
basipterygoid joint, and a smaller secondary palate. In dicyno- 
donts, the interpterygoid vacuity has enlarged so that it 
extends forward between the posterior ends of the vomers, the 
transverse flanges of the pterygoids have been lost, and the 
pterygoids are sutured to one another posterior to the inter- 
pterygoid vacuity (Cluver, 1970). 

3) The articular, as in Venjukovia (Watson, 1948), retains a 
central ridge with two concave depressions on either side of it. 
In dicynodonts, the central ridge is convex in profile, and a 
concave area is present anterior to this (Crompton and 
Hotton, 1967). 

In addition, a number of features seen in the dromasaurs, but not 
known at present in the venjukoviamorphs, are more primitive than 
the condition in dicynodonts. These include: 



30 BREVIORA No. 465 

1) A separate atlas centrum and axis intercentrum are present in 
the dromasaurs. In dicynodonts, these fuse to the axis (Cox, 
1959). However, the atlas centrum and axis may fuse during 
ontogenetic development in the dromasaurs. 

2) No scapular spine or acromion process is present on the 
scapulocoracoid. In dicynodonts, both these features are 
present (Boonstra, 1966). 

3) The shape of the interclavicle and the absence of a sternum in 
the dromasaurs is primitive. In dicynodonts, the interclavicle 
is short and a large sternum is present (Boonstra, 1966). 

4) The iliac blade in the dromasaurs is primitive in having a well- 
developed posterior process. In dicynodonts, the anterior 
ramus of the ilium is more strongly developed than the 
posterior ramus (Boonstra, 1966). 

Despite the primitive structure of the dromasaurs, features are 
present that are advanced over the condition seen in the venjukovia- 
morphs and are similar to the condition seen in dicynodonts. One of 
these is the structure of the jugal. In Otsheria, the jugal is a triradiate 
bone retaining well-developed zygomatic and postorbital branches. 
In Venjukovia, the postorbital branch has been reduced although it 
is still present (Barghusen, 1976). In both dromasaurs and dicyno- 
donts, the postorbital and zygomatic branches of the jugal have 
been reduced or lost. A second advanced feature is the presence of a 
smooth area on the posterior end of the dentary that suggests that 
the adductor muscles extended onto the lateral surface of the 
dentary. In Venjukovia, an external mandibular muscle was present, 
although it had only a small area of insertion on the lateral surface 
of the dentary (Barghusen, 1976). A third advanced feature is the 
structure of the septomaxilla. In Otsheria and Venjukovia, as in 
primitive therapsids generally, the septomaxilla is a large element 
with an exposure on the lateral surface of the face. In dicynodonts 
and dromasaurs, the septomaxilla is reduced; in the dromasaurs, it 
is restricted to the floor of the external narial opening. 

This combination of primitive and derived features demonstrates 
that the dromasaurs are members of a grade of evolution interme- 
diate between the venjukoviamorphs and the dicynodonts, but it 
does not demonstrate that it is a monophyletic group. For this, it is 
necessary to show that features are present in the three genera that 
are advanced over the primitive anomodont condition but are not 



1981 THE DROMASAURS 31 

shared with dicynodonts. The most obvious such feature is the 
dentition, which consists of reduced peglike teeth. This is a clear 
difference from both the primitive anomodont condition and the 
dicynodont condition in which distinct canines are present. An 
additional derived feature that separates dromasaurs from dicyno- 
donts is the presence of a tall, narrow postorbital and a slender, 
rodlike lower temporal bar. In dicynodonts, the postorbital bar is 
low and the lower temporal bar is a flat, beamlike element. 

Also the absence of a contact between the postorbital and 
squamosal is a derived feature different from the condition in both 
dicynodonts and venjukoviamorphs, where the postorbital extends 
along the upper margin of the temporal opening to reach the 
squamosal. However, the only dromasaur in which the posterior 
extent of the postorbital is known is Galeops, so the possibility that 
this is a derived feature present only in that genus cannot be 
discounted. 

Thus the present evidence, although not conclusive, suggests that 
the dromasaurs are a natural group. Within the group, the three 
genera can be placed in a single structural sequence with Galechirus 
being the most primitive and Galeops being the most derived 
member of the sequence. The features that document this series are: 

1) The Tooth Row: In Galechirus, premaxillary teeth are pres- 
ent; in Galepus, premaxillary teeth are absent but the eden- 
tulous region is short; in Galeops, a large edentulous region is 
present. 

2) The Proportions of the Face: In Galechirus, the face is 
relatively long; Galepus and Galeops show a progressive 
reduction in the length of the face (Fig. 12). 

3) The Lower Jaw: In Galechirus, the lower jaw is slender and 
the ventral border is concave; in Galepus and Galeops, a 
progressive decrease in the length of the jaw is seen and the 
ventral margin becomes straight (Fig. 12). 

4) Humerus: In Galechirus and Galepus, the dorsal margin of the 
humerus is straight when seen in anterodorsal view. In 
Galeops, the proximal end of the humerus curves strongly 
dorsally. 

5) Eetepicondylar Foramen: In Galechirus, a well-developed 
ectepicondylar foramen is present; in Galepus, this foramen is 
reduced; in Galeops, an ectepicondylar foramen is absent. 



32 



BREVIORA 



No. 465 






Figure 12. Reconstructions of the skulls of the dromasaurs. A) Galechirus 
sholtzi; B) Galepus jouberti; C) Galeops whaitsi. Not drawn to scale. 



1981 THE DROMASAURS 33 



Since the most derived member of this sequence is the earliest, 
these three genera cannot represent successive stages in a single 
evolving lineage. Rather, they must be regarded as members of 
distinct grades of evolution in a single monopholetic radiation. 

ACKNOWLEDGMENTS 

I would like to thank Dr. R. L. Carroll of McGill University at 
whose suggestion this project was undertaken. Dr. M. A. Cluver 
provided silicone rubber casts of the specimens housed in the South 
African Museum. Drs. M. A. 4 Cluver, A. W. Crompton, and R. L. 
Carroll read the manuscript and made many useful suggestions that 
led to its improvement. Drs. J. A. Hopson and H. R. Barghusen 
freely provided important information on the structure of the 
venjukoviamorphs. I would also like to thank Lillian Mahoney for 
proofreading the manuscript. 

LITERATURE CITED 

Bairp, D. 1951. Latex molds in paleontology. Compass of Sigma Gamma 

Epsilon, 28: 339-345. 
1955. Latex micro-molding and latex-plaster molding mixture. Science, 

122: 202. 
Barghusen, H. R. 1976. Notes on the adductor jaw musculature of Venjukovia, 

a primitive anomodont therapsid from the Permian of the USSR. Ann. S. Afr. 

Mus., 69: 249-260. 
Barry, T. H. 1974. A new dicynodont ancestor from the Upper Ecca (Lower 

Middle Permian) of South Africa. Ann. S. Afr. Mus., 64: 117-136. 
Boonstra, L. D. 1965. The limbs and girdles of the Gorgonopsia of the Tapino- 

cephalus zone. Ann. S. Afr. Mus., 48: 237-249. 
1966. The girdles and limbs of the Dicynodontia of the Tapinocephalus 

zone. Ann. S. Afr. Mus., 50: 1-11. 
Broom, R. 1907. On some new fossil reptiles from the Karroo beds of Victoria 

West, South Africa. Trans. S. Afr. Phil. Soc, 18: 31^2. 
1910. A comparison of the Permian reptiles of North America with those 

of South Africa. Bull. Am. Mus. Nat. Hist., 28: 197-234. 
1912. On some new fossil reptiles from the Permian and Triassic beds 

of South Africa. Proc. Zool. Soc, 1912: 859-876. 
1914. On the origin of mammals. Phil. Trans. Roy. Soc. Lond. B, 206: 



1-48. 



34 BREVIORA No. 465 

Carroll, R. L. 1976. Noteosuchus, the oldest known rhynochosaur. Ann. S. Afr. 

Mus., 72: 37-51. 
Cluver, M. A. 1970. The palate and mandible in some specimens of Dicynodon 

testudirostris Broom & Haughtbn (Reptilia, Therapsida). Ann. S. Afr. Mus., 

56: 133-153. 
1975. A new dicynodont reptile from the Tapinocephalus zone (Karoo 

system, Beaufort series) of South Africa, with evidence of the jaw adductor 

musculature. Ann. S. Afr. Mus., 67: 7-23. 
Cox, C. B. 1959. On the anatomy of a new dicynodont genus with evidence on the 

position of the tympanum. Proc. Zool. Soc. Lond., 132: 231-367. 
Crompton, A. W., and N. Hotton III. 1967. Functional morphology of the 

masticatory apparatus of two dicynodonts (Reptilia, Therapsida). Postilla, 109: 

1-51. 
Hopson, J. A. 1969. The origin and adaptive radiation of mammal-like reptiles 

and nontherian mammals. Ann. New York Acad. Sci., 167: 199-216. 
Kitching, J. W. 1977. The distribution of the Karroo vertebrate fauna. Memoir 

1. Bernard Price Institute for Paleontological research. Johannesburg, Univer- 
sity of Witwatersrand, 131 pp. 
Olson, E. C. 1962. Late Permian terrestrial vertebrates, USA and USSR. Trans. 

Amer. Phil. Soc. N.S., 25: 1-225. 
Romer, A. S. 1956. The Osteology of the Reptiles. Chicago, University of 

Chicago Press, 772 pp. 
Romer, A. S., and L. I. Price. 1940. Review of the Pelycosauria. Geol. Soc. 

Amer. Spec. Pap. No. 28, 538 pp. 
Romer, A. S., and D. M. S. Watson. 1956. A classification of therapsid reptiles. 

Bull. Mus. Comp. Zool., 114: 37-89. 
Watson, D. M. S. 1948. Dicynodon and its allies. Proc. Zool. Soc, 118: 823-877. 



,,,oo. >-* 



3. COMP. l^"*' 

m 1 8 1985 



B R E \$iR) R A 

Miisenm of Comparative Zoology 



US ISSN 0006 9698 



Cambridge, Mass. 30 June 1982 Number 466 



SYSTEMATICS OF THE MEXICANA 

SPECIES GROUP OF THE 

COLUBRID GENUS LAMPROPELTIS, 

WITH AN HYPOTHESIS 

MIMICRY 

William R. Garstka 1 

Abstract. Variation within the mexicana group of kingsnakes. genus Lampropel- 
tis (Fit/inger). is discussed. Three species are recognized: L. mexicana (Garman). L. 
alterna (Broun), and L. rulhveni (Blanchard), which is added to the species group. L. 
leunis (Gunther) is placed in the synonymy of L. mexicana. Qualitative and quantita- 
tive characters of external and internal morphology are used to diagnose the taxa. 
Outgroup comparisons with other Lampropeltis and other colubrids indicate that L. 
mexicana is primitive relative to L. alterna. Evolution within the group and an 
hypothesis of mimicry to explain pattern polymorphisms are discussed. 

L. alterna as interpreted here is restricted to areas north of the Anticline of Arteaga, 
Mexico and occurs throughout the Bolson de Mapimi and north to the foot of the 
Guadalupe Mountains. L. mexicana is a montane and plateau form occurring on and 
to the south of the Anticline of Arteaga. /.. rulhveni has been found in northwestern 
Michoacanand adjacent Queretaro in the transverse volcanic region. Sympatry is not 
recorded for the taxa. 

INTRODUCTION 

The kingsnakes allied with Lampropeltis mexicana (Garman) are 
relatively small (less than one meter) little-known snakes which 
inhabit the Chihuahuan Desert and adjacent areas. They occur from 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02 1 38. Present address: Department of Biological Sciences, University of Alabama in 
Huntsville, Huntsville, Alabama 35899. 



2 BREVIORA No. 466 

northern Michoacan, Mexico, north to perhaps southern New Mex- 
ico. The taxa have been considered closely related and have been 
called variously a species group (Smith, 1942, 1944; Webb, 1961)ora 
species complex (Gehlbach and Baker, 1962; Gehlbach and McCoy, 
1965; Gehlbach, 1967; Tanzer, 1970). The nominal taxa are L. mexi- 
cana (Garman), L. thayeri Loveridge, L. greeri Webb, L. leonis 
Gunther, L. alterna (Brown), and L. blairi Flury. These species have 
been considered as a unit because of the following shared characteris- 
tics: a light-edged red-centered black blotch, mottled speckling, dis- 
tinct head, and long tail (Gehlbach and Baker, 1962). 1 also include in 
this group L. ruthveni (Blanchard) because it clearly fits these crite- 
ria, and is sympatric with L. triangulum. 

The findings reported here suggest that the mexicana group con- 
sists of three species-level taxa which are distinctive in several charac- 
ter suites: L. alterna (including L. alterna and L. blairi), L. mexicana 
(including L. mexicana, L. thayeri, L. greeri, and L. leonis), and L. 
ruthveni (removed from the synonymy of L.t. arcifera). 

The mexicana group is related closely to L. triangulum on the basis 
of shared hemipenial characters (Smith, 1942) and may also show 
close relationship to L. pyromelana and L. zonata when pertinent 
characters are more closely examined in those species. The mexicana 
group has been considered both as ancestral to the remaining species 
of Lampropeltis ( Webb, 1 96 1 ) and as the most derived species (Blan- 
chard, 1921; Smith, 1942; 1944; Tanner, 1953). Data will be presented 
demonstrating that L. alterna is the most derived species in the group, 
and that both it and L. ruthveni are closely related to L. mexicana. 
Although the distribution of all three species appears to be presently 
patchy and perhaps relictual, this is not sufficient justification for 
considering them ancestral. 

Four specimens of Lampropeltis, seemingly related to L. mexi- 
cana, were collected by the author in Coahuila and Nuevo Leon, 
Mexico, during 1975 and 1976. It was not possible to refer three of the 
four specimens to any presently recognized taxon when compared to 
the key in Webb (1961) and the descriptions in Gehlbach (1967). 
Comparison with reference material did not clarify the situation. Of 
the four specimens previously known from the vicinity of Galeana, 
Nuevo Leon (Gehlbach and McCoy, 1965; Gehlbach, 1967), one was 
identified as L. mexicana thayeri, one as L.m. alterna, and two as 
L.m. thayeri X L.m. mexicana intergrades. The only additional 
specimen from Nuevo Leon was the holotype of the obviously closely 



1982 LAMPROPELTIS MEXICANA SPECIES GROUP 3 

related L. leonis. It was clear then that a revision of the goup and 
reidentification of the reference material was necessary. 

One of the above specimens collected by the author (from San 
Lazaro, Coahuila) has a series of 24 laterally truncated body rings; 
thus it should have been classed as L.m. thayeri (Webb, 1961). 
However, in comparison with the type specimen of thayeri and a 
living specimen quite like the type, this Coahuila specimen was 
clearly different. The head was much more triangular, as has been 
noted for altema (Brown, 1901; Gehlbach, 1967); the number of 
ventral scales was nearly 20% greater than reported for thayeri, and 
the iris was silver-grey, like specimens from Texas and unlike the 
gold-brown iris of animals collected in the mountains of Nuevo Leon. 
Living specimens collected in the mountains of Durango, Mexico, 
were similar in these three characters with those from Nuevo Leon, 
and were likewise similar to specimens from Queretaro, except in 
ventral scale number. 

The specimens recently collected in Queretaro, Mexico, posed 
another problem, as they closely resembled the single previous spec- 
imen identified as L. ruthveni (Blanchard, 1921), which had been 
synonymized (Williams, 1978) with L.t. arcifera. These specimens 
share many of the same features of L. mexicana, and indeed are 
distinguishable only on ventral scale number from some L. mexicana 
from Nuevo Leon. They are clearly not L. triangulum nelsoni nor L.t. 
arcifera, with which they are sympatric; both subspecies of L. triangu- 
lum have been collected (Williams, 1978) in the valleys that surround 
the montane area where the ruthveni were collected. Therefore an 
initial hypothesis of three valid taxa, altema, mexicana, and ruth- 
veni, was considered, and characters were analyzed within that 
framework. 

The most striking feature of kingsnakes is their often brilliant color 
and pattern. Color and pattern have been relied upon exclusively in 
diagnoses of the species of Lampropeltis (Blanchard, 1920, 1921; 
Gehlbach, 1967; Smith, 1942; Webb, 1961) and the subspecies of L. 
triangulum (Williams, 1978). The extreme variability of color and 
pattern among individuals in the mexicana group and the similarity 
of the patterns of ruthveni and some mexicana with some subspecies 
of triangulum makes using only this character untenable. 

Color and pattern variability is also well documented in Texas 
altema (Gehlbach and Baker, 1962; Gehlbach and McCoy, 1965; 
Gehlbach, 1967; Tanzer, 1970; Miller, 1979). There is a gradation of 



4 BREVIORA No. 466 

patterns from extremes of more than 1 7 narrow rings with alternating 
dots (alterna) to a low number (9-17) of major bands with no alter- 
nating reduced markings (b/airi). Tanzer (1970) reported a brood 
from an alterna female which contained both blairi and alterna 
offspring. Melanism is also known in both alterna (Miller, 1979) and 
mexicana. 

The morphological characters examined in this study are: 1) lepi- 
dosis, 2) color and pattern, including color of the iris, 3) osteology of 
the skull and vertebrae, and 4) hemipenes. The variation in these 
characters is analyzed in mexicana, ruthveni, and alterna, and each 
taxon is compared with its respective sympatric triangulum popula- 
tion. The three taxa are redefined, and a hypothesis of relationship 
and of selection pressures leading to evolution within the group is 
presented. 

MATERIALS AND METHODS 

Specimens 

Both preserved and living specimens including all age classes were 
examined. Embryo and hatchling specimens were examined and used 
in ontogenetic comparisons with adults. Living snakes were sexed by 
the method of Laszlo (1975) using probes manufactured by J & M 
Specialty Parts. All living specimens in the author's collection will be 
deposited in the Museum of Comparative Zoology. Wild caught 
specimens only were used in the character analysis as captive bred 
animals cannot be considered as statistically independent samples. 

External Morphology 

Scale counts recorded included ventrals (method of Dowling, 
1951), subcaudals, infralabials, supralabials, temporals, and dorsal 
scale rows at mid-body (V method of Peters, 1964). The location of 
the umbilical scar on hatchling specimens was noted. Dissected and 
everted hemipenes of each taxon were examined. Statistical analysis 
of data used methods from Sokal and Rohlf ( 1969). All sample means 
indicated in the text are followed by ± one standard error of the 
mean. 

X-Rays 

Radiographs of preserved specimens were taken on an F.E. Faxi- 
tron X-ray machine using Kodak Industrex R film. Measurements of 



1982 LAMPROPELTIS MEXICAN 'A SPECIES GROUP 5 

five thoracic vertebrae posterior to #20 were taken from each speci- 
men radiographed. Coiling of preserved specimens prevented exami- 
nation of the same five vertebrae on every animal. Measurements and 
terminology of the vertebrae are from Johnson ( 1955a, 1955b) and 
Auffenberg( 1963). Comparisons were made between replicate radio- 
graphs of the same vertebrae, and between measurements from a 
prepared skeleton and radiographs of the same vertebrae to be confi- 
dent of accuracy and precision. Differences were less than 3%. Dor- 
soventral radiographs of heads and measurements of the length and 
width of the skull at various levels were also taken. Individual osteo- 
logical elements of the skull were not examined. Comparisons were 
again made of three sets of duplicate radiographs, and the differences 
were less than 2%. 

Field Work 

During 1972, 1973, and 1975 trips were made to the Chihuahuan 
Desert areas of Texas, principally to Brewster, Presidio, and Val 
Verde counties. During 1974, 1975, 1976, 1979, and 1980 trips were 
made to various localities in Mexico. Collection of living specimens 
provided information on the ecology and extent of distribution of the 
group. 

ANALYSIS OF CHARACTERS 

Lepidosis 

There is a marked difference in the numbers of ventral scales of the 
three taxa (see Table 1 and Fig. 1). A one-way ANOVA shows that 
there is significant heterogeneity of mean ventral scale number 
among taxa (F = 85.3 at 2/ 83 df, p < 0.005). In a Student-Newman- 
Keuls test of differences between ranked pairs of means, all differ- 
ences are significant at the 19r level. In order to be assured that 
distinct populations were sampled, the distributions were tested for 
normality. The individual taxon distributions are not different from 
normal by the Kolmogorov-Smirnov cumulative test (D [a/terna] — 
0.09, critical value = 0.29, a = 0.01; D [mexicana] = 0.07. critical 
value = 0.30, a = 0.01 ; D [ruthveni] = 0. 1 3, critical value = 0.30. a = 
0.01). The total distribution was tested for normality and was found 
to be significantly different from normal (D = 0.27, critical value = 
0.18). 



BREVIORA 



No. 466 



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1982 LAMPROPELT1S WEXICANA SPECIES GROUP 

The variation among the ventral scale means of alterna, mexicana, 
and ruthveni appears as a north to south cline. This variation, how- 
ever, is not clinal within a taxon, and the differences among taxa are 
stable in captive-bred animals. Clinal variation in ventral scale 
number has been documented for Coluber constrictor (Auffenberg, 
1955) and Lampropeltis triangulum (Williams, 1978). In both taxa 
the change is gradual, with counts high in the south, low in the north. 
The effects of differing temperatures of incubation on scale counts 
(Fox et ai, 1961) could explain that situation, although no experi- 
mental evidence was given for either species. 

The geographic distribution of ventral scale numbers within each 
taxon appears random (Table 2). The southernmost specimens of 
alterna from just north of Saltillo, Coahuila (USNM 1 10819) and 
from near San Antonio, Durango (TCWC 36892), have 223 and 218 
ventrals, respectively. The northernmost mexicana specimens, three 
from the Rio Mimbres area of Durango (UCM 21061 and 2 Garstka 
collection) and one from the Anticline of Arteaga near Los Lirios, 
Coahuila (ITESM uncataloged), have 203, 202, 204. and 190. respec- 
tively. The Los Lirios mexicana has the lowest ventral number of any 
mexicana examined. 

The hypothesis that the north to south cline in ventral scale counts 
may be a result of differing incubation temperatures was examined by 
incubating two clutches of mexicana eggs and one clutch of alterna 
eggs in the same container at the same time (alterna: 3 August-23 
October; mexicana: 14 July-25 September; 28 June- 18 September 
1977). The same means of ventral counts are alterna = 218.5 ± 2.50 
(N=2); mexicana = 204.6 ± 1 .45 (N=10), and the difference is signifi- 
cant (t = 8.47, p< 0.001). In an additional sample of captive-bred 
alterna (218.8 ± 0.38, N=40) and captive-bred mexicana (201.6 ± 
7.87, N=7), the difference is also significant (t = 1 8.28, p< 0.00 1 ). The 
conditions and dates of incubation for the latter sample are not 
known. 

Juvenile specimens usually possess an umbilical scar. The number 
of ventral scales anterior to the scar is 1 70.9 ± 1 .48 for 1 1 mexicana 
and 190.0 ± 1.69 for 12 alterna (t= 12.98, p< 0.001). The length of 
the scar is the same in both taxa; the mean lengths in the samples are 
4.5 ± 0.25 scales for mexicana and 5.0 ± 0.37 scales for alterna (t = 
1.25, 0.3 > p > 0.2). No sex differences were noted. The single 
juvenile ruthveni (Kl) 155528) had no visible scar. 



BREVIORA 



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10 BREVIORA No. 466 

The pattern of ventral scale counts in these taxa is clearly differ- 
ent from that of each sympatric triangulum variety (Williams, 1978). 
L. triangulum celaenops and L.t. anmtlata are sympatric with 
alterna, L.t. nelsoni and L.t. arcifera with ruthveni, and there is 
apparently no triangulum sympatric with the montane mexicana. 
The ventral scale counts of Williams' ( 1978) and Quinn's (in prepa- 
ration) samples of celaenops, annulata, arcifera, nelsoni. and a new 
subspecies can serve to distinguish each species of the mexicana 
group from its sympatric L. triangulum subspecies (Table 1 . ) 

There is no clear separation of the taxa in other scale characters. 
The uppermost preoculars in most mexicana and ruthveni extend 
onto the top of the head to or nearly to the frontal, whereas in 
alterna they rarely do so. There is much variability and overlap 
between taxa in the number and arrangement of the temporal scales. 
There is also no differences among taxa in numbers of labial scales, 
subcaudals, or dorsal scale rows, and all specimens examined pos- 
sess a loreal (Table 3). 

Color Pattern 

The variable head and snout markings of this group can be 
grouped into four categories: I) asymmetric spot or blotch markings 
without red, II) snout evenly black to the middle or posterior of the 
parietals, with or without a red infusion, III) a red-centered, three- 
lobed cloverlike marking with the anterior lobe across the frontal or 
usually the prefrontals, and IV) a similar three-lobed marking, 
without red, that can have the anterior lobe either split to form a 
double Y, detached, or enlarged and subrectangular (Fig. 2). None 
of these categories is exclusive to any taxon, but most ruthveni are 
category II, and most alterna are category I. The head markings 
connect to the nuchal blotch only in some mexicana. The nuchal 
blotch of mexicana and alterna is usually light-centered and can be 
split longitudinally. Postocular stripes are usually present in all taxa 
(absent in UCM 21061, a mexicana), but may be reduced or obs- 
cured, as occurs in category II. 

All living specimens examined of alterna have a distinctive silver- 
grey iris; mexicana and ruthveni have a golden-brown iris. Other 
Lampropeltis (getulus, triangulum, pyromelana, and zonata) exam- 
ined also have a golden-brown iris. 



1982 



lAMPROPEl TIS MEXICANA SPECIES (iROlT 



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1982 LAM PROPELTIS MEXICAN A SPECIES GROUP 13 

The number of primary, usually red-centered, body markings var- 
ies considerably and is not by itself indicative of any taxon (Table 
1). The blairi form of alterna has the fewest primary markings 
(9-17). 

The 47 offspring produced in ten broods by one male and two 
female mexicana collected from a single site near San Juanito, 
Nuevo Leon, are variously ringed, blotched, or spotted. All possible 
variants of the dorsal head pattern occur in these offspring, and 
ground color ranges from lemon-yellow through various shades of 
brown, orange, and grey. The blotches vary in number from fewer 
than 20 to more than 40, and in shape from spots to rings extending 
around the body. The number of body bands falls within the ranges 
of all the forms except blairi. In some, alternating reduced markings 
occur in one or several rows. One specimen has red only in the 
nuchal blotch. Thus, neither number, level of truncation (Webb, 
1961) or alternating reduction (Blanchard, 1920, 1921; Smith, 1942) 
of body bands are diagnostic characters. The local variability 
characteristic of mexicana and alterna is not present in any 
triangulum subspecies, nor are alternating reduced markings 
recorded for any L. triangulum. 

Ventral pigmentation ranges from nearly entirely black to nearly 
entirely background color in alterna and mexicana. The ruthveni 
examined most commonly have rings which extend around the 
body; this also occurs with the anterior rings in some mexicana. The 
venter of alterna and mexicana can also be checkered in black and 
background color as in L.t. triangulum and Elaphe guttata (see 
figure in Smith, 1942, pp. 202 203), and some ruthveni have a 
checkered red pattern ventrally. 

A ventrolateral expansion of the first (anteriormost) tail blotch 
with red extending onto (usually across) the subcaudals is character- 
istic of mexicana. This character is not evident in any of the alterna 
examined. The underside of the tail of one specimen of mexicana 
(BMNH 1946-1-4-10) has red infused anteriorly in a longitudinal 
black line. The tails of the ruthveni examined have a ringed condi- 
tion similar to mexicana; however, the first tail ring is not usually 
widened ventrolaterally. The anterior tail rings are not different 
from body rings in any of the L. triangulum subspecies. 

The L. triangulum subspecies sympatric with each taxon can be 
distinguished by a combination of color patterns and scale charac- 
ters from the sympatric member of the mexicana group (Table 1). 



14 



BREVIORA 



No. 466 



The snout color and symmetric body annuli of ruthveni make these 
animals easily distinguishable from the sympatric nelsoni. Similarly, 
the high number of annuli of celaenops and annulata are different 
from the usually few annuli of the ringed or saddled blairi form of 
alterna. In the other cases, some combination of color pattern with 
ventral scale number serves to diagnose members of the mexicana 
group. 

Skull 

Data from dorsoventral radiographs of similarly sized, adult 
alterna (N = 7) and mexicana (N = 12) were compared to quantify 
the observation of a more triangular head shape in alterna. The 
premaxilla to occipital condyle length, the width of the maxilla at 



L. alterna H 
L. mexicana I I 




.72 



.76 



80 



.84 



PRE- -i- POST-ORBITAL WIDTH 



Figure 3. The ratio of pre-orbital and post-orbital width of L. alterna and L. 
mexicana adults. 



1982 LA MPROPELT1S MEXICAN A SPECIES GROUP 15 

the prefrontals, the least width of the frontals, the maximum widths 
across the post-orbitals, across the supratemporals, and across the 
quadrates were measured. The ratio of preorbital to postorbital 
width was used to quantify triangularity of the skull, as that should 
be least influenced by kinesis. The mean ratio of the alterna sampled 
is 0.66 ± 0.007 and of the mexicana sampled is 0.77 ± 0.010. They 
are significantly different (t = 7.343, p < 0.001) (Fig. 3). The single 
ruthveni adult examined for this character (USNM 46558) has a 
ratio of 0.74. 

Additionally, ten juveniles each of alterna and mexicana were 
compared for this character. The mean ratio of the sample of mexi- 
cana juveniles is 0.74 ± 0.007, with one specimen outside the range 
of variation of the adults, in the direction of alterna. In the sample 
of juvenile alterna, five are outside the variational range of adult 
alterna in the direction of mexicana, and the mean is 0.69 ± 0.008. 
Thus the two taxa are more similar as juveniles, and, importantly, 
are more like mexicana than alterna as juveniles. This indicates that 
there is differential growth to the adult stage in alterna. 

Adults of other species of Lampropeltis were also examined in 
outgroup comparison for this character. L. triangulum amaura and 
L.t. polyzona both had a preorbital to postorbital ratio of 0.77, a 
single L. pyromelana had a ratio of 0.73, and a single L. calligaster 
had a ratio of 0.73. Thus, it appears that the lesser triangularity of 
the skull in the mexicana condition is not only the common condi- 
tion for juveniles, but also for other Lampropeltis. 

Vertebrae 

Eight or more individual thoracic vertebrae (Johnson, 1955a, 
1955b; = first precaudal series, Auffenberg, 1963; = lumbar series, 
Bullock and Tanner, 1966) prepared from each of eight adult speci- 
mens of alterna and four adult mexicana were examined. X-rays in 
lateral and dorsoventral aspect of an additional three adult and five 
juvenile alterna and six adult and nine juvenile mexicana were also 
examined. No ruthveni vertebrae were available. None of the com- 
monly used measures (Johnson, 1955a, 1955b) or ratios (Auffen- 
berg, 1963) showed any significant difference between adults of the 
two taxa. The amount of individual variation in qualitative appear- 
ance of the vertebrae is substantial, however, even in the small 
sample examined; because of the lack of descriptions of the range of 



16 



BREVIORA 



No. 466 




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1982 LA MPROPEL T/S M EXICANA SPECIES GROUP 17 

individual variation of colubrid vertebrae, and the importance of 
vertebral characters in the systematics of fossil forms, an analysis of 
the vertebrae follows (Fig. 4). 

The neural spine may be blunt or pointed, and in all individuals of 
mexicana examined it overhangs both anteriorly and posteriorly. In 
lateral X-ray the neural spine of a single alterna, USNM 110819, 
appears to not overhang anteriorly. The zygosphene is crenate to 
convex from above and usually has a broad shallow notch. In ante- 
rior aspect the dorsum of the neural arch is flattened or slightly 
arched and is thinned centrally; the sides are obtusely angled, form- 
ing a dorsolateral articulating surface. The dorsal articulating surfa- 
ces of the anterior zygapophyses are ovoid to reniform and their 
lateral accessory processes can be blunted or pointed, directed later- 
ally or upward, and if upward, may be curved. The processes may be 
rounded to dorsoventrally compressed, and if compressed, may be 
tilted to be parallel to the centrum. The cotyle may be round to 
ovoid and if ovoid, may have accessory flanges or fillets, usually 
ventrally, that make it appear rectangular. The parapophyses may 
or may not project ventral to the cotyle and may be projected ante- 
riorly. The paradiapophysial articulating surfaces are in the form of 
a figure-8 with the upper portion larger and directed anteriorly, and 
the lower surface directed laterally to posteriorly. There is a small 
foramen on the parapophyses lateral to the cotyle on each side 
approximately at its midline. The haemal keel is usually ridgelike, 
though it may be flattened or absent. The subcentrum ridges are 
straight to slightly concave. There are two pairs of subcentrum 
foramina, one pair on either side of the haemal keel and the other 
more lateral and dorsal, just ventral to the lateral flange of the body. 
The condyle is oblique. In posterior view, the neural arch is rounded 
dorsally and flattened ventrally with a central ventral ridge. There 
a*e also lateral minor ridges extending the length of the canal on 
either side approximately one third of the distance superior to the 
base. The ventral exterior articulating surfaces of the posterior 
zygapophyses are ovoid to reniform with the long axis perpendicu- 
lar to the centrum. The medial articulations that match the dorsal 
zygosphene corners, the zygantra, appear as enlarged triangular 
holes lateral to the neural arch. There are small projections poste- 
rior and lateral to the posterior zygapophyses just dorsal to the 
ventral articulating surfaces. 




18 BREVIORA No. 466 

Juvenile vertebrae present quite a different appearance from 
those of adults, which is most noticeable in the proportionally larger 
neural canal. The ratio of length of the centrum to width of the 
neural canal in a sample of three adult mexicana is 2.1 ± 0.10, of 
three adult alterna is 2.3 ± 0.08, and of eight juvenile mexicana is 1.2 
± 0.03. The neural spine is also proportionally higher. The ratio of 
length to height of the neural spine of a sample of three mexicana 
adults is 2.4 ± 0.34 and of three juvenile mexicana is 1 .5 ± 0.09. The 
juvenile alterna sampled had vertebrae insufficiently ossified to 
obtain the measurements for comparison. Since nervous tissues 
have grown most in relation to other tissues at birth, and since 
ossification is incomplete then, these differences are to be expected. 

The thoracic vertebrae of L. triangulum differ from alterna and 
mexicana in the form of the zygosphene which is arched rather than 
flattened (Brattstrom, 1955; Auffenberg, 1963). All of the tricolored 
kingsnakes examined (L. triangulum, L. pyromelana, L. zonata, L. 
alterna, and L. mexicana) share the single foramina lateral to the 
cotyle and the two pairs of subcentrum foramina. L. getulus and L. 
calligaster, however, have two pair of foramina adjacent to the 
haemal keel and a pair of foramina on each side of the cotyle. 

Hemipenes 

Previous workers (Smith, 1942; Gehlbach and Baker, 1962) have 
reported on the hemipenes of alterna and mexicana, and allied the 
taxa with L. triangulum on the basis of overall similarity of gross 
structure. The hemipenes of three adult mexicana, five adult alterna, 
and two subadult ruthveni have been examined. They are uniformly 
longitudinally flounced proximally, then abruptly spinulate, quickly 
grading into slightly spinulate fringed calyces. The sulcus is single 
and terminates laterally on the very slightly bilobed organ. This 
morphology is similar to that of L. triangulum. 

The spination on the hemipenes is clearly different among the 
mexicana group, however (Fig. 5). The spines of mexicana are very 
slightly recurved and are about 0.4 mm long. They appear diamond- 
shaped in cross section and are tightly packed on the organ. The 
spines of alterna and ruthveni are also recurved but are longer, to 
approximately 0.8 mm in alterna and 0.6 mm in ruthveni, and the 
supporting tissue makes them appear more ovoid or flattened in 
cross section. The spines also appear to be fewer in number in these 



1982 LAMPROPELTIS MEXICASA SPECIES GROUP 



19 




L alterna 






L. mexicana 




L. ruthveni 




Figure 5. Hemipenial spines of L. alterna, L. mexicana, and L. ruthveni. Lateral 
(L) and distal ( R) aspects. 



taxa than in mexicana. The spines of both ruthveni examined are 
more strongly recurved than those of both mexicana and alterna. 



Ecology 

The distribution of the entire mexicana group has been character- 
ized as xeric to subhumid (Gehlbach and Baker, 1962) and roughly 
corresponds to the limits of the Chihuahuan Desert as described by 
Morafka( 1977) (Fig. 6). 



20 BREVIORA No. 466 

L. alterna is found within the Chihuahuan Desert, usually in 
association with typically desert floral elements such as Prosopis 
and Larrea (Jameson and Flury,1949; Mechamand Milstead, 1949; 
Worthington, 1974). All known localities are within or north of the 
Bolson de Mapimi. Specimens collected in the Chisos Mountains, 
Brewster County, Texas (Murray, 1939) and near Saltillo, Coahuila, 
Mexico (Smith, 1941) were found in rock crevices, though the spe- 
cies appears to be primarily nocturnal (Miller, 1979). 

L. mexicana seems to be limited, except for CM 59980, to for- 
ested mesic areas peripheral to the desert, but with xerophytic vege- 
tation such as Echinocactus and Ariocarpus. At mexicana localities 
in Nuevo Leon, San Luis Potosi, and Durango, Mexico, the vegeta- 
tion may be termed "mesic to arid oak-pine forest" (J. Henrickson, 
personal communication), with the following dominant plants in 
common at the three localities: Pinus cembroides, Quercus sp., 
Juniperus sp., Buddie ja cor data, and Bouvardia ternifolia. The 
author has collected specimens in rock crevices, under stones in an 
arroyo, and active on the road at night. 

L. ruthveni is presently known from the northeastern portion of 
the trans-volcanic region in a habitat very similar to mexicana. In 
the vicinity of Amealco, Queretaro, Mexico, ruthveni are locally 
abundant in areas with scattered patches of (^wmm-dominated 
woodland along arroyos and on hilltops. Bouvardia is common here 
as well. All recent collections have been of animals either in crevices 
or diurnally active. 

Both mexicana and ruthveni appear to be more mesic species than 
alterna, occurring in more temperate forested areas. However, CM 
59980 was collected in a desert area with stunted Larrea, cacti, and 
short grasses (D. Morafka, personal communication). The elevation 
of this rocky area is over 2,000 m, and while it is surrounded by a 
sandy plain, it is also closely adjacent to mesic upland areas. Many 
similar hilly areas in the vicinity have been deforested in recent times 
(T. Wendt, personal communication) and the area may have been 
woodland as recently as 100 years ago. L. alterna may also occupy 
upland (Worthington, 1974) or riparian habitats within the desert; 
however, it appears to be primarily a desert species. The higher 
ventral scale numbers of alterna may indicate adaptation to a desert 
environment, as Klauber (1941) found a correlation of higher ven- 
tral counts in desert habitats and lower counts in more humid coas- 
tal habitats in 1 2 of 1 3 taxa tested. 



1982 LAMPROPELTIS MEXICANA SPECIES GROUP 21 

DISCUSSION 
Phylogeny 

Underwood (1967) considered Lampropeltis as derived from 
Elaphe on the basis of similarity of a long intrapulmonary bron- 
chus, a series of skull characters, presence of similar scale tubercles, 
similar arrangement of pits on the head, and similar paired pits on 
trunk scales. The characters that define Lampropeltis as a group 
distinct from Elaphe are the entire anal scale and the unkeeled 
dorsal scales. Minton (1976) and Minton and Salanitro (1972) dis- 
cussed the phylogeny of colubrids, based on the immunoelectropho- 
retic analysis of serum proteins and found that both E. guttata and 
L. getulus have the same degree of difference from E. obsoleta. 
Karyological evidence also demonstrates this similarity to Elaphe. A 
karyotype of 2N = 36 (16 macrochromosomes and 20 micro- 
chromosomes) has been reported for L. getulus, ealligaster, and 
alterna, as well as for most Elaphe (E. suhocularis is an exception) 
(Bury, et al. 1970; Baker, et al. 1972). This close relationship 
between Lampropeltis and Elaphe is also substantiated by vertebral 
morphology. The form of the neural spine and the arrangement of 
the vertebral foramina are similar in L. getulus, L. ealligaster, and 
Elaphe. 

All of the tricolored kingsnakes (alterna, mexicana, pyromelana, 
ruthveni, triangulum, zonata) share grossly similar derived hemipe- 
nial (Smith, 1942) and vertebral character states that are unlike 
those of the remaining species in the genus, L. ealligaster and L. 
getulus, which are in turn similar to each other and to Elaphe (Cope, 
1898). 

Captive-breeding experiments (Miller, 1979; Tryon and Garstka. 
in preparation) have produced alterna X me.xieana, pyromelana X 
alterna, and pyromelana X zonata hybrids. None of these species 
will mate with any triangulum subspecies tested in captivity. This 
can be taken as tentative evidence of their relationships. Therefore, 
it appears that all the tricolored kingsnakes share a common ances- 
tor, and that the me.xieana group forms a logical unit within that 
lineage, perhaps most closely related to pyromelana and zonata. 
Within the me.xieana group, alterna is the most derived species, as 
shown by its iris color and head shape. These characters suggest that 
alterna is the most derived species within the genus as well ( Fig. 6). 



22 



BREVIORA 



No. 466 



mexicana 



ruthveni 



zonata 
pyromelana 



getulus 
calligaster 




triangulum 



ELAPHE LAMPROPELTIS 




Figure 6. Hypothesis of relationship within the genus Lampropeltis. Derived 
character states uniting the taxa are: a) paired pits on trunk scales, long intrapulmo- 
nary bronchus; b) entire anal scale, unkeeled dorsal scales; c) single paired foramina 
lateral to hemal keel and cotyle, hemipenes flounced proximally and spinulate dis- 
tally, pattern in three colors including red; d) behavioral traits in male aggression and 
reproduction; e) light-edged, red-centered blotch; f) alternating reduced markings 
possible; and g) grey iris, "triangular" skull. 



1982 LA M PROPEI.T1S M EXICANA SPECIES GROUP 23 

The data presented here suggest that the three taxa hypothesized, 
L. mexicana, L. alterna, and L. ruthveni, are distinct morphological 
species. However, the taxa are allopatric, as are the two apparently 
most closely related species, L. pyromelana and L. zonata. There- 
fore the allocation of species status must still be regarded as an 
hypothesis until appropriate species borders or contact zones are 
established. 

The species borders of mexicana and alterna, that is, the Anticline 
of Arteaga and the uplands of central Durango at the periphery of 
the desert, should be further collected. The presence of alterna at the 
base (USNM 110819) and mexicana (1TESM uncataloged) at the 
top of the Anticline of Arteaga indicate a high probability of a 
contact zone. Southern San Luis Potosi, Guanajuato, and northern 
Michoacan including the Sierra de Santa Rosa (Duges, 1897) and 
the escarpment of the basin of the Rio Lerma should be explored as 
well for a species boundary or area of intergradation as this is 
between the localities of mexicana and ruthveni. Finally, the moun- 
tainous area of northern Durango north of Otinapa and west of 
Santiago Papasquiaro and the upper reaches of the Rios San Juan 
and Verde need investigation. There are no kingsnakes known from 
that large area between the localities of mexicana and pyromelana 
(Taylor, 1940). 

In addition, small sample sizes of some of the Mexican Plateau 
subspecies of triangulum make it difficult to determine the extent of 
sympatry, especially with ruthveni, and make comparisons difficult. 
More collections from that area, and revision of the genus or at least 
of the tricolored group, are necessary before the relationships 
among the various taxa can be better understood. 

Variation and Selection of Color and Pattern 

Many explanations of the significance of color and pattern in 
snakes have been proposed (see review by Jackson et ai, 1976). 
Aposematism, to warn potential predators, has been proposed as an 
explanation of bright colors on venomous snakes (Goodman and 
Goodman, 1976; Gehlbach, 1972; Smith, 1969). Mimicry of coral 
snakes by nonvenomous forms has been discussed extensively 
(Smith, 1969; Gehlbach, 1972; Greene and Pyburn, 1973; Echter- 
nacht, 1973; Hecht and Marien, 1956) as an explanation for the 
brightly colored rings on many snakes. Crypsis on broken back- 



24 



BRFVIORA 



No. 466 




108 106 104 102 100 

Figure 7. Distribution of the species in the mexicana group. State boundaries are 
shown as a dotted line, national borders as a broken line, and the extent of the 
Chihuahuan Desert (after Morafka, 1977) as a solid line. The Anticline of Arteaga is 
the band of mountains extending into the desert across southern Coahuila at about 
latitude 25°N. North of that is the Mapimian portion of the desert, and south of the 
Anticline is the Saladan portion of the desert (Morafka, 1977). Symbols denote only 
localities and not numbers of specimens. 



1982 LAMPROPELTIS MEXICANA SPECIES (.ROUP 25 

grounds (Pough, 1976; Brattstrom, 1955), or lack of crypsis on a 
unicolored background (Camin and Erhlich, 1958) have also been 
cited to explain the presence or absence of a ringed or blotched 
pattern. 

Gehlbach (1972) reported experimental evidence that self-mimicry, 
that is, the coral snake behavior of flipping the tail over, thwarted 
predation by coatimundis (Nasua) and peccaries {Tayassu). This 
effect was enhanced if the rubber models were painted aposemati- 
cally. Evidence of the effectiveness of coral snake mimicry has been 
provided by experiments offering striped and ringed stick models to 
naive captive-raised motmots (Eumomota superciliosa) (Smith, 
1975). The birds responded to red and yellow ringed models by 
avoidance and, in some cases, alarm notes. 

L. mexicana and L. alterna, even though they may be brightly 
colored, do not look very much like coral snakes. Gans (1961) dis- 
cussed an hypothesis of mimicry in Dasypeltis, a small nonveno- 
mous snake that eats eggs of birds large enough to eat it. He 
demonstrated a correspondence of pattern between Dasypeltis and 
the local possible viperid models over its entire range. The pattern of 
the particular similar-sized viper in its range was mimicked. There- 
fore, the distribution of venomous snakes of the same general size 
and habits as Lampropeltis, and occurring within the range of the 
mexicana group, was examined. Crotaline models were considered, 
as the fine dark speckling on the scales lends an illusion of texture to 
the smooth-scaled Lampropeltis. 

Agkistrodon mokeson pictigaster occurs in Texas west through 
the Big Bend region, but west of Val Verde County occurs only in 
the vicinity of permanent water sources. Crotalus lepidus is found 
throughout the range of alterna and is present in the same micro- 
habitat. Crotalus lepidus, with its alternating broken pattern, can 
easily be mistaken for alterna. A similar pattern is also seen on 
Trimorphodon from west Texas. Micrurus fulvius is found in Texas 
west to Val Verde County. 

The variation in Texas alterna could be a function of multiple 
models, the blairi form of alterna being a mimic of Agkistrodon. 
The relative constancy of pattern in alterna outside of the range of 
Agkistrodon could be due to having a single available model in 
those areas, C. lepidus. In recent years Val Verde County, Texas, 
has had increased rainfall, and the blairi form, first collected in 1948 



26 



BREVIORA 



No. 466 









Figure 8. Hypothesis of mimicry in the mexicana group, a) Crotalus lepidus (L) 
with sympatric L. alterna (R); b) Agkistrodon mokeson (L) with sympatric blairi 
morph of L. alterna (R); and c) Crotalus triseriatus (L) with sympatric L. mexicana 

(R). 



1982 LAMPROPEETIS MEXICANA SPECIES GROUP 27 

(Axtell, 1951), is today quite commonly found. Agkistrodon are 
common along the Devil's and Pecos river drainages as well, in 
similar habitat with blairi; Micrurus fulvius is found there as well. 
The sporadic appearance of blairi-Mke animals in Presidio and 
Brewster counties in the Big Bend region is correlated with the 
distribution of Agkistrodon. Both A.m. pictigaster and M. fulvius 
are very conspicuous, brightly colored animals, and though C. lepi- 
dus is also sympatric in Val Verde County, its dull appearance may 
make it a less efficient model. The increased red in the blairi morph 
may be a result of stimulus generalization by predators. 

In San Luis Potosi Crotalus triseriatus aquilis shares the range 
and habitat of mexicana. The pattern match of these animals is 
quite remarkable. In life, C.t. aquilis is gray-brown with black speck- 
ling and a mid-dorsal row of 28 to 40 (mean = 34.5, Klauber, 1952) 
almost rectangular red-brown irregular blotches. Ventrolateral black 
blotches are also present. The W.W. Brown 1923 collection at 
Alvarez for the MCZ contained \0 mexicana and 20 C.t. aquilis. 

The color variability of mexicana in the Nuevo Leon population 
sampled could reflect the local variability of the model, Crotalus 
iepidus, or a multiple model system. In that area C. lepidus varies 
from ochre to blue in background color, with deep reddish-orange 
on the ventral side of the tail, which is also apparent in C.t. aquilis. 
Some mexicana from Nuevo Leon are clearly ringed, like Micrurus. 
Micrurus fulvius is present on the Atlantic versant of the Sierra 
Madre. The mobility of predatory birds, as well as the relative rarity 
of the crotaline model, could affect the occurrence of mimicry in 
mexicana. 

There are two situations in which the mimicry hypothesis is less 
certain. The population of mexicana in Durango, Mexico, is rather 
uniform in appearance and not clearly like any other snake found 
there. Possible models which share the same canyons with mexicana 
are C lepidus, C pricei, and C willardi. The mexicana are most like 
male C. lepidus in that both are yellow-green to gray-green in back- 
ground color with contrasting narrow dark crossbands. The mexi- 
cana, however, have many more bands and the bands are usually 
red-centered. The second situation concerns ruthveni. On the same 
hills in southwestern Queretaro where ruthveni is locally abundant, 
C triseriatus is also abundant, yet ruthveni is a brightly ringed 
animal. Micrurus fit zingeri is sympatric with ruthveni over much of 



28 breviora No. 466 

its range, but appears not to be present at those localities. M. fitzin- 
geri is ringed similarly to ruthveni, with up to 28 triads of more or 
less equal annuli (Smith and Taylor, 1966). Again, the conspicuous- 
ness of the red color may contribute to its stimulus value. 

An alternate prediction can be made (Levene, 1953) that the areas 
showing polymorphism are in some way spatially patchy. The habi- 
tat of mexicana in Nuevo Leon could be considered patchy in that 
there are, in close proximity, areas of pine forest, chaparral, open 
desert, and natural and man-made meadows. There is, however, no 
correlation of observed morph to microhabitat. There is some spa- 
tial heterogeneity in alterna habitat also, with alterna found in both 
riparian and desert situations. However, this does not correlate with 
the distribution of the blairi and alterna morphs. 

Parallel selection processes for crypsis in both the "model" and 
"mimic" could also explain the similarity of pattern in each situa- 
tion. The blairi case, however, with the change of color pattern of 
alterna to the more abundant and conspicuous model Agkistrodon, 
even in the presence of C. lepidus, argues strongly for something 
otherthan parallel selection. 

DESCRIPTIONS AND DIAGNOSES OF THE TAXA 

Lampropeltis alterna ( Brown) 
Ophibolus alternus Brown 1901 
Lampropeltis alterna, Stejnegerand Barbour 1917 
Lampropeltis blairi Flury 1 950 

Lampropeltis mexicana blairi, Gehlbachand Baker 1962 
Lampropeltis mexicana alterna, Gehlbachand Baker 1962 

Type specimen (ho\otyve): Acad. Nat. Sci. Phil. 14977 
Type locality: Davis Mountains, Jeff Davis County, Texas 
Description and Diagnosis: A moderately sized (to about 1 m) 
snake with a very distinct head and overall mottled grey color. The 
pattern is a series of white-edged black blotches or saddles that may 
be red-centered. Alternating reduced markings may be present 
between major markings. The iris of the relatively large eye is silver- 
grey in color. The number of ventral scales is 210-232. The proxi- 
mate spines on the hemipenes are ovoid in cross section and 
approximately 0.7 mm long. 



1982 LAMPROPELTIS MEX1CANA SPECIES GROUP 29 

Distribution: Specimens have been collected over the entire 
Mapimian portion of the Chihuahuan Desert (Morafka, 1977). 
These localities range from latitude 32° N (Worthington, 1974) 
south to latitude 25° N (Tanzer, 1 970). 

Lampropeltis mexicana (Garman) 

Ophibolus triangulus var. mexicanus Garman 1 884 

Oreophis boulengeri Duges 1897 

Coronella mexicana, Gunther 1900 

Coronella leonis Gunther 1 900 

Lampropeltis mexicana, Blanchard 1921 

Lampropeltis leonis, Blanchard 1921 

Lampropeltis leonis, Loveridge 1924 

Lampropeltis thayeri Loveridge 1924 

Lampropeltis greeri Webb 1961 

Lampropeltis mexicana greeri, Gehlbachand Baker 1962 

Lampropeltis mexicana thayeri, Gehlbachand Baker 1962 

Lampropeltis mexicana mexicana, Gehlbachand Baker 1962 

Type specimen (syntypes): MCZ 4652, 4653 
Type locality: near Ciudad San Luis Potosi, Mexico 
Description and Diagnosis: A moderately sized (to about 1 m) 
snake with a slightly distinct head and overall mottled grey to yellow 
or brown color. The pattern is a series of white-edged black 
blotches, saddles or rings that may be red-centered. Alternating 
reduced markings may be present, usually ventrolateral^, between 
major markings. The anterior tail blotch is enlarged ventrolateral^ 
and red extends onto or usually across the subcaudals. The iris of 
the relatively large eye is yellow-brown in color. The number of 
ventral scales falls between 190 and 212. The proximate spines of the 
hemipenes are rhomboidal in cross section and are approximately 
0.4 mm long. 

Distribution: Specimens have been collected mainly from the 
mountains surrounding the Saladan portion of the Chihuahuan 
Desert (Morafka, 1977). The localities have ranged from about lati- 
tude 25° N in the Sierra Madre Oriental (P. Bartlett, personal com- 
munication) south to 21° N (Duges 1897). A single desert locality 
(D. Morafka, personal communication) may be a result of recent 
man-altered changes in the habitat. 



30 



BRF.VIORA 



No. 466 



Lampropeltis ruthveni Blanchard 

Lampropeltis ruthveni Blanchard 1 92 1 

Lampropeltis triangulum arcifera, Williams 1978 (in part) 

Type specimen {\\o\oiypQ): USNM 46558 

Type locality: Potrenaro= Patzcuaro?, Michoacan, Mexico 

Description and Diagnosis: A moderately sized (to about 0.8 m) 
snake with a slightly distinct black head and a uniformly ringed 
pattern. The black rings are light-edged and red-centered and 
extend entirely around the body. The small amount of background 
color is a mottled tan to lime green. The iris is yellow-brown in 
color. The number of ventral scales falls betwen 182 and 196. 

L. ruthveni can be distinguished from the central Mexican milk 
snakes L.t. ne/soni and L.t. arcifera on several features of external 
morphology. First, the head of L. ruthveni is distinct from the neck, 
as in L. mexicana. Second, the ringed pattern of L. ruthveni is 
uniform; the red is not much broader nor is the background color 
much narrower than the black as in L.t. nelsoni. Third, there is no 
tendency in any of the L. ruthveni examined for the black to extend 
along the dorsal midline through the red, as is the case in both L.t. 
nelsoni and L.t. arc if era (Smith, 1942; Williams, 1978). Fourth, the 
black rings of L. ruthveni are bordered with a lighter color, usually a 
pale lime green; this is similar to L. mexicana and is distinct from L. 
triangulum. Finally, the ventral scales of the L. ruthveni examined 
were never greater than 196 (mean = 188), while the range of ven- 
tralsofL./. ne/soni is 203-231 and of L.t. arcifera is 197-217. 

Distribution: The type locality is cited (Blanchard, 1921) as Patz- 
cuaro, Michoacan, but this may be incorrect. The locality with the 
specimen is Potrenaro, Michoacan. The base camp of the collector, 
W.E. Nelson, in August 1892 was Patzcuaro (F. McCullough, per- 
sonal communication). Recent collections have been in the vicinity 
of La Piedad, Queretaro (C. Lieb, J. Dixon, E. Wagner, personal 
communication and personal observation), Morelia (UMSNH) and 
Contepec, Michoacan (D. Armstrong and J. Campbell, personal 
communication) and Tapalpa, Jalisco. All of the recent collections 
have been in rocky, wooded uplands. The range of this species may 
extend entirely across the Mexican Plateau. 



1982 LAMPROPELTIS MEXICANA SPECIES GROUP 31 

SPECIMENS EXAMINED 

Lampropeltis alterna. MEXICO: Coahuila: Cuatrocienegas 
(FMNH 47090); Cruz Verde Mt. c. Saltillo (USNM 1 10819); Puente 
de la Muralla c. Monclova (Garstka coll., 1 spec.). Durango: 26 mi. 
N San Juan del Rio (TCWC 36892). UNITED STATES, TEXAS: 
Brewster Co.: hills N of Study Butte (MCZ 157763). Jeff Davis Co.: 
17.7 mi. N Fort Davis (TCWC 26181). Presidio Co.: 25 mi. W 
Lajitas (UTA 7875). Val Verde Co.: W of Comstock (UTA 2633); 8 
mi. W of Comstock (UTA 2941); 8.5 mi. N of Comstock (UTA 
8690); 32 mi. NW of Comstock (TCWC 33759); 2 mi. W of Com- 
stock (TCWC 30515); N of Comstock (MCZ 157764, 157765; 
Garstka coll., 1 spec); 15 mi. NE Del Rio (TCWC 26179); 1.5 mi. E 
of Langtry (UTA 6680); 7.5 mi. E of Langtry (UTA 6681); N of 
Langtry ( UTA 7 1 88); 6 mi. N of Langtry ( UTA 867 1 ); Langtry Loop 
Road (MCZ 156175); 1 mi. E of Langtry (Garstka coll., 1 spec); 11 
mi. N of Loma Alta (UTA 8668); c. 10 mi. S of Loma Alta (UTA 
7874); 0.5 mi. E of Pecos River on US 90 (UTA 8095); Roadside rest 
c. Pecos River (UTA 8568); Pecos River overlook (TCWC 26180). 
NO SPECIFIC LOCALITY: (UTA 7969, 8179); (Wagner coll., no. 
19). CAPTIVE-HATCHED: Gravid female collected 8.7 mi. N of 
Comstock (TCWC 33761 33763). CAPTIVE-BRED: Male parent 
collected Brewster Co., Christmas Mts.; Female parent collected 
Val Verde Co., c Langtry (MCZ 156271-156273); Both parents 
collected Val Verde Co., c Langtry (MCZ 157724 157728; 156173, 
156174; Garstka coll., 10 spec); Both parents collected no specific 
locality (Texas) (MCZ 156259 156270, 157755 157758, 157760- 
157762, 158326 158335; Garstka coll., 1 spec, UTA 8126-8129. 
7873; Wagner coll., 1,5,7, 19,21 25, 34,44). 

Lampropeltis mexieana. MEXICO: Durango: 42 mi. S of Cd. 
Dgo. (UCM 21061); 23 mi. S of Cd. Dgo. (Wagner coll., Gl. G2); 
Canyon of the Rio Tunal (Garstka coll., 2 spec); Highway 40 at the 
Rio Chico (LACM 107230, 107231); Rancho Sta. Barbara (Hous- 
ton Zoo, 1 spec); no specific locality (San Diego Zoo, 9 spec). 
Nuevo Leon: c. Galeana, Linares-San Roberto highway (ITESM 
2507, 2508); Galeana (TCWC 56823); 5 mi. SE of Galeana (TU 
16483); c Los Lirios (ITESM uncataloged); Ojo de Agua c. Galeana 
(FMNH 30819 30821); c. La Angostura (Garstka coll., 3 spec); no 
specific locality (BMNH 146 1 4 10). San Luis Potosi: Alvarez 



32 BREVIORA No. 466 

(MCZ 19022 19025, 24976 24979; AMNH 77602; USNM 120823); 
c. Armadillo de los Infantes (Wagner coll., 1 spec.); c. Rioverde (KU 
85010); c. Cd. San Luis Potosi (MCZ 4652, 4653); 52 mi. WNW of 
Cd. San Luis Potosi (CM 59980). Tamaulipas: Miquihuana (MCZ 
19551). CAPTIVE-BRED: Both parents collected Mexico, Du- 
rango, no specific locality (MCZ 157754; Garstka coll., 1 spec; 
Wagner coll., G3, G5). Both parents collected Mexico, Nuevo Leon, 
c. La Angostura (MCZ 156274-156277, 157766; also Garstka coll.) 

Lampropeltis ruthveni. MEXICO: Michoacdn: Club Campestre 
at Morelia(UMSNH uncataloged); Potrenaro= Patzcuaro?(USNM 
46558). Queretaro: Canyon of the Rio Galindo c. Amealco (MCZ 
161010-161012; Mexican govt, coll., Agencia Forestal y de la Fauna 
Z-06586, Z-06587; Garstka coll., 5 spec; Wagner coll., 9 spec; 5 
spec, released at site of capture). Jalisco: Mts. W Zacoalco (KU 
155528); Tapalpa (SDMNH 46093); 6.5 mi. E Tapalpa (LACM 
37307). 

ACKNOWLEDGMENTS 

1 would most like to thank E. E. Williams for his patience and 
presence throughout this study. T. Fritts was instrumental in the 
early stages of the study, as were many discussions with D. F. Retes, 
E. Wagner, and others. G. C. Mayer criticized several drafts. D. 
Crews, J. Dixon, M. Scott, and two reviewers also commented on 
the manuscript. Access to specimens was provided by: J. Alvarado, 
Universidad Michoacana de San Nicolas Hidalgo, Morelia, Michoa- 
can, Mexico (UMSNH); P. Bartlett, Instituto Tecnologico y Escu- 
ela Superior de Monterrey, Monterrey, Nuevo Leon, Mexico 
(ITESM); R. Bezy & J. Wright, Los Angeles County Museum 
(LACM); J. Dixon, Texas Cooperative Wildlife Collection (TCWC); 
W. Duellman, University of Kansas (KU); H. Dundee, Tulane Uni- 
versity (TU); T. Fritts, San Diego Museum of Natural History 
(SDMNH); A. G. C. Grandison, British Museum (Natural History) 
(BMNH); W. R. Heyer, United States National Museum (USNM); 
D. Hoffmeister, University, of Illinois Museum of Natural History 
(UIMNH); P. Maslin, University of Colorado Museum (UCM); C. 
J. McCoy, Carnegie Museum (CM); W. Pyburn, University of 
Texas at Arlington (UTA); T. Van Devender, University of Arizona 
(UA); and R. Zweifel, American Museum of Natural History 
(AMNH). Access to specimens was also provided by numerous zoo 



1982 LAMPROPELTIS MEXICANA SPECIES GROUP 33 

and private collections, especially J. Bacon (Zool. Soc. San Diego), 
J. Murphy (Zool. Soc. Dallas), H. Quinn, and B. Tryon (Zool. Soc. 
Houston), E. Wagner (Zool. Soc. Seattle) and D. F. Retes. Botani- 
cal material was identified by J. Henrickson, T. Wendt, and M. 
Donaghue. 

Permission to collect in Mexico was issued by Lie. M. L. Cossio- 
Gabucio, Lie. 1. Ibarrola-Bejar, and Lie. M. Gonzalez-Escamilla, all 
of the Agencia Forestal y de la Fauna. 

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34 BREVIORA No. 466 

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1982 LAMPROPELTIS MEXICANA SPECIES GROUP 35 

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American and Eurasian colubrid snakes. Copeia, 1976(4): 672-678. 
Minton, S. A., and S. K.. Salanitro. 1972. Serological relationships among 

some colubrid snakes. Copeia, 1972 (2): 246-252. 
Morafka, D. 1977. A biogeographical analysis of the Chihuahuan Desert through 

its herpetofauna. Biogeographica vol. 9. The Hague, Junk, vi + 3 1 3 pp. 
Murray, L. J. 1939. Annotated list of amphibians and reptiles from the Chisos 

Mountains. Contrib. Baylor Mus.,24:4-16. 
Peters, J. 1964. Dictionary of Herpetology. N.Y., Hafner, vii + 427 pp. 
Pough, F. H. 1976. Multiple cryptic effects of crossbanded and ringed patterns of 

snakes. Copeia, 1976(4): 834-836. 
Smith, H.M. 1941. Lampropeltis alterna from Mexico. Copeia, 1941(2): 112. 
. 1942. Remarks on the Mexican kingsnakes of the triangulwn group. 

Proc. Roch. Acad. Sci., 8: 196-207. 
. 1944. Snakes of the Hoogstraal expeditions to Northern Mexico. Field 



Mus. Nat. Hist. Ser. Zool., 29: 135-152. 
Smith, H. M., and E. H. Taylor. 1966. The Herpetology of Mexico. Ashton, 

MD., Lund berg, 610 pp. 
Smith, N. G. 1969. Avian predation of coral snakes. Copeia, 1969(2): 402-404. 
Smith, S. M. 1975. Innate recognition of coral snake pattern by a possible avian 

predator. Science, 187(4178): 759-760. 
Sokal, R. R., and F. J. Rohlf. 1969. Biometry. San Francisco, W. H. Freeman, 

xxi + 776pp. 
Stejneger, L., andT. Barbour. 1917. A checklist of North American Amphibi- 
ans and Reptiles. Harvard, Cambridge, Ma. 
Tanner, W. W. 1953. A study of the taxonomy and phytogeny of Lampropeltis 

pyromelana (Cope). Great Basin Nat., 13( 1,2): 47-66. 
Tanzer, E. 1970. Polymorphism in the mexicana complex of kingsnakes with 

notes on their natural history. Herpetologica, 26(4): 4 19-428. 
Taylor, E. H. 1940. A new Lampropeltis from western Mexico. Copeia 1940(4): 

253-255. 
Underwood, G. 1967. A contribution to the classification of snakes. BMNH 

Pub. 654, x+ 179 pp. 
Webb, R. 1961. A new kingsnake from Mexico with remarks on the mexicana 

group of the genus Lampropeltis. Copeia, 1961(3): 326-333. 
Williams, K. 1978. Systematics and natural history of the American milk snake 

Lampropeltis triangulum. Pub. Biol. Geol. Milwaukee Pub. Mus. No. 2, 258 

pp. 
Worthington, R. D. 1974. Western records of the Davis Mountain kingsnake, 

L. m. alterna in Texas. SW Nat., 19(3): 330-33 1 . 






MAR 1 8 1985 

'ARVARD 
UMJVERSiTY 



BREVIORA 

Museum of Comparative Zoology 



US ISSN 0006-9698 



Cambridge, Mass. 30June1982 Number 467 

THREE NEW SPECIES OF THE ANOLIS PUNCTATUS 

COMPLEX FROM AMAZONIAN AND INTER-ANDEAN 

COLOMBIA, WITH COMMENTS ON THE EASTERN 

MEMBERS OF THE PUNCTATUS SPECIES GROUP 

Ernest E. Williams 1 

Abstract. New species of the Anolis punctatus group are described from central 
and eastern Colombia: A. vaupesianus from Comisaria of Vaupes, A. huilae from the 
Departments of Huila and Tolima, A. santamartae from the south slope of the Sierra 
Nevada de Santa Marta. The characters useful in discriminating the eastern members 
of the Anolis punctatus species group are discussed and evaluated, and the species 
themselves are diagnosed. 

INTRODUCTION 

Important new collections from Colombia have yielded Anolis 
that require description. Those dealt with here are allied to Anolis 
punctatus of Amazonia and the Atlantic Forest of Brazil, and, like 
the other recently described members of the punctatus group 
(Williams, 1965, 1974), they occur in areas peripheral to the range of 
that central and widely dispersed species. The three new taxa thus 
fill in blank areas in the known range of the complex. However, like 
many others of the punctatus species group, excepting only the 
central widespread and best-known species punctatus, these are 
species almost without distributions or ecologies. There are so few 
localities from which they are known, so little is known about their 
biology, that they are question marks rather than known compo- 
nents of fauna. They testify primarily to our basic ignorance of 
South American lizards. 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. 



BREVIORA 



No. 467 



I first describe the Amazonian taxa. Since the three best 
specimens are from the Comisaria of Vaupes, I call it: 

Anolis vaupesianus, new species 
(Figs. 1-5 and Map) 

Type: MCZ 156309 (formerly UTA 3626), adult male. 

Type locality: Timbo, a Cubeo village ca. 1 N 70 W on the Rio 
Vaupes, E of Mitii, Vaupes, Colombia, William F. Pyburn coll., 24 
July 1972. 

Paratypes: COLOMBIA: Vaupes. UTA 6850, MCZ 154592 
(formerly UTA 505 1) 1 same locality as type. J. K. Salser, Jr. coll.; 
ICN 4542, Cano Ti near Pamopeta, W. W. Lamar coll.; UTA 10283, 
Wacara, Lois Lores and Marilyn Cathcart coll. Amazonas. MCZ 
79655, lower Rio Apaporis, I. Cabrera coll.; MCZ 112098, Tio 
Miriti-Parana, La Providencia, F. Medem coll. 

Description (paratype variation in parentheses). Head: Head 
scales moderately sized, weakly (sharply) keeled toward tip of snout, 
smooth posteriorly. Ten (8-1 1) scales across snout between second 
canthals. Frontal depression distinct, scales within it smaller than 
those anterior to it (not or not conspicuously so). Eleven (9-10) 
scales border rostral posteriorly. Anterior nasal scale divided (not 
divided), it and inferior nasal scale in contact with rostral. Eight 
(6-7) scales between supranasals. Rostral area swollen, protuberant, 
rostral extending well beyond mental on male (not swollen or 
protuberant, rostral not extending much beyond mental in female). 

Supraorbital semicircles separated by one minute scale (or by one 
or two larger scales or narrowly in contact), separated from the 
supraocular disk by one row of granules. Supraocular disk ill- 
defined (better-defined), grading into surrounding granular scales. 
One elongate supraciliary followed by minute granules. Canthus 
distinct, five (5-6) canthal scales, the second largest. Six (5 7) loreal 
rows counting down from the second canthal, the ventralmost 
largest. 

Temporals and supratemporals granular, the latter grading into 
larger scales around the interparietal. An ill-defined and tapering 
double intertemporal row of somewhat enlarged scales. Inter- 



■This is a specimen referred to as A. punctatus by Greene (1977) 



1982 



THREE NEW COLOMBIAN ANOLIS 




Figure I. Anolis vaupesianus, n. sp. Type $, MCZ 156309. Dorsal view of head. 




Figure 2. Anolis vaupesianus, n. sp. Type $, MCZ 156309. Lateral view of head. 



BREVIORA 



No. 467 




Figure 3. Anolis vaupesianus, n. sp. Type q\ MCZ 1 56309. Ventral view of head. 



(982 



THREE NEW COLOMBIAN ANOLIS 




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J"! 



BREVIORA 



No. 467 




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1982 THREE NEW COLOMBIAN A SOUS 

parietal much larger than ear (larger), separated from the supra- 
orbital semicircles by two (3) rather large scales. Scales surrounding 
interparietal largest anterolateral^', posteriorly grading through an 
area of minute scales into the dorsals that are as large as the 
temporals. 

Suboculars weakly keeled, in contact with supralabials. Seven to 
eight (7-9) supralabials to center of eye. 

Mental semidivided, each half distinctly wider than long, in 
contact with four scales between the keeled sublabials. Three (5) 
sublabials in contact with infralabials. Gulars small, smooth, 
increasing in size laterally. 

Trunk: Dorsals granular, subequal, smaller than the weakly 
keeled ventrals which are subimbricate (juxtaposed), in transverse 
rows. 

Dewlap: Moderate in male, not extending posteriorly much 
beyond axilla. Lateral scales elongate, larger than ventrals, in close- 
packed rows. Edge scales shorter, crowded, subimbricate. Vestigial 
in female. 

Limbs: Largest scales on forelimb unicarinate, on digits weakly 
multicarinate. On hind limbs scales in front of thigh and lower leg 
weakly unicarinate, on back of thigh and lower leg granular. 
Supradigital scales weakly multicarinate. Twenty-five (22 26) la- 
mellae under phalanges ii and iii of fourth toe. 

Tail: Weakly compressed. No dorsal crest. No verticils. Scales 
above weakly carinate. A double middorsal row. Scales behind vent 
at most very weakly carinate, becoming distinctly so posteriorly. 
Postanal scales not well defined in male, absent in female. 

Size: Male type: 82 mm snout to vent; largest male paratype: 78 
mm snout to vent. Female paratype: 75 mm snout to vent. 

Field notes and color in life: F. Pyburn presents the following 
notes on the male holotype: "24 July 1972 I collected a dark green 
anole on the base of a small tree in village (Timbo) about one hour 
before dark. Dorsum of head, body, legs dull uniform green with 
scattered blue flecks; venter grey; dewlap skin black; scales of 
dewlap grey. Lizard turns dull brown when handled. Has fleshy keel 
on nape and along back." 

For MCZ 154592, a male, the notes below are abridged from 
those of J. K. Salser, Jr.: "Anolis sp.; Timbo (Vaupes), Colombia; 
March 20, 1974: very hot noon-time; caught in an aged avocado tree 



8 BREVIORA No. 467 

- scaled tree and jumped from its top in attempt to escape. . . Blue 
dots dorsolateral^ placed in stripes from vertebra down, angling 
down to lower sides; changes from dark green to green-brown 
mottled very quickly; eyes golden, eyelid border yellow..." 

For the female paratype, MCZ 112098 (Fig. 5), from the Rio 
Miriti-Parana, Federico Medem has provided the following infor- 
mation: "Caught 8:30 a.m. on underbrush, one meter high, in open 
country. . . Gular sack small, greenish. Iris golden. Intense green on 
body with lateral small blue spots. Middorsally 16 squarish sand- 
colored spots. Dark transverse bars on tail. Venter light gray with 
dark vermiculations." 

For MCZ 79655, a male, Medem reports the color as: "Dorsally 
green with lateral blue spots; dewlap 'morado' - violet, mulberry- 
colored." 

Comparisons: Very few specimens of A. punctatus are recorded 
from Colombia. The nearest to any of the type series of A. 
vaupesianus are two recently collected specimens in the Paris 
Museum (MHNP-1978-3086-87) from Hameau Kuira, Rio Agara 
Parana, a tributary of the Rio Putumayo, in Amazonas Territory. 
Both are males with strongly keeled ventrals, dorsum an unpat- 
terned dark purplish and dewlap with light skin with minute 
dispersed melanophores and purplish scales. 

The A. vaupesianus most nearly adjacent to these is MCZ 79655 
from La Providencia, Rio Miriti-Parana, a tributary of the Rio 
Caqueta, also in Amazonas. It is unfortunately a female. MHNP- 
1978-3086 has therefore been compared with the topotypic paratype 
UTA 6850, which is of the same size and sex. The dewlap in the 
paratype contrasts strongly with the Amazonas punctatus in its 
intensely black skin and white scales, and the ventral scales in 
vaupesianus are smaller and decidedly more weakly keeled. No 
other scale differences are evident, but the color and pattern of the 
Mitu paratype as preserved are strikingly different from anything 
observed in punctatus: it is brown, strongly blotched with darker, 
the dorsal blotches tending to form transverse series across the back. 
Anteriorly, a dark V, the apex backward, formed of somewhat 
irregular blotches, reaches the middle of the orbit on each side. A 
less well defined V in medial to this, reaching to the tops of the 
orbits. 

This very distinctive color and pattern are not, however, seen in 
the remainder of the type series of vaupesianus which, as preserved. 



1982 THREE NEW COLOMBIAN A NOUS 9 

are not strikingly dissimilar to A. punctatus. At best the distinctive 
pattern of UTA 6850 represents an element of the pattern repertoire 
of A. vaupesianus not present in A. punctatus. It clearly will not 
serve to distinguish most specimens. 

Habitat: The few notes on vaupesianus available give little 
indication of its ecology, except that it is associated with trees. The 
fact that the type was caught in a village probably indicates that it is 
an edge animal, not characteristic of the dark forest. 

The second species was first discovered in the Departamento del 
Huila. It has therefore been called: 

Anolis huilae, new species 
(Figs. 6-9 and Map) 

Type: ICN 3725, an adult male, collected by Helen Chin, October, 
1976. 

Type locality: Herberto Herrera's coffee plantation, Palestina, 
Huila, Colombia. 

Paratypes: COLOMBIA: Huila. KU 169830-31, Parque Ar- 
queologico San Agustin, 3 km SW San Agustin, 1750 m; ICN 3726, 
MCZ 159121-22: Palestina; MCZ 156305: Quebrada La Cascajosa, 
Parque Nacional Natural La Cueva de los Guacharos (northern 
boundary); INDERENA R-0297, -0562, -0665, MCZ 156306, 
156316: Parque Nacional Natural La Cueva de los Guacharos; ICN 
4461-62: 62 km (by road) NW San Isidro de Isnos, 1940 m. Tolima. 
ICN 3732: Amaine; ICN 3727-31, 3733, INDERENA (numbers not 
now available, formerly UVMP 4400, 4404). MCZ 159112-117: 
Cajamarca; Cali (Univ. Valle, Depto. Biol.) UVC 38, 191, ICN 
3735-36, MCZ 159119-120: Juntas; ICN 3734, MCZ 159118: 
Llanitas, 10 km N Ibague. 

[Referred specimen: Meta: ICN 4541 ca. 3-4 km NNE of 
Manzanares.] 

Description (paratype variation in parentheses). Head: Most head 
scales smooth, slightly concave, a few convex, obtusely keeled. Ten 
(8-11) scales across snout between second canthals. A few narrow 
scales in the frontal depression much smaller than surrounding 
scales. Eight (7-9) scales border rostral posteriorly. Anterior and 
inferior nasal scales in contact with rostral. Seven (6-7) scales 
dorsally between nasals. Snout not protuberant in male. 

Supraorbital semicircles broadly in contact with each other and 
with the supraocular disks. Supraocular disks with few (several) 



10 



BREVIORA 



No. 467 




Figure 6. Anolis huilae, n. sp. Paratype $, MCZ 159015. Dorsal view of head. 




Figure 7. Anolis huilae, n. sp. Paratype $, MCZ 159015. Lateral view of head. 



1982 



THREE NEW COLOMBIAN ANOLIS 



I I 




Figure 8. Anolis huilae, n. sp. Paratype $, MCZ 159015. Ventral view of head. 



large smooth scales. One elongate supraciliary on each side followed 
by granules. Canthus blunt, ca. 8 canthal scales, the second and 
third largest. Five (4 5) loreal rows, counting down from the second 
canthal, subequal. 

Temporals slightly smaller than the supratemporals, both granu- 
lar. No differentiated intertemporal line of enlarged scales. The 
scales surrounding the interparietal abruptly larger, those anterior 
and lateral largest (not so in female). Interparietal equal to (smaller 



12 



BREVIORA 



No. 467 




N 
U 

*6 



BO g 



i—iia^—^— 



1982 THREE NEW COLOMBIAN A SOI. IS 13 

or larger than) ear, separated from the supraorbital semicircles by 
2-3 (0 2) scales. Large scales posterior to interparietal followed by 
minute granules on nape, distinctly smaller than the granular 
dorsals. 

Suboculars keeled, in contact with supralabials, continued 
posterior to the eye by two distinct (less distinct) rows of smaller 
scales, anteriorly meeting canthals. Seven (5-7) supralabials to 
center of eye (supralabial five is much the longest in one paratype, 
an obvious anomaly; in the type, the supralabials are nearly 
subequal). 

Mental semidivided, in contact with three ( 1-4) scales between the 
very large, smooth sublabials. Three sublabials (1-4) in contact with 
infralabials. Central throat scales small, swollen, oval, becoming 
abruptly longer laterally alongside the sublabials. 

Trunk: a few middorsal (2-4) rows indistinctly enlarged, weakly 
or distinctly keeled, grading into granular flank scales. Ventrals 
much larger than largest dorsals, smooth, imbricate or subimbricate 
(juxtaposed in female), not in distinct transverse rows (in transverse 
rows). 

Dewlap: Large in male, extending to midbelly (absent in female). 
Scales at edge larger than ventrals, smooth. Lateral scales smooth, 
in double rows separated by naked skin. 

Limbs: Limb scales weakly uni- or multicarinate. Supradigital 
scales multicarinate. Twenty-one (22) lamellae under phalanges ii 
and iii of fourth toe. 

Tail: Slightly compressed, without crest. Enlarged postanals 
present (absent in female). A double row of keeled scale middorsally 
on tail. Scales behind vent smooth. 

Size: The male type is 75 mm in snout-vent length. The largest 
male reaches 82 mm. 

Color in life: Helen Chin reports topotypic animals as 'parrot 
green' in the trees. Her color transparencies show tan and reticulate 
animals after capture. 

Color notes are available for two MCZ specimens. MCZ 156405. 
a male: "Dorsum bright green with yellow spots (seven large spots 
along the spine). Tail banded green and black. A white dot inside a 
black ring on side of neck behind the ear. Eyelid yellow, iris grey. 
Chin grass-green. Dewlap yellow with six green stripes, these 
bordered by numerous black dots. Rest of underparts greyish." 

MCZ 156306. a female: "Dorsum with wide tan stripe from back 



14 BREVIORA No. 467 

of head to halfway down tail, at which point it becomes speckled 
with black dots. Rest of back grey-green, darker next to tan stripe, 
and lighter with dark patches on the sides. Legs banded black and 
light green. Head light green with dark around eyes. Tip of tail 
bronzy. Venter yellow with few small black spots." 

Both these specimens are near-topotypes. Palestina is a settlement 
at the beginning of the 22 km trail leading to the Parque Nacional 
Natural La Cueva de los Guacharos. 

The colors in life of the Kansas specimens from Parque 
Arqueologico de San Agustin are described rather differently 
(although not in the same detail): "At night - dorsum pale green with 
brown reticulations. Dewlap cream with grey-bordered white 
streaks. Iris creamy-grey. Tongue orange." 

Notes are at hand also for two females collected by Pedro Ruiz 
and J. D. Lynch near San Isidro de Isnos (notes by Lynch): 

ICN 4461: "Sleeping on leaf in forest near stream. Iris yellowish. 
Ground color dull green with brown barring; greenish cream spots 
on flanks; tail barred brown and medium brown with cream edges; 
venter dirty cream with brown spots. Throat the same but with 
greenish cast." 

ICN 4462: "Sleeping on Ortega, 3 m above stream at edge of corn 
field. Iris grey brown. Tongue pale orange yellow." 

Dennis M. Harris has provided notes on the series from Tolima: 

"Male: Dorsum pale leaf-green, spotted most vividly on the sides 
by 1-2 mm diameter circular dots of greenish-yellow. Head slightly 
brownish. Eyelid yellow. Nape with a lateral spot of blue-green to 
black, sometimes surrounded by a yellow area, sometimes with a 
light center. Tail barred or not with bands equal in width to the 
green. Venter white to grayish. Dewlap skin pale yellow, scales 
cream, in rows that are edged with black spotting. 

Male may change to a deep brown with vivid yellow spots. It does 
so when handled and possibly as a response to another male. The 
one brown male seen in the field was never a green one, and had a 
vertically flattened body and extended dewlap. (The cream scales of 
the dewlap may become drab greenish in the brown-phase male.) 

Female: Banded or striped, light or dark brown. The stripe is 
crayola brown with dark brown flecks. There is no color change 
except for minor tone changes." 

Color notes have been made available by W. W. Lamar for the 
single animal from Manzanares, Meta: "Dorsum dark gray with 



1982 THREE NEW COLOMBIAN A, XOL/S 15 

numerous light gray-green spots. Mid-dorsum brighter than rest of 
the back. More green toward the center of the flanks. Tops of 
forelegs gray-green with lighter yellow-green elbows and feet. Hind 
legs with 3 light bands above knee. Three broken bands below knee. 
Tail pale green with 11-12 dusty bands. A black spot with an 
aquamarine center on the side of the neck. Yellow eye ring and 
snout aqua-blue-green, darker between the eyes. Venter pale yellow 
with dark gray spots on the sides, brighter yellow near vent, broad 
portion of tail, and inguinal area. A patch of aqua-blue on tail 10-25 
mm from vent, rest of tail dark gray. Dewlap an overall aqua green- 
blue striped with pale green and dotted with black. Spaces between 
stripes are pale yellow. Nostril yellow. Tongue yellow." 

Comparison of these notes for the Meta specimen described 
shortly after death with the detailed account from Huila and Tolima 
shows various discrepancies. Color differences in Anolis, however, 
may be described differently by different individuals or by the same 
person at different times. This, plus the capacity for color change, 
suggest caution in attributing taxonomic significance to color 
differences of the sort seen here. (See also comment below.) There 
are no scale differences. 

Color as preserved: Most male A. huilae are strikingly distinctive 
because of the bold lateral spotting, plus the lines of dark spots on 
the dewlap. Some specimens display a more muted spotting. This is 
especially true of the individual from Manzanares, Meta. The latter 
seems at first distinguishable because the spotting on the dewlap is 
not in lines but quite random. Some specimens from Tolima, 
however, have the spotting on the dewlap irregular also. The Meta 
animal does not, on present evidence, fully match the type series; 
since it is also geographically peripheral (on the east side of the 
Andes, rather than in the Magdalena Valley) I have not made it a 
paratype. With only one specimen and no structural differences it is 
not permissible to describe it taxonomically. 

Habitat: Dennis Harris reports: "found on relatively smooth bark 
tree trunks usually, between 1 and 7 meters. Males almost 
exclusively there, females sometimes on bushes nearby, 1 m high. 
Usually only single male/ tree trunk (25 cm+ diameter). Shrubs 
seem to be only peripheral environments. Not found on very 
smooth-bark guayaba (guava) trees. Not found on trunks with 
heavy moss growth. Common right in town of Cajamarca and along 
edges of road. Males more frequently seen and more of them 



a«i&i^^ 



BREVIORA No. 467 



caught. Only time male and female seen together on tree trunk they 
were copulating. Usually seen head down, out (like frenatus) on 
trunk. At Anaime Prionodactylus argulus found at same site. At 
Llanitas, 10 km from Ibague, Anolis antonii and A. huilae; at 
Juntas, much higher, Phenacosaurus and A. huilae." 

MCZ 156316 was found below a cave-hole of Rupicola rupicola 
among defecated seeds. Apparently a food item, it was in relatively 
good condition and quite recognizable. 

Comparisons: A. huilae, although obviously a member of the 
punetatus species group, is not obviously closely related to any 
other presently described member. The color pattern of males is 
highly distinctive. 

A third species comes from the southern slopes of the isolated 
Santa Marta range. It is to be called: 

Anolis santamartae, new species 
(Figs. 10-13 and Map) 

Type: CAS 113922, adult male. 

Type locality: San Sebastian de Rabago, Sierra Nevada de Santa 
Marta, Cesar, Colombia (10°4" N, 73° 16" W), Borys Malkin coll., 
1 10 April, 1968. 

Paratypes: same data as type: CAS 113924, MCZ 156311. 

Description (paratype variation in parentheses). Head: Moderate- 
ly elongate. Head scales rather large, smooth, flat except for smaller 
bluntly keeled or swollen scales at tip of snout. Six (4-5) scales 
across snout between second canthals. Frontal depression distinct, 
the scales within it larger (some smaller) than those anterior to it. 
Anterior and inferior nasal scales in contact with rostral. Four (5-6) 
scales dorsally between supranasals. Rostral area protuberant in 
male, overhanging lower lip (not so in female). 

Supraorbital semicircles broadly in contact with each other and 
with the supraocular disks. Supraocular disks well defined, com- 
prised of 8 to 10 smooth scales. Remainder of supraocular area with 
large granules. Canthus distinct, four or five canthal scales, the 
second and third (or first and second) largest. Five (4-5) loreal rows, 
the lowermost largest. Temporal and supratemporal scales granular, 
larger at the angle of the mouth an at the angle between temporal 
and supratemporal regions. Scales surrounding the interparietal 
rather abruptly larger, largest laterally, grading into dorsals 



1982 



THREE NEW COLOMBIAN A NOUS 



17 







Figure 10. Anolis santamartae, Type <J, CAS 113922. Dorsal view of head. 







Figure 11. Anolis santamartae. Type $, CAS 113922. Lateral view of head. 



18 



BREVIORA 



No. 467 




Figure 12. Anolis santamartae. Type $, CAS 113922. Ventral view of head. 



posteriorly. Interparietal much larger than ear, broadly in contact 
with the supraorbital semicircles. 

Suboculars weakly keeled, in contact with supralabials, anteriorly 
separated from the canthal ridge by two (1-2) scales, posteriorly 
grading into the temporals. Seven supralabials to the center of the 
eye. 

Mental semidivided (mental region abnormal in MCZ 15631 1), in 
contact with two large scales between the well-developed sublabials. 



1982 



THREE NEW COLOMBIAN ANOLIS 



19 




C/3 
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g, 
P 



60 S 



20 



BREVIORA 



No. 467 




Figure 14. Distribution of the species described herein. 
O: Anolis huilae; and ■ : Anolis vaupesianus. 



A nolis santamartae; 



1982 THREE NEW COLOMBIAN A NOUS 21 

Four (3) sublabials in contact with infralabials. Gulars rather large, 
swollen, increasing in size laterally and anteriorly. 

Trunk: Ca. four to six middorsal rows flat, weakly keeled, 
subimbricate, much larger than flank granules but grading into 
them. Ventrals still larger, smooth or obtusely keeled. Chest scales 
distinctly keeled. 

Dewlap: Large in male, only indicated in female. Scales in close 
set rows, elongate and narrow, except at edge where they are flat 
and smooth. 

Limbs: Scales on limbs uni- to multicarinate, those on digits 
multicarinate. Eighteen (19-21) lamellae under phalanges ii and iii 
of fourth toe. 

Tail: Compressed. Scales weakly to strongly carinate. A double 
middorsal row. No tail crest. No verticils. Scales behind vent 
carinate. Post-anals large (broken up on one side type, absent in 
female.) 

Size: Male type: 50 mm, snout to vent. Male paratype: 55 mm. 
Female paratype: 53 mm. 

Color as preserved. Female paratype: Gray-brown. A light line 
from suboculars to above ear. A narrow dark interorbital bar. A 
series of ovoid or rhomboidal dark blotches paravertebral^. 
Irregular oblique dark streaking on flanks. Limbs irregularly 
banded. Tail very weakly banded. Throat and belly smudged with 
darker. 

Male type: Gray. A light line from suboculars to above ear. 
Another light line from lower edge of ear along lateral edge of 
dewlap onto underside of upper arm. Weak traces of blotching 
dorsally. Irregular oblique dark streaks on flanks. Limbs and tail 
with very weak and obscure markings. Underneath smudged with 
brown. Dewlap skin white, scales very light brown. 

Male paratype: A dark interorbital bar. A light stripe from 
suboculars to above ear, another from below ear to shoulder. 
Middorsum and dorsum of tail obscurely marked with light and 
dark. Flanks with oblique dark streaks. Limbs smudged above. 
Belly with irregular dark markings. Throat with vaguer markings. 

Habitat: San Sebastian is at the border between the Departments 
of Cesar and Magdalena, and the specimens may have been 
collected in either Department. The elevation is at, or above, 2,000 
m. There are no ecological notes. 



22 BREVIORA No. 467 

DISCUSSION 

Comparison of santamartae and the other two new species of the 
punctatus complex with each other and with the other already- 
described members of the complex could be made in tabular form, 
as in Williams (1974). Here, however, I prefer to discuss and 
elevuate the characters on which I recognize the several species. I 
begin with the color differences. 

Color in Life 

For this purpose the few detailed descriptions of color in life of 
Anolis punctatus must be quoted. 

Hoogmoed (1973) has a description for A punctatus 'punctatus' 1 
from Surinam: "Color in life grass-green above with sky-blue dots, 
eyelids purplish with a yellow rim. Chin bluish, throat greenish. 
Belly white to purple-green. In bright sunlight the lizards are purple 
with sky-blue dots. Dewlap in males yellow ochre with rows of white 
scales; in females the scales on the dewlap are yellow. Iris orange- 
brown." 

A parallel description is provided by Dixon and Soini (1975) for 
A. punctatus 'boulengeri' from the Iquitos area in Peru: "The dorsal 
coloration of adults is leaf-green with six to seven very narrow white 
cross-bands confined to the upper body. The lower sides are spotted 
with minute white dots, and the limbs are spotted with minute dark 
brown spots. Occasionally specimens are deep purple dorsally, 
especially after being in captivity for a few hours. The white cross- 
bands and white spots are very distinct when the latter color 
prevails. Male dewlaps are orange to reddish-orange with the 
enlarged lateral scale rows light yellow." 

P. E. Vanzolini has provided me with a translation of the 
unpublished field description of Helmut Sick of an A. punctatus 
'punctatus' from the Atlantic Forest in Brazil (Rio Itaunas, Espirito 
Santo): "General body color alive: grayish sepia, dewlap chrome 
yellow. Freshly killed: moss green, eyelid yellowish green, dewlap 
chrome yellow with stripes composed of whitish green spot, chin 
whitish-green. Underparts greenish-white. Digits and tail with 



■Western specimens of punctatus with keeled ventrals are usually called boulengeri 
O'Shaughnessy whether as subspecies or species. See Williams and Vanzolini (1980) 
for evidence that 'boulengeri' is a morph and not a valid taxon. 



1982 THREE NEW COLOMBIAN ANOLIS 23 

vestiges of dark brown crossbands. Posterior two-thirds of tail 
grayish sepia. Flanks with lines of white spots...." Vanzolini 
himself describes dewlap colors for two additional specimens: 
(Santo Amaro dos Brotas, Sergipe): 'pure yellow'; (Lago Mapori, 
Rio Japura, Amazonas): 'rusty orange.' 

No color descriptions from living specimens are available for A. 
santamartae, A. caquetae, A. deltae, or A. nigropunctatus. The 
color descriptions of A. vaupesianus and huilae have been given 
above. 

There are serious problems in the comparisons of color and 
pattern in the members of this complex. The few good reports are 
for the local populations of widely scattered regions or for 
individual specimens. This would be a difficulty in any taxonomic 
group. In Anolis an additional problem exists: color repertoire may 
be extensive and tends to be especially so in the green species. In the 
latter, the difference between light and dark phases may be extreme, 
and there may be in life intermediate patterns or colors that are only 
briefly or occasionally seen. (This may have been true, for example, 
of the Meta specimen referred to A. huilae above.) Preserved 
specimens compound the dilemma already posed. Pattern or lack of 
pattern may be due entirely to the preservative, or to its strength. 
Specimens done in strong formalin tend to a muddy uniformity; 
many of the older specimens in collections are of this type. It is 
fortunate that none of the specimens considered in this paper are of 
so inferior a quality. Nevertheless, they have been preserved by 
different individuals with different preferences in technique. The 
preserved specimens are therefore only minimally comparable. 

But even the available descriptions of color in life are imperfectly 
comparable. They were not made to be checked against one 
another, and they therefore do not stress quite the same details. 
They were never checked against the same color standards. 

Yet it is quite clear that enough information exists to assure us 
that differences exist. While no possibility exists of entirely avoiding 
error, some plausible conclusions may be drawn. 

Color in anoles is best discussed by breaking it into components. 
It is a fact about Anolis in general that dewlaps are least modified by 
color change and most readily interpretable in preserved specimens. 
This fact quite accords with the view (e.g., Rand and Williams, 
1970; Williams and Rand, 1977) that the dewlap is an intra- and 
interspecific signal which should have high constancy in order 



24 BREVIORA No. 467 

always to convey the same message. (A caveat: this rule may have 
less validity in mainland Anolis than it appears to have in the species 
of the islands.) 

In the present case we have the following evidence of dewlap 
color: 

pimetatus punctatus (Surinam) 

— yellow ochre with white scales (Hoogmoed). The other reports 
from the range of subspecies (Sergipe, Espirito Santo) stress a 
yellow color also. 

punctatus 'boulengeri' 

— orange to reddish-orange (Dixon and Soini). Clearly a deeper 
shade than in punctatus punctatus, but not sharply different. 
Preserved specimens of both taxa show only light skin and 
scales. 

vaupesianus 

— 'black' (Pyburn); 'mulberry red' (Medem) presumably on the 
authority of the collector: 'greenish' in the female; Medem, in 
this case himself the collector. There is an obvious discrepancy 
here, but we are evidently dealing with a dark dewlap, and the 
dewlap skin in all preserved specimens is darkly pigmented. 

From these notes it is clear that vaupesianus is sharply distinct in 
dewlap color from the animals that in size and morphology it most 
resembles, western A. punctatus, and from /I. huilae. The dewlap of 
the latter, with dark spots placed in lines or randomly, is quite 
unlike any of the others. Insufficient information exists for any 
really useful discussion of dewlap color in the other taxa. 

As regards body color, I am very uncertain what can be inferred. 
From the descriptions of Hoogmoed for Surinam and of Dixon and 
Soini for Peru, it would appear plausible that a purple phase exists 
in western A. punctatus which does not occur mA. vaupesianus, but 
Vanzolini has not noticed such a purple phase in the punctatus of 
Amazonia and eastern Brazil punctatus that he has caught. Absence 
of purple may therefore not be fully diagnostic for vaupesianus. 
Other aspects of the color of vaupesianus involve tints and special 
aspects of its color repertoire which again are of uncertain value. 
Size, body, and dewlap color seem safer guides to identification. For 
A. huilae we do not know enough about color repertoire to evaluate 
the colors in life that we do know. The bold flank spotting seen in 
huilae may be useful, like the nape spot, but neither are obvious in 
females and are not mentioned in some of the descriptions of colors 



1982 THREE NEW COLOMBIAN ANGUS 25 

in life. See above for discussion of the specimen from Meta. For 
nigropimctatus, caquetae, santamartae, and deltae, discussions of 
preserved body color are of very dubious value. Are the spots of 
preserved nigropunctatus surely of diagnostic value? There is at 
present no way to tell. The light line in santamartae from suboculars 
to ear seems distinctive. It is less evident in the type than in the other 
darker specimens. It may or may not be readily visible in life. For 
these last four species body color cannot be disregarded but 
determination of specimens must depend upon size, scale characters, 
and locality. Let us look at these features in sequence. 

Size 

A. vaupesianus, A. huilae, and A. nigropunctatus are all within 
the size range of A. p. punctatus (70-80+ mm snout to vent length). 
A. dissimilis, A. caquetae, A. santamartae, and A. deltae are, on 
present evidence, smaller (50-60 mm) (Table 1). (I include in these 
tables and discussions all described members of the punctatus group 
east of the Pacific slopes of the Andes.) 

Table 1. Size of punctatus group anoles from east of the Andes and from the 
inter-Andean valleys. 

Maximum male snout to vent length 
(in mm.) 

punctatus 85 

transversalis 83 

vaupesianus 82 

huilae 82 

jacare 73 

nigropunctatus 72 

deltae 58 

caquetae 57 

dissimilis 56 

santamartae 55 



Scale characters 

The more significant of these may be examined one by one: 
Ventrals: In Anolis it is probable that smooth ventrals are 



26 BREVIORA No. 467 

primitive, keeled ventrals the derived condition. (The argument 
derives from the apparently more frequent correlation of smooth 
than keeled ventrals with other characters considered primitive; this 
is not a strong argument.) Most punctatus group species have 
smooth ventrals, but a few have strong keels on the venter. Table 2 
again lists the conditions of the several species east of the Andes or 
in the inter-Andean valleys. 

Table 2. Ventral scales in punctatus group anoles from east of the Andes and 
from the inter-Andean valleys. 



punctatus 


smooth or weakly to strongly keeled 


vaupesianus 


weakly keeled 


transversalis 


smooth 


huilae 


smooth 


jacare 


smooth 


nigropunctatus 


smooth 


deltae 


smooth 


caquetae 


weakly keeled 


dissimilis 


smooth 


santamartae 


smooth or weakly keeled 



Nostril-rostral relationship: This character is somewhat variable 
within species, but within a narrow range. The punctatus group 
shows only a fraction of the possible range, and each species shows 
only a segment of this restricted range. The relevant conditions and 
the species in which these are seen are listed below: 

1) Circumnasal scale (=nasal) directly in contact with rostral: 
nigropunctatus, jacare. 

2) Circumnasal scale separated from rostral by a small round 
undifferentiated scale: nigropunctatus, dissimilis. 

3) A differentiated anterior nasal scale present and in contact with 
rostral (the anterior nasal is characteristically triangular and 
overlaps posteriorly part of the circumnasal): deltae, huilae. 

4) Anterior plus an inferior nasal in contact with rostral: puncta- 
tus, houlengeri, transversalis, vaupesianus, caquetae, huilae, 
santamartae. 

It is noteworthy that, while a species may exhibit two of these 
four categories, these are always adjacent ones and no species is 



1982 THREE NEW COLOMBIAN ANOLIS 27 

known to range the full gamut. This character will assist in 
discriminating species. 

Snout protuberant or not in males: This character, well known in 
A. punctatus, is clearly present in A. vaupesianus, less obvious in 
nigropunctatus, and absent in the other taxa. It again assists in 
identifying species. 

Scales across the snout between second canthals: The range in the 
punctatus group is moderate, most species ranging between 6 and 9, 
transversalis including, however, values as low as 4, and caquetae, 
nigropunctatus and vaupesianus reaching 10. Very low counts here 
may point to transversalis. Counts otherwise are not diagnostic. 

Scales between the semicircles: the minimum number between the 
semicircles is 0, usually implying broad contact. In the species set as 
a whole the modal conditions are and 1; a count of 2 is relatively 
infrequent. This character cannot be relied on to distinguish species. 

Loreals: The maximum number of loreals (counted down from 
the second canthals) may be as low as 3 (transversalis) or as high as 
7 (vaupesianus). More frequently the count ranges from 4 to 6. 
Again, a very low number of loreals may point to transversalis, but 
in general loreals are not diagnostic. 

Interparietal: A very large interparietal in contact with the 
semicircles is a distinguishing feature of the unique type of caquetae 
and of the type series of santamartae and of dissimilis and deltae. 
An interparietal smaller than the small ear is characteristic of 
nigropunctatus (clearly so only in the type). For the other members 
of the punctatus group the interparietal is not diagnostic. 

Number of supralabials to the center of the eye: The exceptional 
number of 11 is distinctive for A. dissimilis. A range of 6 to 9 is 
characteristic for all the other species and hence does not discrimi- 
nate among them. 

Lamellae under phalanges ii and Hi of fourth toe: Number of toe 
lamellae in Anolis correlates with two factors — perch and size. In 
the known cases, punctatus group anoles tend to high perches— 
usually in the crown — and should have high lamellar counts for 
their size. Correspondingly, the better-known, larger species, known 
to correspond to expectations in perch — punctatus, vaupesianus — 
have counts in the mid- to upper 20s. A. nigropunctatus, A. jacare, 
and A. huilae have relatively low counts for punctatus group 
animals of their size (20-24). 



28 BREVIORA No. 467 

Tail crests: Two species — dissimilis and deltae — have tail crests 
and hence are unique in the group. Others have a double row 
without a crest. 

Species Diagnoses 

What combinations of characters diagnose the punctatus group 
species I have described herein and the others previously described? 
(See also Tables 3 and 4.) 

1) Vaupesianus is distinctive in its black dewlap, strongly 
protuberant snout in males, weakly keeled ventrals, and 
moderate size. (Distribution: Vaupes and Amazonas.) 

2) Huilae is distinctive in its spotted or streaked dewlap, and 
(usually) in the vivid mottling of its flanks and in its moderate 
size (ventrals smooth). (Distribution: the upper Magdelena 
valley in Huila and Tolima, perhaps at high elevations on the 
eastern slope of the Andes in Meta [Manzanares].) 

3) Santamartae is distinctive in the large interparietal broadly in 
contact with the semicircles, plus smooth or weakly keeled 
ventrals, small size, and a light line from supralabials to ear and 
from ear to upper arm. (Distribution: south slope of Santa 

Marta mountains.) 

4) Caquetae is diagnosed by its large interparietal broadly in 
contact with the semicircles, plus keeled ventrals, small size, 
and no light line from suboculars to ear. (Distribution: known 
only from the Upper Apaporis in Caqueta, Colombia.) 

5) Nigropunctatus stands out by its small interparietal, smaller 
than the small ear, an irregularly and weakly punctate pattern 
on the flanks (in males on middorsum also), snout protuberant 
in males, and by moderate size (ventrals smooth). (Distribu- 
tion: from Norte de Santander in Colombia to Tachira and 
Trujillo in Venezuela.) 

6) Deltae has the large interparietal broadly in contact with the 
semicircles and small size of caquetae but has smooth ventrals, 
a tail crest and a short, blunt head. (Distribution: Delta of the 
Orinoco in Venezuela.) 

7) Dissimilis has a very large interparietal in contact with the 
semicircles, small size, smooth ventrals and tail crest like those 
of deltae, but a strikingly elongate head. (Distribution: Madre 
de Dios in Peru.) 



1982 THREE NEW COLOMBIAN ANOL/S 29 

8) Jacare is distinctive in moderate size, male without a protu- 
berant snout, with an immaculate yellow dewlap and a strongly 
reticulate body pattern with often an oblique light streak from 
the throat to above the shoulder; female with unspotted body 
and unmarked throat (ventrals smooth). (Distribution: the 
Sierra Madre de Merida in Venezuela.) 

9) Transversalis is like jacare in moderate size and the lack of a 
protuberant snout in males, but the male has a yellow dewlap 
with streaks and spots, and a body pattern of oblique light lines 
enclosing rows of black spots; and females with bold transverse 
bands on the body and the throat vividly cross-marked with 
black (ventrals smooth). (Distribution: western Amazonia.) 

10) Punctatus has moderate size and a strongly protuberant snout 
in males and a yellow or orange dewlap. (Ventrals smooth or 
keeled). (Distribution: Amazonia, the Guianas, and the Atlan- 
tic Forest of Brazil.) 

The Geography of Difference 

The species of the punctatus group that have been discussed here 
are all those thus far reported that are east of the Andes or inter- 
Andean. (For convenience, santamartae is counted in this group; it 
lies in an eastward-looking valley in the southern part of the isolated 
Sierra Nevada de Santa Marta.) This assemblage, as it turns out, is, 
if not a natural group, clearly as natural as a group which also 
includes taxa west of the Andes, currently referred to the punctatus 
species group. None of the western species are demonstrably close to 
the species here considered. 1 



'I no longer consider A. nigropunctatus to be especially close to A. nigrolineatus 
(Williams, 1965). A. nigropunctatus, as will be suggested below, is probably closer to 
A. jacare, and A. nigrolineatus may be a strict synonym of A. festae Peracca 
(syntypes examined). Both nominal species have the same highly peculiar dewlap 
with an elongate dark spot. There are no scale differences. There is variability in head 
shape. The sole known differences between southern Ecuadorian specimens 
( Machala, El Oro) known from recent large collections by Fitch, Echelle and Echelle, 
and northern populations (Pichincha) known from equally large collections made by 
Kenneth Miyata are smaller size in the northern populations (Miyata, personal 
communication) and a blue iris in the north (Miyata) as compared with the yellow 
iris reported by Fitch el al. (1976) for southern animals. No recently collected 
animals — live or preserved — show the narrow dark middorsal line believed diag- 
nostic for A. nigrolineatus. 



30 



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1982 THREE NEW COLOMBIAN ANOLIS 33 

The eastern and inter-Andean assemblage is, itself, clearly 
morphologically heterogeneous, showing several levels of differen- 
tiation. Yet it is a striking fact that only one of the 1 1 forms here 
called species is widely sympatric with any of the others: A. 
transversalis occurs within the range of A. punctatus and A. 
vaupesianus. It is abundantly distinct not only from the two taxa 
with which it is sympatric, but from all other punctatus group 
species. It differs not only in the striking color characters 
emphasized in the section immediately above, but it stands at one 
extreme in scale characters also (low counts of loreals and scales 
across the snout). 

None of the other nine described forms is known to co-occur. 
Two of them, however, are in their own way sharply distinct. A. 
dissimilis, as its name implies, seemed very distinct when it was first 
made known, but the tail crest, otherwise unique within the 
punctatus group, has since been found in widely allopatric deltae 
(which differs, however, inter alia, in its much shorter head). 

The remaining taxa are allopatric; they do, however, divide into 
low-land and high-land forms, and this, while it is not prima facie 
evidence against conspecificity, makes it a little less likely. 

The low-land species are punctatus, vaupesianus, and caquetae. 
Here it is clear that as regards the first two I have relied on color in 
life. The difference, however, is sharp, and, if there are any 
intermediate populations, we can specify where they must be: in 
Amazonas between the Rios Putumayo and Caqueta. 

It is, of course, crucial to the final allocation of all allopatric taxa 
that are phenetically similar that, when possible, we endeavor to 
discover whether there is a contact zone and to determine what 
happens there. I have expressly chosen not to call distinctive 
populations subspecies merely because they are related. Unless 
variation is clinal and the relevant characters not congruent — in 
which case taxonomic description is not warranted — the alternative 
to the hypothesis of allospecies, which I am using here, is subspecies, 
a highly restrictive hypothesis according to which distinctive 
populations with congruent characters meet in well-defined zones of 
intergradation. I agree with the long-expressed opinion of Wilson 
and Brown (1953) that these restrictive conditions are rarely met. A. 
punctatus and A. vaupesianus are, indeed, distinctive populations 
but closest relatives that are either allospecies, the ends of a cline, or 
supspecies. There is not present evidence of clinal change or of 



34 BREVIORA No. 467 

intergradation. My decision therefore is to tentatively regard the 
two populations as allospecies. 

The case of A. caquetae is different. It is a single specimen 
peculiar in some aspects of color and squamation — in combination 
outside the known limits of other forms. As a single specimen, it is 
not clearly representative of any distinctive population. None of its 
characters is individually unusual within the broad variation of the 
punctatus group. The large interparietal in contact with supra- 
orbital semicircles is not at all usual in the punctatus species group, 
but see A. santamartae. The color as preserved is distinctive but 
suffers the difficulty of preserved color in Anolis, discussed at some 
length above. Clearly, acceptance of A. caquetae as a valid species 
involves risk of error. But if it were to be synonymized, what would 
it be synonymized with? A. punctatus or A. vaupesianus! Or would 
both sink into a variable species to be called A. punctatus! The large 
interparietal in contact with semicircles is rare in A. punctatus (one 
known case in 157 specimens), unknown in A. vaupesianus (but the 
sample is too small for a secure statement). It is closer to A. 
vaupesianus, but it would seem erroneous on present evidence to 
synonymize A. vaupesianus with caquetae. The case here is one of 
inadequate evidence, and, above all, of inadequate search for 
evidence. The fauna of Rios Vaupes, Caqueta, and Putumayo and 
their tributaries is very poorly known. 

The remaining taxa with which we are concerned— y'arare, 
nigropunctatus, huilae, and santamartae — are all animals of high 
elevations. Of these, only jacare is an old name, and huilae and 
santamartae have been described herein. 

Santamartae is the most distinctive of these. In the tendency to 
keeling of the ventrals and the light line from the labials toward the 
ear it is unlike the others. It seems also to be smaller. 

The other three, it must be confessed, are primarily distinctive in 
coloration. A. huilae seems to stand out in this series. The bold light 
spotting and nape spot in males, as well as the contact of 
differentiated anterior and inferior nasals with the rostral, contrast 
with the dark spotting of nigropunctatus and the vermiculation of 
jacare and with the primitive circumnasal scale characteristic of 
both the latter. 

Betweenjacare and nigropunctatus there are no conspicuous scale 
differences other than the smaller interparietal of nigropunctatus 
(but this is sharply evident only for the male type of nigropunctatus, 



1982 THREE NEW COLOMBIAN ANOL1S 35 

the only known male). They differ, however, in color; the sparse 
dark spots of nigropunetatus are quite unlike the bold vermiculation 
of male jacare. I am reinforced in my confidence of this color 
character by examination of a series of 1 1 specimens of jacare from 
three localities in the state of Merida made available to me by James 
R. Dixon. The males seem clearly distinctive; the females are not as 
readily assigned. Between these two, however, and even between 
nigropunetatus and huilae, there might be a contact zone. The 
appropriate intermediate regions remain uncollected. 

I have not here gone beyond a minimal analysis of the phenetic 
similarities of the 10 species of the eastern punctatus species group. I 
do not attempt any history or even any dendrogram of relationship. 
It seems to me totally premature to do so. 

Of the 10 species only three — punctatus, transversalis, and 
jacare — have been known for more than 15 years. Of these, only 
punctatus is widespread and relatively common. A. transversalis is 
widespread but rather scarce in collections, although the collections 
of Pekka Soini (Soini and Dixon) indicate that it must be 
moderately common, at least in the Iquitos region. A. jacare is 
known only from the Merida region in Venezuela where, from the 
experience of Carlos Rivero Blanco and Williams (Williams et ah, 
1970, and the recent collections by James Dixon), it is commoner 
than the infrequent sightings of it and the meager collections 
indicate. 

Anoles of the punctatus group are animals usually of the canopy 
and appear rather infrequently on the trunks or near the ground (A. 
huilae fide Dennis Harris may be an exception since it is, according 
to his report, more of a trunk-ground adapted species like A. 
frenatus [cf. Scott et ah, 1976 for the latter]). Most of them are, 
therefore, not conspicuous lizards, whatever their real abundance. 

Other species may be genuinely rare, rare enough that the 
question of intraspecies and interspecies, even intergeneric compe- 
tition, becomes moot. It is difficult, however, to demonstrate that 
failure to observe is equivalent to real rarity. Many species of Anolis 
also are highly cryptic and behaviorally highly adapted to crypsis, 
almost never visible unless they move. Some, readily capturable 
while sleeping and therefore known to be at least locally common, 
are rarely seen in the daytime, and their diurnal activity and ecology 
are therefore almost unknown. 



36 BREVIORA No. 467 

However, quite apart from any characteristics of the animals that 
may make acquisition of knowledge about them difficult, a primary 
and evident difficulty is failure to explore and to collect. Anolis 
huilae is not only a species just now described, but only very recently 
collected. Dennis Harris is the collector of all the Tolima specimens 
— 23 obtained in one 10-day trip which did not specialize in Anolis 
but sought lizards of all kinds. Such a collection in a brief span 
implies that prior sampling of this area, which has not been totally 
neglected — it is the type locality of Anolis tolimensis and A. ibague - 
has nevertheless been quite inadequate. 

If this is true of areas from which there is moderately good 
representation in old collections, what may still be coming from 
areas that are still altogether or nearly untouched? 

The Department of Narino is one such area. Collections by even 
Ayala's group have been only in the low-lands near Tumaco; there is 
no reason to believe that even near Tumaco there is not much more 
to find. 

An undescribed species is known from the west side of the Sierra 
Nevada de Santa Marta, and one on the south side has been de- 
scribed herein. What occurs on the east side? Are the new species the 
only ones on the west and south sides? Is the fauna of the Choco 
completely known, of Cauca, Antioquia, Bolivar, Santander, and 
Norte de Santander? One can, in fact, call the roster of the 
Departments of Colombia and it will not be possible to say even of 
the relatively well collected areas that the herpetofauna is com- 
pletely known. We in fact do not know enough to estimate the frac- 
tion that is unknown. (However, to shirk the frustrating effort of 
dealing with incomplete and inadequate evidence would be to 
acquiesce in the perpetuation of ignorance. Our obligation is, while 
not avoiding these thorny thickets, to explore carefully and to leave 
the paths that we pursue plainly marked.) 

What has been done in this paper is to attempt — and I do not 
vouch for more than an attempt — to answer the first and most 
elementary (but most fundamental) of biological questions: who? - 
i.e., what is the cast of characters? After we have answered this first 
question we may begin to ask the question where? On the way we 
may begin to answer other questions: what does it do? with whom 
does it interact? Much later, when the picture is much clearer, we 
can try to answer historical questions: how did this system come 
about. On the evidence before us the picture is too incomplete, the 



1982 THREE NEW COLOMBIAN ASOI.IS 37 

unfilled places in the evidence too blatant and glaring, to begin an 
analysis of species group history or even, in any serious sense, of 
inter-group relationship. We do not yet know enough to ask the 
right questions. 

ACKNOWLEDGMENTS 

I am indebted to William F. Pyburn for permission to describe 
the new species collected by him, and to Alan E. Leviton (California 
Academy of Sciences [CAS]), Pedro Ruiz (Instituto de Ciencias 
Naturales, Bogota [ICN]), William E. Duellman (University of 
Kansas Museum of Natural History [KU]), James Dixon (Texas A 
& M University), and Jean Lescure (Museum National de Histoire 
Naturelle, Paris [MNHP]) for permission to examine material in 
their charge. Crucial to the study was newly collected material made 
available by Stephen Ayala, Dennis Harris, and William Lamar. 
Stephen Ayala also prepared the distribution map. Laszlo Meszoly 
in preparing the figures has done me an indispensible service. 



LITERATURE CITED 

Dixon, J. R., and P. SoiNl. 1975. The reptiles of the Upper Amazon Basin. 

Iquitos. Region, Peru. I. Lizards and amphisbaenians. Contrib. Biol. Geol. 

Milwaukee Public Mus. No. 4. pp. 1-58. 
Fitch, H. S., A. F. Echelle, and A. A. Echelle. 1976. Field observations on 

rare or little known mainland anoles. Univ. Kansas Mus. Nat. Hist. Sci. Bull.. 

51: 91-128. 
Greene, H. W. 1977. Lizards of the genus Uracentron (Iguanidae) in east-central 

Colombia. Herpetologica, 33: 256-260. 
Hoogmoed, M. S. 1973. Notes on the herpetofauna of Surinam. IV. The lizards 

and amphisbaenians of Surinam. The Hague. W. Junk. 410 pp. 
Rand, A. S., and E. E. Williams. 1970. An estimation of redundancy and 

information content of anole dewlaps. Amer. Nat.. 104: 99 103. 
Scott, N. J. Jr., D. E. Wilson, C. Jones, and R. M. Andrews. 1976. The 

choice of perch dimensions by lizards of the genus Anolis (Reptilia, Lacertilia. 

Iguanidae). J. Herp.. 10: 75-84. 
Williams, E. E. 1965. South American Anolis (Sauria, Iguanidae): Two new 

species of the punctatus group. Breviora Mus. Comp. Zool. No. 233. pp. 1 15. 
1974. South American Anolis: Three new species related to Anolis 

nigrolineatus and A. dissimilis. Breviora Mus. Comp. Zool. No. 422. pp. I 15. 
Williams, E. E., and A. S. Rand. 1977. Species recognition, dewlap function, 

and faunal size. Amer. Zool., 17: 261 270. 



38 BREVIORA No. 467 



Williams, E. E., O. A. Reig, P. Kiblisky and C. Rivero-Blanco. 1970. Anolis 
jacare Boulenger, a solitary anole from the Andes of Venezuela. Breviora Mus. 
Comp. Zool. No. 353, pp. 1-15. 

Williams, E. E., and P. E. Vanzolini. 1980. Notes and biogeographic com- 
ments on anoles from Brasil. Pap. Avuls. Zool. S. Paulo, 34: 99 1 08. 

Wilson, E. O., and W. L. Brown, Jr. 1953. The subspecies concept and its 
taxonomic application. Syst. Zool., 2: 97-111. 



s*os- 



URRARY 

MAR 1 8 1985 

i iARVARD 
UNIVERSITY 

BREVIORA 

Museum of Comparative Zoology 

IS ISSN 000(3 9(iW 

Cambridge, Mass. 30June1982 Number 468 

A NEW FOREST SKINK FROM PONAPE 

A. Ross KlESTER 1 



Abstract. Emoia ponapea, new species, is described from Ponape in the Caroline 
Islands where it inhabits deep forest. It is distinguished from all other Emoia by the 
presence of 13 premaxillary teeth and a palate intermediate between the alpha and 
beta conditions. 

INTRODUCTION 

A survey of the scincid genus Emoia, undertaken after a collecting 
trip to Ponape in the Caroline Islands and other localities in 
Micronesia, showed that three specimens taken on Ponape repre- 
sent a distinct species of Emoia. In fact, as discussed below, this 
form possesses some characters which make its generic allocation 
somewhat problematic. However, an analysis of the boundaries of 
the genus Emoia is beyond the scope of this paper, and so the new 
taxon is here described as: 

Emoia ponapea, new species 
(Figs. 1-5) 

Holotype: MCZ 121041, forest 1/4 mile above Agricultural 
Station, Kolonia, Ponape Island, Eastern Caroline Islands. Col- 
lected by A. R. Kiester, 28 July 68. 

Paratypes: Eastern Caroline Islands, Ponape Island: MCZ 121042- 
43, same data as for holotype (The skull of 121042 has been 
removed and prepared. This individual was a sexually mature 



'Smithsonian Tropical Research Institute, APO Miami 34002. 



BREVIORA 



No. 468 




Figure I. Head scalation of the holotype (MCZ 121041) ot Emoia ponapea. 
Dorsal view. 






1982 



A NEW FOREST SKINK. 




g> 2 



BREVIORA 



No. 468 




Figure 3. Skull of paratype (MCZ 121042) of Emoia ponapea. Ventral view. 



1982 



A NEW FOREST SKINK 




Figure 4. Skull of paratype (MCZ 121042) of Emoia ponapea. Dorsal view. 



BREVIORA 



No. 468 




Figure 5. Skull of paratype (MCZ 121042) of Emoia ponapea. Lateral view. 



1982 A NEW FOREST SKINK 7 

female.); USNM 138985-86, Nanpil River; USNM 138991, Dolo- 
nier; USNM 139002-05, N end of ridge parallel to Tavensorola 
River; USNM 139006-07, Dolen Eirike; CAS 152222 (formerly 
USNM 138987), Nanpil River. All USNM and CAS specimens 
collected between 13 September 55 and 5 December 55 by J. T. 
Marshall, Jr. See acknowledgments for abbreviations. 

Diagnosis: A member of the genus Emoia, as it is currently and 
broadly construed (Greer, 1974), differing from all other members 
of the genus by the possession of 13 premaxillary teeth. It is 
distinguishable externally by the combination of the following 
characters: interparietal present; subdigital lamellae of the 4th toe 
38-46; midbody scale rows 30-32; middorsal scale rows 48-54 
counting from the nuchals to the anterior insertion of the hind limb 
and 52-60 counting to the point directly above the vent; body form 
small (snout-vent length less than 50 mm) and distinctly slender; 
head narrow and relatively pointed; coloration without metallic 
blues or greens. 

DESCRIPTION OF HOLOTYPE 

The range of paratypical variation is given in parentheses after the 
description of the holotype character. 

General Appearance: Body small with a snout-vent length of 46 
mm (21 mm to 49 mm for the paratypes), form distinctly slender and 
gracile. Tail long, over 1.5 times the snout- vent length. Limbs well 
developed, overlapping easily when adpressed to the body; hindlimb 
length 23 mm, forelimb length 13 mm. Five digits on each foot. 

Head and Head Scalation (Figs. 1-2): Head narrow and some- 
what depressed. Rostral twice .as long as deep; supranasals 
separated, much wider anteriorly than posteriorly; prefrontals 
moderately to narrowly separated by frontonasal and frontal; 
supraoculars 4; frontoparietals fused; interparietal single, distinct, 
and large; one pair of parietals in contact behind interparietals; two 
pairs of temporals corresponding to the primary and secondary 
temporals of Taylor (1935) with only the secondary temporals 
placed toward the top of the head; supraciliaries 9-9; loreals two; 
single pair of nuchals; 6th supralabial subocular; nasal divided 
below nostril; eyelid window undivided; postmental contacts first 
two infralabials on each side. 



8 BREVIORA No. 468 

Ear: Tympanum sunk below level of epidermis. A single larger 
anterior lobule with two smaller ones below (a single smaller one on 
one side of one of the paratypes). Vertical (longest) diameter of ear 
opening 1.1 mm. 

Body Scalation: Scales smooth. Midbody scale rows 30 (30-32). 
Middorsal scale rows 48 (48 -54) counting from the first scale 
posterior to the nuchals to the anterior insertion of the hind leg and 
53 (52-60) counting posterior to a point directly above the vent. 
Subdigital lamellae on the 4th toe of the hind foot rounded with 42 
on both sides (38-46). Preanals not enlarged. 

Coloration: Dorsum with a central tan stripe with irregular edges 
extending from the snout to the end of the tail. A small number of 
black spots on the stripe on the head (some of the paratypes show 
black spotting on the stripe on the back as well). A poorly defined 
laterodorsal stripe on either side of this beginning behind the eye 
and extending one third of the way down the tail. These stripes are 
formed by two irregular rows of black dashes on the tan color. Sides 
reddish brown becoming spotted over cream color toward the 
venter. Limbs reticulated brownish red over cream above. Chin, 
throat, venter, and undersides of limbs and tail all immaculate 
cream. 

DESCRIPTION OF SKULL 

Skull MCZ 121042, paratype (Figs. 3-5): General appearance 
very delicate and somewhat narrow (as compared, say, to a similarly 
sized E. cyanura). Premaxillary teeth 13. Secondary palate moder- 
ately well developed. Palatines meet extensively along the midline 
and send pointed projections posteriorly between the anterior 
portions of the pterygoids. One of the projections is folded slightly 
under the other. Pterygoids intermediate in condition between the 
alpha and beta configurations of Greer and Parker (1968) and Greer 
(1974), with the anterior portion showing distinct expansion toward 
the midline without achieving a strongly recurved process. 

Lower Jaw: The Meckelian canal is as the second of the two 
conditions described by Greer (1974) for the Leiolopisma group 
of lygosomine genera: canal closed with no suture evident. 



1982 A NEW FOREST SKINK 9 

DISTRIBUTION AND ECOLOGICAL OBSERVATIONS 

At present this species is known only from the island of Ponape in 
the Caroline Islands. Ponape is also known as Ascension Island in 
some of the older literature. 

All three MCZ specimens were collected in the late morning on 
the floor of a mature forest in an area where sun flecks occurred. No 
other emoias were seen in this deep forest habitat. E. cyanura, E. 
caeruleocauda, and E. boetigeri were found only in more open areas 
towards the coast. Ecological notes by J. T. Marshall, Jr. ac- 
companying the other specimens indicate that they were also col- 
lected on the floor of the forest. This species bears a similar 
ecological relationship to its congeners as E. parkeri does to the 
other emoias in the Fiji Islands (Brown et al., 1980). 

Two of the USNM specimens are hatchlings. Marshall notes that 
the eggs were collected on 27 October 55 in a rotted palm stump in 
the rain forest and that they hatched on 6 November 55. A clutch 
size of two is typical for most species in this genus (Greer, 1968). 

DISCUSSION 

The current state of confusion in the genus Emoia prevents any 
accurate assessment of the relationships of Emoia ponapea. A 
thorough revision of the genus would be necessary to understand 
the relationships of any but the most closely related of its species. 
Lacking this, a comparison with the smaller emoias found in 
Micronesia may be useful. E. cyanura and E. caeruleocauda, both 
also found on Ponape, have true beta palates. This character is 
considered derived in the genus (Greer, 1974), as is the very high 
subdigital lamellae count of cyanura itself. Thus, since E. ponapea 
has a palate intermediate between the alpha and beta conditions, it 
is unlikely that it is derived from any of the members of the cyanura 
group directly. Within Micronesia a possible candidate for a related 
species could be E. mivarti. However, there are notable differences 
from this species as well. E mivarti has the interparietal fused with 
the frontoparietal and is a much stouter animal. None of these 
species seems closely related to E. ponapea; a search for its relatives 
will have to extend outside of Micronesia. 



10 breviora No. 468 

The most unusual character of E. ponapea is the possession of 13 
premaxillary teeth. All other members of the genus Emoia have 1 1, 
as do all of the genera regarded as related to Emoia (the members of 
Group II of Greer, 1974). The only other leiolopismid genus with 13 
premaxillaries is Carlia, which is clearly unrelated on other grounds 
(Greer, 1974). For the genus Emoia this character state must be 
derived. This character and the intermediate condition of the palate 
argue that E. ponapea is rather different from the rest of the genus. 
Just how different it really is will have to be determined by future 
work. 

ACKNOWLEDGMENTS 

I wish to thank W. C. Brown, Allen Greer, the late T Preston 
Webster, and Ernest E. Williams for much help and discussion on 
matters related to this paper. The illustrations are by Lazlo Meszoly. 
This work was supported by the Evolutionary Biology Committee 
of Harvard University, The Society of Sigma Xi, and Ernest E. 
Williams. For permission to examine specimens in their care, I 
thank A. G. C. Grandison, British Museum (Natural History); H. 
Marx, Field Museum of Natural History; A. Leviton, California 
Academy of Sciences (CAS); Ernest E. Williams, Museum of 
Comparative Zoology, Harvard University (MCZ); R. Stebbins, 
Museum of Vertebrate Zoology, University of California at Berk- 
eley; and the late J. Peters and W. R. Heyer, United States National 
Museum (USNM). 

LITERATURE CITED 

Brown, W. C, J. C. Pernetta, and D. Waiting. 1980. A new lizard of the 

genus Emoia (Scincidae) from the Fiji Islands. Proc. Biol. Soc. Wash., 93: 

350-356. 
Greer, A. E. 1968. Clutch size in the genus Emoia. Copeia, 1968: 417-418. 
1974. The generic relationships of the scincid lizard genus Leiolopisma 

and its relatives. Austr. Jour. Zool. Suppl. Ser. No. 31, pp. 1 67. 
Greer, A. E., and F. Parker. 1968. Geomyersia glabra, a new genus and species 

of scincid lizard from Bougainville, Solomon Islands, with comments on the 

relationships of some lygosomine genera. Breviora Mus. Comp. Zool. No. 302, 

pp. 1-17. 
Taylor, E. H. 1935. A taxonomic study of the cosmopolitan scincoid lizards of 

the genus Eumeces, with an account of the distribution and relationships of 

its species. Kans. Univ. Sci. Bull., 23: 1-643. 
Webster, T. P. 1969. Aspects of the morphological and ecological variation in 

the cyanura group of the lizard genus Emoia (Sauria: Scincidae) in the Solomon 

Islands. Honors Thesis, Harvard University. 



MUS. COMP. ZOOL 
LfBRARY 



BRE S'O R A 

UNIVERSITY 

seum of Comparative Zoology 



.11 



US ISSN 0006-9698 



Cambridge, Mass. 30 June 1982 Number 469 

CATALOG OF THE PRIMARY TYPES OF 

BOMBYLIIDAE (DIPTERA) 

IN THE ENTOMOLOGICAL COLLECTIONS OF 

THE MUSEUM OF COMPARATIVE ZOOLOGY, 

WITH DESIGNATIONS OF LECTOTYPES 

Neal L. Evenhuis 1 

Abstract. One hundred and nine holotypes, lectotypes. and syntypes of Bomby- 
liidae deposited in the entomological collections of the Museum of Comparathe 
Zoology are listed. Lectotypes are designated for 27 taxa based on material in MCZ. 

INTRODUCTION 

This catalog lists the primary types of 109 taxa of Bombyliidae 
located in the entomological collections of the Museum of Compara- 
tive Zoology, Harvard University, Cambridge (MCZ). Fifty-seven 
of these taxa are represented by holotypes, 1 7 by syntype series, and 
35 by lectotypes. All but one of the types listed here are deposited in 
the general collection of the MCZ. The unique male holotype of 
Epibates harrisi Osten Sacken is deposited in the Thaddeus W. 
Harris collection, which is maintained separately from the general 
collection. Twenty-seven new lectotype designations are recorded 
herein including two by Jack C. Hall, University of California, Riv- 
erside. Of the remaining 25 new lectotypes, three are designated in 
Hall and Evenhuis (1981). The format used here follows that of 
Radovsky et al. (1976) with modifications. Information presented in 
brackets [ ] is data additional to or corrected from data recorded in 
the original description. The present combination of each taxon 



'Department of Entomology, Bishop Museum. P.O. Box I9000-A, Honolulu. 
Hawaii 96819. 



2 BREVIORA No. 469 

listed, if it differs from the original combination, is also included 
after the remarks section under each taxon concerned. 

The majority of species recorded here were described by Loew in 
his "Centuries," Osten Sacken in his "Western Diptera," and C.W. 
Johnson in various publications. Most of the types or type series in 
the collections carry a red label with a type register number placed 
there by Nathan Banks when he curated the collection. Recent addi- 
tions to the type collection carry a similar label with a type number 
placed there by various curatorial technicians since Banks. Banks' 
type labeling has posed problems because he sometimes failed to 
label all of the syntypes in a syntype series or labeled more than the 
original description of that particular taxon was based upon (this is 
especially true for many of Loew's species). In compiling this 
catalog, all original descriptions were checked to make sure the 
correct number of syntypes were located and labeled. Discrepancies 
in the number of specimens found versus the number of specimens 
upon which that species was based are noted in the remarks section 
under each taxon concerned. In some cases, syntypes of certain taxa 
were found to be located in other museums. These museums include 
the United States National Museum of Natural History, Washing- 
ton, D.C. (USNM), the American Museum of Natural History, New 
York (AMNH), the Snow Collection at the University of Kansas, 
Lawrence (SNOW) and the Zoologisches Museum of the Museum 
fur Naturkunde der Hurnboldt-Universitat, Berlin (ZMHU). This 
information is supplied in the remarks section under each taxon 
concerned. 

Some syntypes of the bombyliid taxa described by Osten Sacken 
in "Biologia Centrali-Americana" are deposited in the MCZ. Lecto- 
types for most of these species were designated by Painter and Paint- 
er (1962) from material in the British Museum (Natural History), 
where in most cases the majority of the syntypes of these particular 
taxa are housed. Specimens of these species represented in the MCZ 
carry labels signifying that the lectotype for that species is in the 
BMNH. 

SPECIES-GROUP INVENTORY 

adusta Loew, Anthrax SYNTYPES 2 9 12672 

1869 Berliner Entomol. Z. 13: 26 pin 

CUBA: [no further data] Gundlach 



1981 BOMBYLIIDAE TYPES IN MCZ 3 

The specimens in the MCZ are left here as syntypes pending revision of the genus. 
Present name: Villa adust a (Loew) 

agassizi Loew, Exoprosopa HOLOTYPE 9 12635 

1869 Berliner Entomol. Z. 13: 16 pin 

UNITED STATES: California: [no further data] Agassiz 

albicapillus Loew, Bombylius LECTOTYPE 9 12686 

1872 Berliner Entomol. Z. 16: 78-79 pin 

UNITED STATES: [California]: San Francisco [no date] 

H. Edwards 
Lectotype designated by Evenhuis in Hall and Evenhuis (1980: 6). 

albidipennis Loew, Geron HOLOTYPE S 12713 

1869 Berliner Entomol. Z. 13: 174 pin 

UNITED STATES: California: [no further data] Agassiz 

aldrichi Johnson, Phthiria SYNTYPES 2 #, 1 $ 7555 

1903 Psyche 10: 184 pin 

UNITED STATES: Idaho: Caldwell 24.IV. 1901 C.W. John- 
son Collection 

These specimens are left as syntypes pending revision of the genus. 
Present name: Phthiria (Poecilognathus) aldrichi Johnson 

alpha Osten Sacken, Anthrax LECTOTYPE 9 1 2660 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 239-40 
UNITED STATES: Wyoming: Cheyenne 21. VIII. O. Sacken 
Four (three male, one female) of the five specimens on which Osten Sacken based 
his description of alpha have been located at the MCZ. The fifth syntype is deposited 
in AMNH. The best preserved of the MCZ material, the female, is here designated 
lectotype. 

Present name: Poecilanthrax alpha (Osten Sacken) 

amabilis Osten Sacken, Ploas LECTOTYPE $ 12703 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 261-62 
UNITED STATES: California: Yosemite 5.VI. O. Sacken 

Osten Sacken based his description of amabilis on two female and one male. The 
male is here designated lectotype. 

Present name: Conophorus amabilis (Osten Sacken) 



4 BREVIORA No. 469 

arenicola Johnson, Anthrax lateralis LECTOTYPE $ 7548 

1908 Psyche 15: 15 pin 
UNITED STATES: New Jersey: Clementon 16.V. 1917 

C.W. Johnson Collection 

Lectotype male and allolectotype female designated herein by J.C. Hall. 

Present name: Villa lateralis arenicola (Johnson) 

arizonicus Banks, Systropus SYNTYPES 2 $ 13484 

1909 Entomol. News 20: 18 pin 
UNITED STATES: Arizona: Palmerlee [no date] N. Banks 

Collection 

These specimens are left as syntypes pending revision of the genus. 

atratula Loew, Ploas HOLOTYPE $ 12698 

1872 Berliner Entomol. Z. 16: 79-80 pin 

UNITED STATES: California: [no further data] H. Edwards 
Present name: Conophorus atratulus (Loew) 

atriceps Loew, Bombylius LECTOTYPE $ 12688 

1863 Berliner Entomol. Z. 7: 301 pin 

UNITED STATES: Florida: [no further data] O. Sacken 

Lectotype designated by Evenhuis in Hall and Evenhuis (1980: 15). 

aurifer Osten Sacken, Bombylius LECTOTYPE? 12691 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 249-50 
UNITED STATES: California: Webber Lake 26. VII. 

O. Sacken 

Lectotype designated by Evenhuis in Hall and Evenhuis (1980: 18). 

banksi Johnson, Dipalta HOLOTYPE $ 13485 

1921 Occas. Pap. Boston Soc. Nat. pin 

Hist. 5: 12-13 
UNITED STATES: Virginia: Great Falls 12.IX. Collection 

N. Banks 

barbatus Osten Sacken, Anastoechus LECTOTYPE S 12719 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 252-53 
UNITED STATES: Wyoming: Cheyenne 21. VIII. O. Sacken 

Lectotype designated by Hall in Hall and Evenhuis (1981: 153-54). 



1981 BOMBYLIIDAE TYPES IN MCZ 5 

bifurca Loew, Exoprosopa HOLOTYPE $ 12636 

1869 Berliner Entomol. Z. 13: 15-16 pin 

UNITED STATES: California: [no further data] Agassiz 

Loew's description of bifurca was apparently based on a single specimen because 

there is no range of measurements recorded. There are two female specimens in the 

MCZ labeled as types. The one in the best condition is considered to be the holotype. It 

fits Loew's description and measurements and carries a label "bifurca m." in Loew's 

handwriting. The other specimen is denuded, smaller than the holotype, and carries a 

label "bifurca" in Banks' handwriting. The latter specimen probably does not belong to 

the original type series. 

bigradata Loew, Anthrax HOLOTYPE [$] 12655 

1869 Berliner Entomol. Z. 13: 23-24 pin 

CUBA: [no further data] Gundlach 

Original description incorrectly cites sex of type as "<3". 

borealis Johnson, Phthiria HOLOTYPE $ 27054 

\9\Q Psyche 11:229 pin 

UNITED STATES: Maine: Fort Kent 7.VIII. 1910 
C.W. Johnson Collection 

Present name: Phthiria (Poecilognathus) borealis Johnson 

brevicornis Loew, Sparnopolius SYNTYPES 2 9 12723 

1872 Berliner Entomol. Z. 16: 79 pin 

UNITED STATES: Texas: [no further data] Belfrage 

Though Loew gives no range of measurements, two specimens are labeled as types 
in the MCZ. They are left here as syntypes until their type status can be clarified. 

cachinnans Osten Sacken, Bombylius LECTOTYPE S 12693 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 250 
UNITED STATES: California: Sonoma County 2.IV.-9.V. 
R. Osten Sacken 

Male and female on same pin. Lectotype designated by Evenhuis in Hall and 
Evenhuis(1980:29). 

calvus Loew, Geron SYNTYPES 2 $ 1 27 1 1 

1863 Berliner Entomol. Z. 7: 303-04 pin 

UNITED STATES: New York [no further data] 
Loew's description does not include a range of measurements though two speci- 
mens are labeled as types in the MCZ. Until this discrepancy can be clarified, they are 
left as syntypes. 



6 BREVIORA No. 469 

candidulus Loew, Systoechus LECTOTYPE $ 12716 

1863 Berliner Entomol. Z. 7: 302 pin 

UNITED STATES: Wisconsin: [no further data] O. Sacken 

Loew's description cites a range of measurements though only one male was 
located in the MCZ. The male is here designated lectotype. 

canuta Hall, Thevenemyia HOLOTYPE S 31339 

1969 Univ. Calif. Publ. Entomol. pin 

56: 28-29 Fig. 8 
UNITED STATES: Arizona: Reef 10.X. C.W. Johnson 
Collection 

capito Osten Sacken, Pantarbes LECTOTYPE S 12720 

1877 Bull. U.S. Geo!. Geogr. Surv. pin 

Terr. 3: 256 
UNITED STATES: California: Sonoma County 2.IV.-9.V. 

R. Osten Sacken 
Lectotype male designated herein by J.C. Hall. 

cervinus Loew, Aphoebantus HOLOTYPE? 12684 

1872 Berliner Entomol. Z. 16: 76-77 pin 

UNITED STATES: Texas: [no further data] Belfrage 

ceyx Loew, Anthrax LECTOTYPE S 12664 

1869 Berliner Entomol. Z. 13: 19 pin 

UNITED STATES: Virginia [no further data] 

Loew's description cites a range of measurements though only one male was found 
in the MCZ. It is here designated lectotype. 

Present name: Poecilanthrax demogorgon (Walker) 

contigua Loew, Argyromoeba HOLOTYPE $ 12679 

1869 Berliner Entomol. Z. 13: 30-31 pin 

UNITED STATES: Virginia: [no further data] Loew 
Present name: Anthrax argyropygus argyropygus 
Wiedemann 

coquilletti Johnson, Phthiria SYNTYPES 2 $, 4 $ 7554 

1902 Can. Entomol. 34: 240-41 pin 

UNITED STATES: New Jersey: 1 #, 4 9, Riverton 4. VII. 1891; 
1 S, Jamesburg4.VII.1891 C.W. Johnson Collection 



1981 BOMBYLIIDAE TYPES IN MCZ 

Three syntypes are in USNM; one syntype is in SNOW; these specimens are left as 
syntypes pending generic revision. 

Present name: Phthiria (Poecilognathus) coquilletti Johnson 

cubana Loew, Exoprosopa SYNTYPES I $, 1 9 12641 

1869 Berliner Entomol. Z. 13: 14-15 pin 

CUBA: [no further data] Gundlach 
Two syntypes are also located in ZMHU. These specimens are left as syntypes 
pending generic revision. 

curta Loew, Anthrax HOLOTYPE S 12654 

1869 Berliner Entomol. Z. 13: 22 pin 

UNITED STATES: California: [no further data] Agassiz 
Present name: Hemipenthes curta (Loew) 

cyanoceps Johnson, Phthiria HOLOTYPE S 27053 

1903 Psvc/^ 10: 184 pin 

UNITED STATES: Massachusetts: Cohasset 8.IX. 

C.W. Johnson Collection 
Present name: Phthiria (Poecilognathus) cyanoceps Johnson 

decora Loew, Exoprosopa HOLOTYPE $ 12630 

1869 Berliner Entomol. Z. 13: 13 pin 

UNITED STATES: Wisconsin: [no further data] Kennicot 

delila Loew, Argyromoeba HOLOTYPE $ 12674 

1869 Berliner Entomol. Z. 13: 28 pin 

UNITED STATES: California: [no further data] Agassiz 
Present name: Anthrax delila (Loew) 

diagonalis Loew, Anthrax HOLOTYPE? 12656 

1869 Berliner Entomol. Z. 13: 21 pin 

UNITED STATES: California: [no further data] Agassiz 
Present name: Paravilla diagonalis (Loew) 

dodrans Osten Sacken, Exoprosopa SYNTYPES 2 $ 12634 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 234-35 
UNITED STATES: [Colorado]: Colorado Springs [no date] 
O. Sacken 

These specimens are left here as syntypes pending generic revision. 



8 BREVIORA No. 469 

dorcadion Osten Sacken, Exoprosopa LECTOTYPE $ 12631 

1877 Bull. U.S. Geo I. Geogr. Surv. pin 

Terr. 3:231-33 
UNITED STATES: Maine [no further data] 

Osten Sacken's original description of dorcadion is based on syntypes from 
"Summit Station, Central Pacific Railroad, California (July 17); Webber Lake, 
Sierra Nevada, California (July 26); Shasta district, California (H. Edwards); 
Washington Territory (the same); Georgetown, Colorado (August 12); Twin Lake 
Creek, Colorado (W.L. Carpenter); White Mountains, New Hampshire (H.K. Mor- 
rison); Maine — 2 specimens from Denver, Colorado (Uhler)." The MCZ has all but 
the New Hampshire and Shasta district specimens. The Summit Station specimen 
also carries the label "Sierra Nevada." All the specimens are females except for the 
one male from Denver, Colorado. The female specimen from Maine is in the best 
condition and is here designated lectotype. Three syntypes are also deposited in 
AMNH. 

doris Osten Sacken, Exoprosopa SYNTYPE $ 12632 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 235-36 
UNITED STATES: Nevada: Humboldt Station 29.VII. 
O. Sacken 

One syntype (headless) is deposited in AMNH. These specimens are left as syn- 
types pending generic revision. 

edwardsii Loew, AHocotus HOLOTYPE [S] 12704 

1872 Berliner Entomol. Z. 16: 81-82 pin 

UNITED STATES: California: [no further data] H. Edwards 

Original description incorrectly states sex as "9". 
Present name: Paracosmus edwardsii (Loew) 

egerminans Loew, Phthiria LECTOTYPE [9] 12708 

1872 Berliner Entomol. Z. 16: 80-81 pin 

UNITED STATES: California: [no further data] H. Edwards 

Original description incorrectly cites sex of type as "(5". Loew's description cites a 
range of measurements though only one female was found in the MCZ. It is here 
designated lectotype. 

eremita Osten Sacken, Exoprosopa HOLOTYPE S 12637 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 236 
UNITED STATES: California: Shasta District [15. VII.] 
H. Edwards 



1981 BOMBYLIIDAE TYPES IN MCZ 9 

euplanes Loew, Argyromoeba HOLOTYPE 9 12681 

1869 Berliner Entomol. Z. 13: 30 pin 

CUBA: [no further data] Gundlach 
Present name: Anthrax euplanes (Loew) 

fenestrata Osten Sacken, Ploas LECTOTYPE $ 12702 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 260-61 
UNITED STATES: California: San Raphael 12.IV. O. Sacken 

Osten Sacken based his description of fenestrata on two males and one female. Of 
these syntypes, the best preserved (the female) is here designated lectotype. 
Present name: C onophor us fene stratus (Osten Sacken) 

flaviceps Loew, Anthrax LECTOTYPE $ 12663 

1869 Berliner Entomol. Z. 13: 18-19 pin 

MEXICO: Tamaulipas [no further data] 

Of the two female syntypes of this species in the MCZ, the best preserved is here 
designated lectotype. 

Present name: Poecilanthrax flaviceps (Loew) 

fraudulently Johnson, Bombylius HOLOTYPE S 7552 

1907 Psyche 14: 99-100 pin 

UNITED STATES: Massachusetts: Provincetown 27.VI.1904 
C.W. Johnson Collection 

fuliginosa Loew, Anthrax HOLOTYPE 3 12661 

1869 Berliner Entomol. Z. 13: 20 pin 

UNITED STATES: California [no further data] 

Specimen rubbed. 
Present name: Poecilanthrax fuliginosus (Loew) 

fur Osten Sacken, Argyromoeba LECTOTYPE $ 12678 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 244-45 
UNITED STATES: Texas: Dallas [no date] Boll 

Osten Sacken based his description of fur on one male and two female syntypes. 
The best preserved of these (the male) is here designated lectotype. 
Present name: Anthrax limatulus fur (Osten Sacken) 

gazophylax Loew, Exoprosopa HOLOTYPE $ 12629 



10 BREVIORA No. 469 

1869 Berliner Entomol. Z. 13: 12-13 pin 

UNITED STATES: California: [no further data] Agassiz 
Head missing. 
Present name: Ligyra gazophylax (Loew) 

gibbus Loew, Lordotus HOLOTYPE $ 12724 

1863 Berliner Entomol. Z. 7: 303 pin 

MEXICO: [Tamaulipas]: Matamoros [no further data] 

The label data cites the locality as "Matamoras." 

haemorrhoicus Loew, Bombylius HOLOTYPE S 12696 

1863 Berliner Entomol. Z. 7: 300 pin 

CUBA: [no further data] [Gundlach] 

Original description cites collector as "Riehl," label data cites "Gundlach." 
Present name: Heterostylum haemorrhoicum (Loew) 

harrisi Osten Sacken, Epibates HOLOTYPE S 26400 

1877 Bull. U.S. Geo!. Geogr. Surv. pin 

Terr. 3: 273 
[No locality label] H. Gray 
Hall (1969: 38) proposed a lectotype male (MCZ Type No. 31340) and allolecto- 
type female for harrisi under the assumption that the original type specimen was 
lost. 1 The type male is, in fact, present in the T.W. Harris Collection [in MCZ] as 
described by Osten Sacken. 

Present name: Thevenemyia harrisi (Osten Sacken) 

humilis Osten Sacken, Phthiria HOLOTYPE S 12710 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 264 
UNITED STATES: California: Sonoma County 4. VII. 

O. Sacken 

incanus Johnson, Bombylius HOLOTYPE 3 7553 

1907 Psyche 14: 97-98 pin 

UNITED STATES: Massachusetts: Provincetown 2. VI. 1904 

C.W. Johnson Collection 
Present name: Bombylius (Zephyrectes) incanus Johnson 



'Though neotypes should be proposed for lost or destroyed types. Hall mistakingly 
proposed a lectotype and allolectotype which, therefore, have no validity no- 
menclatorially. 



1981 BOMBYLIIDAE TYPES IN MCZ 11 

johnsoni Painter, Geron HOLOTYPE <$ 27266 

1932 Trans. Am. Entomol. Soc. pin 

58: 155 
UNITED STATES: Massachusetts: Horseneck Beach 30. 
VII. 1913 C.W. Johnson Collection 

lancifer Osten Sacken, Bombylius LECTOTYPE S 12692 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 251 
UNITED STATES: California: [no further data] Sacken 

Lectotype male designated by Evenhuis in Hall and Evenhuis (1980: 57). 

lepidotoides Johnson, Chrysanthrax HOLOTYPE S 7550 

1919 Psyche 26: 12 pin 

UNITED STATES: New Jersey: Iona 16. VI. 1902 
C.W. Johnson Collection 

limbata Loew, Ploas LECTOTYPE $ 12700 

1869 Berliner Entomol. Z. 13: 31 pin 

UNITED STATES: New Mexico [no further data] 

Specimen badly rubbed; right wing glued to point below specimen; head, left wing, 
left fore- and hind- and right mid- and hindlegs beyond coxae missing. Loew's 
description cites a range of measurements though only one female was found in the 
MCZ. It is here designated lectotype. 

Present name: Conophorus limbatus (Loew) 

luctifer Osten Sacken, Epibates HOLOTYPE S 12728 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 271-72 
[CANADA: British Columbia]: Vancouver Island [no date] 

G.R. Crotch 
Present name: Thevenemyia luctifera (Osten Sacken) 

macer Loew, Systropus LECTOTYPE? 12726 

1863 Berliner Entomol. Z. 7: 305 pin 

UNITED STATES: Pennsylvania [no further data] 

Of the three syntypes of macer (1 male, 2 females) in the MCZ, the best preserved 
of these (a female) is here designated lectotype. 

macropterus Loew, Geron HOLOTYPE 3 12712 



12 BREVIORA No. 469 

1869 Berliner Entomol. Z. pin 

13: 172-73 
UNITED STATES: New York: Geneseo [no further data] 
Head and thorax damaged; abdomen beyond third segment missing. 
Present name: Geron (Empidigeron) calvus Loew 

magnus Osten Sacken, Epibates HOLOTYPE $ 12731 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 272-73 
[CANADA: British Columbia]: Vancouver Island [no date] 

G.R. Crotch 
Present name: Thevenemyia magna (Osten Sacken) 

marginatus Osten Sacken, Epibates HOLOTYPE $ 12730 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 272 
UNITED STATES: California: San Francisco [no date] 

H. Edwards 
Present name: Thevenemyia marginata (Osten Sacken) 

metopium Osten Sacken, Bombylius HOLOTYPE $ 12690 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 249 
UNITED STATES: California: [San Francisco] 19.IV 

O. Sacken 

modestus Loew, Leptochilus SYNTYPES 1 (5, 1 $ 12685 

1872 Berliner Entomol. Z. 16: 77-78 pin 

UNITED STATES: Texas: [no further data] Belfrage 

The specimens are left here as syntypes pending revision of the genus. 
Present name: Epacmus modestus (Loew) 

molitor Loew, Anthrax HOLOTYPE? 12652 

1869 Berliner Entomol. Z. 13: 26-27 pin 

UNITED STATES: California: [no further data] Agassiz 
Head glued to thorax. 
Present name: Villa molitor (Loew) 

mucorea Loew, Anthrax HOLOTYPE 9 12651 



1981 BOMBYLIIDAE TYPES IN MCZ 13 

1869 Berliner Entomol Z. 13: 27 pin 

UNITED STATES: Nebraska: [no further data] [F.V. Hayden] 

Collector recorded in Osten Sacken ( 1903: 95). 
Present name: Villa mucorea (Loew) 

muricatus Osten Sacken, Epibates HOLOTYPE S 12729 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 272 
UNITED STATES: California: Sierra Nevada [no date] 

H. Edwards 
Present name: Thevenemyia muricata (Osten Sacken) 

mus Osten Sacken, Triodites LECTOTYPE $ 12683 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 246-47 
UNITED STATES: Utah: Salt Lake City I. VIII. O. Sacken 
Of the three syntypes of mus (1 male, 2 females) located in the MCZ, the best 
preserved of these (the male) is here selected as lectotype. 

Present name: Aphoebanlus mus mus (Osten Sacken) 

nigricauda Loew, Anthrax HOLOTYPE? 12650 

1869 Berliner Entomol. Z. 13: 24 pin 

UNITED STATES: Massachusetts: [no further data] Scudder 
Head missing. 
Present name: Villa fulviana nigricauda (Loew) 

nigripennis Loew, Ploas LECTOTYPE 9 12697 

1872 Berliner Entomol. Z. 16: 80 pin 

UNITED STATES: California: [no further data] H. Edwards 

Loew's description cites a range of measurements though only one female was 
found in the MCZ. It is here designated lectotype. 

Present name: Conophorus nigripennis (Loew) 

notata Loew, Phthiria HOLOTYPE 9 12706 

1863 Berliner Entomol. Z. 7: 113-14 pin 

UNITED STATES: California: [no further data] Agassiz 
Present name: Phthiria (Poecilognathus) loewi Painter 

nubifera Loew, Exoprosopa LECTOTYPE 3 12639 

1869 Berliner Entomol. Z. 13: 16 pin 



14 BREVIORA No. 469 

CUBA: [no further data] [Poey] 

Original description cites collector as "Gundlach ," label data cites "Poey." Loew's 
description cites a range of measurements though only one male was found in the 
MCZ. The male is here designated lectotype. 

obesula Loew, Ploas SYNTYPES 3 3 12699 

1872 Berliner Entomol. Z. 16: 80 pin 

UNITED STATES: California: [no further data] H. Edwards 

Loew's original description of obesula was based on an unstated number of males 
and females. Only three males were found in the MCZ. They are left here as syntypes 
until the genus is revised. 

Present name: Conophorus obesulus (Loew) 

obsoleta Loew, Argyromoeba HOLOTYPE [$] 12675 

1869 Berliner Entomol. Z. 13: 29 pin 

UNITED STATES: Missouri [no further data] 

Original description incorrectly states sex of type as "<3". 
Present name: Anthrax limatulus limatulus Say 

occidentalis Johnson, Spogostylum SYNTYPES 2 $ 7547 

1913 Bull. Am. Mm. Nat. Hist. pin 

32:56 
UNITED STATES: 1 9, Colorado: Denver 5.VIII.1897; 1 $, 
Washington: Seattle [no further data] C.W. Johnson 
Collection 
These specimens are left here as syntypes pending generic revision. 
Present name: Anthrax analis Say 

oreas Osten Sacken, Systoechus LECTOTYPE S 12718 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 254 
UNITED STATES: California: Webber Lake 22. VII. 
O. Sacken 

Lectotype designated by Hall in Hall and Evenhuis (1981: 147). 

palliata Loew, Anthrax HOLOTYPE? 12657 

1869 Berliner Entomol. Z. 13: 20-21 pin 

UNITED STATES: Illinois: [no further data] Osten Sacken 
Present name: Paravilla palliata (Loew) 



1981 BOMBYLIIDAE TYPES IN MCZ 15 

parva Loew, Exoprosopa LECTOTYPE S 12640 

1869 Berliner Entomol Z. 13: 17 pin 

CUBA: [no further data] Gundlach 

Though Loew's description cites a range of measurements, only one male was 
located in the MCZ. It is here designated lectotype. 

Present name: Neodiplocampta (Neodiplocampta) parva (Loew) 

parvicornis Loew, Anthrax HOLOTYPE S 12658 

1869 Berliner Entomol. Z. 13: 23 pin 

UNITED STATES: Illinois: [Chicago] [no date] Osten Sacken 
Present name: Rhynchanthrax parvicornis (Loew) 

pauper Loew, Argyromoeba HOLOTYPE? 12677 

1869 Berliner Entomol. Z. 13: 29-30 pin 

UNITED STATES: Illinois [no further data] 
Present name: Anthrax pauper (Loew) 

pertusa Loew, Anthrax HOLOTYPE $ 12659 

1869 Berliner Entomol. Z. 13: 18 pin 

UNITED STATES: New Mexico: [10.IV.] [no further data] 
Present name: Thyridanthrax pertusa (Loew) 

planus Osten Sacken, Lordotus HOLOTYPE 3 12725 

1877 Bull. U.S. Geol. Geogr. pin 

Surv. Terr. 3: 258-59 
UNITED STATES: California: Marin County [no date] 
O. Sacken 

proboscidea Loew, Anthrax HOLOTYPE # 12653 

1869 Berliner Entomol. Z. 13: 17-18 pin 

MEXICO: Sonora: [no further data] Schott 
Present name: Lepidanthrax proboscideus (Loew) 

pulchellus Loew, Bombylius LECTOTYPE S 12687 

1863 Berliner Entomol. Z. 7: 300 pin 

UNITED STATES: Illinois [no further data] 
Lectotype designated by Evenhuis in Hall and Evenhuis ( 1980:79). 

quinquenotata Johnson, Phthiria HOLOTYPE $ 7556 



16 BREVIORA No. 469 

1903 Psyche 10: 185 pin 

UNITED STATES: Colorado: Grand Junction 25.V.1900 

C.W. Johnson Collection 
Present name: Oligodranes quinquenotata (Johnson) 

ravus Loew, Bombylius HOLOTYPE $ 12694 

1863 Berliner Entomol. Z. 7: 301-02 pin 

MEXICO [no further data] 
Present name: Bombylius (Zephyrectes) ravus Loew 

rufula Osten Sacken, Ploas LECTOTYPE S 12701 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 261 
UNITED STATES: California: San Geronimo 19.IV. 

O. Sacken 
Present name: Conophorus rufulus (Osten Sacken) 

sagata Loew, Anthrax LECTOTYPE $ 12671 

1869 Berliner Entomol. Z. 13: 21-22 pin 

MEXICO: [Tamaulipas]: Matamoros [no further data] 

Head missing; label data cites locality as "Matamoras". Loew's description cites a 
range of measurements, though only one male was found in the MCZ. It is desig- 
nated here as lectotype. 

Present name: Hemipenthes sagata (Loew) 

scolopax Osten Sacken, Phthiria SYNTYPES [2(5, 1$] 12789 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 263-64 
UNITED STATES: Colorado: Manitou 17-18.VIII. O. Sacken 
The original description of scolopax was based on one male and three females. 
Two male and one female syntypes are located in the MCZ. These specimens are left 
here as syntypes pending generic revision. 

Present name: Phthiria (Poecilognathus) scolopax Osten Sacken 

scrobiculata Loew, Anthrax LECTOTYPE 3 12648 

1869 Berliner Entomol. Z. 13: 24-25 pin 

UNITED STATES: Illinois [no further data] 
Of the two male syntypes of scrobiculata in the MCZ, the best of these is here 

designated lectotype. 

Present name: Villa scrobiculata (Loew) 



1981 BOMBYLIIDAE TYPES IN MCZ 17 

seminigra Loew, Hemipenthes HOLOTYPE [#] 12673 

1869 Berliner Entomol. Z. 13: 27-28 pin 

CANADA: Saskatchewan: [no further data] Kennicot 
Head missing; original description incorrectly cites sex of type as "9" 
Present name: Hemipenthes seminigra Loew 

semirufus Loew, Bombylius LECTOTYPE S 12695 

1872 Berliner Entomol. Z. 16: 78 pin 

HAITI: [Geremie]: [no date] P.R. Uhler 

The location of Geremie is cited from the label data. It was omitted from the original 
description. Of the two syntypes of semirufus (one male, one female) in the MCZ, the 
best preserved (the male) is here designated lectotype. 

Present name: Heterostylum semirufum (Loew) 

serpentina Osten Sacken, Dipalta LECTOTYPE? 12646 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 237 
UNITED STATES: California: Mt. Shasta District [no date] 
O. Sacken 

Osten Sacken based his description of serpentina on one male and two female 
syntypes. The best preserved of these (a female) is here designated lectotype. 

shawii Johnson, Anthrax HOLOTYPE^ 7549 

1908 Psyche 15: 14-15 pin 

UNITED STATES: New Hampshire: Hampton 27.VIII. 1906 
S.A. Snow 

The holotype also carries a type label reading "holotype no. 599". ' 
Present name: Villa shawii (Johnson) 

sima Osten Sacken, Exoprosopa LECTOTYPE [<$] 12628 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3:231-33 
UNITED STATES: Nevada: Humboldt Station: Central Paci- 
fic Railroad [no further data] 
Osten Sacken's original description of sima incorrectly cites the sex of the three 
syntypes as female. They are, in fact, all males. The best preserved is here designated 
lectotype. 



'This number probably pertains to Johnson's personal collection prior to MCZ 
accession (Woodley, personal communication). 



18 BREVIORA No. 469 

slossonae Johnson, Spogostylum HOLOTYPE $ 7546 

1913 Bull. Am. Mus. Nat. Hist. pin 

32:55 
UNITED STATES: Kentucky: Cumberland Gap VII. 1876 G. 

Dimmock 
Present name: Anthrax aterrimus (Bigot) 

sordida Loew, Exoprosopa SYNTYPES [2$] 12642 

1869 Berliner Entomol. Z. 13: 14 pin 

MEXICO: 1 9, Tamaulipas: [no further data]; 1$, [Tamauli- 

pas]: Matamoros [no further data] 
Loew's original description of sordida cites an unspecified number of males and 
females. Only two males were found in the MCZ; label data cites locality of the latter 
syntype as "Matamoras." These specimens are left here as syntypes pending generic 
revision. 

stellans Loew, Argyromoeba HOLOTYPE [9] 12676 

1869 Berliner Entomol. Z. 13: 28-29 pin 

UNITED STATES: Oregon [no further data] 

Right wing broken; original description incorrectly cites sex of type as "$". 
Present name: Anthrax stellans (Loew) 

stenozona Loew, Anthrax LECTOTYPE 9 12649 

1869 Berliner Entomol. Z. 13: 25 pin 

UNITED STATES: Illinois [no further data] 

A lectotype is here designated for the one female in MCZ as Loew's description 
cites a range of measurements, implying a series of specimens. 
Present name: Villa stenozona (Loew) 

subauratus Loew, Geron SYNTYPES 2 S, 1 $ 12714 

1863 Berliner Entomol. Z. 7: 304-05 pin 

UNITED STATES: Pennsylvania: [no further data] Osten 

Sacken 
The specimens are left here as syntypes until the genus is revised. 

subvarius Johnson, Bombylius LECTOTYPE 9 7551 

1907 Psyche 14: 98-99 pin 

UNITED STATES: Pennsylvania: Lehigh Gap 26.VI.1901 

H.L. Viereck 
Lectotype female designated herein. 
Present name: Bombylius atriceps Loew 



1981 BOMBYLIIDAE TYPES IN MCZ 19 

sulphurea Loew, Phthiria HOLOTYPE $ 12705 

1863 Berliner Entomol. Z. 7: 113 pin 

UNITED STATES: New Jersey: [no further data] Glover 
Present name: Phthiria (Poecilognathus) sulphurea Loew 

titubans Osten Sacken, Exoprosopa HOLOTYPE [<J] 12633 
1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 233-34 
UNITED STATES: [Colorado]: Denver [no date] O. Sacken 

Original description incorrectly states sex of type as "$". 

trabalis Loew, Exoprosopa HOLOTYPE? 12638 

1869 Berliner Entomol. Z. 13: 13-14 pin 

MEXICO: [Veracruz]: Jalapa [no further data] 
Present name: Exoprosopa anthracoidea Jaennicke 

validus Loew, Bombylius LECTOTYPE S 12689 

1863 Berliner Entomol. Z. 7: 300 pin 

UNITED STATES: Virginia: [no further data] Osten Sacken 

Lectotype designated by Evenhuis in Hall and Evenhuis (1980: 90). 

virgata Osten Sacken, Toxophora SYNTYPES 1 6\ 1 $ 12727 

1877 Bull. U.S. Geol. Geogr. Surv. pin 

Terr. 3: 266-67 
UNITED STATES: Georgia [no further data] 

Osten Sacken based his description of virgata on two males and two females from 
Georgia and Texas. A male and a female from Georgia are in the MCZ. The Texas 
specimens have not been located. These specimens are left here as syntypes pending 
generic revision. 

vitripennis Loew, Geron SYNTYPES 2 $ 12713 

1869 Berliner Entomol. Z. 13: 173 pin 

UNITED STATES: "Middle States" [no further data] 

No locality labels on type specimens. Until the genus is revised they are left as 
syntypes. 

vulgaris Loew, Systoechus LECTOTYPE 3 12717 

1863 Berliner Entomol. Z. 7: 302-03 pin 

UNITED STATES: Nebraska: [no further data] [F. V. Hayden] 
Lectotype designated by Hall in Hall and Evenhuis (1981: 151); original descrip- 
tion incorrectly states collector as "Osten Sacken." Osten Sacken (1903: 95) gives the 
correct collector. 



20 BREVIORA No. 469 

webberi Johnson, Villa HOLOTYPE $ 12680 

1919 Psyche 26: 11 pin 

UNITED STATES: Massachusetts: Lunenberg 3. VI. 1914 [no 

collector] 
Present name: Hemipenthes webberi (Johnson) 

willistoni Osten Sacken, Pantarbes HOLOTYPE $ 12722 

1887 Biologia Cent rali- Americana pin 

1: 153 
UNITED STATES: Arizona: [no further data] O. Sacken 

xanthomeros Marston, Anthrax HOLOTYPE $ 32555 

1970 Smithson. Contrib. Zool. pin 

43: 52-53 PI. 3g 
BRITISH HONDURAS: Benque Viejo [no date] Father 
Stanton 

GENUS-GROUP INVENTORY 

(indexed by present combination) 

Anastoechus: barbatus 

Anthrax: bigradata, contigua, delila, euplanes, fur, obsoleta, occi- 

dentalis, pauper, slossonae, stellans, xanthomeros 
Aphoebantus: cervinus, mus mus 
Bombylius (Bombylius): albicapillus, atriceps, aurifer, cachinnans, 

fraudulentus, lancifer, metopium, pulchellus, subvarius, validus 
Bombylius (Zephyrectes): incanus, ravus 
Chrysanthrax: lepidotoides 
Conophorus: amabilis, atratulus, fenestratus, limbatus, nigripennis, 

obesulus, rufulus 
Dipalta: banksi, serpentina 
Epacmus: modestus 
Exoprosopa: agassizi, bifurca, cubana, decora, dodrans, dorcadion, 

doris, eremita, nubifera, sima, sordida, titubens, trabalis 
Geron: albidipennis, calvus, johnsoni, macropterus, subauratus, 

vitripennis 
Hemipenthes: curta, sagata, seminigra, webberi 
Heterostylum: haemorrhoicum, semirufum 
Lepidanthrax: proboscideus 



1981 BOMBYLIIDAE TYPES IN MCZ 21 

Ligyra: gazophylax 

Lordotus: gibbus, planus 

Neodiplocampta (Neodiplocampta): parva 

Oligodranes: quinquenotata 

Pantarbes: capito, willistoni 

Paravilla: diagonalis, palliata 

Paracosmus: edwardsii 

Phthiria (Phthiria): egerminans, humilis 

Phthiria (Poecilognathus): aldrichi, borealis, coquilletti, cyanoceps, 

loewi, scolopax, sulphurea 
Poecilanthrax: alpha, ceyx, flaviceps, fuliginosus 
Rhynchanthrax: parvicornis 
Sparnopolius: brevicornis 
Systoechus: candidulus, oreas, vulgaris 
Systropus: arizonicus, macer 

Thevenemyia: canuta, harrisi, luctifera, magna, marginata 
Thyridanthrax: pertusa 
Toxophora: virgata 
Villa: adjusta, lateralis arenicola, molitor, mucorea, fulviana nigri- 

cauda, scrobiculata, shawii, stenozona 



ACKNOWLEDGMENTS 

I wish to thank Mr. Jack C. Hall for reviewing the manuscript of 
this paper and Drs. Pedro Wygodzinsky of the American Museum 
of Natural History and H. Schumann of the Museum fur Natur- 
kunde der Humboldt-Universitat, Berlin for their help in locating 
Osten Sacken and Loew types, respectively, in their museums. I also 
wish to express my sincere gratitude to Mr. Norman E. Woodley for 
his assistance during my visit to the MCZ and for his comments and 
suggestions concerning the manuscript of this study. I thank Ms. 
Margaret K. Thayer for her help in obtaining for me the initial loans 
of some of the type material in the MCZ. 

Travel for this study was supported by National Science Founda- 
tion Grant DEB 79- 1 1 92 to the University of California, Riverside. 

LITERATURE CITED 

Banks, N. 1909. A new species of Systropus (Bombyliidae). Entomol. News, 20: 18. 



22 BREVIORA No. 469 

Cole, F.R. 1917. Notes on Osten Sacken's group "Poecilanthrax," with descriptions 

of new species. J. N.Y. Entomol. Soc, 25: 67-80. 
Hall, J.C. 1969. A review of the subfamily Cylleniinae with a world revision of the 

genus Thevenemyia Bigot (Eclimus auct.) (Diptera: Bombyliidae). Univ. Calif. 

Publ. Entomol., 56: 1-85. 
Hall, J.C, and N.L. Evenhuis. 1980. Part 13, Number 1. Family Bombyliidae, pp. 

1-96. In Griffiths, G.C.D. (ed.), Flies of the Nearctic Region. Vol. V. Stuttgart, E. 

Schweizerbart'sche. 
1981. Part 13, Number 2. Family Bombyliidae, pp. 97-184. In Griffiths, 

G.C.D. (ed.), Flies of the Nearctic Region. Vol. V. Stuttgart, E. Schwei- 
zerbart'sche. 
Johnson, C.W. 1902. New North American Diptera. Can. Entomol., 34: 240-42. 
. 1903. Descriptions of three new Diptera of the genus Phthiria. Psyche, 10: 

184-85. 
. 1907. A review of the species of the genus Bombylius of the eastern United 

States. Psyche, 14: 95-100. 
1908. Notes on New England Bombyliidae, with a description of a new species 

of Anthrax. Psyche, 15: 14-15. 
. 1910. Some additions of the dipteran fauna of New England. Psyche, 17: 

228-35. 

1913. Insects of Florida. I. Diptera. Bull. Amer. Mus. Nat. Hist., 32: 37-90. 

. 1919. New species of the genus Villa (Anthrax). Psyche, 26: 1 1-13. 

1921. New species of Diptera. Occas. Pap. Boston Soc. Nat. Hist., 38: 57-99. 



Loew, H. 1863a. Diptera Americae septentrionalis indigena. Centuria tertia. Berliner 

Entomol. Z., 7: 1-55. 
. 1863b. Diptera Americae septentrionalis indjgena. Centuria quarta. Berliner 

Entomol. Z., 7: 275-326. 
. 1869a. Diptera Americae septentrionalis indigena. Centura octava. Berliner 

Entomol. Z., 13: 1-52. 
. 1869b. Diptera Americae septentrionalis indigena. Centuria nona. Berliner 

Entomol. Z., 13: 129-186. 
. 1872. Diptera Americae septentrionalis indigena. Centuria decima. Berliner 



Entomol. Z., 16:49-115. 

Marston, N. 1970. Revision of New World species of Anthrax (Diptera: Bombyliidae) 
other than the Anthrax albofasciatus group. Smithson. Contrib. Zool., 43: 
1-148. 

Osten Sacken, C.R. 1877a. Art. XIII. Western Diptera: Descriptions of new genera 
and species of Diptera from the region west of the Mississippi and especially 
from California. Bull. U.S. Geol. Geogr. Surv. Terr., 3: 189-354. 

1887b. Diptera. pp. 129-216. In Godman, F.D. and O. Salvin (eds.), Biologia 

Centrali-Americana. Zoologia-Insecta-Diptera. Vol. I. London, Taylor & Fran- 
cis, 378 pp. 

. 1903. Record of my life work in entomology. Cambridge, Mass., University 



Press, John Wilson & Sons, 204 pp. 
Painter, R.H. 1932. A monographic study of the genus Geron Meigen as it occurs in 
the United States (Diptera: Bombyliidae). Trans. Amer. Entomol. Soc, 58: 
139-67. 



1981 BOMBYLIIDAE TYPES IN MCZ 23 

Painter, R.H., and E.M. Painter. 1962. Notes on and redescriptions of types of 
North American Bombyliidae (Diptera) in European Museums. J. Kans. 
Entomol. Soc., 35: 1-164. 

Radovsky, F.J., G.A. Samuelson, and W.A. Steffan. 1976. Catalog of entomological 
types in the Bishop Museum. Introduction. Pac. Insects, 17: 1-5. 



"OS. CCMP. ZOOL 
LfDRARY 

MAR 1 8 1985 

BRE V°'f^ R A 

Museum of Comparative Zoology 

IS ISSN 0006 9698 

Cambridge. Mass. 30 June 1982 Number 470 

SYSTEMATIC IMPLICATIONS OF 
INNERVATION PATTERNS 
IN TELEOST MYOTOMES 

Quentin Bone 1 and R. Dana Ono 2 

Abstract. The peripheral innervation patterns of the red and white myotomal 
muscles from over 230 species representing more than 125 families of teleosts were 
studied. A distributed, multiple innervation pattern of teleost red superficial 
myotomal muscles was found without exception in all groups examined. There is 
variability in the innervation patterns of the white myotomal muscles, however \ 
terminally innervated pattern seems to be present in the basal groups of teleosts. 
while a trend toward distributed innervation occurs in the Neoteleostei. Stomii- 
formes possess a rather different distributed pattern which we suggest is the early, 
transitional stage from terminal to distributed innervation patterns in teleosts There 
appears to be a distinct functional difference in the distributed and terminal 
innervation patterns. The innervation of the white myotomal libers should be 
considered a taxonomicallv useful character in elucidating familial relationships 

INTRODUCTION 

In all fishes, there are two main muscle fiber types in the 
myotomes, usually readily visible to the naked eye when the fish is 
sectioned transversely. Small-diameter, well-vascularized red mus- 
cle fibers normally form a thin superficial layer covering the much 
more numerous, larger-diameter, poorly-vascularized white muscle 
fibers that make up the major portion of the myotome. In some 
fishes, but not in all, other myotomal fiber types are present. These 
minor myotomal components will not concern us here. In all fishes. 



'The Laboratory, Citadel Hill. Plymouth I'll 2PB, Devon. F.ngland. 

: Museum of Comparative Zoology. Harvard University. Cambridge. Massachusetts 

02138. 



BREVIORA 



No. 470 



— a «s 




« M E 



1982 



MYOTOME INNERVATION 



OSTARIOPHYS 




Clupeocephala 
E lopocephala 



Figure 2. Hypothetical relationship of the Teleostei modified from Fink and 
Weitzman (1982) and Fink and Fink (1981) to show the extent of the distributed 
innervation pattern in teleosts. ^™ = distributed innervation in all.<=i = distributed 
innervation in some members. 



the red muscle fibers are multiply innervated in a distributed 
fashion, so that each muscle fiber receives a number of motor 
terminations along its length (Fig. 1A). In most teleosts, the white 
muscle fibers are similarly innervated (Figs. ID, 3, 6), but in some 
an entirely different innervation pattern is found. In such fishes the 
white muscle fibers are innervated only at their myoseptal ends, so 
that innervation is focal and terminal (Figs. IB, 4, 5). 

Although this mode of innervation of white muscle fibers is rare 
in teleosts (Barets, 1961; Bone, 1964), investigations on other fish 
groups such as the Halecomorphi, Ginglymodi, Brachiopterygii, 



BREVIORA 



No. 470 



*~mw% < .-**. -» 




v 7 2 in 



"^^.^A*. 



v *» y 









r 



* 

* 






•M*^P- 



^ 






Hi * i 




Figure 3. Distributed innervation pattern of motor endplates in white myotomal 
muscle fibers of Morone labrax as revealed by acetylcholinesterase ( AChE) studies. 
(Mag. X90). 





Figure 4. Terminal innervation pattern of motor endplates in white myotomal 
muscle fibers of the clupeomorph, Sprattus sprattus, as revealed by AChE studies. 
(Mag. X90). 



1982 



MYOTOME INNERVATION 




Figure 5. Silver impregnated axons of terminal innervation pattern in the white 
myotomal muscle fibers of the alepocephalid, Xenodermichthys copei. Winkelmann 
and Schmitt technique. (Mag. X190). 




Figure 6. Silver impregnated axons of the distributed inner\ation pattern in the 
white myotomal muscle fibers of the gymnotid, Eigenmannia virescens. Winkelmann 
and Schmitt technique. (Mag. X228). 



6 BREVIORA No. 470 

Chondrostei (Acipenseridae, Polydontidae), hagfish, Elasmobran- 
chiomorpha, and Latimeria chalumnae have shown that terminal 
innervation is universal in non-teleost groups (Bone, 1964 and 
unpublished; Ono, unpublished; Sakharov and Kashapova, 1979). 
In addition, terminal innervation is found in the Dipnoi, in the 
myotomal musculature of adult urodeles, and in urodele and anuran 
larvae (Best and Bone, 1973; Bone, unpublished; Ono, unpub- 
lished). 

The wide distribution of the terminal innervation pattern of white 
muscle fibers led to a preliminary attempt (Bone, 1970) to link the 
occurrence of the terminal pattern in teleosts with the systematic 
position of the families in which it was found. This attempt was 
unsuccessful since relatively few families were examined. In this 
study, we report the results of a more detailed survey of the 
innervation pattern of white muscle fibers in different teleost groups 
that suggests that this character may be of interest to systematists. 

MATERIALS AND METHODS 

The innervation pattern of white muscle fibers was studied either 
by supravital staining with methylene blue in teleost Ringer 
solution, or by silver staining of 10% unbuffered formalin and 
Bouin-fixed material using the methods of Palmgren (1948) and 
Winkelmann and Schmitt (1957). During the last several years, we 
have collected marine fish material from the Caribbean, the Indian 
Ocean, the North Atlantic Ocean, and the Bay of Biscay. The 
freshwater fish material was collected by us from North American 
and European waters. In addition, we have been much assisted by 
other workers who have provided specially fixed or museum 
material from the waters of South America, Africa, Japan, New 
Zealand, and several deep-sea localities. We owe a particular debt of 
gratitude to Dr. P. H. Greenwood (BMNH) and to Mr. J. B. 
Badcock and Mr. N. Merrit (IOS) who have generously provided 
material from their collections. 

TELEOST GROUPS AND THEIR 
INNVERVATION PATTERNS 

The status of the different groups of teleosts is still subject to 
review, and different authors have adopted different systems of 
classification to indicate their views of the relationships of the 



1982 MYOTOME INNERVATION 7 

various groups. In this paper we have adopted the classification 
scheme of Rosen (1973) and Fink and Weitzman (1982). A complete 
checklist showing the teleosts examined is given in Table 1. 

1) Osteoglossomorpha 

Nine genera from this group were examined. With the exception 
of the two Hiodon species, all possess the distributed pattern of 
white muscle innervation. 

2) Elopomorpha 

We have examined 12 genera from this group representing the 
Elopiformes, Anguilliformes, and Notacanthiformes. All possess 
the terminal innervation pattern. Both the adults and leptocephali 
of Elops sp. and Albula sp. have the terminal pattern of white 
muscle innervation. 

3) Clupeomorpha 

Twenty-one species of clupeids belonging to 16 genera were 
studied. All possess the terminal innervation pattern with the sole 
exception of Denticeps, in which the white myotomal muscle has a 
distributed innervation. 

4) Ostariophysi 

We have examined 32 genera of ostariophysans representing the 
Gonorynchiformes, Characiformes, Cypriniformes, and Siluri- 
formes. Some ostariophysans possess the terminal pattern (some 
siluroids and all gonorynchids), but the majority have the distrib- 
uted pattern of innervation. 

5) Protacanthopterygii 

There is considerable uncertainty about the limits of this group, 
which has been reduced from its original size (Fink and Weitzman, 
1982). The 16 species representing 14 genera listed in Table 1 are 
provisionally placed in this group. 

6) Stomiiformes 

Fifteen genera representing the two Infraorders Gonostomata 
and Photichthya (Weitzman, 1974) were studied. Two innervation 
patterns are found in these two lineages of Stomiiformes. While the 
Gonostomata possess the usual distributed innervation pattern 
found in other teleosts, members within the Photichthya have the 
white zone of the myotome innervated in a different way. Terminal 
endplates are seen on many fibers which are probably innervated at 



8 BREVIORA No. 470 

both ends. In addition, some axons pass from the myosepta into the 
midregions of the myotome to form a sparsely distributed innerva- 
tion pattern. Thus, the Photichthya possess a third type of 
innervation pattern (Fig. 1C), distinct from either the terminal or 
distributed patterns, apparently combining elements of both. 

7) Eurypterygii 

All 125 species representing 99 genera examined in the Aulopi- 
formes, Myctophiformes, Paracanthopterygii, and Acanthopterygii 
possess the distributed innervation pattern (see Table 1 for list of 
eurypterygians examined). 

DISCUSSION 

All non-teleost fishes possess terminally innervated white muscle 
fibers (Ono, unpublished). It is reasonable to suppose therefore that 
this pattern is the primitive condition and that the distributed 
pattern of innervation is derived. The universal occurrence of the 
distributed pattern in the Eurypterygii supports this view. 

Two questions arise. First, is there a functional advantage in the 
distributed pattern that has led to its universal appearance in 
"higher" teleosts? Secondly, can white muscle innervation prove 
useful in dealing with the controversial questions of the limits and 
interrelationships of different teleost groups? 

Consideration of the habits of fishes possessing the two types of 
innervation pattern, for example, Hiodon and Heterotis or Albula 
and Chanos, shows no obvious correlation between habitat and 
innervation pattern, nor any obvious differences in locomotor 
ability. But studies of fishes swimming under controlled conditions 
have revealed a notable difference in locomotor ability between 
fishes where the white muscle is innervated terminally, and where it 
is innervated in the distributed manner. Few fishes have yet been 
studied in this way (where activity in different zones of the myotome 
is monitored by electromyography as the fish swims at different 
speeds), but results are consistent, and are probably generally 
applicable. In all fishes studied, the red muscle fiber zone of the 
myotome is active during slow cruise swimming that can be 
maintained indefinitely. In fishes where the white muscle fibers are 
terminally innervated, these are only active during bursts of rapid 
swimming, and are rapidly exhausted (Bone, 1966; Bone et ah, 



1982 MYOTOME INNERVATION 9 

1978). Where the white musele fibers are innervated in the 
distributed manner, they operate quite differently, for they are 
active not only during bursts of rapid swimming, but also during 
cruise swimming at intermediate speeds (Hudson, 1973; Johnston et 
ah, 1977; Bone et ai, 1978). 

Electromyographic records from the white muscle /one during 
cruise swimming in such fishes are different from those obtained 
during rapid swimming, and it appears that where the fibers receive 
distributed innervation, they can operate in two ways propagating 
action potentials only during rapid swimming (Bone et ai, 1978). 
Extracellular recordings cannot provide other than suggestive 
evidence on this point, and definite proof of the hypothesis that 
contraction of the same fast muscle fiber in such fishes can either 
follow local potentials or propagated action potentials awaits 
intracellular investigation. 

However, the results of these electromyographic studies clearly 
show that fishes with the distributed pattern of innervation of the 
white muscles are able to recruit these fibers to give a wider range of 
sustainable cruising speeds than can be obtained when the white 
fibers are terminally innervated and are used only during burst 
swimming. Evidently, this ability to swim for long periods over a 
wide speed range could confer a significant advantage to fishes in 
some particular habitats. For example, it may be advantageous to 
fishes living in streams where flow varies and the fishes are required 
to keep stationary. This ability to swim over a wide speed range 
would be insignificant to benthic fishes, however, at least in the 
adult stage due to the relatively stable flow regime of the water 
column. 

We conclude that the distributed pattern of innervation was 
derived in teleostean evolution when adults or larvae were pelagic, 
and has been retained in those groups where the adults today are 
relatively sedentary, so that the habits of the adults today are not 
correlated with the innervation pattern. 

The first attempt to use the innervation pattern as a systematic 
character made the simple assumption that this derivation from the 
ancestral terminal pattern occurred only once during teleost 
phylogeny (Bone, 1970). The present more complete survey 
(unfortunately still lacking data on several important species) 
demonstrates that this assumption can no longer by justified. Figure 



10 BREVIORA No. 470 

2 shows a current view of the relationships among teleost groups 
(Fink and Weitzman, 1982), and indicates points where transition 
from terminal to distributed innervation is assumed to have taken 
place. In this view of the relationships of the groups, the change 
presumably took place independently on at least eight occasions. 

In general, our survey has shown that the more primitive groups 
possess the terminal innervation pattern, as expected. Thus in the 
Osteoglossomorpha, the Hiodontoidei is a phylogenetic relict in the 
sense that the two living Hiodon species form a lineage that has 
retained a large number of primitive features (Greenwood, 1970); 
only these members of the group possess the terminal pattern. 

Both the Elopomorpha and Clupeomorpha (with the sole 
exception of Denticeps) also possess the terminal pattern. The 
freshwater Denticeps retains many primitive characters, (Green- 
wood, 1968), but is unique among clupeomorphs in having achieved 
the distributed innervation pattern. Since a wider sustainable speed 
range might be a distinct advantage in the fluviatile freshwater 
environment, we examined other African freshwater clupeids, but 
all five genera studied possessed terminally innervated white muscle 
fibers. 

In the Ostariophysi, all Gonorynchiformes and some Siluriformes 
have the terminal innervation pattern, and if the scheme of 
ostariophysan relationships proposed by Fink and Fink (1981) is 
accepted, this implies that convergent origin of the distributed 
pattern has occurred in all lines except that leading to the 
Gonorynchiformes (Fig. 2). Certainly, although the phylogenetic 
position of Siluriform families is uncertain, the innervation pattern 
is terminal in the most primitive, the relict Diplomystidae. 

Perhaps the most interesting group with respect to muscle 
innervation pattern is the Stomiiformes. This group has been less 
thoroughly studied than the Ostariophysi, and despite recent work 
by Fink and Weitzman (1982), and an earlier study by Weitzman 
(1974), relationships within the group are still uncertain. It is 
notable that the two major sister groups forming the Stomiiformes, 
the Gonostomata and Photichthya (Weitzman, 1974), have different 
innervation patterns. In the Gonostomata, innervation is dis- 
tributed, but in the Photichthya, innervation of the white fibers is 
mixed. Relatively few axons course through the white portion of the 
myotome, and in addition there are axons terminating on the fiber 



1982 MYOTOME INNERVATION 1! 

ends as in the terminal pattern. Perhaps in this group of teleosts, we 
witness the distributed innervation pattern at an early stage in its 
development. It would be interesting to see whether the mixed 
pattern of adult photichthyans develops from an initially terminal 
pattern in the larval stage. 

Few studies involving the nervous system in fishes have been used 
in a comparative fashion to solve problems of classification. On the 
whole, our survey suggests that the innervation of white myotomal 
fibers, despite evident convergence in its origin, should be con- 
sidered a systematically useful character, and may prove particu- 
larly helpful in elucidating interrelationships within the Siluriformes 
and the Stomiiformes, respectively. 

ACKNOWLEDGMENTS 

We express our gratitude to Drs. P. H. Greenwood, William L. 
Fink, Karel F. Liem, Ian A. Johnston, as well as to Mr. Julian B. 
Badcock and Ms. Sara V. Fink for providing helpful comments and 
specimens. R. D. O. was partially supported with funds from a 
Raney Award (American Society of Ichthyologists and Herpeto- 
logists), National Science Foundation Grant BNS-79 15308, and 
National Institutes of Health Predoctoral Fellowship GMO-7117 
during the tenure of this project. 

LITERATURE CITED 

Barets, A. 1961. Contribution a l'etude des systemes moteurs lent-et rapid du 

muscle lateral des teleosteens. Arch. Anat. Morphol. Exp., 50 (suppl.): 91 187. 
Best, A. C. G., and Q. Bone. 1973. The terminal neuromuscular junctions of 

lower chordates. Z. Zellforsch., 143: 495-504. 
Bone, Q. 1964. Patterns of muscular innervation in the lower chordates. Int. Rev. 

Neurobiol., 6: 99-147. 
1966. On the function of the two types of myotomal muscle fibre in 

elasmobranch fish. J. Mar. Biol. Ass. U.K.. 46: 321 349. 
1970. Muscular innervation and fish classification, pp. 369 377. //; A. 



de Haro (ed.), I Simposio Internacional de Zoofilogenia, Fac. Ciencias. Univ. 

Salamanca, 492 pp. 
Bone, Q., J. Kiceniuk, and D. R. Jones. 1978. On the role of different fibre 

types in fish myotomes at intermediate swimming speeds. Fi.-.h. Bull.. 

76: 691 699. 
Fink, S. V., and W. L. Fink. 1981. Interrelationships of the ostariophysan 

fishes (Teleostei). Zool. J. I. inn. Soc. 72(4): 297 353. 



12 BREVIORA No. 470 



Kink. W. I ... \\n S. H. Weitzman. 19X2. Relationships of the Stomiiform fishes 

(Teleostei) with a description of Diplophos. Bull. Mus. Comp. Zool., 150: 31 93. 
Greenwood, P. H. 1968. The osteology and relationships of the Denticipitidae, 

a family of clupeomorph fishes. Bull. Br. Mus. Nat. Hist. (Zool.)., 16: 213 273. 
1970. On the genus, Lycoptera. and its relationships with the family 

Hiodontidae (Pisces, Osteoglossomorpha). Bull. Br. Mus. Nat. Hist. (Zool.)., 

19: 257 285. 
Hudson, R. C. L. 1973. On the function of the white muscles in teleosts at 

intermediate swimming speeds. J. Exp., Biol.. 58: 509-522. 
Johnston, 1. A., W. Davidson, and G. Goldspink. 1977. Energy metabolism 

of carp swimming muscles. J. Comp. Physiol., 114: 203 216. 
Palmgren, A. 1948. A rapid method for selective silver staining of nerve fibres 

and nerve endings in mounted paraffin sections. Acta Zool., 29: 378-392. 
Rosen, D. E. 1973. Interrelationships of higher euteleostean fishes, pp. 397 513. 

In P. H. Greenwood, R. S. Miles, and C. Patterson (eds.), London, Academic 

Press, 536 pp. 
Sakharov, D. A., and L. A. Kashapova. 1979. The primitive pattern of the 

vertebrate body muscle innervation: Ultrastructural evidence for two synaptic 

transmitters. Comp. Biochem. Physiol., 62(A): 771 776. 
Weitzman, S. H. 1974. Osteology and evolutionary relationships of the 

Sternoptychidae with a new classification of stomiatoid families. Bull. Am. Nat. 

Hist.. 153(3): 329-478. 
Winkelmann, R. K.., and R. W. Schmitt. 1957. A simple silver method for 

nerve axoplasm. Proc. Staff Meeting Mayo Clinic. 32: 217 222. 






1982 



MYOIOMl IWI KY \ I I()\ 



13 



Table 
Teleosts. 



Patterns of Innervation in the White MwHomal Muscle Fibers ol 



List of Teleosts Examined for 

Innervation Pattern 



White Myotomal 
Muscle Kibers 



Terminal 
Pattern 



Distributed 

Pattern 



Osteoglossomorpha 
Notopteridae 

Notopterus chitala 
Xenomystus nigri 

Osteoglossidae 

Heterotis niloticus 
Osteoglossum bicirrosum 

Pantodontidae 

Pantodon bucholzi 

Mormyridae 
Mormyrus sp. 

Mormyrops engystoma 
Gnathonemus petersii 

Hiodontidae 
Hiodon tergisus 
Motion alosoides 



Elopomorpha 

Elopiformes 

Elopidae 
El ops sp. 

Albulidae 
Alhitla sp. 

Anguilliformes 

Anguillidae 
Anguilla sp. 

Muraenidae 
Gymnothorax sp. 

Congridae 

Conger sp. (2) 
Paraconger sp. 

Muraenesocidae 
Muraenesox sp. 

Ophichthidae 
Ophichthus sp. 



14 



BREVIORA 



No. 470 



Table 1. continued 



List of Teleosts Examined for 
Innervation Pattern 



White Myotomal 
Muscle Fibers 



Terminal 
Pattern 



Distributed 
Pattern 



Eurypharyngidae 

Eurypharynx pelecanoides 

Notacanthiformes 

Halosauridae 
Halosaurus sp. 
Halosauropsis sp. 

Notacanthidae 

Polyacanthonotus sp. 

Clupeomorpha 

Clupeidae 
Clupea sp. 
Harengula sp. (2) 
Sardinella sp. 
Pellonula atzeliusi 
Opisthopterus sp. 
Limnothrissa miodon 
Thrissocles sp. 
Euplatygaster sp. 
Cynothrissa memo 
Poecilothrissa congicae 
Stolothrissa tanganicae 
Microthrissa sp. 
Opisthopterus sp. 
Alosa pseudoharengus 
Alosa aestivalis 
Alosa sp. 
Sprattus sprattus 

Engraulidae 

Anchoa mil chilli 
Anchoa sp. 

Chirocentidae 

Chirocentrus dorab 

Denticipitidae 

Demiceps clupeoides 

EUTELEOSTEI 

Ostariophysi 

Gonorynchiformes 



+? 



1982 



myotome; innervation 



\5 



Table 



continued 



List of Teleosts Examined for 
Innervation Pattern 



White Myotomal 
Muscle Fibers 



Terminal 
Pattern 



Distributed 
Pattern 



Chanidae 

Chanos chanos 

Kneriidae 
Kneria mittei 

Phractolaemidae 

Phraetolaemus ansorgei 

Gonorynchidae 

Gonorynchus gonorynchus 

Characiformes 

Characidae 

Crenuchus spilurus 
Hyphessobrycon flammeus 
Hyphessobrycon serpae X 

Hyphessobrycon collistus 
Hyphessobrycon pulchrspinnis 
Astyanax mexicanus 

Lebiasinidae 

Nannostomus nannostomus 

Gasteropelecidae 
Gasteropelecus sp. 

Hemiodontidae 
Hemiodus sp. 

Cypriniformes 

Cyprinidae 

Notropis hudsonius 
Cyprinus sp. 

Gyrinocheilidae 

Gyrinocheilus aymonieri 

Catostomidae 

Catostomus catostomus 

Cobitidae 

Noemacheilus sp. 

Siluriformes 

Siluroidei 



16 



BREVIORA 



No. 470 



Table I. continued 



White Myotomal 
Muscle Fibers 



List of Teleosts Examined for 
Innervation Pattern 



Terminal 
Pattern 



Distributed 
Pattern 



Bagridae 

Parauchenoglanis macrostoma 

Siluridae 

Kryptopterus bicirrhis 

Malapteruridae 
Malapterurus electricus 

Pangasiidae 

Pangasius sutchi 

Chacidae 
Chaca chaca 

Mochokidae 
Synodontis sp. 

Aspredinidae 

Bunocephalus sp. 

Callichthyidae 
Cory dor as sp. 
Hoplosternum sp. 
Dianema sp. 

Loricariidae 
Ancistrus sp. 

Ictaluridae 
Ictalurus sp. 

Diplomystidae 
Diplornystes sp. 

Ariidae 
Arius sp. 

Doradidae 
Doras sp. 

Pimelodidae 

Sorubim limas 
Pimelodella sp. 

Gymnotoidei 

Rhamphichthyidae 
Eigenmannia virescens 



1982 



MYOTOME INNERVATION 



17 



Table 1. continued 



White Myotomal 
Muscle Fibers 



List of Teleosts Examined for 
Innervation Pattern 



Terminal 
Pattern 



Distributed 
Pattern 



Protacanthopterygii 

Salmoniformes 

Esocidae 
Esox niger 
Esox americanus 

Umbridae 
Umbra limi 
Dallia peel oralis 

Salmonidae 
Salmo truita 
Salmo sp. 

Retropinnidae 
Retropinna sp. 

Galaxiidae 
Galaxias sp. 

Osmeridae 

Osmerus tnordax 

Plecoglossidae 

Plecoglossus altivelis 

Argentinidae 
Argentina sp. 

Opisthoproctidae 
Opisthoproctus sp. 

Alepocephalidae 
Alepoeephalus sp. 
Xenodermichthys copei 
Bathylaco nigricans 

Searsiidae 
Searsia sp. 

Stomiiformes 

Gonostomata 

Gonostomatidae 

Cyclothone obscura 
Gonostoma elongatum 
Maurolicus sp. 



18 



BREVIORA 



No. 470 



Table 1. continued 



White Myotomal 
Muscle Fibers 



List of Teleosts Examined for 
Innervation Pattern 



Terminal 
Pattern 



Distributed 
Pattern 



Sternoptychidae 
Argyropelecus sp. (2) 
Sternoplyx sp. 

Photichthya 

Chauliodontidae 
Chauliodus sp. 

Stomiatidae 
Stomias sp. 
Macrostomias longibarbatus 

Astronesthidae 
Astronesthes sp. 

Melanostomiatidae 
Melanostomias sp. 
Eustomias sp. 
Echiostoma barbatwn 

Malacosteidae 
Malacosteus sp. 
Photostomias sp. 

Idiacanthidae 
Idiacanthus sp. 

EURYPTERYGII 

Aulopiformes 

Aulopodidae 
Aulopus sp. 

Synodontidae 
Sy nodus sp. (2) 

Giganturidae 
Gigantura sp. 

Bathypteroidae 

Batbypterois sp. (2) 

Myctophiformes 

Myctophidae 
Myciophum sp. 
Diaphus sp. 



1982 



MYOTOME INNERVATION 



19 



Table I. continued 



List of Teleosts Examined for 
Innervation Pattern 



White Myotomal 
Muscle Fibers 



Terminal 
Pattern 



Distributed 
Pattern 



Paralepididae 
Paralepis sp. 

Omosudidae 
Omosudis sp. 

Evermannellidae 
Coccorella sp. 

Scopelarchidae 
Scopelarchus sp. 

Paracanthopterygii 

Gadiformes 

Moridae 
Antimora sp. 

Macrouridae 

Nematonurus sp. 

Lophiiformes 

Lophiidae 
Lophius sp. 

Antennariidae 

Antennarius hispidus 
Antennarius scaber 

Acanthopterygii 

Atheriniformes 

Exocoetidae 

Parexocoetus sp. 
Cypselurus sp. 
Hemiramphus sp. (2) 

Belonidae 
Be lone sp. 
Platybelone sp. 
Tylosurus sp. 

Cyprinodontidae 
Belonesox belizanus 

Scomberesocidae 
Scomberesox sp. 



20 



BREVIORA 



No. 470 



Table 1. continued 



List of Teleosts Examined for 
Innervation Pattern 



White Myotomal 

Muscle Fibers 



Terminal 
Pattern 



Distributed 
Pattern 



Atherinidae 

Menidia menidia 
Atherinomorus sp. 

Beryciformes 

Polymixiidae 
Polymixia lowei 
Polymixia japonica 

Holocentridae 

Hulocentrus sp. (3) 

Cetomimidae 
Cetomimus sp. 

Melamphaeidae 
Melamphaes sp. 

Dactylopteriformes 

Dactylopteridae 
Dactylopterus sp. 
Zeiformes 

Zeidae 

Zeus faber 

Sygnathiformes 

Aulostomidae 
Aulostomus sp. 

Fistulariidae 

Fistularia sp. (2) 

Sygnathidae 
Nerophis sp. 
Hippocampus sp. 

Scorpaeniformes 

Triglidae 
Trigla sp. 

Cottidae 

Coitus cognathus 

Perciformes 

Centrarchidae 

Pomoxis nigromaculatus 
Lepomis gibbosus 



1982 



MYOTOME INNERVA I IO\ 



21 



Table 



continued 



White Myotomal 
Muscle Fibers 



List of Teleosts Examined for 
Innervation Pattern 



Terminal 
Pattern 



Distributed 
Pattern 



Priacanthidae 
Priacanthus sp. 

Carangidae 
Caranx sp. (3) 
Oligoplites sp. 
Seriola sp. 

Lutjanidae 

Lutjanus sp. (2) 
Ocyurus sp. 

Plectorynchidae 
Gaterin sp. 

Pentapodidae 
Monotaxis sp. 

Serranidae 

Epinephelus sp. (3) 
Peirometapun sp. 
Serranus sp. 
Cephalopholis sp. 

Pomadasyidae 
Haemulun sp. (2) 
Anisotremus sp. 

Percichthyidae 
Morone lahrax 

Sphyraenidae 
Sphyraena sp. 

Grammistidae 
Rypticus sp. 

Scaridae 

Sparisoma sp. (3) 

Percidae 

Eihevsiuma olmstedi 

Stromateidae 
Strumateid sp. 

Istiophoridae 
Makaira sp. 



22 



BREVIORA 



No. 470 



Table 1. continued 



White Myotomal 
Muscle Fibers 



List of Teleosts Examined for 
Innervation Pattern 



Terminal 
Pattern 



Coryphaenidae 
Coryphaena sp. 

Sciaenidae 

Cynoscion regalis 
Equetus sp. 

Cichlidae 
Julidochromis sp. 

Belontiidae 

Trichogaster trichopterus 

Channidae 

Channa micropeltes 

Mullidae 

Mullus surmuletus 
Mulloidichthys sp. 

Kyphosidae 
Kyphosus sp. 

Chaetodontidae 
Chaetodon sp. (6) 
Pomacanthus sp. 

Pomacanthodes sp. 

Pomacentridae 
Abudefduf sp. 

Cepolidae 

Cepola rubescens 

Centropomidae 
Centropomus sp. 

Acanthuridae 

Adanthurus sp. (4) 
Zanclus sp. 

Scombridae 

Scomber sp. (2) 
Thunnus sp. 
Euthynnus sp. 

Gerreidae 
Gerres sp. 
Eugerres sp. 



Distributed 
Pattern 









1982 



MYOTOME INNERVATION 



23 



Table 1. continued 



List of Teleosts Examined for 
Innervation Pattern 



White Myotomal 
Muscle Fibers 



Terminal 
Pattern 



Distributed 
Pattern 



Siganidae 
Siganus sp. 

Labridae 

Halichoeres sp. 
Lepidaplois sp. 
Anampses sp. 
Lachnolaimus maximus 

Gobiidae 

Periophthalmus koelreuteri 

Ephippidae 

Chaetodipterus sp. 

Tetraodontiformes 

Balistidae 
Batistes sp. 
Xanthichthys sp. 
Melichthys sp. 
Cantherhines sp. 
Balistapus sp. 

Ostraciotidae 

Acanthostracion sp. (2) 
Lactophrys sp. 

Diodontidae 
Diodon sp. (2) 
Chilomycterus sp. 

Tetradontidae 
Sphoeroides sp. 



In all the fishes listed above, the red myotomal muscle fibers had the distributed 
pattern of innervation. 



MUS. COMP. Zovl 
LfRRARY 



B R E Olt) R A 

UNIVERSITY 

Museum of Comparative Zoology 



US ISSN 0006 9698 



Cambridge, Mass. 30June1982 Number471 

ARTHUR LOVERIDGE — A LIFE IN RETROSPECT 

Ernest E. Williams 1 

On February 16, 1980, at the age of almost eighty-nine, Arthur 
Loveridge, former Curator of Reptiles and Amphibians at the 
Museum of Comparative Zoology, died on the island of St. Helena 
in the South Atlantic after a short illness. 

In 1924 A. Lawrence Lowell, then President of Harvard, wrote to 
the Immigration Department in Boston in these terms: 

"This is to inform you that Arthur Loveridge, Esq.. formerly of 
the Manchester University Museum, National Museum of Wales, 
and latterly Director of the British East African Museum in 
Nairobi, a gentleman standing high in his chosen field, is due on the 
steamer Laconia, arriving in Boston on or about May 1st. 

"On March 14th of this year Mr. Loveridge was appointed by the 
Faculty of the Museum of Comparative Zoology in Harvard 
University to the position of Associate in Zoology in the Harvard 
University Museum, where he will exercise his profession during the 
coming years as an officer of Harvard University. 

"Since I am informed that the British quota is full, I am anxious 
that you should know in advance that Mr. Loveridge is a teacher, 
scientist and author of high professional standing, and that he 
comes here already appointed to a University position in Harvard. 

"Any kindness you may show him in expediting his entry will be 
very greatly appreciated by me." 

It is obvious that President Lowell's plea was effective. It is 
known that Glover Allen, then Curator of Mammals, met Loveridge 



'Museum of Comparative Zoology. Harvard University, Cambridge. Massachusetts 
02138. 



2 BREVIORA No. 471 

at the boat and drove him to Cambridge, where he was to spend 
thirty-three years (till 1957). At first he was Thomas Barbour's 
assistant, reorganizing the Museum's herpetological collection and 
then, when Barbour was appointed Director of the Museum, 
continuing to supervise with surpassing care the expansion of one of 
the world's great collections of reptiles and amphibians. From 1931 
he had the volunteer assistance of Benjamin Shreve. 

Thomas Barbour brought Loveridge to the New World, but 
Loveridge's heart remained in the continent of Africa where he had 
spent almost ten years. There he had made the reputation that 
brought the Harvard appointment. There he had met his wife, and 
there he had indulged to the full his passion for collecting and for 
general natural history that had been his since childhood — indulged 
it despite (or by means of) service with the East African Mounted 
Rifles, the Nairobi Museum, and the Game Department in 
Tanganyika (now Tanzania). 

Loveridge's association with Harvard was no bolt from the blue; 
it was a tie that had gradually strengthened. In the Museum Report 
for 1919-1920, there is a mention of a first gift from Loveridge. His 
name recurs in 1921 1922, and then, in 1923-1924, there is 
Barbour's comment: "This year has been eventful in that during its 
course the Arthur Loveridge African collection was received and 
Mr. Loveridge arrived to assist in a general overhauling of the study 
series." 

Harvard got a bargain in Loveridge. Harvard bought Loveridge's 
collection, but with it came a Curator — one the collection des- 
perately needed. The other side of Loveridge's passion for collecting 
was a passion for order and for tidiness: what he brought back or 
had brought back to him had to be as perfectly classified, ticketed, 
and put away as human power could manage. 

For a while he surely had the best of his two worlds. On the one 
hand, he was in charge of a major but crowded, ill-labeled, ill- 
organized gathering of collections. He was able to transform it into 
a model of collections as he felt they should be — taking frogs, 
snakes, and lizards out of tanks and putting them in fine glass- 
stoppered bottles with labels written in hard pencil in his own neat 
hand and carefully arranging them within trays, each with neatly 
typed labels. His collection, when he finished, was a thing of beauty 
(and fiercely kept so). 



1982 LOVERIDGE'S LIFE IN RETROSPECT 3 

For the other part, in the first years he was repeatedly able to go 
back to Africa and, doing what he most wanted to do, simultane- 
ously enrich the Harvard collections and provide for himself the 
study material he needed. Clearly this had been part of the 
understanding that went with the Harvard appointment. He did 
general collecting, not only herpetological collecting. He had always 
done so, and museum workers in that day were always, whatever 
their specialty, general collectors. He did sometimes collect large 
mammals, but this, I am told, was not to his preference. He did get 
to Africa and to parts of it he had not seen before. In the years 
between 1924 and 1940 he was away from Cambridge four times 
(1925-1926, 1928-1929, 1933 1934, and 1938-1939). On each 
occasion he was away a full year. In terms of his additions to the 
Harvard collections, this was his prime time. 

The first years were active years in many ways. These were years 
of affluence for the Museum. Barbour's money immensely aug- 
mented collections that Louis Agassiz had been at feverish pains to 
acquire. Although Loveridge's African expeditions were undoub- 
tedly the greatest source of additions to the herpetological 
collections during these years, Barbour did not cease, so long as he 
was able, to encourage and directly finance every sort of acquisition 
from any part of the world. This flood of material was certainly 
Loveridge's joy. 

Loveridge was something of a public figure in the first years. He 
routinely gave lectures, wrote articles for "Fauna," "Frontiers," and 
"Natural History" and in 1928 gave a series of twenty lectures for 
the Boston Society of Natural History on Boston's WBET entitled 
"Tales from Tanganyika." He made "Who's Who" in 1938. 

There is much to indicate that the world changed for Loveridge 
after the 30s. The Depression had come; if its impact was not 
immediate, it was fundamental. The concomitant diminution of 
Barbour's fortune meant that the flood of specimens began to come 
to an end. (Loveridge once showed me how plainly this change was 
demonstrated on our species cards.) It was later in this period of 
diminished affluence that Loveridge refused to take more than two 
of a series, offered by Vanzolini, of a species not represented in the 
MCZ collections; "Bottles," he is reported to have said, "are 
precious." For some time the momentum of previous activity 
continued. By 1942, the number of species and subspecies in the 



4 BREVIORA No. 471 

collection surpassed 6,000. The Department had to be enlarged, and 
a new room (the old Aquarium) was taken over for snakes. 

But already in the previous year (1941), Barbour's report as 
Director had begun to take a mournful tone: "Increased taxes are 
going to make it difficult or impossible for the Museum to expect to 
receive the private assistance which it has received in the past." 

In 1942 Shreve left for the army. In 1942-1943 only 400 specimens 
were catalogued; 140 of these were exchanges. Loveridge's own 
collecting suffered also. There was to be only one more African trip. 

Decreased curating and collecting did not impair Loveridge's 
productivity, however. His previous work had been primarily 
reports of collections and faunal studies. He now began revisions, 
and in a popular vein began the series of books that gave him wider 
fame. "Many Happy Days I've Squandered" (1944) was the first. 
"Tomorrow's a Holiday" succeeded it in 1947, then "I Drank the 
Zambesi" (1953), and "Forest Safari" (1956). All included accounts, 
highly entertaining, of his African experiences. 

The end of the Second World War had brought some bonuses. 
Shreve came back. W. H. Stickel, Sergeant Beck, Captain Jarvo, an 
Australian, Gunner Tovell, and others sent to Loveridge material 
from the Pacific area collected during their service. Loveridge had 
written a little book, "Reptiles of the Pacific World" (100,000 copies 
were printed for the Armed Forces, and it has recently been 
reprinted), and these collections were its rewards. Loveridge 
dutifully reported on the collections and on similar material ob- 
tained by the National Museum. 

(It is curious that the most massive of all the acquisitions since 
Loveridge were the result of these activities peripheral to Lov- 
eridge's major interest. Correspondence with Fred Parker in 1960 
was initiated by a request for Loveridge's Australian and New 
Guinean papers and has resulted in the MCZ's now huge Solomons 
and New Guinea collections.) 

Loveridge remained in charge of the collections for almost ten 
years more. He made the last African expedition of his Harvard 
career, that to Nyassaland and Tete in 1948-1949. Thereafter, the 
entire period was devoted to his intended summary of East African 
herpetology, most of it to the series of revisions that he had begun 
earlier, in 1940, with some snake genera, and that culminated with 
the East African Check List published in the year of his retirement, 
1957. One paper on "The Cryptodira of Africa" was in collaboration 






1982 LOVERIDGE'S LIFE IN RETROSPECT 5 

with the man who was to succeed him — myself — published again in 
1957. 

Loveridge left Cambridge in 1957, immediately after his retire- 
ment, for the island of St. Helena in the South Atlantic. Although 
he did visit England, and I once saw him in the British Museum, and 
although we maintained a correspondence and he even published on 
material he sent to the Museum, he never returned to the United 
States. It is not known why — perhaps because the collection was no 
longer his in the special sense that it had been for thirty-three years. 

The Boston Globe of July 21, 1957 headed its four column 
account of Loveridge's retirement with the statement: "Retiring 
Curator to Avoid Work Temptation." If that was genuinely 
Loveridge's intention, it did not turn out that way. He did make his 
retirement home at Varney's on St. Helena, but it often seemed that 
he was only a little less active in these final years than he had been in 
the MCZ. 

He kept up an intense interest in both African herpetology and 
the Museum and in collecting: his letters of 1958 are full of 
impatience to get the tubes to collect St. Helena spiders. His 
correspondence, in fact, began on the boat to the island, and he was 
soon to start numbering his letters. There were already 2,472 in early 
1965, and they were nearing 7,000 at the time of his death. (He was a 
punctilious writer, always answering a letter, but always insisting 
also that his letters be answered before he would write again.) 

He travelled also, not infrequently to England, and at least once 
collected again in Africa — a small collection — Chsmaeleo, Mabuya, 
and frogs from Mau Narok at 9,000 ft. in Kenya, donated to the 
MCZ. He received specimens from correspondents also and 
published on some of them, reporting Hemidactylus mercatorius as 
new to Ascension Island and describing new amphisbaenids 
collected by Ionides in Tanganyika. His most interesting paper from 
his "exile" on St. Helena may be unique in herpetology: his own 
report, published at his own expense, on "The status of new 
vertebrates described or collected by Loveridge." 

His wife died suddenly on St. Helena in 1972. His son Brian 
joined him on the island four years later. In another four years 
Loveridge himself was dead. 

What of the man behind the Curator? He was, of course, a very 
special individual in his own right, but he was also one of a breed 
that is now extinct because the times have made its life style no 
longer viable. 



6 BREVIORA No. 471 

Born in Penarth, Glamorgan, Wales, 28 May 1891, Loveridge was 
thirty-three when he came to the Harvard Museum. He had already 
been Curator in Nairobi and served in museums in Wales and 
England. As he reports in "Many Happy Days I've Squandered." he 
had decided to become a Museum Curator at the age of ten. He tells 
in the preface of that book of "the acquiescence of a kindly father." 
However, the Harvard Archives contains his application in 1914 for 
the newly created post of Curator at the Nairobi Museum. This 
reveals that he had to "serve time" for two years as apprentice in the 
family business of ship furnishing, and that only then was he 
allowed to take a year's course in Zoology and Botany in the 
University College of South Wales on the way to appointments first 
at the Manchester University Museum and then in the Temporary 
Museum in Cardiff. 

It was while he was in the latter post, and, in addition to his 
regular duties, making a card index of the whole British Fauna 
(about, he reports, 23,000 cards), that he serendipitously received 
knowledge of an open position in Africa. Although he already had a 
private collection of "nearly 250 jars of preserved reptiles and over 
300 glass topped drawers containing birds' eggs, insects and other 
specimens," he was always avid for more. When he heard about a 
civil engineer from British East Africa due home on leave who "had 
in his youth shown a fondness for snakes," he tried to inveigle the 
man into collecting for him. Utilizing a joint interest in stamps and 
bribing him with duplicates of these, Loveridge extorted a promise 
to pickle lizards and snakes. An inquiry six months later produced 
an apologetic reply which included the news that the East Africa 
and Uganda Natural History Society needed a curator for a new 
museum that would have government support. "Why don't you 
apply for the post and then you can collect your own bally snakes" 
was the advice. Loveridge applied at once, was accepted, and 
arrived in Nairobi in mid-1914. 

The first World War very soon cast its shadow over Africa; it did 
not interrupt Loveridge's career as a naturalist. Although he joined 
the local forces shortly after his arrival and after six months' 
training was on active duty, it is often difficult, from his account of 
the next four years, to be conscious that a war was on. The 
occasional moment of danger was memorable for him because of 
the capture of a rare animal. His story of the capture of his first 
Boulengerula boulengeri is characteristic: "This rare Caecilian was 



1982 I ()\ FRIDGE'S LIKE IN RETROSPECT 

obtained under rather unusual circumstances during the East 
African campaign. We were busily engaged in 'digging in' under an 
unpleasant shellfire, when it was unearthed by one of my fellow 
troopers in the Mounted Rifles. He humourously called out that one 
of my snakes had escaped and that, if I did not come over and take 
charge of it at once, he would run his bayonet through it. Needless 
to say, when I saw what it was, I very gladly took charge of it." The 
same total devotion to natural history made him- as he himself 
recounts — badger first his sergeant and then the general in the midst 
of his staff for permission to obtain bottles of pickled snakes from 
an abandoned German house. 

He got the snakes; this was no isolated incident. One oi his 
periodical summaries of his activities preserved in the Harvard 
Archives mentions: "Travelling through German East Africa from 
north to south provided exceptional opportunities for collecting . . . 
All necessary preservatives and pickling jars were 'found' in 
captured German towns." (The 'found' is in quotation marks in his 
own typescript.) 

We have here the image of a man wholly devoted to an a\ ocation 
that he made his vocation and who found his life "one long holiday. 
Gratefully I confess to being one of the favored few whose waking 
thoughts in the morning consist of the pleasant planning of the day's 
work." It was he who also wrote: "Probably only a zoologist can 
look at an uncaught cobra and feel the joy a child feels on Christmas 
morning." 

It is this spirit of Loveridge that is well-caught in the cartoon that 
I have chosen to illustrate this memorial of a life. It is a man 1 would 
have liked to have known. It is not, I think, the man I knew. 

I came into Loveridge's ambience late, in 1947, after the Second 
World War, and while I was working on my thesis. Visiting the 
Museum, I was able to re-identify one or two turtles and so gained 
his confidence. This began a cordial relationship. Eventually, after 1 
came to Harvard, I was able to call him "Arthur." (The first level o\ 
intimacy was "Loveridge" rather than "Mr. Loveridge.") 

The man I knew was stiffish. Some called him "Sir Arthur." This 
was probaby mere Englishness plus a firm insistence on standards 
that he did not allow to be relaxed, not for himself, not for anyone. 
Romer called him the "Demon Curator," and this was the aspect 
that most of us knew, who knew him late in his career. 

He could be kind and very helpful. Many of his correspondents. 



8 BREVIORA No. 471 

the visitors to the Department, the young questioner, and even some 
of the anatomists in search of specimens for study knew his kindness 
and assistance well. The more demanding might get short shrift, and 
for poseurs and frauds — so he regarded Ivan Sanderson — he had no 
kindness at all. His review of a book of Sanderson's, and of its 
gentler reviewers, is classic vitriol. 

His tidiness extended to classification. He preferred clarity, was 
unhappy with complication, was impatient of subtlety. He wanted 
problems solved cleanly, once and for all. Therefore he was very 
much a lumper. He was so much a lumper that many of our species 
cards record the species name with an interval between the genus 
and species name — an interval for the eventual insertion of the 
species name of which Loveridge was sure the taxon in question 
could only be a subspecies. (He was very insistent also that 
subspecies be readily recognizable from museum material. He 
reportedly gave K. P. Schmidt the chance to sort out unlabelled 
MCZ specimens into subspecies that K. P. was describing: K. P. 
flunked. It is known that he gave a similar test to Vanzolini on the 
subspecies of Arnphisbaena fuliginosa that Vanzolini passed hand- 
somely. Vanzolini is now not certain that one of his subspecies is 
valid.) 

This fervor for lumping and tidiness sometimes caused disagree- 
ments. I was invited into collaboration with him on the Cryptodira 
of Africa. As he told me, this was partly because I knew turtles, but 
also because I could read German and translate type descriptions. 
That the collaboration succeeded is evidenced by a thickish volume, 
but there were moments of discord. My discussions were too 
theoretical and too verbose, and my taxonomy too splitting. 
Loveridge told me that he had lost a year of his scheduled program 
because of me, and a well-known footnote (softened at the advice of 
his wife) testifies to our taxonomic disagreement. (It was character- 
istic of him that this did not impair a good relationship.) 

I, and others of my time, knew Loveridge only in the Curator- 
facet of his life. We knew him after his last field trip. That part of his 
life had ended. 

But more, I think, was gone by then than just the opportunity for 
year-long field trips. The world had changed. The British Empire 
was diminished if not extinct. Africa had changed, Harvard had 
changed. His chosen profession as naturalist-curator was no longer 
highly regarded at Harvard, or elsewhere. Africa was not the same 



1982 



LOVERIDGE'S LIFE IN RETROSPECT 




Cartoon of Arthur Loveridge from the newspaper East Africa. 



10 BREVIORA No. 471 

land in which Salimu, his favorite No. 1 Boy, had chosen to be 
naturalist-servant to a naturalist-master. 

It is notable that he did not choose to go back to Africa upon his 
retirement. There is a story that his wife chose isolated St. Helena 
because they had once stopped there on their way to Africa in one of 
the two months of the year when the climate is pleasant. But surely, 
even if that were true, the choice involved more than that. Africa 
was no longer the Africa he had loved. 

The man of the later years was not, at least on the surface, the 
man one would expect to write a book with the title "Many Happy 
Days I've Squandered." The man seemed sterner and more prim, 
more New England — as though he had acquired the characteristics 
of the New England breed for whom the verb "to squander" borders 
on obscenity. 

Loveridge's aspect as Demon Curator provoked as much 
astonishment as appreciation. Romer, the new Director after 
Barbour, regarded Loveridge with a respect not unmixed with 
amusement. Their psychologies were nearly antithetical, and 
Romer' s appreciation of Loveridge's value was very incomplete. It is 
true that Loveridge could not have been quite happy with Barbour's 
exuberant and insouciant carelessness, but at least Barbour and 
Loveridge were of one mind about collecting and collections. 
Romer marked the first of the transitions to another museum style. 

The element of fanaticism in Loveridge's neatness quite naturally 
evoked legends. There is a tale that there was in the Department a 
drawer labelled "string too short to use." Neatness and routine were 
at times extreme. Shreve's work counter had to be cleared at 4:30 
when he left. Books had to be put back. Loveridge told both Carl 
Gans and Vanzolini that gaps on the book shelves were to him like 
teeth that had been knocked out. Even the chairs had to be in 
correct positions under the counter. I was reprimanded one 
Monday, when, working over a weekend, I left all three chairs 
improperly aligned. 

Loveridge's fanatic passion for his collection astonished his 
colleagues; clearly he did not fit too well in the new world at 
Harvard. But fanatic attention to detail is a good thing in a curator, 
and certainly the Museum was well served by Loveridge's devotion. 
The organization of the herpetology collection was his and his 
alone; the task that confronted him when he first arrived must have 
been fabulous, and the order he achieved remains an achievement as 



1982 LOVERIDGE'S LIKE IN RETROSPECT 1 1 

great as his African contributions. When I took over the collection, 
all was in perfect shape. There were few curatorial tasks to do: only 
the one collection from the Riu-Kiu Islands that had not yet been 
wholly identified and put away. 

Loveridge served in a University Museum, but he was in no sense 
an academic. He belonged to another generation and another life 
style — he was pre-eminently a collector-naturalist. It is interesting to 
put him in context in the succession of herpetological curators at the 
MCZ. Agassiz must be counted here, but he was clearly unique — a 
European emigre, professor, builder and acquirer of collections, 
intellectual parent to whole generations of natural historians in the 
United States, he is not at all comparable to anyone else. 

Of Garman, the next in line, we know too little, not much beyond 
Barbour's unsympathetic remarks and some plaintive autobio- 
graphical notes of his own. Before the MCZ, he apparently had an 
irregular career; at the MCZ, fide Barbour, he remained isolated 
and apart. Clearly in his day he was useful and respected. He began 
the MCZ's West Indian interest which Barbour so much cultivated 
after him. 

Barbour had, in contrast, the full academic panoply, a doctoral 
degree, and, at least late in life, professorial status. For all that, he 
was throughout his life the Wealthy Amateur, never quite willing to 
go very deep, never quite serious enough to be professional. He 
could be pontifical and very disparaging of others, but many of the 
criticisms could have been turned against him. He had notable 
proteges — G. K. Noble and E. R. Dunn — and was, within American 
herpetology, for a while something of a father figure with equal 
colleagues but no admitted superior. 

Loveridge was very different. British always and a Briton of the 
Empire, he was a man who, without inherited wealth, had chosen, 
very stubbornly, a poorly remunerative career — the career of Bates 
and Wallace, the naturalist-collector. He had chosen also a 
continent. His eyes and his interests turned eastward toward Atrica; 
he was hardly part of American herpetology. For all that in 
Cambridge he was physically close to his American colleagues, it 
was hardly different from what it might have been had he been 
across the sea. If ever he was further west than New York, I have no 
record or report of it. 

In a sense that Barbour was not, Loveridge was a professional. 
Without interest in theory or in biology beyond field and museum 



12 BREVIORA No. 471 

natural history, he was totally professional in what he did — 
completely dedicated to competence in that rather narrow area. He 
never formally taught, and his own formal instruction was limited. 
In herpetology, like -hrs'predecessors (and like myself), he was self- 
taught. (The new curator at MCZ is the first to have had formal 
specific instruction in herpetology.) What he chose to be, he was par 
excellence — Curator-Collector-Naturalist. 

With Loveridge's departure — and his literal departure from 
Cambridge for St. Helena came only a few weeks after his formal 
retirement as Curator -the Department settled into quite another 
style. I was the first curator to be fully a product of Academia — not 
only the holder of the conventional degrees, but one who needed 
them for a living. The world has changed post-Loveridge; the pure 
naturalist-collector is, when he exists at all, an anachronism. 

Arthur and Mary Loveridge's one son, Brian, was schooled at 
Harvard, and had early gone to England for his career. Apart most 
of their lives, Brian joined his father on St. Helena and was building 
a home there near Varney's when his father died. Brian has said of 
his father that his work was his life. That is an affirmation that is 
also a tribute, and the tribute that Arthur Loveridge would have 
most wanted. 

In one of the entryways to the MCZ there is a plaque on the wall 
honoring Alexander Agassiz and with the Latin motto, "Omnia 
quae hie vides monumentum." The Herpetology Department might 
very reasonably display a similar motto in Loveridge's honor. 1 he 
collection's order and style have his imprint. I have added somewhat 
to that collection, but he provided the solid base. 

His taxonomic work is now history; his revisions are now revised. 
It would disappoint him bitterly that this is true, but his passion for 
simplicity is now judged to have gone too far. He was concerned in 
his "Status" paper to learn — certainly with a twinge of heart — which 
of his species had been synonymized. He would view with dismay, if 
not distaste, the sibling species that are now commonplace. He 
would not understand the concerns and disputes of modern 
taxonomists, nor care to. In this sense time has passed him by, but 
his own collections and the collections he so diligently curated are 
his enduring monument. 



,nw * ^OMP, 200L 

'MAR 1 8 1Q«S 

HARVARO 

BREV I'TTR A 

Muse mm of Comparative Zoology 

US ISSN ()()( Hi 9698 
Cambridge, Mass. 30 June 1982 Number 472 



FISHES OF THE SUBORDER LABROIDEI 

(PISCES: PERCIFORMES): 

PHYLOGENY, ECOLOGY, AND 

EVOLUTIONARY SIGNIFICANCE 

Leslie S. Kaufman 1 and Karel F. Liem 2 

Abstract. We postulate that the Pomacentridae, Cichlidae. Embiotocidae. 
Labridae, Odacidae, and Scaridae comprise a monophyletic assemblage: the 
Labroidei. Four groups within the Labroidei can be defined as monophyletic 
assemblages on the basis of shared derived characters: the Pomacentridae. Cichlidae. 
Embiotocidae, and Labridae (which includes the Scaridae and Odacidae). The 
Pomacentridae is considered the primitive sister group of all other Labroidei; the 
Cichlidae is a sister group of embiotocids and labrids, and the Embiotocidae is a 
sister group of the Labridae. Labroids are characterized by (I) united or fused fifth 
ceratobranchials resulting in the formation of one functional unit; (2) a true 
diarthrosis between upper pharyngeal jaws and the basicranium without an 
intervening part of the transversus dorsalis anterior muscle; and (3) the presence of 
an undivided sphincter oesophagi muscle forming a continuous sheet. It is proposed 
that ( 1 ) the ecological and functional versatility of the trophic apparatus is correlated 
with a characteristic structural design, and that (2) this design has contributed to the 
dominant position of labroids in diurnal communities of tropical marine and lentic 
fresh waters. 

INTRODUCTION 

Liem and Greenwood (1981) have recently reviewed the compara- 
tive functional morphology of the pharyngeal jaw mechanism in 
acanthopterygian fishes. On the basis of functional considerations 
they have proposed that the Cichlidae, Embiotocidae, Labridae. 
Odacidae, and Scaridae comprise a monophyletic assemblage. 



'-Museum of Comparative Zoology, Harvard University. Cambridge, Massa- 
chusetts 02138. 



2 BREVIORA No. 472 

Pharyngognathy, as expressed in the Cichlidae, has been correlated 
with several functional and ecological attributes which distinguish 
cichlids from most other Acanthopterygii. First, cichlids can 
mechanically process a broader range of food types (Liem, 1974). 
Second, they have greater feeding versatility (Liem and Osse, 1975; 
Liem, 1980) and hence a broadened fundamental niche (sensu 
Hutchinson, 1958, 1965). Finally, cichlids exhibit an extremely high 
species diversity, perhaps because extinction rates in changing 
environments are decreased (Liem, 1982). To test this hypothesis, 
evolutionary patterns in the Cichlidae must be compared with those 
of related fishes both more and less specialized with respect to 
pharyngeal jaw morphology. Such an analysis requires a thorough 
knowledge of phylogenetic relationships (Lauder, in preparation). 
In this paper we offer a revised hypothesis of the phylogenetic 
relationships of pharyngognath acanthopterygians first proposed by 
Liem and Greenwood (1981). New morphological evidence offered 
here and by Stiassny (1981, personal communication) requires the 
inclusion of the Pomacentridae in the monophyletic assemblage 
Pharyngognathi {sensu Liem and Greenwood, 1981), and a re- 
arrangement of the Embiotocidae as the sister group of the Labridae 
instead of the Cichlidae. The resulting scheme of classification is as 
follows: 

Suborder Labroidei 

Family Pomacentridae 

Family Cichlidae 

Family Embiotocidae 

Family Labridae (including Labridae, Odacidae, Scaridae) 
The proposed phylogenetic relationships provide a basis for 
assessing patterns of change in the feeding apparatus, historical 
consequences of new feeding mechanisms, and the evolution of 
coral reef fish communities. 

MATERIALS AND METHODS 

Morphological studies were conducted with the aid of a Wild-M5 
dissecting microscope and camera lucida. Clearing and staining 
followed the techniques of Taylor (1967). Scanning electron 
microscopy was conducted on an AMR- 1000, and x-ray cineradiog- 
raphy of feeding labrids and cichlids was carried out using the 
Siemens Cineradiographic Unit at 150 frames sec" , The following 
material was examined: 






1982 LABROID PHYLOGENY 3 

Pomacentridae: AbudefcluJ taurus MCZ 42755. Amphiprion xanihurus M( / 
14852. A. percula MCZ 33399, Dascyllm trimaculala MCZ 14837. p. alhisella MCZ 
51671. Eupomacentrus planifrons MCZ 44745. /.. avapukensus MCZ 43961. 
Pomacentrm littoralis MCZ 5794. Chromis atrilobatus MCZ 44640. 

Cichlidae: "Haplochromis" leuciscus MCZ 49517. 

Embiotocidae: Rhacochilus vacca MCZ 57708. Damalichthys vacca MCZ 54333. 
Cymatogaster aggregata MCZ 57707. Phanerodon furcatus MCZ 54334. Embiotoca 
jacksoni MCZ 54332. 

Labridae: Tautogolabrus adspersus uncat.. Tautoga onitis uncut.; Halichoeres 
bivittatum, Scarus croicensis, Sparisoma viride, all MCZ acq. 1981-002-6. 

Caribbean reef fishes were studied in Salt River Canyon and 
Tague Bay, St. Croix, U.S. Virgin Islands, and also studied at the 
Discovery Bay Marine Laboratory, Jamaica. W.I. Ecological 
classifications of reef fishes were based in part on observations made 
from the NULS-1 Hydrolab during mission 81-8. 

ANALYSIS OF CHARACTERS 
Definition of the Labroidei 

We postulate that the Pomacentridae, Cichlidae, Embiotocidae. 
Labridae, Odacidae, and Scaridae comprise a monophyletic lineage, 
the Labroidei. A cladogram defining this group and expressing 
relationships among its major clades has been derived on the basis 
of three investigations: Liem and Greenwood (1981), Stiassny's 
analysis of the phylogenetic relationships of the Cichlidae (in which 
extensive out-group comparisons are described, Stiassny. 1981 and 
personal communication), and this study (Fig. 1). 

All Labroidei share the following three derived characters: (1) 
junction or fusion of the two fifth ceratobranchial bones into a 
single unit, (2) diarthrosis between the upper pharyngeal jaws and 
the basicranium (Fig. 2 A F: APU; Stiassny, personal communica- 
tion), and (3) the presence of the sphincter oesophagi muscle as a 
continuous sheet, with no dorsal subdivision (Fig. 2; Stiassny 
personal communication). Within Perciformes. fused or joined 
lower pharyngeal jaws also occur among the Anabantidae (all), 
Kyphosidae (Girella tricuspidata), and Sciaenidae (Pogonias chro- 
mis and Aplodinotus grunniens). The pharyngeal jaw morphology 
and biting mechanisms of these fishes differ appreciably, however, 
from those of the Labroidei (Liem and Greenwood. 1981). Other 
acanthopterygians show some form of articulation between the 
upper pharyngeal jaws and the basicranium (e.g.. Sparidae. 



BREVIORA 



No. 472 




Figure 1. Cladogram illustrating interrelationships of the major labroid clades. 
Bars represent shared derived characters: (1) United or fused fifth ceratobranchials; 
(2) True diarthrosis between upper pharyngeal jaws and basicranium; (3) Undivided 
sphincter oesophagi muscle; (4) Strong sheet of connective tissue joining lower jaw 
with a ligament which inserts on the ceratohyal bone; (5) Nipple-like bony process 
on ventral surface of lower pharyngeal jaw; (6) Pharyngo-cleithral articulation of 
characteristic form; (7) Obliquus posterior dominant muscle to lower pharyngeal 
jaw; levator externus 4 and obliquus posterior vertically aligned on fourth 
epibranchial, separated by oblique aponeurosis or tendon; (8) Transverus dorsalis 
muscle subdivided into four parts; (9) Premaxillae and maxillae functionally 
decoupled; (10) Cartilagenous cap on anterior border of epibranchial 2; ( 1 1) Micro- 
branchiospinae of characteristic form present on outer faces of second, third, and 
fourth gill arches; (12) As and A« portions of adductor mandibulae complex lacking 
major structural association; insertion of large ventral division of A; onto angulo- 
articular; (13) Head of epibranchial 4 distinctly expanded; (14) Intra-uterine 
development of young with strongly modified vascularized median fins; (15) Mus- 
cular sheet joining Ai and A2.3 portions of adductor mandibulae; (16) Levator 
posterior dominant muscle to the lower pharyngeal jaw, forming a force couple with 
the pharyngocleithralis muscle; (17) Toothplates of fourth pharyngobranchials 
absent (either lost or fused with pharyngobranchial 3), first pharyngobranchials 
absent or reduced; (18) Fourth epibranchials highly modified, articulating with 
upper pharyngeal jaws; (19) True pharyngo-cleithral articulation functioning as 
sliding and hinge joint; (20) Levator externus 4 is a continuous muscle joining 
prootic region to muscular process on lower jaw; (21) Predisposition for insertion of 
levator posterior muscle on lower pharyngeal jaw; (22) Loss of second pharyngo- 
branchial toothplates; (23) First three branchial adductor muscles cover antero- 
dorsal faces of the epibranchials; (24) Ligament connecting postmaxillary process of 
maxilla with anterior border of palatine and ectopterygoid; (25) Tooth rows 
arranged radially across the lower pharyngeal jaw, teeth located directly over the 
symphysis between left and right fifth ceratobranchials. LPJ toothplate composed of 
an anterior, small-toothed field and a posterior, large-toothed pavement replaced by 
addition along the rear margin of the LPJ. 



1982 LABROID PHYLOGENY 5 

Gerreidae, Pogonias, Aplodinotus), but only in the Labroidei is 
there a true diarthrosis. In other perciforms a portion of the 
transversus dorsalis muscle or its aponeurosis passes between the 
apophyses of the upper pharyngeal jaws and basicranium (Stiassny, 
1981). 

Synapomorphies Characterizing the Pomacentridae 

The damselfishes can be defined on the basis of four characters. 
(1) Stiassny (1981: 286) observed that "A strong sheet of connective 
tissue originates from the dorsal border of the bony ridge on the 
medial face of the lower jaw [dentary] and merges with a cylindrical 
ligament that passes posteriorly and inserts onto the ceratohyal 
bone." In other acanthopterygians she examined this ligament was 
wanting. The remaining three characters concern the structure of 
the lower pharyngeal jaw (LPJ). (2) The LPJ's of all pomacentrids 
we have examined bear on their ventral surfaces a pair of small 
nipplelike processes, which serve as the insertion sites for the 
pharyngohyoideus muscle. These processes are absent in all other 
acanthopterygians examined. (3) In primitive acanthopterygians 
there is no contact between the fifth ceratobranchial and the 
cleithrum. In most pomacentrids, however, the muscular processes 
of the LPJ abut upon the cleithrum and slide along it by means of 
articular facets. Two such facets may be present (e.g., Pomacentrus 
littoralis, MCZ 5794): a dorsal facet lying parellel to the dorso- 
ventral plane, and a ventral facet which is curved slightly outwards 
from this plane and may provide the LPJ with some lateral freedom 
of movement. Labrids, in contrast, have a true pharyngo-cleithral 
joint. The degree of pharyngo-cleithral articulation varies con- 
siderably among pomacentrids. Even when the two bones are 
closely related, the nature of the articulation differs from that seen 
in the Labridae (Liem and Greenwood, 1981). This difference is also 
reflected in the unique and complex shape of the muscular processes 
of the LPJ in pomacentrids, a feature related to their peculiar 
musculature (Figs. 2, 3). In some pomacentrids (e.g., Mierospatho- 
dori) pharyngo-cleithral articulation appears to have been lost as 
part of a general reduction of the pharyngeal apparatus. (4) In 
pomacentrids, as in all more primitive perciforms, the fourth levator 
externus muscle (Fig. 2 A-F: LE 4 ) and levator posterior (LP) insert 



BREVIORA 



No. 472 




RD TOP 



Figure 2. Dorsal aspect of the branchial musculature viewed from posterior to 
elucidate the muscles surrounding the esophagus and posterior branchial arches. 
A) Pomacentrus littoralis; B) Abudefduf taurus; C) Tautogolabrus adspersus; 
D) Amphiprion xanthurus; E) " Haplochromis" leuciscus; F) Embiotoca jacksoni. 



Abbreviations: AD, adductor branchialis; APU, apophysis of upper pharyngeal 
jaw (third pharyngobranchial); CB% fifth ceratobranchial (lower pharyngeal jaw, 
LPJ); EB, epibranchial; ES, esophagus; LE, levator externus muscle; LI, levator 
internus muscle; LP, levator posterior muscle; OD, obliquus dorsalis muscle; OP, 
obliquus posterior muscle; PB, pharyngobranchial; RD, retractor dorsalis muscle; 
SE, sphincter oesophagi muscle; TDS, transversus dorsalis anterior muscle; TDP, 
transversus dorsalis posterior muscle. 



[982 



LABROID PHYLOGENY 




a. < i 



8 BREVIORA No. 472 

on the dorsal aspect of the fourth epibranchials (Liem, 1974). 
However, the pomacentrid LE4 (Fig. 2A,B,D) splits near its 
insertion site on the fourth epibranchial (EB 4 ). The larger head 
inserts near the insertion site of LP, while a smaller medial head 
joins an oblique aponeurosis that runs anteromedially along the 
border of EB4. The obliquus posterior (OP) is the major muscle 
operating the posterior region of the LPJ. The OP muscle runs 
between the broad dorsal flange of each muscular process and the 
flange along the EB 4 just below the insertion site of LE 4 (Fig. 2A, B, 
D). Here the LE4 and OP muscles are separated by the oblique 
aponeurosis. Thus the sites of origin and insertion for OP in 
pomacentrids are the same as the other acanthopterygians, but the 
extremely close relationship between LE 4 and OP on the fourth 
epibranchial is a derived character. The aponeurosis is present in all 
pomacentrids examined, and it clearly separates the fibers of the 
LE 4 from those of the OP. Nevertheless, the insertions for the two 
muscles are extremely close together, a condition which seems to 
foreshadow the muscular sling of cichlids and other Labroidei. 
There is a broad flange on EB 4 to receive LE 4 , and the posterior 
flanges on EB 4 and the fifth ceratobranchial (CBs) are vertically 
aligned. Between them, and possibly contributing to the mechanical 
linkage between the two, is the stout dorsal portion of CB 4 . Thus, 
while the muscles of the LPJ in pomacentrids still originate and 
insert on the same bones as in non-labroid acanthopterygians, the 
geometry of these insertions has been modified in a characteristic 
fashion. The resulting condition may represent a primitive counter- 
part to the cichlid muscular sling (Liem, 1974). 

Synapomorphies Characterizing the Cichlidae 

Six characters clearly distinguish the Cichlidae as a monophyletic 
assemblage: (1) The transversus dorsalis muscle is subdivided into 
four parts (Liem and Greenwood, 1981; Fig. 2E:TDA). (2) The 
premaxillae and maxillae of all cichlids are functionally decoupled; 
i.e., premaxillary protrusion can be regulated independently from 
motion of the maxillae by means of multiple mechanical pathways 
(Liem, 1978, 1979). Four additional characters have been described 
by Stiassny ( 1 98 1 ): (3) There is an extensive cartilaginous cap on the 
anterior border of EB2 (Cichla ocellaris being the exception). (4) 
Microbranchiospinae of characteristic form are present on the outer 



1982 LABROID PHYLOGENY 9 

faces of the second, third, and fourth gill arches. (5) The A: and A u 
portions of the adductor mandibulae complex have lost a major 
structural association, and there is an insertion of a large ventral 
division of A: onto the angulo-articular. (6) The head of EB4 is 
distinctly expanded. 

Synapomorphies Characterizing the Embiotocidae 

Two characters are considered synapomorphies for the Embio- 
tocidae. ( 1 ) All embiotocids are fully viviparous fishes, which deliver 
large, well-developed young. The males have a small intromittent 
organ derived from anal fin rays. The young are usually closely 
packed in the ovarian sacs which function in the fashion of a uterus. 
Monophyly of the embiotocids can be established on the basis of the 
highly specialized mode of "intra-uterine" development with its 
associated structural, physiological, and behavioral features, invol- 
ving spatulate extensions of the webs of median and caudal fins, 
each with a rich blood supply (Webb and Brett, 1972). (2) Stiassny 
(1981) has found a small sheet of parallel muscle fibers arising from 
the anteromedial region of the Ai part of the adductor mandibulae 
muscle and inserting upon the dorsal aponeurosis of A2+3 part of this 
muscle. 

Synapomorphies Characterizing the Labridae 

The families Labridae, Odacidae, and Scaridae have been 
recognized as close relatives within the Labroidei (Greenwood et ai, 
1966). Here the three groups are recognized as a single family, the 
Labridae, since the monophyletic nature of this assemblage is 
strongly indicated by both morphological and functional characters. 
(1) The levator posterior muscle (Fig. 2C:LP) is the dominant 
muscle of the LPJ, forming a force couple with the pharyngo- 
cleithralis externus muscle (Liem and Greenwood, 1981). (2) The 
toothplates of the fourth pharyngobranchials are absent (either lost 
or fused with pharyngobranchial 3; Stiassny, 1981), while the first 
pharyngobranchials are absent or reduced. (3) The fourth epi- 
branchials are highly modified and of characteristic form, articulat- 
ing with the upper pharyngeal jaws (Yamaoka, 1978, 1980; Gobalet, 
1978). (4) There is some form of physical contact between the LPJ 
and the cleithrum in all members of the clade (Liem and 
Greenwood, 1981). 



10 BREVIORA No. 472 

The posterior face of the muscular process on each side of the 
LPJ bears an articular facet, which fits against a small fossa on the 
cleithrum. The mechanics of this joint are quite complex. Cine- 
radiography of the LPJ in Tautogolabrus adspersus during mastica- 
tion shows that there is a biphasic pattern involving both a hingelike 
and a sliding movement. The dynamics are reminiscent of those in 
the human temporo-mandibular joint. At rest and during the bite 
the LPJ is in close contact with the cleithrum and exhibits the 
pattern of a hinge-joint. During protraction the LPJ slides down 
and forward, using the anterior face of the cleithrum as a track. The 
morphology of the pharyngocleithral joints of odacids and scarids is 
basically similar to that of labrids, though the support system is 
stronger, and it is unknown whether the LPJ disarticulates during 
protraction. The condition in odacids and scarids overlaps broadly 
with that of the Labridae. Crypt otomus roseus is a scarid with a 
labridlike LPJ; both it and Nicholsina denticulata have many 
wrasselike features (see Gobalet, 1980 for comparative discussion of 
Nicholsina). Pseudodax mollucanus is a labrid with a scaridlike LPJ 
(Bleeker, 1862). 

Interrelationships Among the Labroidei 

The relationships proposed here differ from those of previous 
investigators (Greenwood et al., 1966; Nelson, 1967; Liem and 
Greenwood, 1981) in three important ways. The Pomacentridae is 
considered the primitive sister group of other Labroidei. The 
Embiotocidae is postulated to be the sister group of the Labridae, 
and not the Cichlidae. Finally, the Labridae, Scaridae, and 
Odacidae are united in a single family, Labridae, to reflect striking 
similarities in their morphological as well as functional specialized 
features (Fig. 1). 

Chief evidence for the proposed primitive sister-group relation- 
ship of the Pomacentridae to other Labroidei is found in the 
osteology and myology of the LPJ. In the Cichlidae, Embiotocidae, 
and Labridae, the LE 4 and (in general) LP muscles insert on the 
LPJ. The LE4 and LP muscles of Pomacentridae insert on the 
fourth epibranchials, which represents the primitive condition in the 
Perciformes. Aerts and Verraes (1982) have demonstrated that the 
LE4 of the cichlid Haplochromis elegans (and presumably, other 
cichids as well) is actually a composite of LE4 plus a large medial 
head of the obliquus posterior muscle, the two fusing during 



1982 LABROID PHYLOGENY 11 

development. This condition is never fully developed in Poma- 
centridae (Fig. 2A, B, D: OP, LE4). Obliquus posterior (OP) is the 
dominant muscle to the LPJ while LE 4 remains a separate muscle, 
though it is large, and is aligned with the OP. The result is that the 
two muslces together bridge the gap between prootic and LPJ, 
meeting across the fourth epibranchial, where they are separated by 
an aponeurosis (Fig. 2A, B, D: OP, LE 4 ). 

Four derived characters establish sister-group relationship be- 
tween the Embiotocidae and Labridae. (1) These groups have lost 
the second pharyngobranchial toothplates (Nelson, 1967; Stiassny 
1981). (2) The first three branchial adductor muscles cover the 
anterodorsal faces of the epibranchials (Stiassny, 1980 and Fig. 2C, 
F: ADi + :). (3) A ligament connects the postmaxillary process of the 
maxilla with the anterior border of the palatine and ectopterygoid 
(Stiassny, 1980). (4) Tooth rows are arranged radially across the 
LPJ, with teeth located directly over the symphysis between left and 
right fifth ceratobranchials. The LPJ toothplate is composed of two 
fields: an anterior field (often lost in durophages), and a posterior 
field bearing the large cardinal teeth used in crushing or grinding. 
New cardinal teeth are added only to the rear margin of the LPJ, 
forming a conveyor-like pavement with greatest wear toward the 
front (Fig. 3; see also Embiotocidae: Damalichthys vacca; Labridae: 
Pseudodax mollueanus; all scarids). 

The phylogeny proposed in Figure 1 assumes that similarities in 
pharyngeal dentition between the Labridae and Embiotocidae on 
the one hand, and Pomacentridae on the other, are homoplasies. In 
cichlids the LPJ toothplate is divided into left and right regions, 
with the largest teeth arranged in two main rows parallel to the 
symphysis, and there are no • teeth located directly over the 
symphysis. This is clearly the primitive condition, since (a) the LPJ 
of labroids is derived from two separate fifth ceratobranchial bones, 
and (b) this condition is displayed in those non-labroid perciforms 
possessing fused or joined fifth ceratobranchials. In pomacentrids, 
as well as in embiotocids and labrids, the teeth cross the plate in a 
radial series and there are teeth located directly over the symphysis. 
As a sole synapomorphy for pomacentrids, embiotocids, and labrids 
this character seems too tentative. It appears independently in the 
Beloniformes, another group with fused lower pharyngeal jaws, 
which is clearly unrelated to the Labroidei (Collette, 1966). 
Pharyngocleithral joints appear in both the Pomacentridae and the 



12 BREVIORA No. 472 

Labridae, but are clearly dissimilar in form. Thus in terms of the 
specialized osteological and myological characters discussed above, 
the pharyngeal jaws of the Cichlidae, Embiotocidae, and Labridae 
resemble each other much more closely than any one of these groups 
resembles the Pomacentridae. 

Independent investigations (Stiassny, 1981; Liem and Green- 
wood, 1981, and this study) have resulted in very similar hypotheses 
of labroid phylogenetic relationships. Stiassny (1981, personal 
communication) has based her studies primarily on soft-tissue 
characters, while we have concentrated on functional and osteo- 
logical characters. The relationships postulated in Figure 1 deviate 
drastically from previous schemes. This is mainly because important 
new osteological, dental, and myological evidence has emerged. 

DISCUSSION 

Ecology of the Labroidei 

The new phylogenetic scheme of the Labroidei has important 
implications for our perception of the ecology and functional 
morphology of this group. The gradal nature of former classifica- 
tions obscures relationships and thereby masks evolutionary se- 
quences, ecological diversity, and changes in functional patterns. To 
illustrate this point we will briefly discuss some of the implications 
of the new phylogeny for ecological concepts as they pertain to the 
Labroidei. 

Labroidei as defined here unites at least 1,470 species (some 5 to 
10 percent of living fishes) that are extremely diverse ecologically. 
Nevertheless, the majority of labroids occur within one general type 
of environment: warm, slow-moving water with abundant habitat 
structure. Tropical marine reefs are densely populated by pomacen- 
trids and labrids. These are joined by embiotocids on temperate 
Pacific reefs. All four clades figure prominently in aquatic 
macrophyte forests such as grass beds, kelp beds, algal reefs, or 
heavily vegetated pond and stream edges. Relatively few labroids 
are abundant in pelagic, soft-bottom, or strictly lotic assemblages. 

Modes of life exhibited by marine labroids differ characteris- 
tically from those of sympatrically occurring non-labroid and 
functionally intermediate forms. We illustrate these patterns with 
data from a coral reef fish assemblage observed in Salt River 
Canyon, St. Croix, U.S. Virgin Islands (Table 1). The 137 species 



1982 LABROID PHYLOGENY 13 

Table 1. Relationship between functional morphology of the pharyngeal jaws 
and potential anti-predator mechanisms for 137 species of coral reef fishes observed 
during visual censuses in Salt River Canyon, St. Croix, U.S. Virgin Islands 
(Kaufman and Ebersole, in preparation). 



Potential 




Functional 


Morphology of 


Pharyngeal Jaws 


Anti-Predator 










Mechanisms 




3 rimitive 


Intermediate 


Labroid 


Non-territorial 


O 


= 52 


O = 


10 


O = 5 


Non-schooling 


E 


= 33.74 


E = 


18.09 


E = 15.16 


Not heavily armed 


X : 


= 9.88 


X : = 


3.62 


X : = 6.81 




o 


= 16 


o = 


17 


O=0 


Heavily armed 


E 


= 16.62 


E = 


8.91 


E = 7.47 




X : 


= 0.02 


X : = 


7.35 


X : = 7.47 


Territorial or 





= 1 


O = 


10 


O = 26 


Schooling 


E 


= 18.64 


E = 


9.99 


E = 8.37 


Not heavily armed 


X : 


= 16.69 


X : = 


0.00 


X : = 37.13 




X 2 


= 88.98. p 


< .001 







O = observed frequencies for numbers of species in each category. 

E = expected frequencies. 

X : = Chi-square value within cell. 

X 2 — total Chi-square. 

observed during two series of replicated visual censuses (Kaufman 
and Ebersole, in preparation) were divided into three categories 
according to pharyngeal jaw functional morphology: (1) primitive, 
with pharyngeal jaws unspecialized for mastication; (2) intermediate 
forms exhibiting some, but not all of the features found in Labroidei 
(as discussed below in greater detail); and (3) labroids. The species 
were also placed into three other categories related to strategies for 
avoiding predation (refuging): (1) non-territorial and non-habitually 
schooling species, both unarmed; (2) strongly territorial and 
habitually schooling species, both unarmed; and (3) species armed 
with frank defensive mechanisms (toxin, venom, dermal armor, 
enlarged spines), or which live inside corals, sponges, invertebrate 
tests, and boreholes. It was postulated that trophic mechanisms and 
refuging strategies would be interrelated. Table 1 provides evidence 
of highly significant relationship between "pharyngeal jaw" and 
"anti-predation" categories (X 2 = 88.98; p < .001 ). A detailed list of 
the species and their categorizations can be obtained from the 
authors. 



14 BREVIORA No. 472 

The data in Table 1 suggest that there are characteristic modes of 
life for coral reef labroids. Most of them both feed and refuge on the 
reef. They are not heavily armed. They are, for the most part, either 
territorial or schooling. Territorial labroids defend a general area as 
a multipurpose territory rather than occupying one specific hole or 
cavity as do many inquiline gobies, blennies, or jawfishes (Opis- 
thognathidae). The residents exclude potential competitors for 
food, hiding places, and mates, as well as species that threaten the 
integrity of the territory (Low, 1971; Thresher, 1976; Kaufman, 
1977, 1979; Potts, 1977; Williams, 1979, 1980; Ebersole, 1977; 
Lobel, 1980). Even the schooling and planktivorous labroids are 
strongly reef-associated, utilizing benthic cover as the ultimate 
means of escape from predators. In summary, members of the 
Labroidei generally rely on the reef for refuge, exploiting those 
foods which are, or can be made available without travelling over 
long distances. There is an exception to this pattern. Adults of the 
larger species (such as the huge Caribbean parrotfishes Scarus 
guacamaia and S. coelestinus) sometimes forage or migrate to and 
from the reef as individuals. For these fishes, large size alone may be 
a sufficient deterrent against predators. 

Other reef-dwelling fishes exhibit modes of life that contrast with 
those of the labroids. The non-labroid category in Table 1, including 
such fishes as berycoids, apogonids, serranids, and lutjanids, 
consists of both diurnal .and nocturnal predators which stalk small 
soft-bodied prey. The intermediate pharyngeal jaw category is 
dominated by two groups: armed fishes that feed on the reef by day 
and rest on the reef at night, and unarmed fishes that feed off-reef at 
night and shelter on the reef during the day. The heavily armed 
plectognaths, chaetodontids, and acanthurids comprising the bulk 
of the first group together exploit almost as broad a range of foods 
as the labroids. Individually, however, they exhibit functional 
limitations related to gape, mouth position, jaw mobility and 
pharyngeal jaw mechanics that should seriously limit feeding 
versatility in comparison to that of similar-sized pomacentrids or 
labrids. This is reflected in what is known of their diet (e.g., Randall, 
1967; Hobson, 1974; Reese, 1975). Relatively few of these species 
enhance their own local food supply by defending feeding territories 
(possibly Acanthurus sohal, Vine, 1974; Chaetodon trifascialis 
[formerly Megaprotodon strigangulus], Reese, 1975). The nocturnal 



1982 LABROID PHYLOGENY 15 

off-reef predators (e.g., Pomadasyidae) prey chiefly on small 
benthic invertebrates. 

We postulate that feeding versatility was a chief factor in shaping 
the characteristic modes of life exhibited by marine labroids; i.e., 
schooling or territorial behavior with a strong reliance on the reef 
for both food and shelter. Trophic mobility can be one key to 
survival when spatial mobility is limited by a high risk of predation. 
The antithesis of this strategy, rarely exhibited by labroids, is to 
reduce the risk of predation by adopting some active defensive 
mechanism. 

Labroids play a disproportionate role in determining the dis- 
tribution and abundance of benthic organisms in tropical marine 
hard-bottom communities (Randall, 1961, 1974; Ogden and Lobel, 
1978; Brock 1979). In part this is due to broad-spectrum feeding 
capabilities (hard-shelled invertebrates, coral rock, coral, algae). 
Many labroids locally manipulate the substratum and its occupants 
to suite their own needs (Brawley and Adey, 1977; Kaufman, 1977, 
1979; Wellington, 1981). This constitutes a patchy disturbance to 
sessile invertebrates (Kaufman, 1977; Connell, 1978) and could be a 
principal factor regulating food abundance for other reef organisms. 

Labroid Phylogeny and the Evolution of 
Acanthopterygian Feeding Mechanisms 

When the cichlid pharyngeal jaw mechanism was first described, 
it appeared to represent an abrupt breakthrough in the acantho- 
pterygian feeding mechanism, radically different from anything 
known in the cichlids' presumed ancestors (Liem, 1974). Subsequent 
radiation seemed to involve little modification of the basic feeding 
mechanism (Fryer and lies, 1972; Greenwood, 1974). The new 
hypothesis on the genealogical relationships of the Labroidei 
presented here requires that these views be revised considerably. 

One erroneous hypothesis was that the pharyngeal jaw complex is 
unique to cichlids; present evidence rejects such a hypothesis (Liem 
and Greenwood, 1981). Many features present in cichlids are 
present in Labroidei. Second, there was thought to be a large 
morphological and functional gap between cichlids and their 
primitive (ancestral) counterparts; there is not. The apparent gap 
was an artifact of insufficient data, now bridged by primitive 
labroids and certain non-labroid perciforms. Aerts and Verraes 
(1982) have shown that during the ontogeny of a cichlid (Astatoti- 



16 BREVIORA No. 472 

lapia elegans) the LE 4 splits into a lateral and a medial head. The 
medial head of LE 4 unites with the medial head of the OP, thus 
establishing a functionally as well as a structurally uninterrrupted 
muscle between the prootic and the muscular process of the LPJ, 
i.e., a compound LE 4 . The characteristic arrangement of LE 4 and 
OP in pomacentrids (e.g., Pomacentrus lit (oralis, Fig. 2) resembles 
those early ontogenetic stages in cichlids before the compound 
muscle is formed. The Pomacentridae is clearly primitive, however, 
in terms of the origins and insertions of LE 4 and OP. Thus the 
Pomacentridae is intermediate between the more derived labroids 
(cichlids, embiotocids, and labrids) and other perciforms. 

Other perciforms which approach the labroid condition in one or 
more respects are morphologically (and perhaps phylogenetically) 
intermediate between labroids and primitive perciforms. The 
anabantoids, Kyphosidae, and Sciaenidae have fused or joined 
pharyngeal jaws in some members. The fifth ceratobranchials of the 
Gerreidae and Pomadasyidae (especially Anisotremus surinamen- 
sis) have no true bony junction, but are in some species very tightly 
bound together by strong ligaments. Certain Sciaenidae (e.g., 
Pogonias chromis, Aplodinotus grunniens) and Gerreidae have an 
articulation between the upper pharyngeal jaws and the basicranium 
although it is not as well developed as in labroids. The shell-cracker 
centrarchid Lepomis microlophus and the molluscivorous Carangi- 
dae {Trachinotus spp.) have broad, hypertrophied lower pharyngeal 
jaw elements which meet closely at the midline (Kaufman and Ono, 
in preparation). Nearly all of these morphologically intermediate 
forms feed habitually on hard-shelled benthic invertebrates in 
addition to a wide variety of other organisms, both hard and soft. 
Selection favoring a broader, more inclusive diet could have been a 
major factor in the early evolution of labroids. This hypothesis can 
not be tested without first developing a better picture of perciform 
phylogeny. However, the presence of so many intermediate forms 
(one of which may represent the primitive sister group of the 
Labroidei) and the intermediate characteristics displayed in Poma- 
centridae, indicate that advanced acanthopterygian pharyngeal jaws 
are the result of a series of morphological changes. There was no 
single "adaptive breakthrough" {sensu Simpson, 1944, 1953; Liem, 
1974). 



1982 LABROID PHYLOGENY 17 

In summary, this new model of labroid relationships will permit 
us to examine the nature of evolutionary change in a structurally 
complex mechanical system. Judging from the great ecological 
diversity of labroids, it seems that'their specialized pharyngeal jaw 
apparatus has greater structural potential and functional flexibility 
than that of its more primitive counterparts. The more precise 
labroid phylogenetic scheme will allow us to determine if there is a 
general relationship between design versatility and historical pat- 
terns of morphological change. 

ACKNOWLEDGMENTS 

We owe much to the useful comments made by William and Sara 
Fink, Humphry Greenwood, and Dana Ono. Melanie Stiassny 
contributed generously from her own observations on labroid 
systematics. Karsten Hartel aided in locating study material. Ed 
Seling and Al Coleman assisted in electron microscopy and 
photography. William B. Hamner, Jr. made important contribu- 
tions to studies of the pharyngeal jaw in labrids, and helped in 
collecting specimens; William McFarland contributed many embio- 
tocid specimens. We thank the staffs of the West Indies Marine 
Laboratory and the Discovery Bay Marine Laboratory for assist- 
ance in specimen collection and field observations. This work was 
supported by National Science Foundation Grant DEB-79-00955 to 
K. Liem; National Oceanographic and Atmospheric Administra- 
tion Grant NULS-1 Mission 81-8 to L. Kaufman and J. Ebersole; 
and National Science Foundation Grant OCE 79-12674 to J. B.C. 
Jackson. This paper is contribution No. 261 from the Discovery Bay 
Marine Laboratory, Discovery Bay, Jamaica, W.I. 

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18 BREVIORA No. 472 

Brock, R. E. 1979. An experimental study of the effect of grazing by parrotfishes 

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Soc. Lond. 189: 93-125. 
1980. Adaptive significance of intra- and interspecific differences in the 

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1982 LABROID PHYLOGENY 19 

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Vine, P. J. 1974. Effects of algal grazing and aggressive behavior of the fishes 
Pomacentrus lividus and Acanthurus sohal on coral reef ecology. Mar. Biol., 24: 
131-136. 

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Embiotoca lateralis. J. Fish. Res. Bd. Canada, 29: 1525-1542. 

Wellington, G. M. 1981. The role of competition, niche diversification and 
predation on the structure and organization of a fringing coral reef in the Gulf of 
Panama. Ph.D. Dissertation, University of California, Santa Barbara. 

Williams, A. H. 1979. Interference behavior and ecology of the threespot dam- 
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1980. The threespot damselfish: a noncarnivorous keystone species. 

Amer. Natur., 116: 138-142. 

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1980. Some pharyngeal jaw muscles of Calotomus japonicus (Scaridae, 

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_;,.;> AV/^O 






LIBRARY 



B R E 



MAR 1 8 1985 

SR) R A 




II 



seiim of Comparative Zoology 



US ISSN 0006 9698 



Cambridge, Mass. 



29 April 1983 



Number 473 



THE INTERRELATIONSHIPS OF PELYCOSAURS 

Donald Brinkman 1 and David A. Eberth 2 

Abstract. An analysis of 17 characters forms the basis of a hypothesis of 
relationships of seven pelycosaur genera. These genera include representatives from 
most of the higher taxa recognized by Romer and Price ( 1940). In agreement with the 
phylogeny proposed by Romer and Price, Edaphosaurus is considered to be more 
closely related to Dimeirudon than is Ophiacotton. In contrast to the phylogeny 
proposed by Ronur and Price, Ophiacudun is considered more closely related to 
Dimeirudon than are Varanops and Aerosaurus. Three character-states that are 
interpreted as being derived are shared by Casea, Varanops, and Aerosaurus, 
suggesting that these three genera are members of a clade distinct from the clade 
including Ophiacodon, Edaphosaurus, and Dimeirudon. 

INTRODUCTION 

Pelycosaurs occupy a central position in amniote evolution. As a 
paraphyletic taxon within the clade including mammals, pelyco- 
saurs have played an important role in considerations of the origin 
of mammals. Also, pelycosaurs include some of the most primitive 
known reptiles, and are an important element in consideration of 
the early evolution of amniotes. Thus, an understanding of pelyco- 
saur interrelationships has implications for many broader problems 
of reptile diversification. 

Pelycosaurs were the subject of a detailed monographic study by 
Romer and Price ( 1940), and as a result are one of the best under- 
stood groups of Paleozoic reptiles. Romer and Price used evolu- 
tionary systematics in their study of pelycosaurs (Fig. 1A). In line 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 

02138. Present Address: Tyrrell Museum of Paleontology, Box 7500. Drumheller. 

Alberta, Canada. TOJ OYO. 

"Department of Geology, University of Toronto, Toronto, Ontario, Canada, M5T 

IR7. 



BREVIORA 



No. 473 




EOAPHOSAURIOAE 



LuPEOSAURIDAE 



NIT03AURIDAE 



SPHENACODONTIA v. EDAPHOSAURIA 

EOTHYRIDIDAE \ / OPHI ACODONTIDAE 





Figure I. Hypotheses of pelycosaur interrelationships. A) Phylogeny of pely- 
cosaurs presented by Romer and Price (1940); B) Cladogram showing the inter- 
relationships of pelycosaurian families presented by Reisz ( 1980). A is from Romer 
and Price, 1940; B is from Reisz, 1980. 



with this approach, taxa were established on the basis of phenetic 
similarity. This is reflected in the diagnoses, which consist of gene- 
ralized descriptions that encompass the anatomy of most of the 
members of the group. The taxonomic rank assigned to a group was 
based on both the diversity within the assemblage and the morpho- 
logical distance between that and other groups. Ecological interpre- 
tations also played a significant role in their study of pelycosaur 
evolution, and a scenario uniting the morphological and ecological 
interpretations was developed. 



1983 PELYCOSAUR INTERRELATIONSHIPS 3 

Some aspects of the phylogeny proposed by Romer and Price 
were brought into question by Langston (1965), who proposed a 
close relationship between Oedaleups and Eothyris, and argued that 
these genera were close to the ancestry of caseids. This implied that 
caseids were not closely related to the edaphosaurids, as was sug- 
gested by Romer and Price. 

Recently, Reisz (1980) reviewed pelycosaur interrelationships 
using a cladistic analysis of the characters available to Romer and 
Price. Reisz concluded that many of the families recognized by 
Romer and Price were monophyletic, but the interrelationships of 
the families that Reisz proposed (Fig. IB) differed from those sug- 
gested by Romer and Price. Also, the relationships that Reisz rec- 
ognized did not conform to the scenario of pelycosaur evolution 
proposed by Romer and Price. 

During the study of a new pelycosaur from El Cobre Canyon, 
New Mexico, it became clear that much additional morphological 
information could be brought to bear on the problem of pelycosaur 
interrelationships, and a review of the group was undertaken. The 
results of this review are intended to provide a testable hypothesis of 
pelycosaur interrelationships, and to serve as a framework in which 
detailed taxonomic and morphological revisions of individual gen- 
era and families can be interpreted. 

MATERIALS AND METHODS 

The central problem in a cladistic analysis is establishing the 
polarity of character-states. Out-group comparison has been consid- 
ered the most powerful tool for this (Watrous and Wheeler, 1981). 
The interrelationships of the groups used in determining the polarity 
of character-states within pelycosaurs are shown in Figure 2A. 
These relationships are based on Carroll ( 1970), except for the posi- 
tion of Diadectes, which, following Heaton (1980), is placed with 
Limnoscelis and Tseajaia in the Diadectomorpha. Also, following 
Kemp ( 1980), pelycosaurs are considered to be the sister-group of all 
other reptiles. 

In using out-group comparison to interpret polarities, two princi- 
ples are used. One is the principle of parsimony: the hypothesis of 
polarities that requires the least number of evolutionary events to 
account for the distribution of character-states will be accepted. 



BREVIORA 



No. 473 



PRIMITIVE OUT-GROUPS 



IN-GROUP 



SISTER-GROUP 



Anthrocosoria 



Diadectomorpho 



Pelycosauno ail other reptiles 







primitive sister- 

out-groups Pelycosauna group 



primitive 

out-groups Pelycosauno 



sister- 

group 





primitive 

out-groups Pelycosauno 



sister- 
group 



primitive 

out-groups Pelycosauno 



sister- 
group 




primitive 

out-groups Pelycosauna 





1983 PELYCOSAUR INTERRELATIONSHIPS 5 

This principle is the basis tor the interpretation of polarities for the 
character-states whose distribution is shown in Figure 2B-E. 

The second principle is the co-occurrence of primitive character- 
states. According to this principle, primitive character-states tend to 
occur together. Using this principle, the character-state that occurs 
in the outgroup that, on the basis of other evidence, is thought to be 
more primitive is accepted as the primitive character-state. This is 
the basis for the interpretation of polarities for the character-states 
whose distribution is shown in Figure 2F and G. It shoud be noted 
that implicit in this principle is the assumption that reversals are less 
likely to occur than are independent acquisitions. The use of this 
principle, therefore, limits the extent to which the results presented 
here can be used to test this assumption. 

Characters of uncertain polarity will not be used to support rela- 
tionships. However, such characters may provide corroboration of 
relationships proposed on the basis of derived character-states, since 
the distribution of uncertain character-states will be either consis- 
tent with the cladogram or will require that the occurrence of parallel 
evolution be hypothesized. 

One of the problems in the study of the evolution of groups 
represented by fossil material is the incorporation of data both from 
well-known animals and from animals represented by incomplete 
specimens. This is especially true in the study of the evolution of 
early reptiles, where material is rare and, in many cases, animals are 
represented by single fragmentary specimens. Often animals repre- 
sented by such material have been assigned to a taxon erected on the 
basis of well-known animals, and discussions of the ev olution of the 
group have focused on the interrelationships of the higher taxa. 



Figure 2. The interpretation of polarities ol character-states by outgroup compari- 
son. A) Cladogram showing the interrelationships of the taxa used as out-groups 
lor interpreting the polarity of character-states within pelycosaurs; Diadectomorpha 
includes Diadevtes, Linmoscelis. and Tseijaia; B G) Interpretation of polarities of 

character-states. In B E. this is based on parsimony, in F and G, this is based on the 
principle of co-occurrence ol primitive character-states. ■ represent the derived 
in-group character-states; □ represent the primitive in-group character-states; repre- 
sent character-states not present in the in-group. 



6 BREVIORA No. 473 

In order that information from both fragmentary and well-known 
animals can be incorporated into the analysis of pelycosaur inter- 
relationships, this study will be separated into two steps. The first 
will be to construct a cladogram showing the interrelationships of 
the most completely known genera using all available morphological 
features. This will provide a framework in which data from the less 
well known genera can be interpreted. The results of this part of the 
study are presented here. The second part will be the inclusion of the 
less well known animals in the analysis. Through this, the clad- 
ogram will be tested and a more detailed understanding of pelyco- 
saur interrelationships will be obtained. 

By using genera as the basis for considering pelycosaur interrela- 
tionships, all morphological diversity known to be present within 
higher taxa will be incorporated in a single analysis of diversifica- 
tion of pelycosaurs. Also, assumptions about pelycosaur evolution 
are minimized. Some currently recognized higher taxa are almost 
certainly monophyletic and could be treated as units in a discussion 
of pelycosaur interrelationships. Others, such as the Ophiacodon- 
tidae, as currently defined may be either polyphyletic or paraphyletic. 
By treating each genus as an independent entity, this potential 
source of error is avoided. 

The pelycosaurs considered here are Casea, Varanops, Aero- 
saurus, Ophiacodon, Edaphosaurus, Dimetrodon, and Sphenacodon. 
The restriction of the study to these genera was necessitated by the 
accessibility of material for direct observation and the fragmentary 
nature of available specimens of other genera. Only characters that 
were actually observed were utilized in the analysis, and many of the 
reconstructions presented here are modified from previously pub- 
lished reconstructions on the basis of observations made during the 
course of this work. This required that a large amount of material be 
examined^and supplying a complete list of specimens studied is not 
possible. To facilitate future work, the specimens that showed par- 
ticular structures most clearly are listed in Table 1, and the speci- 
mens that are the basis for the modifications in the reconstructions 
are listed in the figure legends. In all the features considered here, 
Sphenacodon and Dimetrodon are indistinguishable (Eberth, 1981). 
Thus, illustrations from only one of these genera are included here. 



1983 



PELYCOSAUR INTFRRELAI IONSH1PS 



Table 1. List of 


specimens p 


reserving the 


structures d 


escribed in the text. 








Ophia- 


Edapho- 


Dime- 


Character 


Casea 


Varanups 


codon 


saurus 


trodon 


Supraoccipital 


UC698 





MCZ 1366 


MCZ 1762 


MCZ 1347 


Tabular 


UC698 





MCZ 1426 


MCZ 1762 


MCZ 1347 


Basipterygoid 












tubercula 


UR 1011 





MCZ 4920 


MCZ 1531 


MCZ 1697 


Stapes 


UC698 


P 12841 


MCZ 1350 


MCZ 1762 


MCZ 1347 


Quadrate ramus 












of pterygoid 


UC698 


P 12841 


MCZ 1350 


MCZ 1762 


MCZ 5950 


Basisphenoid shelf 


UC 698 


UR 2423 


MCZ 4820 


MCZ 1531 


MCZ 1697 


Frontal 


UC 656 


UR 2423 


MCZ 1366 


MCZ 4309 


MCZ 4430 


Maxilla 


UC698 


UR 2423 


MCZ 1366 


MCZ 1762 


MCZ 1347 


Premaxilla 


UC698 


MCZ 1926 


MCZ 1366 


MCZ 1680 


MCZ 4982 


Quadratojugal 


UC656 


U R 2423 


MCZ 1366 


MCZ 1762 


MCZ 6173 


Pterygoideus process 





MCZ 1926 


MCZ 1203 


MCZ 1370 


MCZ 3244 


Angular 


UC698 


UR 2423 


MCZ 1203 


MCZ 1370 


MCZ 7539 


Vertebrae 


UC 883 


MCZ 1926 


MCZ 5912 


MCZ 1754 


MCZ 5210 


Humerus 


UC656 


UR 695 


MCZ 1486 


MCZ 3417 


MCZ 8708 


Postparietal 


UC698 








MCZ 1762 


MCZ 1347 


Paroccipital process 


VC 656 


UR 2423 


MCZ 1426 


MCZ 1762 


MCZ 1347 



Abbreviations: UC. UR, and P: 
History; MCZ: specimens housed 



specimens housed in the Field Museum of Natural 
in the Museum of Comparative Zoology. 



CHARACTER ANALYSIS: I. FEATURES FOR WHICH 
POLARITY CAN BE INTERPRETED 

Ventral Margin of Skull 

In Dimetrodon, Sphenacodon (Fig. 3F), Edaphosaurus (Fig. 3E), 
and Ophiacodon (Fig. 3D), the ventral margin of the cheek region 
of the skull is concave (i.e, bowed upward). 

In Casea (Fig. 3B), Varanops (Fig. 3C), and Aerosaurus (Lang- 
ston and Reisz, 1981), the ventral margin is convex and extends 
below the level of the tooth row. 

The ventral margin of the cheek is convex in Limnoscelis (Romer, 
1946), Diadectes (Fig. 3 A), and anthracosaurs (Panchen, 1970). In 
Hylonomus, the cheek margin is straight (Carroll, 1964). Thus, 
three character-states can be recognized: the presence of a concave 
ventral cheek margin, a convex ventral cheek margin, and a straight 



BREVIORA 



No. 473 



ventral cheek margin. They have the distribution shown in Figure 
2C, with the derived character-state present in pelycosaurs being the 
presence of a concave cheek margin. The condition seen in Hylu- 
nomus is a separate, derived character-state. 




Figure 3. The skulls in right lateral view of A) Diadectes; B) Casea; C) Vara- 
nops; D) Ophiacodon; E) Edaphosaurus; and F) Sphenacodon. Drawing of Dia- 
dectes based on MCZ 1739. MCZ 1736, and MCZ 2086; Casea based on UC 698 and 
UC 656; Varanops based on MCZ 1926 and UR 2423; Ophiacodon based on UC 671; 
Edaphosaurus based on MCZ 1762, MCZ 1764, and USNM 299844; Sphenacodon 
from Eberth (1981). 



Abbreviations: U frontal; j, jugal; la, lacrymal; max, maxilla; na, nasal; pf, post- 
frontal; pmx. premaxilla; po, postorbital; prf, prefrontal; pt, pterygoid; q, quad- 
rate; qj, quadratojugal; sq, squamosal; st, supratemporal. 



1983 PELYCOSAUR INTERRELATIONSHIPS 9 

Premaxilla 

In Dimetrodon, Sphenacodon (Fig. 3F), Edaphosaurus (Fig. 3E), 
and Ophiacodon (Fig. 3D), the anterior margin of the premaxilla 
slopes posteriorly from the anterior termination of the tooth row 
giving a convex outline to the snout. 

In Casea and Varanops (Fig. 3B -C), the anterior margin of the 
premaxilla first extends anteriorly from the anterior termination of 
the tooth row, giving a forwardly projecting rostrum. Langston and 
Reisz (1981) argue that the forward sloping premaxilla seen in 
Aerosaurus specimen UCMP 40096 is a result of displacement of 
the element, and they reconstruct the skull with a posteriorly sloping 
premaxilla. However, apart from the slope of the premaxilla, there 
is no direct evidence that the premaxilla has been displaced, and no 
evidence that the structure of this part of the skull was different than 
in Varanops. 

In Diadectes, the dorsal process of the premaxilla is vertical (Fig. 
3A). Limnoscelis is like Varanops, Casea, and Aerosaurus in having 
a forwardly projecting rostrum (Romer, 1946). Anthracosaurs, with 
an elongate, flattened face, are not directly comparable to either 
condition seen in pelycosaurs. The structure of the premaxilla in 
Hylonomus and Paleothyris is not known for certain. In Romeria, 
the dorsal ramus of the premaxilla is nearly vertical (Heaton, 1979), 
a condition not specifically similar to either of the character-states 
present in pelycosaurs. Thus, these character-states have the distri- 
bution shown in Figure 2E, with the presence of a forwardly sloping 
dorsal ramus of the premaxilla being the primitive condition. 

Frontals 

In Dimetrodon and Sphenacodon (Fig. 4E), the frontal has a 
laterally directed lappet that is greater in mediolateral width than is 
the posterior end of the frontal. Edaphosaurus was figured as hav- 
ing a broad flange extending laterally along the entire anterior half 
of the frontal by Romer and Price (1940). However, isolated front- 
als show that the lateral flange illustrated by Romer and Price is 
formed in part by the prefrontal, and that a lateral lappet like that of 
Dimetrodon and Sphenacodon is present (Fig. 4D). 

In Ophiacodon (Fig. 4C), the frontal is without a strongly devel- 
oped lateral lappet. A small projection extends between the base of 



10 



BREVIORA 



No. 473 






1983 PELYCOSAUR INTERRELATIONSHIPS 11 

the prefrontal and postfrontal, but the width of this is much less 
than the width of the posterior end of the frontal. 

In Casea (Fig. 4A), Varanops (Fig. 4B), and Aerosaurus (Lang- 
ston and Reisz, 1981), frontal lappets are absent. 

Frontal lappets are absent in Limnoscelis (Romer, 1946), Dia- 
dectes (Lewis and Vaughn, 1965), anthracosaurs (Panchen, 1970), 
and Paleothyris (Carroll, 1969). Thus, these character-states have 
the distribution shown in Figure 2B, with the absence of a frontal 
lappet being the primitive character-state. 

Maxilla 

In Dimetrodon, Spenacodon (Fig. 3F), Edaphosaurus (Fig. 3E), 
and Ophiacodon (Fig. 3D), the maxilla does not extend posterior to' 
the orbit and does not meet the quadratojugal. In Varanops (Fig. 
3C) and Aerosaurus (Langston and Reisz, 1981), the maxilla 
extends posterior to the orbit and meets the quadratojugal, exclud- 
ing the jugal from the ventral margin of the skull. In cross section, 
the jugal slopes laterally so that a ridge is present at the contact of 
the jugal and maxilla. In Casea the maxilla meets the quadratojugal, 
excluding the jugal from the ventral margin of the skull (Fig. 3B). A 
ridge is present along the contact of the maxilla and jugal as in 
Aerosaurus and Varanops. 

Thus two structural patterns are present: the Dimetrodon pattern 
in which the jugal enters the ventral margin of the skull and no ridge 
is present at the contact of the jugal and maxilla, and the ( aranops 
pattern in which the jugal is excluded from the ventral margin of the 
skull by a contact between the maxilla and quadratojugal and a 
ridge is present along the contact of the maxilla and jugal. Diadectes 
(Fig. 3A), Limnoscelis (Romer, 1946). Paleothyris (Carroll, 1969). 



Figure 4. The skulls in dorsal view of A) Casea: B) Varanops; C) Ophiacodon; 
D) Edaphosaurus; and E) Sphenacodon. Reconstruction of Casea based on LJC 656 
and UC 698; Varanops based on MCZ 1926 and UR 2423: Ophiacodon based on 
MCZ 1366; Edaphosaurus based on MCZ 1762 and USNM 299844; and Sphena- 
codon from Eberth (1981 ). 

Abbreviations: f, frontal; j, jugal; la, lacrymal; na, nasal; op, opisthotic; p. pari- 
etal; pf, postfrontal; prf, prefrontal; so, supraoccipital; sq. squamosal; tab. tabu- 
lar. 



12 BREVIORA No. 473 

and anthracosaurs (Panchen, 1970) are like Dimetrodon in these 
features. Thus, these character-states have the distribution shown in 
Figure 2B, with the absence of a contact between the maxilla and 
quadratojugal and the absence of a ridge along the contact of the 
maxilla and jugal being the primitive character-state. 

Quadratojugal 

In Dimetrodon and Sphenacodon (Fig. 3F), the quadratojugal is 
a small bone sitting on the posterolateral corner of the quadrate and 
is without an anterior zygomatic process. 

Edaphosaurus was reconstructed with a large quadratojugal 
(Romer and Price, 1940), but specimen MCZ 1762 shows that a 
small quadratojugal like that of Dimetrodon was present (Fig. 3E). 

In Ophiaeodon (Fig. 3D), Varanops (Fig. 3C), Aerosaurus (Lang- 
ston and Reisz, 1981), and Casea (Fig. 3B), the quadratojugal 
extends forward from the posterior corner of the skull forming the 
ventral border of at least the posterior half of the cheek. 

In Diadeetes (Fig. 3 A), Limnoseelis (Romer, 1946), anthraco- 
saurs (Panchen, 1970), and Paleothyris (Carroll, 1969), the quadra- 
tojugal is a large element extending well anteriorly. Thus, these 
character-states have the distribution shown in Figure 2B, with the 
presence of a large quadratojugal being the primitive condition. 

Quadrate Ramus of Pterygoid 

In Dimetrodon (Fig. 5F), Sphenacodon, and Edaphosaurus (Fig. 
5E), the quadrate ramus of the pterygoid is a vertical sheet with a 
rounded ventral edge. In Ophiaeodon (Fig. 5D), Varanops (Fig. 
5C), and Casea (Fig. 5B), a shelf extends medially from the ventral 
edge of the quadrate ramus, flooring the lateral most part of the 
tympanic cavity. The quadrate ramus of the pterygoid in Diadeetes 
has a similar shelf (Fig. 5A), as does Limnoseelis (Romer, 1946). In 
anthracosaurs, a tympanic shelf is not present, but the quadrate 
ramus of the pterygoid is little differentiated from the more anterior 
part of the bone (Panchen, 1970), and is not comparable to the 
quadrate flange of pelycosaurs in structural details. Paleothyris is 
like Dimetrodon and Edaphosaurus in not having a medially 
directed tympanic flange. Thus, these character-states have the dis- 
tribution shown in Figure 2G, with the presence of a medially 



1983 



PELYCOSAUR INTERRELATIONSHIPS 



13 




Figure 5. Ventral view of the posterior half of the skulls of A) Diadectes; 
B) Casea; C) Varanops; D) Ophiacodon; E) Edaphosaurus; F) Dimetrodon. Re- 
construction of Diadectes based on MCZ 1739 and MCZ 2042; Casea based on UC 
698; Varanops based on P 12841; Ophiacodon based on MCZ 1366 and UC 1638; 
Edaphosaurus based on MCZ 1762; and Dimetrodon based on MCZ 1365, MCZ 
2168. and MCZ 5950. 



Abbreviations: bo, basioccipital; bsp, pasisphenoid; ec, ectopterygoid; ept, epi- 
pterygoid;j, jugal; op, opisthotic; pt, pterygoid; q, quadrate; sq, squamosal. 



14 BREVIORA No. 473 

directed shelf flooring the tympanic cavity being the primitive 
character-state and the absence of this flange being a derived 
condition. 

Suprauccipital 

In Dimetrodon, Sphenacodun (Fig. 6D), Edaphusaurus (Fig. 
6C), and Aerosaurus (Langston and Reisz, 1981), the supraoccipital 
has an area dorsal to the posttemporal fenestra forming the dorsal 
margin of the posttemporal fenestra, termed the lateral process by 
Romer and Price (1940). In Ophiacodon (Fig. 6B), the supraoccipi- 
tal is without an ossified lateral process. Romer and Price inter- 
preted the absence of a lateral process as a result of the tendency for 
poor ossification in that genus, and reconstructed cartilaginous lat- 
eral processes. Reisz (1980) accepted this hypothesis and showed a 
lateral process of the supraoccipital bordering the posttemporal 
fenestra in his reconstruction of the occiput of Ophiacodon. While 
cartilaginous lateral processes of the supraoccipital may have been 
present, these would not have been visible in posterior view since the 
tabular of Ophiacodon, as illustrated by Romer and Price (1940), 
contacts the opisthotic and has a finished ventral surface that would 
have formed the dorsal edge of the posttemporal fenstra. In Casea 
(Fig. 6A), the arrangement of the supraoccipital and tabular is like 
that of Ophiacodon: the supraoccipital is not exposed above the 
posttemporal fenestra when seen in occipital view, and the tabular 
contacts the paroccipital process, excluding the supraoccipital from 
the margin of the posttemporal fenestra. 

In Diadectes,' the elements of the occiput are fused. In Linmosce- 
lis, the relationships of the tabular are like those of the Ophiacodon 
(Romer, 1946). Paleothyris is without lateral processes on the 
supraoccipital, and the dorsal margin of the posttemporal fenestra is 
formed by .the tabular (Carroll, 1969), as in Ophiacodon. However, 
in contrast to that genus, Paleothyris has a large posttemporal 
fenestra that is bounded medially by the supraoccipital, and thus is 
not distinctly like either condition seen in pelycosaurs. Thus, these 
character-states have the distribution shown in Figure 2E, with the 
exclusion of the supraoccipital from the border of the posttemporal 
fenestra by the tabular being the primitive character-state, and the 
presence of a lateral process of the supraoccipital forming the dorsal 
border of the posttemporal fenestra being a derived character-state. 



1983 



PELYCOSAUR INTERRELATIONSHIPS 



15 



Basipterygoid Pr< > t esses 

The basipterygoid processes of Dimetrodon (Fig. 7G), Sphena- 
codon, and Edaphosaurus (Fig. 7F) extend anteriorly, ventrally, 
and laterally from the lateral surface of the basisphenoid. Each 
articular surface is divided into two areas, a flat anteroventrally 
facing area and a dorsally facing area at nearly 90 degrees to this. 




Figure 6. Occipital view of the skulls of A) Casea; B) Ophiacodon; C) Edapho- 
saurus; and D) Sphenacodon. Drawing of Casea based on UC 698 and UC 656; 
Ophiacodon based on MCZ 1366. MCZ 1426, and MCZ 1 121; Edaphosaurus based 
on MCZ 1762; and Sphenacodon from Eberth (1981). 



Abbreviations: bo, basioccipital; eo, exoccipital; op, opisthotic; par, parietal; 
pp. postparietal; q, quadrate; qj, quadratojugal; sq, squamosal; tab, tabular. 



16 



BREVIORA 



No. 473 



In Ophiacodon, the basipterygoid processes extend ventrally (Fig. 
7E). The long axis of the articular surfaces are anteroposteriorly 
oriented and the surface curves smoothly from its ventral to its 
lateral aspect. 

In Casea (Fig. 7C), the basipterygoid processes extend laterally 
from the side of the cultriform process and have strongly curved 
articular surfaces. Aerosaurus (Langston and Reisz, 1981) is similar 
to Casea, but differs in that the articular surfaces are more elongate 
mediolaterally. 

Thus, two distinct characters can be recognized in the structure of 
the basipterygoid process. One of these is the curvature of the artic- 
ular surfaces: the curved articular surfaces such as are seen in Aero- 
saurus being one character-state and two flat articular surfaces set at 




Figure 7. The basisphenoid in ventral view of A) Archeria. B) Diadectes, 
C) Casea, D) Varanops, E) Ophiacodon, F) Edaphosaurus, and G) Dimetrodon. 
Drawing of Archeria based on MCZ 8736; Diadectes based on MCZ 3277; Casea 
based on UC 698; Varanops based on UR 2423; Ophiacodon based on UC 1638 and 
MCZ 4820; Edaphosaurus based on MCZ 1762; and Dimetrodon based on MCZ 
1697. 



Abbreviations: bs s, basisphenoid shelf; bs w, basisphenoid wings; bst tub, basi- 
pterygoid tubercula. 



1983 PELYCOSAUR INTERRELATIONSHIPS 17 

nearly 90 degrees to each other such as in Dimetrodon being a 
second character-state. In Diadectes (Fig. 7B), Archeria (Fig. 7A), 
and Eocaptorhinus (Heaton, 1979), the basipterygoid processes are 
curved. Thus, for the character of the curvature of the articular 
surfaces, the character-states have the distribution shown in Figure 
2B, with the presence of curved articular surfaces being the primitive 
character-state and the flat articular surfaces of Dimetrodon, Sphe- 
nacodon, and Edaphosaurus being a derived character-state. 

The second character is the orientation of the basipterygoid pro- 
cesses: the laterally oriented basipterygoid processes such as is seen 
in Aerosaurus being one character-state, anteroventrally oriented 
processes such as is seen in Dimetrodon being a second character- 
state, and ventrally oriented basipterygoid processes with antero- 
posteriorly oriented articular surfaces such as is seen in Ophiacodon 
being a third character-state. In all the outgroups, the basipterygoid 
processes extend ventrolaterally from the side of the cultriform pro- 
cess as in Dimetrodon. Thus, for the character of the orientation of 
the basipterygoid processes, the character-states have the distribu- 
tion shown in Figure 2B, with the presence of ventrolaterally 
oriented basipterygoid processes being the primitive character-state. 
The laterally orientated basipterygoid process seen in Casea and 
Aerosaurus is one derived character-state, and the ventrally oriented 
basipterygoid process with anteroposteriorly oriented articular sur- 
faces of Ophiacodon is a second derived character-state. 

Shelf Between Basisphenoid Wings 

In Dimetrodon (Fig. 7G), Sphenacodon, and Edaphosaurus, the 
basisphenoid wings extend posteriolaterally from the base of the 
basipterygoid tubercula with a smooth trough between them. In 
Ophiacodon, the arrangement of the basisphenoid wings is similar, 
but a shelf (bs.s., Fig. 7E) extends between the base of the wings, 
roofing over the anterior end of the trough. This is also the case in 
Varanops (Fig. 7D) and Aerosaurus (Langston and Reisz, 1981). In 
Casea (Fig. 7C), a basisphenoid shelf is present, although its poste- 
rior edge, rather than being straight, is V-shaped with the lateral 
ends of the V raised. 

In Limnoscelis (Romer, 1946), a shelf is present roofing the ante- 
rior portion of the trough between the basisphenoid wings. In Dia- 
dectes, this shelf is hypertrophied and extends to the posterior ends 



18 



BREVIORA 



No. 473 



of the basisphenoid wings so that, in ventral view, the shelf appears 
to occupy the position of the wings (Fig. 7B). The anthracosaur 
Archeria is like Limnoscelis in having a shelf covering the anterior 
portion of the trough between the basisphenoid wings (Fig. 7A). 
Paleothyris is without such a shelf. 

Thus, these character-states have the distribution shown in Figure 
2F, with the presence of a shelf between the bases of the basis- 
phenoid wings being the primitive character-state, and the absence 
of the shelf being a derived condition. 

Stapes 

The stapes of Dimetrodon (Fig. 8E) consists of a triangular plate 
oriented anteromedially with a posteromedially oriented footplate 
(fp.. Fig. 8E) directed about 90 degrees from the medial edge of the 
triangular plate. The wide end of the triangle is the dorsal process 
(dp.. Fig. 8E), and the narrow end is the quadrate process (qp.. Fig. 
8E). The dorsal process articulates in a socket on the ventral surface 
of the paroccipital process. 

The stapes of Edaphosaurus (Fig. 8C) differs from that of Dime- 
trodon only in proportions, the dorsal process being relatively wider 




Figure 8. The left stapes in posterior view of A) Casea; B) Varanops; C) Ophia- 
codon; D) Edaphosaurus; E) Dimetrodon. Drawing of Casea based on UC 698; 
Varanops based on UR 2423; Ophiacodon based on MCZ 1366; Edaphosaurus based 
on MCZ 1762; and Dimetrodon based on MCZ 1347. 



Abbreviations: dp, dorsal process; fp, footplate; qp, quadrate process. 



1983 PELYCOSAUR INTERRELATIONSHIPS 19 

and the quadrate process shorter. The dorsal process articulates in a 
socket on the paroccipital process as in Dimetrodon. 

In Ophiacodon (Fig. 8C), the stapes is more rodlike than Dime- 
trodon. The dorsal process of the stapes is narrower, although it still 
articulates in a socket on the paroccipital process, and the footplate 
is oval, rather than circular in end view as in Dimetrodon and 
Edaphosaurus. 

In Casea (Fig. 8A), the stapes consists of a footplate and a single 
triangular flange of bone extending laterally from this. One corner 
of the flange approaches the paroccipital process and presumably 
represents the base of the unossified dorsal process. The other 
corner is directed towards the stapedial pit of the quadrate. A dis- 
tinct socket in the paroccipital process for the dorsal process of <the 
stapes is not present. 

The stapes of Varanops (Fig. 8B) differs from that of Casea in the 
presence of a V-shaped notch in its lateral edge. The edges of this 
notch are damaged, and therefore the notch may be articicial. In 
other regards, the stapes is directly comparable to that of Casea. As 
in Casea, no socket is present on the paroccipital process for the 
dorsal process of the stapes. 

The stapes of Aerosaurus is not known, but the paroccipital pro- 
cess shows no articular surface for the dorsal process of the stapes 
(Langston and Reisz, 1981). 

The stapes of Paleothyris (Carroll, 1969) appears most similar to 
the rodlike laterally directed stapes in captorhinids (Heaton, 1979), 
in which the dorsal process does not articulate in a socket on the 
paroccipital process. The stapes of Diadectes, as described by Olson 
(1966) is not readily comparable to that of any other tetrapod. A 
stapes is not known in Limnoscelis or anthracosaurs. In the early 
temnospondyle Greererpeton, the stapes is similar to that of Vara- 
nops and Casea in the presence of a triangular lateral process that 
does not have an ossified dorsal head articulating in a socket on the 
paroccipital process (Carroll, 1980). Since Greererpeton and pelyco- 
saurs are both without an otic notch, probably primitively so (Lom- 
bard and Bolt, 1979; Carroll, 1980), the similarity of the stapes in 
these animals can be used as evidence that the stapes of Casea and 
Varanops is primitive in not having a socket on the paroccipital 
process that receives the dorsal process of the stapes. 



20 



BREVIORA 



No. 473 



Prearticular 

In Dimetrodon (Fig. 9C), Sphenacodon, and Edaphosaurus, the 
prearticular underlies the pterygoideus process of the articular, a 
distinctive twisting of the bone being present as it passes under the 
process. 

In Ophiacodun (Fig. 9A), the prearticular underlies the medial 
portion of the articular but does not show the twisting seen in 
Dimetrodon. This is also the case in Varanops (Fig. 10) and Aero- 
saurus (Langston and Reisz, 1981). The condition in Casea is 
unknown. 




ptpr 



Figure 9. The right lower jaw in medial view of A) Ophiacodon; B) Edapho- 
saurus; C) Dimetrodon. From Romer and Price, 1940. 



Abbreviations: ang, angular; ar, articular; prear, prearticular; pt pr, pterygoi- 
deus process; sur ang, surangular. Arrows indicate the position of sections shown in 
Figure 1 1. 



1983 



PELYCOSAUR INTERRELATIONSHIPS 



21 





pt p 
C 



Figure 10. The lower jaw of Varanops in A) ventral view; B) medial view of 
articular region; and C) section through articular and pterygoideus process. Draw- 
ings based on MCZ 1926. 

Abbreviations: pt pr, pterygoideus process; ret ar pr, retroarticular process. 



Didaectes is without the twisting of the prearticular, as are cap- 
torhinids (Heaton, 1979), and anthracosaurs (Panchen, 1970). Thus, 
these character-states have the distribution shown in Figure 2B, 
with the absence of a twisted prearticular being the primitive 
character-state. 

Angular 

In Dimetrodon (Fig. 11E) and Sphenacodon the angular is a 
vertically oriented plate that supports the prearticular and surangu- 
lar by its dorsal edge (Fig. HE), and the articular by its medial 
surface. The posterior edge of the angular is notched, forming the 
reflected lamina of the angular. 

The angular of Edaphosaurus (Fig. 1 ID) and Ophiacodon (Fig. 
1 1C) is like that of Dimetrodon in being a vertically oriented plate. 
They differ in that they are without a posterior notch. 



22 BREVIORA No. 473 



D E 




Figure 1 1 . Sections through the postdentary bones of the lower jaw at the posi- 
tion just posterior to the dentary (marked by an arrow for the jaws shown in Figure 
9). A) Varanops; B) Casea; C) Ophiacodon; D) Edaphosaurus; and E) Dime- 
trodon. Drawing of Varanops based on UR 2423; Casea based on UC 698; Ophia- 
codon. Dimetrodon and Edaphosaurus from Romer and Price, 1940. 

Abbreviation: ang, angular. 

In Casea (Fig. 1 1 A), Varanops (Fig. 1 1 B), and Aerosaurus (Lang- 
ston and Reisz, 1981), the angular is a troughlike bone. In Vara- 
nops, a keel is present on its ventral edge, but in no position along 
the angular does the bone form an extended vertical plate. 

The angular in Limnoscelis and Diadectes is like that of Casea 
and Varanops in being troughlike. In Paleothyris, a keel is present 
in the region of the angular, but the bone does not form an extended 
vertical plate. Thus, these character-states have the distribution 
shown in Figure 2B, with the absence of an extended vertical plate 
being the primitive character-state. 

Pterygoideus Process of Articular 

In Dimetrodon (Fig. 9C), Sphenacodon, and Edaphosaurus (Fig. 
9C), a pterygoideus process is present as a distinct process on the 
articular. In primitive species of Dimetrodon, and in Sphenacodon 
and Edaphosaurus, this is located medial to the glenoid. 

In Ophiacodon (Fig. 9 A) a distinct pterygoideus process is not 
present on the articular. A flange on the prearticular just anterior to 
the articular may be functionally equivalent to the pterygoideus 
process of Dimetrodon. 

In Varanops (Fig. 10) and Aerosaurus (Langston and Reisz, 
1981), a pterygoideus process is not present on the articular, 



1983 PELYCOSAUR INTERRELATIONSHIPS 23 

although a well-developed flange on the prearticular just anterior to 
the articular is present. The condition in Casea is unknown. 

In Diadectes, the articular does not have a distinct pterygoideus 
process. The articular extends medially from the inner surface of the 
jaw (Romer, 1956; Fig. 107C), but the medially projecting portion is 
located beneath the medial condyle of the gleniod and is not a 
distinct process. No flange is present on the prearticular anterior to 
the articular. This is also the case in anthracosaurs (Panchen, 1970). 
The internal surface of the articular is not known in Paleothyris. In 
captorhinids, a pterygoideus process is absent or poorly developed, 
although a medial flange formed by the prearticular just anterior to 
the articular is present (Heaton, 1979). Thus, these character-states 
have the distribution shown in Figure 2D, with the absence of a' 
pterygoideus process being primitive for tetrapods, the presence of a 
flange on the prearticular just anterior to the articular being primi- 
tive for reptiles, and the presence of a pterygoideus process formed 
by the articular being a derived feature within pelycosaurs. 

Neural Arch 

In Dimetrodon (Fig. 12F) and Sphenacodon, the neural arch has 
a pit in its lateral surface above the level of the transverse process. In 
Edaphosaurus boanerges (Fig. 12E), the neural arch is without pits, 
although in an undescribed species of Edaphosaurus from near 
Garnett, Kansas, Sphenacodon-Wkt pits are present along the length 
of the column (Reisz et al, 1982). Varanops (Fig. 12C) and Aero- 
saurus (Langston and Reisz, 1981) are like Dimetrodon in having a 
pit in the lateral surface of the neural arch. Ophiacodon (Fig. 12D) 
and Casea (Fig. 12B) are without a pit in the lateral surface of the 
neural arch. 

The neural arches of Limnoseelis, Diadectes, anthracosaurs, and 
Paleothyris are without pits. Thus, these character-states have the 
distribution shown in Figure 2B, with the absence of pits in the 
neural arch being the primitive character-state. 

Transverse Processes 

In Dimetrodon (Fig. 12F) and Sphenacodon, the longest trans- 
verse processes are in the area of the posterior cervicals and anterior 
dorsals. They extend laterally a distance about equal to the width of 
the centrum. 



24 



BREVIORA 



No. 473 








Figure 12. Anterior dorsal vertebrae in lateral and anterior view, and a cross 
section of an anterior dorsal centrum of A) Labidosaurus; B) Casea; C) Varanops; 
D) Ophiacodon; E) Edaphosaurus; and F) Dimetrodon. Drawing of Labidosaurus 
based on MCZ 8724; Casea based on P 12841; Varanops based on MCZ 1926; 
Ophiacodon based on MCA 5912; Edaphosaurus based on MCZ 1359; and Dime- 
trodon based on MCZ 5216. 






Abbreviations: pit, pit in the lateral surface of the neural arch; vent flange, ventral 
flange supporting the transverse process. 



1983 PELYCOSAUR INTERRELATIONSHIPS 25 

In Edaphosaurus, the longest transverse processes are shorter than 
the width of the centrum and the transverse processes are braced by 
a vertical flange of bone ventrally. 

Ophiacodon (Fig. 12D) has short transverse processes directed 
more ventrally than in Edaphosaurus and Dimetrodon. The ventral 
flange forms a complete web of bone connecting the transverse 
proces to the anterior edge of the centrum. 

Varanops (Fig. 12C), Aerosaurus (Langston and Reisz, 1981), 
and Casea (Fig. 12B) have long laterally directed transverse pro- 
cesses. Limnoscelis, Diadectes, anthracosaurs, and Paleothyris all 
have short transverse processes, although only in captorhinids (Fig. 
12A) are these supported by a web of bone that extends to the 
anterior edge of the centrum. Thus, for the character of the trans- 
verse processes, the character-states have the distribution shown in 
Figure 2D, with the presence of short transverse processes con- 
nected to the anterior edge of the centrum by a web of bone being 
the primitive character-state for reptiles, and the long transverse 
processes being a derived character-state present within pelycosaurs. 

Centrum 

In Dimetrodon (Fig. 12F) and Sphenacodon, the vertebrae in the 
cervical and anterior to mid-dorsal regions of the vertebral column 
have well-developed keels. 

In Edaphosaurus (Fig. 12E), keels are present only on the cervical 
vertebrae. The dorsal vertebrae have broadly rounded ventral 
surfaces. 

In Ophiacodon (Fig. 12D), the cervical vertebrae are keeled. The 
mid-dorsal vertebrae are wedge-shaped in cross section but have a 
pair of ridges ventrally, rather than a single keel. The posterior 
dorsal and lumbar vertebrae are circular in end view. 

In Varanops (Fig. 12C), the cervical and mid-dorsal vertebrae are 
keeled. 

In Casea the most anterior cervical vertebrae are unknown. All 
the more posterior vertebrae are without keels (Fig. 12B). In cross 
section, the centra have flatter ventral surfaces than in Edapho- 
saurus. 

The centra of Diadectes and Limnoscelis are not specifically sim- 
ilar to any pelycosaur. In cross section, they are generally wedge- 
shaped, but are without a distinct keel. The vertebrae of Protero- 



26 



BREVIORA 



No. 473 



2°ndge 







Figure 13. The humerus in distal ventral view of A) Casea; B) Aerosaurus; 
C) Varanops; D) Ophiacodon; E) Eiiaphosaurus; and F) Dimetrodon. Drawing of 
(Ta.ypfl from Williston (1911), drawing of Aerosaurus based on UC 464; Varanops 
based on UR 695; Ophiacodon based on MCZ 1486; Edaphosaurus based on MCZ 
3417; Dimetrodon based on MCZ 1304. 



Abbreviations: delt, deltopectoral crest; ect, ectepicondyle; ect f, ectepicondyl- 
car foramen; ent f, entepicondylar foramen; pect fossa, pectoralis fossa; sup, supi- 
nator process; 2° ridge, secondary pectoralis ridge. 



1983 PELYCOSAUR INTERRELATIONSHIPS 27 

gyrinus an anthracosaur with a protoreptilian vertebral pattern 
(Holmes and Carroll, 1977), are without keels. This is also the case 
in captorhinids (Fig. 12A). Thus, the absence of a distinct keel can 
be considered a primitive feature. 

Deltopectoral Crest 

In Sphenacodon and Dimetrodon (Fig. 13F), the deltopectoral 
crest has a bulbous distal end and a sharp knife-edge base. In Ophia- 
codon and Edaphosaurus, the distal end of the deltopectoral crest is 
bulbous, but the proximal end is a broad ridge (Fig. 13 D-E). 

In Varanops (Fig. 13C), Aerosaurus (Fig. 13B), and Casea (Fig. 
13 A), a secondary ridge is present extending from the distal end of 
the deltopectoral crest to a more medial position on the proximal 
end of the humerus. In Casea and Aerosaurus, this is a sharp crest 
that results in the presence of a distinct fossa at the base of the 
deltopectoral crest. In Varanops this is a low, rounded ridge. 

In captorhinids (Holmes, 1977), the crest is absent. The tuberosity 
is located on the lateral edge of the proximal end of the humerus. In 
Diadectes, the deltopectoral crest is like that of Ophiacodon, 
although it forms a more obtuse angle with the proximal end of the 
humerus. Thus, these character-states have the distribution shown 
in Figure 2E, with the presence of a deltopectoral crest formed by a 
single broad ridge extending from the tuberosity to the proximal 
end of the bone being a primitive reptilian condition, and the pres- 
ence of a secondary ridge extending from the deltopectoral tuber- 
osity to the proximal end of the bone being a derived condition. 

Results 

The distribution of the character-states for which polarity can be 
interpreted, and the cladogram that requires the fewest number of 
reverseals or convergent evolutionary events to explain this distribu- 
tion, is shown in Figure 14. 

CHARACTER ANALYSIS: II. FEATURES FOR WHICH 
POLARITY CANNOT BE INTERPRETED 

As argued in the materials and methods section, characters for 
which polarity of the character-states cannot be interpreted are of 
use in providing a test of the caldogram presented on the basis of 
other characters. Three such characters are considered here. 



28 



BREVIORA 



No. 473 




Casea Varanops Aerosaurus Ophiacodon Edaphosaurus Dimetrodon Sphenocodon 







Figure 14. Hypothesis of relationships of selected genera of pelycosaurs based on 
the characters described in the text for which polarity can be interpreted. □ indicate 
primitive character-states, ■ and (ED indicate derived character-states. For description 
of characters and character-states see Table 2. 






1983 PELYCOSAUR INTERRELATIONSHIPS 29 

Table 2. The characters that form the basis for the cladogram shown in Figure 
14. The numbers refer to the characters shown in Figure 14. 

1) humerus, deltopectoral crest: 

primitive character-state: the presence of a single ridge leading from the distal end of 
the deltopectoral crest to the proximal end of the humerus; 

derived character-state: the presence of a secondary ridge leading from the distal end 
of the deltopectoral crest to a more medial position on the proximal end of the 
humerus with a fossa at the base of the crest. 

2) maxilla: 

primitive character-state: no contact between the maxilla and the quadratojugal; 
derived character-state: the maxilla contacts the quadratojugal with the bones raised 
to form a ridge along the contact of the maxilla and jugal. 

3) basipterygoid process: 

primitive character-state: basipterygoid process directed anteroventrally; 

derived character-state #1 (indicated by ■ in Figure 14): basipterygoid process 

directed laterally and are mediolaterally elongate; 
derived character-state #2 (indicated by m in Figure 14): basipterygoid processes 

directed ventrally and articular surface oriented anteroposteriorly. 

4) cheek margin: 

primitive character-state: cheek margin convex; 
derived character-state: cheek margin concave. 

5) premaxilla: 

primitive character-state: anterior margin of the premaxilla extends anteriorly from 

the anterior termination of the tooth row; 
derived character-state: anterior margin of the premaxilla slopes posteriorly from the 

anterior termination of the tooth row. 

6) stapes: 

primitive character-state: dorsal process of stapes not articulating in socket on the 
paroccipital process; 

derived character-state: dorsal process of stapes articulating in a socket on the paroc- 
cipital process. 

7) angular: 

primitive character-state: angular without an extended ventral plate in the region 

anterior to the articular; 
derived character-state: angular with an extended ventral plate in the region of the 

articular. 

8) basipterygoid articular surfaces: 

primitive character-state: articular surfaces curved; 

derived character-state: articular surfaces differentiated into two flat areas at right 
angles to each other. 

9) shelf between basisphenoid wings: 

primitive character-state: shelf between basisphenoid wings present; 
derived character-state: no shell between basisphenoid wings. 



30 BREVIORA No. 473 

Table 2. Continued 

10) frontal: 

primitive character-state: frontal without a lateral lappet; 
derived character-state: frontal with a lateral lappet. 

1 1 ) quadratojugal: 

primitive character-state: quadratojugal extends anteriorly forming ventral margin of 

skull along posterior half of cheek; 
derived character-state: quadratojugal restricted to ventro-lateral corner of skull. 

12) prearticular; 

primitive character-state: prearticular not twisted; 
derived character-state: prearticular twisted. 

13) pterygoideus process: 

primitive character-state: pterygoideus process not formed by articular alone; 
derived character-state: pterygoideus process formed entirely by articular. 

14) quadrate ramus of pterygoid: 

primitive character-state: quadrate ramus of pterygoid with a medially directed tym- 
panic flange along its ventral edge; 
derived character-state; quadrate ramus of pterigoid with rounded ventral edge. 

15) supraoccipital: 

primitive character-state: supraoccipital without a lateral process, dorsal border of 

posttemporal fenestra formed by tabular; 
derived character-state: supraoccipital with a lateral process that forms the dorsal 

border of the posttemporal fenestra. 

16) neural arch: 

primitive character-state: neural arch without pits; 

derived character-state: neural arch with pits on its lateral surface at the level of the 
zygapophyses. 

17) transverse processes: 

primitive character-state: transverse processes supported by a web of bone extending 

to the anterior edge of the centrum; 
derived character-state: transverse processes extend laterally, ventral flange not 

extending to the anterior edge of the centrum. 



Postparietal 

In Dimetrodon, Sphenacodon (Fig. 6D), and Edaphosaurus (Fig. 
6C), the postparietal is a single median element that broadly overlies 
the supraoccipital. 

The Ophiacodon, a postparietal was not observed, but the 
supraoccipital shows that the postparietal did not greatly overlap 
that bone. 



1983 PELYCOSAUR INTERRELATIONSHIPS 31 

In Casea (Fig. 6A), the postparietal is a paired element. 

The postparietal of Varanops and Aerosaurus is unknown. 

Limnoscelis is like Dimetrodon and Edaphosaurus in having a 
single postparietal (Romer, 1946). In anthracosaurs, as in primitive 
tetrapods generally, the postparietal is a paired element (Panchen, 
1970). 

Two explanations for this distribution of character-states are 
equally possible: that Limnoscelis is apomorphic in having a single 
postparietal and a paired postparietal is primitive for reptiles, or 
that a single postparietal is primitive for reptiles and Casea is 
derived in having a paired postparietal. In view of this, the presence 
of single or paired postparietals must be considered character-states 
for which polarity cannot be determined. 

Paroccipital processes 

The paroccipital processes in Dimetrodon and Sphenacodon are 
rodlike structures sloping ventrally and posteriorly. In cross section, 
they are triangular with their height less than twice their width. In 
Edaphosaurus, the processes are like those in Dimetrodon, but they 
are shorter and extend more directly laterally (Fig. 6C). In Ophia- 
codon, the process is short and laterally directed (Fig. 6B). It is 
rodlike, although its distal end is unossfied so that, as preserved, it 
does not reach the cheek. In Varanops (Fig. 4B) and Aerosaurus 
(Langston and Reisz, 1981), the paroccipital process is platelike: in 
lateral view its height is more than three times its width. In Casea, 
the platelike nature of the paroccipital process is exaggerated (Fig. 
6A). 

Thus, two distinct character-states can be recognized: one present 
in Varanops and Casea in which the paroccipital process is platelike, 
and one seen in Ophiacodon, Edaphosaurus, Dimetrodon, and 
Sphenacodon in which the paroccipital process is rodlike. The 
paroccipital process of Limnoscelis, as described by Romer ( 1946), 
is not directly comparable to either pelycosaur condition. The 
paroccipital process of Paleothyris is unossified (Carroll, 1969). 
Thus, there is no basis for interpreting the polarity of these 
character-states. 

Supinator Process 

In Dimetrodon and Sphenacodon, the supinator process is elon- 
gate proximodistally, and its distal end curves distally (Fig. 13F). 



32 BREVIORA No. 473 

In Edaphosaurus (Fig. 13E), an ectepicondylar foramen is pres- 
ent. Romer and Price ( 1940) interpreted this as a result of the devel- 
opment of a bony connection between the ectepicondyle and the 
distal end of the supinator process. Without this connection, the 
supinator process would have the morphology of that of Dime- 
trodon. 

In Ophiacodon (Fig. 13D), the supinator process is a narrow 
triangular process that projects laterally. 

Casea (Fig. 13 A) and Varanops (Fig. 13C) are like Dimetrodon in 
the development of their supinator process. The morphology of the 
process in an adult humerus of Aerosaurus is not known. 

The structure of the supinator process is variable in early tetra- 
pods. In anthracosaurs, this is represented by a crest running the full 
length of the humerus (Panchen, 1970). In Limnosce/is and Dia- 
dectes, it is a narrow laterally directed process located distal to the 
radial condyle (Romer, 1956). In its shape, it is like that of Ophia- 
codon. but in Ophiacodon, as in other pelycosaurs and in contrast 
to Diadectes and Limnoscelis, the supinator process is located dis- 
tally and is separated from the radial condyle by a distinct groove. 
The supinator process of Paleothyris is located near the distal end of 
the humerus and is separated from the radial condyle by a groove 
(Carroll, 1969). Its shape does not compare directly with either 
pelycosaur condition. This distribution of character-states does not 
allow polarity of the character in pelycosaurs to be interpreted. 

Results 

The characters for which polarity cannot be interpreted can be 
separated into two groups: those whose distribution does not 
require hypothesizing the occurrence of parallel evolution, and 
those whose distribution requires a more complicated hypothesis of 
evolution of the character. In the first group are the structure of the 
paroccipital process and the presence of paired postparietals. The 
structure of the paroccipital process is in accordance with the sepa- 
ration of Ophiacodon, Edaphosaurus, Dimetrodon, and Sphena- 
codon from Aerosaurus, Varanops, and Casea, and thus is in 
agreement with the separation of pelycosaurs into these two groups. 

In the second group is the structure of the supinator process. If 
the supinator process of Ophiacodon is autapomorphic, these 
character-states are consistent with the cladogram; if primitive, then 



1983 PELYCOSAUR INTERRELATIONSHIPS 33 

the proximodistally elongate supinator process must have evolved 
independently in the clade including Casea and Varanops and the 
clade including Edaphosaurus, Dimetrodon, and Sphenacodon. 
Thus, this character is not necessarily in conflict with the clado- 
gram, but requires a more restrictive hypothesis of polarities to 
account for the distribution of the character-states. 

DISCUSSION 

These results differ from those of Reisz (1980) in three features: 1) 
Edaphosaurus and Dimetrodon are interpreted as members of a 
clade more derived than Ophiacodon; 2) Varanops and Aerosaurus 
are interpreted as being more primitive than the clade including 
Ophiacodon, Edaphosaurus, Dimetrodon, and Sphenacodon; and 
3) Casea, Varanops, and Aerosaurus are interpreted as being 
members of a single clade distinct from the clade including Ophia- 
codon, Edaphosaurus, and Dimetrodon. 

The more primitive position of Ophiacodon relative to Edapho- 
saurus and Dimetrodon is the same as that in the phylogeny pro- 
posed by Romer and Price (1940). The alternate interpretation of 
relationships of these genera proposed by Reisz (1980) was based 
primarily on cranial features, including the structure of the frontal, 
supratemporal, and the proportions of the skull. The structure of 
the frontal of Edaphosaurus was incorrectly interpreted by Romer 
and Price (1940). As discussed above, the frontal of Edaphosaurus is 
directly comparable to that of Dimetrodon, and the frontals in these 
animals share features that can be interpreted as being derived with 
respect to Ophiacodon. Thus, the frontal supports a relationship 
between Edaphosaurus and Dimetrodon, rather than between Ophi- 
acodon and Dimetrodon. The structure of the supratemporal is 
poorly known in most genera. The element that Romer and Price 
identified as the supratemporal in Casea is better interpreted as the 
proximal end of the squamosal. This is also the case in Edaphosau- 
rus. Thus, this character cannot be used at present in interpreting 
pelycosaur interrelationships. The proportions of the skull of Eda- 
phosaurus are not known with sufficient certainty to be used in 
interpreting relationships. 

The position of Varanops and Aerosaurus is the most striking 
difference from both the cladogram of Reisz (1980) and the phyl- 
ogeny of Romer and Price (1940). This difference is based on the 



34 BREVIORA No. 473 

interpretation that Ophiacodon and Dimetrodon share derived 
cranial features not present in Varanops and Aerosaurus, and that 
Casea, Varanops, and Aerosaurus share derived cranial features not 
present in Ophiacodon. Necessarily, we interpret the derived verte- 
bral features shared in Dimetrodon and Varanops, the features that 
Romer and Price used to unite these genera, as convergent. 

The suggestion that Casea, Varanops, and Aerosaurus are mem- 
bers of a single clade is the least well supported of the relationships 
proposed above. This reflects the problems inherent in determining 
relationships of primitive members of clades. Greater certainty 
about the primitive character-state of various features in pelyco- 
saurs will come from a better understanding of the interrelation- 
ships of early reptiles and the use of an out-group that is more 
closely related to pelycosaurs than are diadectomorphs in the 
analysis of polarities. This will allow better separation of characters 
that are primitive for pelycosaurs from those that are derived for the 
clade including Casea and Varanops. 

ACKNOWLEDGMENTS 

The authors are greatly indebted to the scientific and curatorial 
staff of the Field Museum of Natural History and the American 
Museum of Natural History for permission to study specimens in 
their collections. Grants from the K. P. Schmitt Fund of the Field 
Museum enabled the authors to visit the Field Museum. Versions of 
this manuscript were read by J. Hopson, H. Barghusen, R. Reisz, 
M. Heaton, F. Jenkins, Jr., H. Sues, and G. Meyer. Their critical 
comments substantially improved the manuscript. 

Publication costs of this study were covered in part by a grant 
from the Wetmore Colles Fund. 



LITERATURE CITED 

Carroll, R. L. 1964. The earliest reptiles. J. Linn. Soc. (Zool.), 45: 61-83. 

1969. A Middle Pennsylvanian captorhinomorph and the interrelation- 
ships of primitive reptiles. J. Paleontol., 43: 151-170. 

1970. The ancestry of reptiles. Phil. Trans. Roy. Soc. Lond. B, 257: 

267-308. 

1980. The hyomandibular as a supporting element in the skull of 

primitive tetrapods pp. 293-317. In A. L. Panchen (ed.). The Terrestrial 



1983 PELYCOSAUR INTERRELATIONSHIPS 35 

Environment and the Origin of Land Vertebrates, London and New York, 

Academic Press. 
Eberth, D. 1981. The skull of Sphenacodon ferocior and comparisons within the 

subfamily Sphenacodontinae. MSc thesis. Department of Paleontology, Berke- 
ley, University of California. 
Heaton, M. H. 1979. Cranial anatomy of primitive captorhinid reptiles from the 

Late Pennsylvanian and Early Permian, Oklahoma and Texas. Oklahoma 

Geol. Surv. Bull. No. 127, 84 pp. 
1980. The Cotylosauria: A reconsideration of a group of archaic tetra- 

pods, pp. 479-551. In A. L. Panchen (ed.). The Terrestrial Environment and the 

Origin of Land Vertebrates, London and New York, Academic Press. 
Holmes, R. 1977. The osteology and musculature of the pectoral limb of small 

captorhinids. J. Morphol., 152: 101-140. 
Holmes, R., and R. L. Carroll. 1977. A temnospondyl amphibian from the 

Mississippian of Scotland. Bull. Mus. Comp. Zool., 147: 489-511. 
Kemp, T. S. 1980. Origin of mammal-like reptiles. Nature, 283: 378-380. 
Langston, W. 1965. Oedaleops campi (Reptilia: Pelycosauria). A new genus and 

species from the Lower Permian of New Mexico and the family Eothyrididae. 

Bull. Texas Mem. Mus., 9: 1-46. 
Langston, W., and R. Reisz. 1981. Aerosaurus wellesi, new species, a varanop- 

seid mammal-like reptile (Synapsida: Pelycosauria) from the Lower Permian of 

New Mexico. J. Vertebr. Paleontol., 1: 73-96. 
Lewis, G. E., and P. P. Vaughan. 1 965. Early Permian vertebrates from the Cutler 

Formation of the Placerville area, Colorado. Contrib. Paleontol. Geol. Surv. 

Prof. Pap. No. 503-C, pp. 1-46. 
Lombard, R. E., and J. R. Bolt. 1979. Evolution of the tetrapod ear: an analysis 

and reinterpretation. Biol. J. Linn. Soc, 11: 19-76. 
Olson, E. C. 1966. The middle ear — morphological types in amphibians and rep- 
tiles. Am. Zool., 6: 399^119. 
Panchen, A. L. 1970. Batrachosauria, part A, Anthracosauria. In O. Kuhn (ed.), 

Handbuch der Palaoherpetologie, Stuttgart, Germany and Portland, USA, 

Gustav Fischer Verlag, 84 pp. 
Reisz, R. 1980. The Pelycosauria: A review of phylogenetic relationships, pp. 

553-592. In A. L. Panchen (ed.). The Terrestrial Environment and the Origin of 

Land Vertebrates, London and New York, Academic Press. 
Reisz, R., M. J. Heaton, and B. R. Pynn. 1982. Vertebrate fauna of the Late 

Pennsylvanian Rock Lake Shale near Garnett, Kansas: Pelycosauria. J. Paleon- 
tol., 56: 741-750. 
Romer, A. S. 1946. The primitive reptile Limnoscelis restudied. Am. J. Sci., 244: 

151-188. 
1956. The Osteology of the Reptiles. Chicago and London, University of 

Chicago Press, 772 pp. 
Romer, A. S., and L. I. Price. 1940. Review of the Pelycosauria. Geol. Soc. Am. 

spec. Pap. No. 28, 538 pp. 
Watrous, L. E., and Q. D. Wheeler. 1981. The out-group comparison method 

of character analysis. Syst. Zool., 30: I— II. 
Williston, S. W. 1911. American Permian Vertebrates. Chicago, University of 

Chicago Press, 145 pp. 



■W M "S; COMP. zoo,. 

MAR 1 8 S85 



BREVIty 




usenoi of Comparative Zoology 



US ISSN 0006-9698 



Cambridge, Mass. 29 April 1983 Number 474 

RUTHIROMIA ELCOBR1ENSIS, A NEW PELYCOSALR 
FROM EL COBRE CANYON, NEW MEXICO 

David A. Eberth 1 and Donald Brinkman 2 



Abstract. Ruthiromia elcobriensis, a new genus and species of pelycosaur from 
the Permo-Carboniferous redbeds of El Cobre Canyon, New Mexico, is based upon 
an articulated portion of the vertebral column, partial appendicular skeleton, and 
associated vertebral and cranial fragments. Ruthiromia is tentatively included within 
the Varanopseidae on the basis of postcranial characters. It is distinguished from all 
other varanopseids by its larger size, pinched-in lumbar centra, and massively 
developed delto-pectoral crest. 

INTRODUCTION 

The Permo-Carboniferous, Cutler formation redbeds of El Cobre 
Canyon, New Mexico, have yielded a unique assemblage of fossil 
tetrapods. El Cobre Canyon, near Abiquiu, New Mexico, is a large 
box canyon that has been sporadically prospected by collectors and 
paleontologists since the late 1870's. The fossil fauna and flora, 
known from both the canyon walls and floor, was most recently 
reviewed by Fracasso (1980). 

In 1965 A. Lewis and S. Olsen collected the larger part of the 
postcranial skeleton and disarticulated cranial fragments of a 



'Department of Geology. University of Toronto, Toronto, Ontario. Canada, M5T 

IR7. 

"Museum of Comparative Zoology. Harvard University. Cambridge. Massachusetts 

02138. Present Address: Tyrrell Museum of Paleontology. Box 7500. Drumheller. 

Alberta. Canada. TOJ OYO. 



2 breviora No. 474 

pelycosaur from the west wall of the canyon. The specimen, referred 
to Ophiacodun navajovicus, remained largely unprepared in the 
collections at the Museum of Comparative Zoology at Harvard 
University until the spring of 1980 when the authors undertook a 
study of the specimen (MCZ 3150). Preparation and study of the 
material has shown that while it is clearly a pelycosaur, it is 
unassignable to Ophiacodon. It is here described as a new genus and 
species, a primitive member of the family Varanopseidae. 

SYSTEMATIC PALEONTOLOGY 

Class: Reptilia 

Order: Pelycosauria 

Family: Varanopseidae 

Genus: Ruthiromia gen. nov. 

Type species: Ruthiromia ekobriensis gen. et sp. nov. 

Etymology: Named in honor of Ruth Romer. 

Diagnosis: A large varanopseid distinguished from all other 
members of the family in the following features: linear measure- 
ments approximately 30 percent greater than those of Varanodon, 
previously the largest known varanopseid; the first eight centra 
directly anterior to the pelvis deeply pinched-in laterally, just below 
the level of the notochordal pit, giving the centra an hour-glass 
appearance when seen in cross section; each of the eight centra 
displays a broad rounded ventral surface; the delto-pectoral crest of 
the humerus is massively developed and continues distally as a sharp 
ridge below the area of pectoralis muscle insertion; the neck of the 
astragulus represents 35 percent of the total height of that element. 

Ruthiromia ekobriensis differs from Varanops brevirostris in 
having a keeled angular in the region of the articular; a large medial 
process of the articular positioned farther forward than in the latter 
genus; and a low bladed ilium. Ruthiromia ekobriensis differs from 
Aerosaurus greenleeorum in not exhibiting the excavated anterior 
lip in its anterior dorsal vertebrae. Ruthiromia ekobriensis differs 
from Aerosaurus wellesii in retaining a short paroccipital process 
and two sacral ribs. 



1983 A NEW PELYCOSAUR 3 

Ruthiromia elcobriensis gen. et sp. nov. 
(Figs. 2-8,9-5) 

Etymology: Named in reference to El Cobre Canyon, the type 
locality. 

Diagnosis: Same as for the genus. 

Holotype (MCZ 3150): An articulated, partial vertebral column, 
pelvis and right hind limb found in close association with disarticu- 
lated elements. All material is considered to represent a single indi- 
vidual. Material includes: an occipital fragment including supra- 
occipital and opisthotic; a left quadrate fragment in articulation 
with a partial quadratojugal and squamosal; the posterior portion 
of the left mandible, the atlas centrum and axis intercentrum; a 
cervical rib; four disarticulated dorsal vertebrae; one disarticulated 
neural arch; one partial neural spine; articulated posterior axial 
elements including the last seven presacral vertebrae, two sacral 
vertebrae and five caudal vertebrae; right and left partial scapulaco- 
racoids; a complete right pelvic girdle; a partial left pelvic girdle 
including pubis, ischium and a partial ilium; a right humerus; a 
right femur; a left proximal femur; a right tibia, fibula, tarsus, and 
pes. 

Type Locality and Age: Cutler Formation, Lower Permian. Col- 
lected from the fluvial redbeds in the west wall of El Cobre Canyon. 
Specific collecting site unknown. 

METHODS 

Although comparisons of Ruthiromia elcobriensis with other 
pelycosaur taxa have been performed in line with phviogenetic sys- 
tematic principles, a few words on polarity determinations within 
the pelycosaurs are in order. Following Brinkman and Eberth 
(1983), we accept a very primitive position for the varanopseids 
within the paraphyletic taxon Pelycosauria (Fig. 16). Thus any pely- 
cosaur taxon may appear closely related to the varanopseids, 
caseids, and eothyridids simply by exhibiting primitive characters. 
To guard against the use of symplesiomorphy in establishing the 
affinities of Ruthiromia elcobriensis, we have evaluated character 
state polarities within the context of what we consider to be a gener- 
ally accepted view of amphibian, reptile relationships. In this paper 



4 BREVIORA No. 474 

we adopt the classification of Carroll (1969) modified by Heaton 
( 1980) (Fig. 1 ). It is bv reference to the Diadectomorpha (including 
Tseajaia. Limnoscelis, and Diadectes) as the closest sister taxon of 
reptiles and the relative distribution of character states in reptiles 
that we determine character polarity in primitive pelycosaurs and 
specifically Ruthiromia elcobriensis. 

DESCRIPTION AND COMPARISON 

Skull: A small fragment oi the left side of the occiput is preserved 
(Fig. 2A,B). The tall anteroposteriorly flattened paroccipital process 
extends directly laterally beneath the posttemporal fenestra and 
terminates in a convex, thin edge as in I aranops and Aerosaurus 
wellesii. No trace of a suture between the opisthotic and supraoccip- 
ital is visible. In posterior view, unfinished lateral edges of the 
supraoccipital suggest that the absence of lateral processes is a result 
o\ incomplete preservation. The anterior surface of the occipital 
fragment displays a small gap between the anterior surface of the 
supraoccipital and an anteromedial continuation o( the opisthotic. 
No notch for the trigeminal is visible however, and it appears that 
only the corner of the braincase in this region is perserved. 

As in pelycosaurs and primitive reptiles generally, the quadrate is 
quite tall (Fig. 3A.B.C). Both condyles are oriented with their long 
axes anteromedially, the medial condyle being the more elongate. A 
saddle-shaped concavity occurs between the two condyles. Frag- 
mentary bone lateral to the posterior edge of the element probably 
represents the squamosal and a portion of the quadratojugal. Distal 
extremities are not preserved. No stapedial recess is discernible on 
the medial surface of the quadrate. 

The posterior portion of the left mandible ( Fig. 4 A,B,C), includes 
the articular, part of the surangular, angular and possibly prearticu- 
lar and measures 7.7 cm long. The condylar recesses of the articular 
are in the horizontal plane. The articular has a large triangular, 
medially projecting process whose medial apex is in line with or 
slightly ahead of the posterior margin of the Meckelian trough and 
well ahead of the condylar recesses. This is closely comparable to 
the condition seen in Aerosaurus wellesii. In I aranops the medial 
apex of the process is located just slightly anterior to the condylar 
recesses. In the area of the articular, the angular is keeled and com- 
pares quite well with A. wellesii and I aranodon. 1 aranops differs in 



1983 



A NEW PELYCOSAUR 



ANTHRACOSAURIA 



DIADECTOMORPHA 



PROTOROTHYRIDIDAE 
ALL OTHER 6 REPTILES 




Figure 1. Cladogram depicting relationships of taxa closely involved in the 
amphibian, reptilian transition. Derived from Carroll ( 1969) and modified by Heaton 
(1980). 





Figure 2. R. ehobriensis, MCZ 3150. A. Posterior view of left opisthotic/supraoc- 
cipital fragment. B. Anterior view of the same. Scale equals 1 cm. 

Abbreviations: op, opisthotic; st, supratemporal. 



having an angular that is gently rounded in this portion of the jaw. 
The posterior limit of a medial fenestra is preserved at the anterior 
most portion of the mandibular fragment and appears to be 
bounded by the angular and the prearticular. 

Axial Skeleton: The vertebral column is known from an articu- 
lated series of 14 vertebrae and five disarticulated vertebrae. All 
regions are represented, but only the atlas is present from the cervi- 
cal region. The centra are clearly larger than their counterparts in 
other varanopseid taxa (Table 1) and appear to be relatively shorter 
than those of Varanops and A. greenleeorum (Table 2). 



BREVIORA 



No. 474 






Figure 3. R. elcobriensis, MCZ 3150. A. Anterior view of left quadrate squamo- 
sal, quadratojugal fragment. B. Medial view of the same. C. Ventral view of the same. 
Scale equals 1 cm. 



Abbreviations: q, quadrate; sq, squamosal. 



1983 



A NEW PELYCOSAUR 





Figure 4. R. elcobriensis, MCZ 3150. A. Dorsal view of left mandibular fragment. 
B. Medial view of the same. C. Lateral view of the same. Scale equals 1 cm. 

Abbreviations: a, angular; ar, articular; cr, condylar recesses of articular; mp, 
medial process of the articular; mt, Meckelian trough; sa. surangular. 



BREVIORA 



No. 474 



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1983 A NEW PELYCOSAUR 



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1983 



A NEW PELYCOSAUR 



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12 breviora No. 474 




Figure 5. R. elcobriensis, MCZ 3150. A. Anterior same and right cervical rib. 
Scale equals 1 cm. 

Abbreviations: ai, axis intercentrum. 



A small block of matrix contains the atlas centrum and the axis 
intercentrum as well as a spatulate cervical rib (Fig. 5A,B). The atlas 
centrum is fully ossified and is exposed ventrally. The axis intercen- 
trum is a large crescentic element fused to the posterior surface of 
the atlas centrum. Although the atlas centrum is visible ventrally, it 
is the axis intercentrum that is more prominent, contributing to 
more than 50 percent of the ventral surface. The condition in 
Varanops is like that in Ruthiromia. The condition in Varanodon, 
A. green/eeorum, and A. wellesii is unknown. 

The head of the cervical rib exhibits separate capitular and tuber- 
cular processes. The shaft is expanded anteroposteriorly as a spatu- 
late convex blade. 

A disarticulated anterior dorsal vertebra includes the centrum 
and partial neural arch and displays an anterolaterally projecting 
diapophysis that extends 6.8 mm from the unexpanded neural arch 
(Fig. 6). The ventral surface of the centrum is sharply keeled. This is 
the typical morphology of all varanopseids with the exception of A. 
greenleeorum. There, the anterior lip of the centrum is deeply 
recessed to receive the intercentrum and the ventral surfaces are 
never as strikingly keeled. The floor of the neural canal is not ossi- 
fied, resulting in a deep excavation which extends nearly to the 
mid-height of the centrum as in Varanops and A. greenleeorum. The 
articulating surface of the zygapophysis is slightly inturned and 
forms an angle of 20 degrees to the horizontal. 

The three remaining disarticulated dorsal vertebrae (Fig. 7) can 
be arranged relative to one another but position in the column is 



1983 A NEW PELYCOSAUR 13 




Figure 6. R. elcobriensis, MCZ 3150. Left lateral view of anterior dorsal vertebra 
and a cross section through the mid-length. Scale equals 1 cm. 



uncertain. All three display a small pit on the lateral surface of the 
centrum wall at mid-height. The more anterior of the three is keeled, 
although not as sharply as the anterior dorsal vertebra just de- 
scribed. The ventral surface of the centrum displays three closely set 
ridges extending anteroposteriorly. In Varanops comparable ridges 
occur on the 22nd presacral vertebra (Fig. 9). 

The second vertebra retains a laterally projecting diapophysis that 
extends 7 mm from the neural arch. The ventral surface is gently 
rounded, however, and shows no trace of a keel or ridges. 

The third vertebra differs from the preceding two in that the 
ventral surface of the centrum is more broadly rounded. 

Neural arch excavations are present on the third vertebra and 
extend anteroventrally and open onto the surface of the neural arch 
between the diapophyses and the anterior zygapophyses. The dia- 
pophysis projects posterolaterally and measures 5 mm from the 
surface of the neural arch. The articulating surface of the zyga- 
pophvsis appears nearly horizontal. The orientation of the diapo- 
physis and the smoothly rounded ventral edge of the centrum 
suggest that this vertebra was a lumbar. 

Although the last seven presacral vertebrae are in articulation 
(Fig. 8), only the first four of the series are preserved well enough to 
merit discussion. As in the preceding vertebrae, each displays a 
small pit in the lateral wall of the centrum at mid-height. The lateral 



14 



BREVIORA 



No. 474 









Figure 7. R. elcobriensis, MCZ 3150. Left lateral views and cross sections of three 
dorsal centra arranged with the anterior most to the left. Scale equals 1 cm. 



walls of the centra are strongly concave and the ventral surfaces are 
broadly rounded giving the centra an hour-glass shape in cross sec- 
tion. This condition is termed pinched-in. No other varanopseid 
displays this condition in the posterior dorsal centra. Varanops dis- 
plays broadly rounded ventral surfaces on its last four presacral 
vertebrae ( Fig. 9), but there is no indication of pinching-in or pitting 
of the lateral walls, of the centra. A posterior dorsal vertebra of an 
immature A. wellesii (UCMP 20096?) displays a pit in the lateral 
wall of the centrum, but lacks the pinched-in condition. 

The neural arches of the last three presacral vertebrae are deeply 
excavated at the level of the articular surfaces of the zygapophyses. 

All neural spines in this articulated series are imperfectly pre- 
served and show only the primitive condition of being laterally 
compressed. The centra of all of the vertebrae posterior to the third 
vertebra of the series are either inaccessible or poorly preserved. 

A disarticulated neural arch with a complete neural spine is 
poorly preserved but shows that the spine is relatively lower in 
height than those of Varanops and A. wellesii (Table 2). 



1983 



A NEW PELYCOSAUR 



15 




Figure 8. R. elcubriensis, MCZ 3150. Right lateral view of articulated lumbar, 
sacral, proximal caudal vertebrae and pelvis. Scale equals 1 cm. 

Abbreviations: cv, caudal vertebrae; Iv, lumbar vertebrae. 




Figure 9. Varanops hrevirostris, MCZ 1926. Ventral view and cross sections of 
last eight presacral vertebrae. Anteriormost to the right. Scale equals 1 cm. 



The neural arches of the sacral vertebrae are deeply excavated 
above the articular surfaces of the zygapophyses. Two poorly pre- 
served sacral ribs are present. Aerosaurus wellesii has three sacral 
ribs and appears to be unique in this feature within the varanopseids 
(Langston and Reisz, 1981). 



16 BREVIORA No. 474 




Figure 10. H. elcobriensis, MCZ 3150. Lateral vieu of right scapulocoracoid 
fragment. Scale equals 1 cm. 

Abbreviations: g, glenoid; trie cor, process for the coracoid head of the M. triceps. 



The first caudal vertebra is distorted and thus appears wider than 
the more posterior caudals. The neural spines of the caudal verte- 
brae are smaller than those of the sacral or presacral vertebrae and 
appear almost triangular in lateral aspect. All of the five articulated 
caudals display excavated neural arches. 

Appendicular Skeleton: Fragmentary right and left scapulocora- 
coids are preserved. Both display at least a portion of the glenoid 
and a well-developed process for the origin of the coracoid head of 
the M. triceps (Fig. 10). In primitive fashion the glenoid is not raised 
above the surface of the coracoid and continues back onto the tri- 
ceps process. Similar conditions exist elsewhere in all known 
varanopseids. 

The right humerus is well preserved (Fig. 1 1) and has proportions 
conformable with those present in Varanops (Table 2). Varanodon 
and A. wellesii have a proportionately narrower proximal ends. The 
proximal end of the humerus displays a stout M. latissimus tubercle. 
This tubercle is poorly ossified in Varanops and A. wellesii but is 
well ossified in A. greenleeorum and most other pelycosaurs. The 
proximal surface in the region of the deltopectoral crest exhibits two 
well-developed parallel ridges with a depression between them. Such 
a condition exists in A. greenleeorum and elsewhere in pelycosaurs 
only in caseids (Brinkman and Eberth, 1983). A feature of the 
humerus not seen in other varanopseid taxa is that of a massively 
developed delto-pectoral crest that continues distally as a sharp 
ridge. This distal extension is so prominent that the entire region 
appears almost square in anterior aspect. Romer and Price (1940) 
distinguished A. greenleeorum partly on the basis of a well- 



1983 



A NEW PELYCOSAUR 



17 





lat tub 




d.p.cr. 



Figure II. R. ekubriensis, MCZ 3150. A. Proximal dorsal view of left humerus. 
B. Proximal ventral view of the same. C. Proximal view ot the same. D. Distal view 
of the same. Scale equals 1 cm. 



Abbreviaiinns: dpcr. delto-pectoral crest; lat tub. latissimus tubercle. 



18 BREVIORA No. 474 

developed deltopectoral crest. Direct comparison indicates that A. 
greenleeorum lacks the distal ridge. A strong ectepicondyle occurs 
distally and is well separated from the partially preserved supinator 
process. Much of the entepicondyle is missing, and the position of 
the entepicondylar foramen is uncertain. Distoventrally, a large 
radial condyle is preserved. 

The right pelvis is well preserved and has been completely pre- 
pared laterally and ventrally. The medial surface of the ilium has 
also been prepared. The pelvis is quite primitive in general structure, 
having a posteriorly projecting iliac blade and a solid puboischiadic 
plate. The ilium (Fig. 12A,B) is triangular in lateral aspect. It 
becomes wider and longer ventrally where it forms the dorsal por- 
tion of the acetabulum. The elongate, triangular posterior process is 
low as in A. wellesii and Varanops but appears longer exhibiting 
lower base/ length and neck/ length proportions than in those two 
species (Table 2). A small anterior projection of the ilium is poorly 
preserved. In medial aspect (Fig. 12B) a ridge bisects the length of 
the posterior process and appears conformable with that present in 
all other varanopseids. The pubis is primitively short and ends ante- 
riorly in a slightly thickened knob (Fig. 12D). In lateral aspect, the 
dorsal margin of the pubis is thickened and appears concave. A 
poorly preserved bump on the lateral surface, directly anterior to 
the acetabulum, may be a pubic tubercle. A pubic tubercle is 
unknown in other varanopseid taxa, but is found in the ophiaco- 
dontids Clepsydrops and Varanosaurus as well as caseids and eda- 
phosaurids. The widespread occurrence of a pubic tubercle in 
pelycosaurs as well as many cotylosaurs suggests that it is primitive. 

The ischium is complete and also displays a slightly thickened 
dorsal margin in lateral aspect (Fig. I2C). The ischiadic plate is thin 
and meets its counterpart of the left side in the mid line. The ischium 
appears conformable with the known morphology of that same ele- 
ment in other varanopseids and ophiacodontids. 

The right femur is complete (Fig. 13) although a small portion of 
the proximal end is broken off and remains in articulation with the 
acetabulum, obscuring the morphology of the latter. The proximal 
end of the left femur is also preserved. Measurements and propor- 
tions are listed in Tables 1 and 2 respectively. No significant dif- 
ferences in femoral proportions between Ruthiromia and other 
varanopseid taxa are apparent. In anterior aspect, the proximal end 



1983 



A NEW PELYCOSAUR 



19 




Figure 12. R. elcobriensis, MCZ 3150. A. Lateral view of right ilium. B. Medial 
view of the same. C. Ventrolateral view of right ischium. D. Ventrolateral view of 
right pubis. Scale equals 1 cm. 

Abbreviation: pt, pubic tubercle. 





Figure 13. R. elcobriensis, MCZ 3150. A. Dorsal view of right lemur. B. Ventral 
view of the same. C. Posterior view of the same. D. Anterior view of the same. Scale 
equals I cm. 



Abbreviations: ac, adductor crest; 4th t, 4th trochanter; it, internal trochanter. 



20 BREVIORA No. 474 

of the right femur displays a large, tall internal trochanter well set 
off from the thickened proximal surface of the femur. Varanops (the 
only varanopseid where this condition can be compared) does not 
exhibit such a well-developed internal trochanter. The intertrochan- 
teric fossa is extensive, occupying about a third of the length of the 
femur as in most non sphenacodontid pelycosaurs. A large knob 
marks the point of attachment of the M. pubo ischio femoralis 
internus. The distinct, although small, fourth trochanter is present 
just distal to the intertrochanteric fossa. A prominent, well-ossified 
adductor crest extends across the ventral surface of the element to 
the fibulad edge. The crest is relatively larger than in any other 
varanopseids and appears to be a primitive feature. In cross section 
the shaft appears tear drop-shaped, in contrast to Varanops where a 
cross section appears oval. In dorsal view, the two well-developed 
ossified tibial condyles are well separated by a deep intercondylar 
fossa. Their dorsal surfaces are essentially flat, and their articular 
surfaces are oriented ventro laterally. 

In dorsal view (Fig. 14) the right tibia and fibula have straight 
external and concave internal edges. The fibula is slightly shorter 
than the tibia (Table 1). The lateral femoral cotyle of the tibia is not 
strongly compressed. Sharp ridges are present on the lateral and 
medial edges of the distal half of the bone. The proximal end of the 
fibula is damaged, so the extent of the dorsal tubercle is unknown. 
The distal end is wide mediolaterally and compressed dorsoven- 
trally. Similar overall conditions are seen to exist in all vara- 
nopseids. 

Eight elements are present in the tarsus: the astragulus, calca- 
neum, a centrale and five distal tarsals (Fig. 15). The astragulus 
(Fig. 15A,B) is L-shaped and supports both the tibia and fibula. The 
neck of the astragulus is elongate and contributes to 35 percent of 
the height of the element; it thus appears more like that of a sphena- 
codontid than all other known varanopseids. It is unclear at present 
whether this is a primitive or derived feature. In contrast to sphena- 
codontids however, no tubercle is present on the ventral surface of 
the astragulus medial to the arterial groove. The calcaneum (Fig. 
15A,B) is primitive in being wide and having a convex lateral edge 
when seen in dorsal view. In contrast to the varanopseid genera A. 
wellesii and Varanops, the element is much taller than it is wide. 

The single centrale (Fig. 15C) is a small element with a limited 
area of finished bone on its dorsal surface. The largest of the distal 



1983 



A NEW PELYCOSAUR 



21 





Figure 14. R. elvobriensis, MCZ 3150. A. Dorsal view of right tibia and fibula. 
B. Ventral view of the same. Scale equals I cm. 



tarsals is the fourth. In dorsal view (Fig. 15C) it is essentially pen- 
tagonal in contrast to the rectangular fourth distal tarsal of A. 
wellesii and Varanops. The fourth distal tarsal also shares an equal 
contact with the astragulus. The first and third distal tarsals are 
subequal in size and the second is slightly smaller than these. This is 
also the case in Varanops and A. wellesii. The metatarsals increase 
in length from first to fourth (Table 1). The first metatarsal is sup- 
ported entirely by the first distal tarsal. In dorsal view, the second 
and third articulate with the distal tarsal of the same number. In 
ventral view (Fig. 15D) each of these metatarsals is supported by 
both the distal tarsal of the same number and the laterally adjacent 
distal tarsal. The fourth metatarsal is longer than the combined 
length of the first metatarsal and digit. This condition is seen devel- 
oped to a greater degree in Varanops and to a lesser degree in A. 
wellesii and occurs elsewhere within the pelycosaurs only in Varano- 
saurus and Ophiacodon mirus. The first toe has a clawlike terminal 
phalanx. The remaining toes are incompletely preserved or not 
present. 

In overall appendicular proportions (Table 2), Ruthiromia elvo- 
briensis appears more primitive than any other varanopseid in hav- 
ing a humerus and femur of subequal lengths and a lower tibia 
femur ratio. 



22 



BREVIORA 



No. 474 




Figure 15. R. elcobriensis, MCZ 3150. A. Dorsal view of right astragalus, calca- 
neum complex. B. Ventral view of the same. C. Dorsal view of right pes. D. Ventral 
view of the same. Scale equals 1 cm. 

Abbreviation: c, centrale. 



DISCUSSION 

Ruthiromia elcobriensis is considered a pelycosaur on the basis of 
the combined, derived characters (within reptiles) of excavated neu- 
ral arches, keeled mid-dorsal vertebrae, a strongly developed pro- 
cess for the coracoid head of the triceps and a keeled angular in the 
region of the articular. These constitute a suite of characters which 
is found only within the pelycosaurian families Varanopseidae and 
Sphenacodontidae. 

The Varanopseidae is currently defined by cranial characters 
(Langston and Reisz, 1981), yet shows a unique combination of two 
derived, postcranial characters: excavated neural arches and the 



1983 A NEW PELYCOSAUR 23 

presence of a shallow fossa on the proximal surface of the deltopec- 
toral crest (Brinkman and Eberth. 1983). Ruthiromia elcobriensis is 
tentatively assigned to the Varanopseidae on the basis of displaying 
these same characters. Following the cladistic classification of 
Brinkman and Eberth (1983, Fig. 16) for the better known pelyco- 
saur genera, the derived character of excavated neural arches is 
thought to have developed either two (Fig. 16A) or three (Fig. 16B) 
times. All varanopseid and sphenacodontid taxa, where postcranial 
material is available, show excavated neural arches. Within the eda- 
phosaurids an undescribed specimen (E2) from the Pennsylvanian 
of Kansas shows this condition (R. Reisz, personal communica- 
tion). Within this context and assuming parsimony, four equally 
correct positions denoting propinquity of descent of Ruthiromia 
elcobriensis may be depicted in a cladogram (Fig. 16C). 

Position 1 places Ruthiromia elcobriensis as a member of the 
varanopseids in the clade including Varanopsj Aerosaurus and 
Casea. This clade is united by the derived morphology of the prox- 
imal surface of the humerus in the region of the delto-pectoral crest. 
As described above, Ruthiromia elcobriensis shares this same 
morphology. 

Relationships indicated by positions 2, 3, and 4 each necessitate 
reversals of those characters at nodes (c), (c,d,e) and (c,d) respec- 
tively. The clade including Ophiacodon, Edaphosaurus, and Dime- 
trodon is defined by the shared presence of a rodlike paroccipital 
process with a socket on the ventral surface for contact with a 
distinct dorsal process of the stapes, a concave ventral margin of the 
cheek and a single postparietal (node c). R. elcobriensis has a nar- 
row paroccipital process that shows no evidence of a socket along its 
ventral edge for contact with a dorsal stapedial process. A relation- 
ship indicated by position 2 therefore involves the reversal of this 
character. Other regions are not preserved in the holotype. Sim- 
ilarly, Ruthiromia elcobriensis does not show the derived characters 
of the twisted prearticular and ventromedial projecting pterygoideus 
process which partially define the clade consisting of Edaphosaurus 
and Dimetrodon (node d). In this case, a total of three reversals 
(reversal at both c and d) is necessary to explain the relationship 
indicated by position 4. Finally, the relationship indicated by posi- 
tion 3 necessitates the reversals listed above as well as reversals of 
the autapomorphic characters that define the Edaphosauridae and 



24 



BREVIORA 



No. 474 



OTHER REPTILES VARANOPSLIDAI 



EDAPHOSAURIDAE 
CASEIDAE 0PHIAC0DONTIDAE EDAPHOSAURUS E2 



Mill Vtl IIIKINI 1IIAI 




OTHER REPTILES VARANOI' M I IHI 



OPHIACODONTIDAE 



EDAPHOSAURIDAE 

EDAPHOSAURUS 



si'iii ssi i .; int] 




Dim P REPTll.l s VARANIil'SI I MM 



EDAPHOSAURIDAE 
OPHIACODONTIDAE EDAPHOSAURUS E2 



'-.PHI Mi nMo',1 I MAI 




Figure 16. Cladograms depicting relationships of pelycosaur families. Adopted 
and modified from Brinkman and Eberth (1983). A&B. Two equally parsimonious 
hypotheses lor the appearance and distribution of excavated neural arches in pelyco- 
saurs. Solid bars represent those points where excavated neural arches are exhibited. 
Dashed bar represents point of character loss. C. Four equally correct positions for 
Ruthiromia on the basis of the derived character of excavated neural arches. 






1983 A NEW PELYCOSAUR 25 

are not seen in Ruthiromia elcobriensis (elongate neural spines with 
lateral tubercles, low position of the jaw articulation relative to the 
tooth row). Clearly, the most parsimonious course to take is to 
tentatively place Ruthiromia elcobriensis among the Varanopseidae 
as a primitive member of that family. The tentative nature of this 
placement reflects both the lack of diagnostic cranial material in the 
type and the employment of a combination of derived postcranial 
characters, none of which can serve alone as an autapomorphic 
character of the family Varanopseidae. 

Ruthiromia appears to be more closely allied with A. wellesii in 
sharing the unique character of the forwardly placed medial process 
of the articular and the keeled angular. 

Ruthiromia shares only primitive characters with Varanops and 
appears similar in retaining a laterally projecting paroccipital pro- 
cess and in having only two sacral vertebrae. 

The morphology of the lumbar vertebrae and more specifically 
the position in the column where keeled vertebrae first appear is 
more difficult to assess. The seven articulated presacral vertebrae of 
Ruthiromia show no signs of possessing keels and all retain a gently 
rounded ventral surface. The inclusion in the holotype of one disar- 
ticulated lumbar (posterior dorsal) with a similarly unkeeled ventral 
surface places the minimum location for the transitional vertebra at 
a position nine vertebrae in front of the pelvis. In addition, a signifi- 
cant difference in diapophysis orientation between the transitional 
vertebra and the isolated lumbar (lateral vs. postero lateral respec- 
tively) warrants placement of the transitional vertebra even further 
forward in the column. Varanops develops a fully keeled vertebra 
six vertebrae ahead of the pelvis (Fig. 9). The fifth is transitional, 
displaying three parallel ridges along the ventral surface. The only 
fully prepared vertebrae known from A. wellesii that can be placed 
in the column with certainty are those nine and ten positions ahead 
of the pelvis. These show keeled ventral surfaces. No idea concern- 
ing the initial position of keeled vertebrae can be determined in 
either A. greenleeorum or Varanodon, but A. greenleeorum does 
not appear to develop strong keels of the sort seen in the one known 
anterior dorsal of Ruthiromia anywhere in its column. Ruthiromia 
similarly does not share with A. greenleeorum the condition of the 
deeply recessed anteroventral lips of the anterior dorsals. Keeled 
lumbar vertebrae are considered an advanced character state (Romer 



26 breviora No. 474 

and Price, 1940). Ruthiromia appears more primitive than either 
Varanops or A. wellesii in displaying a greater number of unkeeled 
vertebrae in the presacral region of the vertebral column. 

In summary, Ruthiromia elcobriensis represents a varanopseid 
which is very primitive postcranially, with some unique vertebral 
specializations. Cranially, it shows some of the same advanced spe- 
cializations seen in A. wellesii and perhaps Varanodon. 

ACKNOWLEDGMENTS 

The authors wish to express their gratitude and appreciation to 
Robert Reisz for his many valuable comments and criticisms as well 
as the technical assistance that he provided during the preparation 
of this manuscript. Although his help must be regarded as a crucial 
factor in the production of this paper, the authors assume full 
responsibility for any errors that might be present. 

Travel and research expenses were covered, in part, by a Schmitt 
Fund research grant provided by the Field Museum of Natural 
History, Chicago. 

LITERATURE CITED 

Brinkman, D., and D. Eberth. 1983. The interrelationships of Pelycosaurs. 
Breviora Mus. Comp. Zool., No. 473. 

Carroll, R. L. 1969. The origin of reptiles, pp. 1-44. In A.d'A. Bellairs el al. 
(eds.), Biology of the Reptilia. New York, Academic Press, 373 pp. 

Fracasso, M. A. 1980. Age of the Permo-Carboniferous Cutler Formation ver- 
tebrate fauna from El Cobre Canyon, New Mexico. J. Paleontol., 44: 156-163. 

Heaton, M. J. 1980. The Cotylosauria: A reconsideration of a group of archaic 
tetrapods, pp. 497-551. In. A. L. Panchen (ed.). The Terrestrial Environment 
and the Origin of Land Vertebrates. Systematics Assoc. Spec. Vol. No. 15, 
London, Academic Press, 633 pp. 

Langston, W. Jr., and R. R. Reisz. 1981. Aerosaurus wellesii, new species, a 
varanopseid mammal-like reptile (Synapsida: Pelycosauria) from the lower 
Permian of New Mexico. J. Vertebr. Paleontol., 1(1): 73-96. 

Olson, E. C. 1965. New Permian vertebrates from the Chickasha Formation in 
Oklahoma. Oklahoma Geol. Surv. Circ. No. 70, 70 pp. 

Romer, A. S., and L. I. Price. 1940. Review of the Pelycosauria. Geol. Soc. 
Amer. Spec. Pap. No. 28, 538 pp. 



STjr 



MUS. COMP. ZOOL 
LIHRARY 

MAR 18 1985 

?VARD 



B R E V I O R A 

Museum of Comparative Zoology 



US ISSN 0006 9698 



Cambridge, Mass. 29 April 1983 Number 475 

NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. 

I. ANOLIS CALIMAE, NEW SPECIES, FROM THE 

CLOUD FOREST OF WESTERN COLOMBIA 

Stephen Ayala, 1 Dennis Harris, 2 and Ernest E. Williams- 1 



Abstract. Anolis calimae, new species, is described from the cloud forest of 
western Colombia in Departmento Valle del Cauca. Referable to the Anolis punctatus 
species group, it is distinctive in its coloration, in the absence of an interparietal scale. 
and in the apparent trend to reduction of the elongate anterior supraciliary scale usual 
in Anolis. 



INTRODUCTION 

In a recent paper (Williams, 1982), the description of three new 
species was made the occasion of a summary of the eastern members 
of the punctatus species group. Several new species must be described 
before a similar summary will be possible for the western punctatus 
group. The first is here described, a small species and initially recog- 
nized only from a single specimen collected in cloud forest near Lake 
Calima. Even with one specimen, its striking color pattern and dis- 
tinctive habitus made it obvious that it is a new species. Subsequently, 
three additional specimens have been obtained, one from the original 
locality, two from Television Tower Mountain near Cali. These four 
specimens have been divided between the Museum of Comparative 



1 120 Vista View Place, Petaluma, California 94952. 

-"Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109. 

"Museum of Comparative Zoology, Harvard University, Cambridge. Massachusetts 

02138. 



2 BREVIORA No. 475 

Zoology and the lnstitutode Ciencias Naturales(ICN, Bogota). Still 
more recently a Museo La Salle (MLS, Bogota) specimen from a 
third locality has been recognized as belonging to this species. At the 
suggestion of Fernando Castro, we call it after the place of its first 
discovery: 

Anolis calimae, new species 
(Figs. 1-6) 

Type: MCZ 158392, adult male. 

Type locality: San Antonio, Television Tower Mountain, Depto. 
Valle del Cauca, Colombia, (3° 28'N 76° 40' W) 1,800 m elevation, 
Dennis Harris, Humberto and Fanny Carvajal, coll., 23 January 
1980. 

Paratypes (all from Valle): MCZ 1 58393, adult female, same data 
as type; ICN 3678, adult female, approx. 1 km below Lake Calima 
dam, (3° 50'N 76° 32'W) Dennis Harris, coll.. 18 January 1980; ICN 
3679, adult male, same place as ICN 3678, William Duellman and 
Fernando Castro, coll., 17 March 1979. MLS 122: Mares, 3 km N of 
San Antonio (3° 30'N 76° 40'W). 

Diagnosis. A small green cloud forest species of the punctatus 
group distinguished by its short body, legs and tail, consistent 
absence of the interparietal scale, a blunt and weak canthal ridge with 
poorly differentiated canthal scales, only a short supraciliary scale 
followed by granules or granules only, small, low number of loreal 
scales(4 5), smooth ventrals and very short stubby toes with 15 to 17 
lamellae under phalanges ii and iii of fourth toe. Dewlap present in 
both sexes, small in female and with larger scales. 

Description. Head. Rather short. Head scales small, flat, very 
slightly wrinkled. Seven to ten scales across the snout between the 
second canthals. Frontal depression shallow, the scales within it as 
large or larger than some of those anterior to it. Four to seven scales 
border rostral posteriorly. Circumnasal scales of each side separated 
from rostral by one elliptical scale which lies above the suture 
between rostral and first supralabial. Six scales between supranasals 
dorsally. Snout elongate, slightly protuberant, extending slightly 
beyond mental. 

Supraorbital semicircles separated in both males by two rows of 
large scales, as large as the scales of the semicircles, in the females in 
contact or separated by one row of small scales or granules. Supraor- 



1983 



ANOLIS CAl'LIMAE 




Figure I. Anolis calimae. new species, in life. Female paratype above, male type 
below. 



bital scales in contact with or barely separated from the supraocular 
disk of each side, which consists of six to ten enlarged, slightly 
wrinkled scales, remainder of the supraocular area granular or sub- 
granular. One or two short supraciliaries on each side followed by 
granules, or only granules present. Canthus blunt, canthals small, 
poorly differentiated, the first or first and second largest. Loreal rows 
four or five, the lowermost slightly larger. 

Temporals granular. An indistinct intertemporal double line of 
slightly enlarged scales. Supratemporals granular laterally, becoming 
larger and flattened toward the interparietal area, which is bounded 
by weak ridges. No differentiated interparietal, but a zone of enlarged 
scales, largest anteriorly and laterally, posteriorly grading slowly into 
the dorsal granules. Ear opening small, elliptical, but larger than any 
scale in the interparietal area. Occiput with small blunt median knob, 
obvious in males, not evident in females. 

Suboculars in contact with supralabials. Seven supralabials to the 
center of the eye. 



BREVIORA 



No. 475 




Figure 2. Anulis calimae, new species. Male type, MCZ 158392. Dorsal view of 
head. 



Mental semidivided, each part slightly or distinctly wider than 
deep. Four granules are in contact with the mental between the large 
first sublabials. Two to five sublabials in contact with the infralabials. 
Gular scales subgranular, very little enlarged laterally where they 
border the sublabials. 

Trunk. Dorsal scales granular, convex, subequal. Ventrals much 
larger, smooth, juxtaposed or subimbricate, rounded or slightly 
pointed in males, broader, squarish in females, in tranverse rows. 
Lateral chest scales not keeled. Males may develop a low nuchal crest 
when aroused. 



1983 



ANOLIS CAtLIMAE 




Figure 3. Anulis calimae, new species. Male type, MCZ 158392. Lateral view of 
head. 



Dewlap. Present in both sexes. Large in male, reaching anterior 
abdomen, scales small, smaller than ventrals, weak, in rows, one scale 
wide, separated by naked skin, edge scales larger, somewhat crowded, 
imbricate, smooth; small to moderate in females, scales larger, as 
large or larger than ventrals, in close-packed rows, smaller and much 
more numerous along the edge. 

Limbs and digits. Upper arm scales granular. Lower arm scales 
granular to larger and unicarinate. Leg scales granular posteriorly, 
larger and uni- to multicarinate anteriorly. All supradigital scales 
multicarinate. Toes relatively short and stubby. Digital dilations 
moderately wide. Fifteen to 18 scales under phalanges ii and iii of 
fourth toe. 

Tail. Compressed, short, ca. 1.5 X body length, possibly prehen- 
sile. No tail crest. Two weakly keeled middorsal rows, ventral rows 
larger and more strongly keeled, verticils not evident. Scales poste- 
rior to vent smooth. Large postanals in males, none in females. Tail 
base prominently swollen to accommodate hemipenes in adult males. 

Size. Anoiis calimae is a small but somewhat robust species. Sizes 
of the four recent specimens are: 59 (type), 58, 55, and 58 mm 
snout-vent length respectively. 

Color. This is a green anole with a considerable capacity for rapid 
pattern and color change. It may be almost uniform green or yellow- 
green with little or no pattern, or it can have three prominent broad 



BREVIORA 



No. 475 




Figure 4. -Anvil's calimae, new species. Male type. MCZ 158392. Ventral view of 
head. 



black bands across the back and sides (apparently more prominent in 
the males), with pale yellow or cream colored spots in the dark bands. 
At another state of excitement, the sides and back are mostly green- 
grey with scattered small black spots on the sides and neck and a few 
dark brown crossbars on the vertebral line. Females especially may 
show alternating sets of narrow, light yellow and dark brown-black 
spots along the midline. There is no prominent dark band across the 
head between the eyes. The large male dewlep is unpigmented: pale 
yellow-green with salmon pink near anterior edge and white or pale 



1983 



ANOLIS CAULIMAE 




8 BREVIORA No. 475 

yellow scales. The small female dewlap is pigmented: blue with yellow 
or white scales. The tail is banded: broad dark bands in the male and 
narrow bands in the female. The lining of the throat is pale. The iris of 
1CN 3678 (a female) was golden yellow above and below a zone of 
orange; that of the holotype male, MCZ 158392, was orange. 

Preserved specimens are greenish-grey with few to many small but 
prominent dark (and light in the male) spots on the back and sides, 
elongate dark and light spots along the dorsal midline and a darker 
brown, unpatterned head. The belly is pale, with small grey spots 
under the chin. The dewlap has rows of white scales over white skin 
(males) or pigmented skin (females). The peritoneal lining is heavily 
pigmented. 

Habitat and reproduction. The two recent collection sites are 
about 50 km apart in the same cloud forest region (tropical premon- 
tane wet forest) in the western Colombian cordillera. All four speci- 
mens were collected at night while they were sleeping in exposed sites 
40 to 150 cm above the ground, the type and first paratype on a fern 
leaf and a low shrub, in a cool forested region at about 1,800 m 
elevation. The two Lake Calima paratypes were on exposed twigs, in 
a somewhat warmer, more densely vegetated area also subject to 
frequent rains and cool fogs at 1,300 m elevation. Other anoline 
species known to occur in the same sites or in the same general area 
are A. ventrimaculatus, A. eulaemus, A.fraseri, A. antonii, Phenaco- 
saurus heterodermus and an undescribed punctatus group anole 
known at present from a single specimen. 

The specimens were in reproductive condition at the time of cap- 
ture: the males with enlarged testes (6. 1 x 4.0 mm approx.), and the 
females with a single oviducal egg on one side and an enlarged, 
yolking follicle on the other. 

Etymology. The name calimae refers to the site where William 
Duellman and Fernando Castro collected the first specimen. Lake 
Calima is in turn named after the Calima Indians who inhabited the 
region centuries ago. 

Comparisons. A. calimae is a very distinctive anole. No previously 
described member of the punctatus group is known to lack an inter- 
parietal scale. This condition is unusual in any group of Anolis, but it 
is curious that it is known as a moderately common variation in two 
species of the aequatorialis species group {A. ventrimaculatus and A. 
gemmosus) which, like calimae, are inhabitants of Andean cloud 



1983 



ANOLIS CAULIMAE 




, ft £ 




10 breviora No. 475 

forest. It is possible, since there are only four specimens of calimae 
thus far known, that absence of an interparietal will be found to be 
inconstant in this species too. 

The wholly granular supraciliary margin found in one specimen of 
calimae is unique, not only for the puncatatus group, but for the 
genus Anulis. The alternative and commoner condition (in three of 
the four calimae) of one elongate but short supraciliary scale is 
known elsewhere (e.g., in the tigrinus species group); the extreme 
shortness of the scale in the three specimens may, however, be indica- 
tive of a strong trend toward de-differentiation of this scale, which 
then culminates in the completely granular margin. 

The presence of a dewlap in both sexes, and the fact that it is 
smaller and differently pigmented in the female, may be a primitive 
feature of calimae and occurs erratically in a number of Anolis 
species groups. In the punctatus group the condition is known in 
chocorum and in transversalis, and in these species, as in calimae, is 
associated with a more or less marked difference in body color and 
pattern between the sexes. 

Unique to calimae is the larger size of the dewlap scales in the 
female. The reverse or equal-sized scales is typical for Anolis species 
in which the female retains a dewlap. 

Unusual also and requiring confirmation by additional material is 
the apparent difference between males and females of calimae in the 
size and number of scales between the supraorbital semicircles. 

From the two other previously described punctatus group species 
of western Colombia A . calimae is as distinctive in color as in scales. 
Both these species are also Pacific lowland forms: Anolis chocorum 
(also a larger species, reaching 79 mm snout-vent length) has the 
dorsum uniform green or with oblique rows of dark green blotches on 
the flanks, never with prominent black bands enclosing yellow or 
cream spots. The male dewlap is orange laterally, green basally; or the 
female dewlap is green with pale yellow or gray at the base. The rows 
of scales in the dewlaps are three to four scales wide. Anolis chloris 
(about the same size as A. calimae, ca. 55 mm snout-vent length ) is 
grass-green in color, with the potentiality of turning a dark olive 
green which then may have diagonal darker bars, but again there are 
never the black bands of calimae. The dewlap color in males is yellow 
or white, never green, the rows of scales in the dewlap are one scale 
wide and there is no dewlap in the female. In contrast to A. calimae, 
both A. chocorum and A. chloris turn blue or purple in preservative 



1983 ANOLIS CAULIMAE 11 

rather than the greyish green of A. calimae. The occiput knob is not 
seen in A. chocorum or A. clitoris. 

There has been no evident close relative for A. calimae. However, 
an undescribed species from the same region, known only from a 
juvenile male, seems closer than any other, although still sharply 
distinct. 



ACKNOWLEDGMENTS 

We are grateful to the collectors of the species here described. 
William E. Duellman, Humbertoand Fanny Carvajal, and Fernando 
Castro. Ayala's work in Colombia was sponsored by grants from the 
U.S. Public Health Service (N1A1D AI-A2I5I 1), Tulane University 
(N1AID AI-10050), and COLCIENC1AS, the Colombia National 
Science Institute. The drawings are by Laszlo Meszoly. 



LITERATURE CITED 

Williams, E.E. 1982. Three new species of the Anulis punctatus complex from 
Amazonian and inter-Andean Colombia, with comments on the eastern members 
of the punctatus group. Breviora Mus. Comp. Zool. No. 467, pp. 1 38. 



MU3. COMP, ZOOL 

Linr?ARY 

MAR 1 8 1985 

ARVARD 

B R E VtO R A 

Museum of Comparative Zoology 

US ISSN 0006 9698 



Cambridge, Mass. 29 April 1983 Number 476 

TOWNSEND'S UNMAPPED NORTH ATLANTIC 
RIGHT WHALES (EUBALAENA GLACIALIS) 

William E. Schevill 1 and Karen E. Moore 2 

Abstract. Townsend's detailed maps of worldwide whale distribution from 19th 
century American whaling logbooks (1935, Zoologica, N.Y., 19: 1-50) omitted his 
North Atlantic records for Euhalaena. We have read 12 of his 15 sources and have 
mapped the right whales therein recorded. 

INTRODUCTION 

Very little is known about the distribution of right whales (Euha- 
laena glacialis) in the North Atlantic Ocean. Among others, J. A. 
Allen (1908), Collett (1909), G. M. Allen (1916), Thompson (1928), 
Slijper, van Utrecht, and Naaktgeboren (1974), Reeves, Mead, and 
Katona (1978), Schevill, Moore, and Watkins (1981), Kraus and 
Prescott (1981), Reeves and Brownell ( 1 982), and Watkins and Sche- 
vill (in press) have discussed sightings and catches in particular areas. 
Townsend ( 1 935), in his study of "the distribution of certain whales as 
shown by logbook records of American whaleships," examined log- 
books from 1,665 voyages, worldwide, mostly of the 19th century. 
Out of the 8,415 Eubalaena taken, he found only 35 in the North 
Atlantic, recorded on 1 5 voyages, and did not bother to plot these on 
his charts by latitude, longitude, and month as he did the rest of his 
total of 53,877 whales of six species (see his pp. 10 and 18). Our note is 
a response to this omission, and like Townsend's paper is based only 



'Museum of Comparative Zoology. Cambridge. MA 02 1 38 and Woods HoleOceano- 

graphic Institution, Woods Hole, MA 02543. 

: Woods Hole Oceanographic Institution, Woods Hole, MA 02543. 



2 breviora No. 476 

on Yankee whaling logbooks. Thus it does not deal with the historic 
Eubalaena fishery of the eastern North Atlantic, well-established in 
the Bay of Biscay by the 12th century, and continuing intermittently 
along the western coast of Europe from the Mediterranean to the 
Barents Sea (where this whale got the name of nordkaper) and 
Iceland; this fishery pretty well stopped in the 1920's, and now 
Eubalaena is very rarely seen in those waters. 

Because so little is known, the location and season of even Town- 
send's few North Atlantic right whales are important. We have con- 
sulted 12 of his 15 sources (Table 1), but failed to find the other three 
logbooks, which he credits with one North Atlantic right whale each 
(sloop Greyhound, 1753; ship Governor Troup, 1868-70; bark Sea 
Ranger, 1879-84). The Providence Public Library has a partial 
journal by William H. Tilton, a boatsteerer/ third mate of Sea 
Ranger during this voyage, but it does not tell where whales were 
encountered. The 1 2 logbooks that we read are listed by Townsend as 
recording the other 32 Eubalaena glacialis. 

Our results do not entirely agree with Townsend's. For one thing, 
as he explains on page 16, his counts of whales include "not only the 
whales turned into oil ['saved', as the whalers said], but also those 
killed. . . but subsequently lost." In our logbook reading we have 
tried to separate these categories, and since we are interested more in 
whale distribution than in oil production, we have also counted 
whales sighted (and identified) but not struck. The logs do not always 
specify the numbers sighted if greater than 1 or 2. When they said 
merely "right whales," we counted it as 2+; the sum of 5+ and 4+ is 
given conservatively as 9+, though the actual count might have been 
appreciably higher. We assume that these old-time whalers were 
competent at recognizing the species of interest — probably much 
more reliably than most seafarers of today; sometimes sightings were 
logged simply as "whales," and these we did not count. We cannot 
explain our discrepancies with Townsend, and can only say that we 
have reported what we found in our reading. We failed to find 
mention of the single Eubalaena each that he recorded for Jireh 
Swift and Emma Jane, and of the 3 he ascribed to Endeavor. 



1983 



NORTH ATLANTIC RIGHT WHALES 



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breviora No. 476 



FINDINGS 



The 1 2 voyages summarized (Table I ) for Eubalaena glacialis in the 
North Atlantic were made by 1 1 whaleships over a 45-year span ( 1 853 
to 1898). The highest and lowest latitudes reached by these are well 
within the known range of this species. No Eubalaena were recorded 
in the months of January, April, September, October, and December. 
The records from November through March are all south of N. lat. 
32°, and those from May through August are between N. lat. 46° and 
62°, but there is no detailed evidence for migration patterns. We find 
the whales in high latitudes in summer and lower in winter, but there 
are no helpful hints of routes to and fro. The wint«r records are right 
along the shore on both sides of the Atlantic (Cintra Bay, N. lat. 
23° in western Africa, and N. lat. 31° at the Georgia-Florida border 
in North America); on each coast the ship sometimes anchored and 
whaled from the boats. 

Six voyages (Ansel Gibbs in 1 868, Daniel Webster and A. J. Ross 
in 1877, A. R. Tucker in 1880, Palmetto in 1886, and Canton in 
1897) reported whales in June, July, and August to the eastward of 
Cape Farewell, Greenland, between N. lat. 59° and 62°, W. long. 32° 
and 38°, at the southern entrance to Denmark Strait. They logged 12 
whales saved, 7 struck but lost, and at least 16 more sighted; Town- 
send recorded 16 whales, but without specifying the region (3 of these 
voyages whaled also in other areas). These 35 whales were encoun- 
tered in 26 ship-weeks of hunting in five years, which implies a 
consistently good hunting ground. This ground is not referred to by 
name in these logs. We found no hint that bowheads were seen or 
taken, although they might have been expected in these waters. (For a 
recent report, see Jonsgard, 1982, Fig. 1.) As pointed out by J. A. 
Allen (1908: 281-288 and especially 286), since the 17th century 
whalers have distinguished bowhead (Balaena mysticetus) from 
nordkaper, or right whale in American usage (Eubalaena glacialis), 
both on morphology and oil yield, as well as on behavior. While the 
Ross was on these Denmark Strait grounds near N. lat. 60° and W. 
long. 35°, the whales were specified as "Wright Whales" and seven or 
more were sighted, of which four were saved. After leaving these 
grounds, the Ross went through the Canaries, and on 24 November 
"Saw A Wright Whale" in "Senter Bay" (Cintra Bay) on the African 
coast. Therefore we conclude that only Eubalaena glacialis were 
found on these voyages. 



1983 



NORTH ATLANTIC RIGHT WHALES 



70° N 



4> ** 

' VII-68 




Figure 1 . Distribution of North Atlantic right whales (Eubalaena glacialis) recorded 
in American whaling logbooks between 1857 and 1897. The dots show where whales 
were encountered at sea, and the coastal encounters are indicated by arrows. Months 
are indicated bv roman numerals. 



Although these whaleships departed and returned to port (usually 
New Bedford) in all seasons, there is no mention of right whales seen 
near the New England or Canadian coasts. Before the mid 1700's 
right whales were caught in Massachusetts waters from small shore- 
based boats, mostly in the winter and spring (Macy, 1835). From 
1955 to the present, right whales have been observed locally in all 
seasons (Schevill, Moore, and Watkins, 1981; Watkins and Schevill, 
in press). It seems highly unlikely that these 19th century whalers 
would have ignored such valuable animals wherever seen, and so it 
may be that Eubalaena were then very scarce in these coastal waters, 
although they had been abundant there from the 16th well into the 
18th centuries. 



6 breviora No. 476 

Against the background of scanty and often imprecise data on the 
seasonal and geographic distribution of Eubalaena glacialis, even 
these few additional points are significant. We have mapped about 60 
whales logged on 9 voyages by 8 ships during the 40 years from 1857 
to 1 897. More than half of these were in the Denmark Strait area in 
June, July, and August. More than 1 3 of the remainder were at Cintra 
Bay in November, February, and March. 

A question about the North Atlantic Eubalaena has been whether 
there was more than one stock in the population. Did the relative 
abundance that we report for Denmark Strait a hundred years ago 
hint that right whales from both sides of the North Atlantic sum- 
mered together, indicating one general population for this ocean? Or 
might there have been two stocks, with Cintra Bay perhaps being in 
the winter range of the nordkapers of the Barents Sea? This is the sort 
of problem for which radio tagging and tracking are needed. There 
are still at least several hundred Eubalaena in the North Atlantic. 

ACKNOWLEDGMENTS 

We would like to acknowledge help from Stuart C. Sherman, who 
told us where to find the logbooks, Richard Kugler and Virginia 
Adams of the Old Dartmouth Historical Society, Paul Cyr of the 
New Bedford Public Library, and Lance Bauer of the Providence 
Public Library; Colleen Hurter helped with bibliothecal matters. 
William A. Watkins and Richard H. Backus helpfully commented on 
the manuscript. Support for this study was from the Oceanic Biology 
Program of the Office of Naval Research, Contract No. N000 14-82- 
C0019 NR083-004. This is Contribution No. 5241 from the Woods 
Hole Oceanographic Institution. 

LITERATURE CITED 

Allen, G. M. 1916. The whalebone whales of New England. Mem. Boston Soc. Nat. 

Hist. 8(2): 107-322, plates 8-16. 
Allen, J. A. 1908. The North Atlantic right whale and its near allies. Bull. Am. 

Mus. Nat. Hist., 24: 277-329, plates 19-24. 
Collett, R. 1909. A few notes on the whale Balaena glacialis and its capture in recent 

years in the North Atlantic by Norwegian whalers. Proc. Zool. Soc. Lond., 1909: 

91-98, plates 25-27. 
JonscjAkd, A. 1982. Bov/head (Balaena mysticetus) surveys in the Arctic Northeast 

Atlantic waters in 1980. Rep. Int. Whal. Commn., 32: 355-356. 



1983 NORTH ATLANTIC RIGHT WHALES 7 

Krai's, S. D., and J. H. Prescott. 1981. Distribution, abundance and notes on 

the large cetaceans of the Bay of Fundy, summer and fall 1980. Final Report to 

the U.S. Department of Commerce, NOAA, National Marine Fisheries Service, 

87 pp. 
Macy, O. 1835. The History of Nantucket. Hilliard, Gray, and Co., Boston, 300 

pp. Reprinted by Research Reprints Inc., New York, 1970. 
Reeves, R. R. and R. L. Brownell, Jr. 1982. Baleen whales. Eubalaena glacialis 

and allies, pp. 415-444. In J. Chapman, G. Feldhamer (eds.). Wild Mammals of 

Nurih America. The Johns Hopkins University Press. 
Reeves, R. R., J. G. Mead, and S. Katona. 1978. The right whale, Eubalaena 

glacialis, in the western North Atlantic. Rep. Int. Whaling Comm., 28: 303-312. 
Schevill, W. E., K. E. Moore, and W. A. Watkins. 1981. Right whale, Euba- 
laena glacialis, sightings in Cape Cod waters. Ref. No. 81-50, Woods HoleOcean- 

ographic Institution, 16 pp. 
Slijper, E. J., W. L. van Utrecht, andC. Naaktgeboren. 1964. Remarks on the 

distribution and migration of whales, based on observations from Netherlands 

ships. Bijdragen tot de Dierkunde, Aflevering 34: 3-93. 
Thompson, D'Arcy W. 1928. On whales landed at the Scottish whaling stations 

during the years 1908-1914 and 1920-1927. Fishery Board Scotl., Sci. Invest. 3. 

39 pp. 
Townsend, C. H. 1935. The distribution of certain whales as shown by logbook 

records of American whaleships. Zoologica, 19(1): 1-50, plates 1-4. 
Watkins, W. A., and W.E. Schevill. In press. Observations of right whales (Eubalaena 

glacialis) in Cape Cod waters. Fishery Bull. 

POSTSCRIPT 

While this paper was in press, we were enabled by the courtesy of 
Professor Howard E. Winn of the University of Rhode Island to 
consult a collection of about half the worksheets used in preparing 
the maps of Townsend's 1935 compilation. These had been pre- 
served at the New York Zoological Society; for each logbook they 
tally the date, location, and species of each whale taken. The log- 
books are of course the primary sources, but these tallies tell us 
something of the compilation, and account for occasional discrep- 
ancies. Thus we have learned that the particular Emma Jane log- 
book read by us is incomplete; it ends on 1 January 1880. The tally 
shows that the voyage continued into August 1881, and that this 
schooner did take a right whale on 1 5 February 1 880 off Brunswick, 
Georgia, near where Golden City took her 2 whales of 1876 and 
1882, also in the latter part of February. The tally for Endeavor 
1854-1856 confirms our failure to find in her logbook any right 
whale captures during either North Atlantic passage, and shows that 



8 breviora No. 476 

the 3 whales published as from that ocean were actually taken in the 
Sea of Okhotsk. The Governor Troup tally for 1868-1870 tells us 
that the whale listed for the North Atlantic was taken in the South 
Atlantic, 20 miles east of Tristan da Cunha in about 37° 15'S. We 
still have not seen her actual logbook. The Canton tally for 
1897-1898 counts 1 of the 2 whales that we list as struck and lost in 
July; the other 2 cited as North Atlantic turn out to have been taken 
in the South Atlantic in November. The sheets for Jireh Swift are 
missing, as are those for Sea Ranger and the sloop Greyhound; we 
hope that someone will find our three missing logbooks. 



MUft COMP. zooc 
LIBRARY 

MR 1 8 885 

REV r^R A 

.iseum of Comparative Zoology 



US ISSN 0006 9698 



Cambridge, Mass 7 September 1984 Number 477 

NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. 

II. ANOLIS PROPINQUUS, ANOTHER NEW SPECIES 

FROM THE CLOUD FOREST OF WESTERN COLOMBIA. 

Ernest E. Williams 1 

Abstract. A. propinquus, new species, from the cloud forest of Departamento 
Valle in western Colombia resembles sympatric A. calimae Ayala, Harris, and 
Williams in the absence of an interparietal scale, but differs by its uniform coloration, 
distinctly keeled head scales, elongate supraciliary scales, and blue rather than yellow 
dewlap. 

Another western punctatus group species has languished un- 
recognized in the Museum of Natural History, the University of 
Kansas (KU), since its collection in 1974. Known only from a male 
near hatchling, it is unremarkable as regards color, but it is 
structurally nearly as distinctive an animal as the recently described 
A. calimae with which it occurs. 

Because it is another member of an anole fauna that is just 
beginning to be known, it receives the Latin name that means 



'neighbor': 



Anolis propinquus, new species 
(Figs. 1-3) 



Type: KU 169833, male juvenile. 

Type locality: Rio Calima, 1.5 km W Lago Calima, Valle, 
Colombia, W. E. Duellman, coll., 15 September 1974. 

Diagnosis. Another green cloud forest species of the punctatus 
group somewhat similar to A. calimae and, like that species, lacks a 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. 



BREVIORA 



No. 477 




Figure 1. Anolis propinquus, new species. Holotype, K.U 169833. Dorsal view of 
head. 



1984 



ANOL1S PROPINQUUS. NEW SPECIES 




Figure 2. Anolis propinquus, new species. Holotype, K.U 169833. Lateral view of 
head. 



parietal eye, but distinguished among other features by keeled head 
scales, an elongate supraciliary scale which occupies half the 
supraciliary margin, a higher number of loreal rows (7) and a higher 
lamellar count under phalanges ii and iii of the fourth toe (25), and 
its blue rather than yellow dewlap. 

Description. Head. Blunt, short. Head scales rather small, most 
scales unicarinate, the keels irregular in direction. Twelve scales 
across snout between second canthals. Frontal depression very 
shallow, the scales within it larger than those anterior to it. Six 
scales border rostral posteriorly. Anterior nasal scale weakly 
differentiated, narrow, separated from the rostral-first supralabial 
suture by one scale. Eight scales between supranasals dorsally. 

Supraorbital semicircles separated by three rows of scales, the 
lateral ones almost as large as the scales of the semicircles and 
keeled, the middle row minute, granular. Supraocular disk mod- 
erately developed, containing ca. 21 keeled scales, those next to the 
semicircles largest, grading laterally into smaller but still keeled 
scales, two rows of subgranular scales separating the disk on each 
side from the supraciliaries. Anterolateral corner of supraocular 
area with subgranular scales, posterolateral corner with minute 
granular scales. One elongate supraciliary extending half the 
supraciliary distance and followed by granules. Canthus blunt, of 
ca. seven scales, the first, second and third larger. Seven loreal rows 
below second canthal. 



BREVIORA 



No. 477 




Figure 3. Anolis propinquus, new species. Holotype, KU 169833. Ventral view of 
head. 



Temporal scales granular, a triangle of subgranular intertemporal 
scales. Supratemporal scales granular, minute, becoming abruptly 
larger toward the interparietal area. Ear small, but larger than any 
presumptive interparietal. Enlarged scales of interparietal area 
largest laterally, all abruptly larger than the minute nape scales 
which are distinctly smaller than the dorsals. No parietal eye. 

Suboculars weakly keeled, separated from the supralabials by one 
row of scales. Seven to eight supralabials to the center of the eye. 

Mental divided, each half a little wider than long. Two small 
scales posteriorly in the notch between the mentals. Two much 
larger rectangular scales lateral to them, between the trapezoidal 
first sublabials. Three sublabials in contact with the infralabials on 
each side. 

Throat scales swollen, rather elongate, minute posteriorly, 
becoming larger anteriorly and laterally. 



1984 ANOLIS PROPINQUVS, NEW SPECIES 5 

Trunk. Dorsal scales granular, convex, subequal. Ventrals not 
much larger, swollen, smooth, juxtaposed or subimbricate, in 
transverse rows. An umbilical scar still detectable. 

Dewlap. Retracted in the unique type, not readily visible. Large, 
scales crowded, swollen, imbricate, raised into series of rows or 
ridges, a little smaller than the ventrals. 

Limbs and Digits. Anterior arm and leg scales unicarinate 
except multicarinate at the knee. Supradigital scales multicarinate. 
Ca. 25 lamellae under phalanges ii and iii of fourth toe. 

Tail. Weakly compressed. No dorsal crest. One row of keeled 
scales middorsally, not differentiated from those lateral to it. No 
evident verticils. Enlarged postanals present (#). 

Size. The snout-vent length of the unique type is 41 mm. 

Color. As preserved, the specimen shows no pattern: it is dark 
above, light below and the dewlap skin appears to be dark. W. E. 
Duellman provides notes of color in life: "Dorsum green. Venter 
pale bluish green. Dewlap pale blue. Iris dark brown. Mouth bluish 
black, tongue pink." 

Habitat. The only notes on habitat or ecology are Duellman's: 
"Sleeping on herb ca. 0.5 m above ground at night." 

Comparisons. The most pertinent comparison of A. propinquus 
is with sympatric and just-described A. calimae (Ayala et a!., 1983) 
(Table 1). The unique type is a near hatchling, but the dewlap is 
already well indicated, extending posterior to the axilla and the high 
number of lamellae under the fourth toe (25) indicates a species of at 
least moderate size. 

It is again astonishing to find the interparietal reduced (i.e., a 
parietal eye lacking). In contrast to A. calimae, the supraciliary 
scales are unusually well developed and the head scales very 
distinctly keeled. The blue dewlap is also distinctive. 



ACKNOWLEDGMENTS 

I am grateful to William E. Duellman, Curator at the Museum of 
Natural History, University of Kansas for the opportunity to 
examine and describe this peculiar animal. The drawings are by 
Laszlo Meszoly. 



BREVIORA 



No. 477 



Table 1. Comparison of Anolis calimae and A. propinquus. 





calimae 


propinquus 


scales across snout 


wrinkled 


unicarinate 


number between 


7-10 


12 


second canthals 






circumnasal/ rostral 


separated by one round 


a weakly differentiated 


scale 


scale 


subtriangular anterior 
nasal scale 


scales between 


2 large squarish scales 


3 rows of scales, the 


supraorbital 


in males, none or one row 


lateral ones large and 


semicircles 


of small scales in females 


keeled, the middle minute 


scales of supraocular 


smooth or wrinkled 


keeled 


disk 






differentiated 


one short or none 


one very elongate 


supraciliaries 




C/2 supraciliary margin) 


loreal rows 


4-5 


7 


interparietal 


not differentiated 


not differentiated 


scales between 


not determinable 


not determinable 


interparietal and 






semicircles 






scales between 





1 


supraoculars and 






supralabials 






supralabials to center 


7 


7 


of eye 






trunk scales 


uniform, granular 


uniform, granular 


ventrals 


larger than dorsals, 


larger than dorsals, 



dewlap 
dewlap scales 

dewlap skin 
postanal scales 



smooth, juxtaposed 
or subimbricate 

large in <$, 
smaller in 9 

lateral scales smaller 

in males than in females, 

smaller than ventrals 

unpigmented in $, 
pigmented in 9 

very large in $, 
absent in 9 



smooth, juxtaposed 
or subimbricate 

large in juvenile male 
only male known 

pigmented in $ 
large in $ 



1984 



ANOLIS PROPINQUUS, NEW SPECIES 



Table 1. Comparison of Anolis calimae and A. propinquus. 



calimae 



propinquus 



scales posterior 


to 


smooth 


vent 






tail 




compressed 


tail crest 




not a crest, but 2 
middorsal rows 



smooth 



slightly compressed 

not a crest but a 
single middorsal row 



LITERATURE CITED 



Ayala, S., D. Harris, and E. E. Williams. 1983. New or problematic Anolis 
from Colombia. 1. Anolis calimae, new species from the cloud forest of western 
Colombia. Breviora Mus. Comp. Zool., No. 475, pp. 1-1 1. 



■\ <-\f- 



>HV'- ,: 



Mu S. COMP. 200L 
LIBRARY 

MAR181985 

BREV I u ^ A 

Museum of Comparative Zoology 

US ISSN 0006-9698 

Cambridge, Mass. 7 September 1984 Number 478 

NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. 

III. TWO NEW SEMIAQUATIC ANOLES 

FROM ANTIOQUIA AND CHOCO, COLOMBIA. 

Ernest E. Williams 1 

Abstract. Two new semiaquatic anoles from Colombia, partly sympatric, and 
sometimes syntopic, prove to belong to distinct lineages despite convergence in habits 
and habitat. The larger of the two — A. maculigula, new species — belongs to the alpha 
section of the genus Anolis and the eulaemus subgroup of the A. aequatorialis species 
group. It is confined, so far as known, to the Departamento Antioquia and 
apparently to larger streams. The smaller species, A. rivalis, new species, is a beta 
anole of the lionotus species group and tends to prefer smaller streams and extends 
beyond the known range of A. maculigula in Antioquia and in Choco. 

In 1968 Norman J. Scott, collecting on a tributary of the Rio 
Arquia in western Antioquia, Colombia, obtained a large anole of 
aquatic habits which he thought resembled A. aquaticus. It proves, 
however, to be a new species belonging to the alpha section of the 
genus. Syntopic with this species was a smaller form which Scott 
recognized as a member of the lionotus species group (beta section). 
This also proves to be new. These specimens were deposited in the 
collections of the Los Angeles County Museum (LACM). 

Since these first collections, additional material of both species 
has been obtained in another area of western Antioquia some 70 km 
farther north by Juan Manuel Renjifo and Vladimir Corredor 
(material in the Instituto de Ciencias Naturales, Bogota [ICN]). 
Specimens of the larger species have been collected at Urrao, 
Parque Las Orquideas, ca. 50 to 60 km east of the type locality, by 



1 Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. 



BREVIORA 



No. 478 



Horatio Echeverri for Marco Serna (collection of Colegio San Jose, 
Medellin (CSJ). Specimens of the smaller species have been 
collected in the Choc6 independently by Philip Silverstone (MCZ) 
and Charles Myers (ICN). A further Choc6 specimen of the smaller 
species has been found in the collections of the San Diego Natural 
History Society (SDSNH) collected by C. B. Perkins. 

The larger new anole may be named, with reference to its heavily 
spotted throat: 

Anolis maculigula, new species 
(Figs. 1-3) 

Holotype: LACM 42150, adult male. 

Type locality: Quebrada San Lorenzo, tributary of the Rio 
Arquia near the small town of Belen (6° 15'N 76° 39'W), about 10 to 
15 km upstream from the junction of the Rio Arquia with the Rio 
Atrato, western Antioquia, Colombia, N. J. Scott coll., 17 April 
1968. 

Paratypes: Antioquia: LACM 42142, 42144-49, 42151-54, same 
data as the type; ICN 5917, Camp Pegadorcito (Ingeominas), (about 
6° 42'N 76° 27'W,) on the height of the Cordillera Occidental 
between Frontino on the south and Dabeiba on the north, Rio 




Figure 1. Anolis maculigula, new species. Type, LACM 42150. Dorsal aspect of 
head. 



1984 



TWO NEW SEMIAQUATIC ANOLES 




_2 



BREVIORA 



No. 478 




Figure 3. A. maculigula, new species. Type, LACM 42150. Throat pattern. 



Amparrad6, 805 m elevation, Juan M. Renjifo and Vladimir 
Corredor coll., 16 September 1981; ICN 5918, at the first stream 
beyond Pegadorcito, J. M. Renjifo and V. Corredor coll., 14 
September 1981; CSJ 308-309, Urrao, Parque Las Orquideas, 
Horatio Echeverri coll., 21 May 1981; CSJ 431, 445, 447, Urrao 
(Rio Calles), Marco A. Serna and Horatio Echeverri coll., 10-12 
July 1983. 



1984 TWONEWSEMIAQUATIC ANOLES 5 

Diagnosis. An alpha anole of moderate size related to A. 
eulaemus Boulenger and, like the latter, having small head scales 
and narrow digital dilations, but differing in having fewer scales 
between the supraorbital semicircles and between the semicircles 
and the interparietal, differing also in color, habits and habitat. 

Description. Head. Head scales small, pustulose, swollen, 
keeled. Twelve to 19 scales between the second canthals. Eight to 11 
scales bordering rostral posteriorly. Anterior nasal separated from 
rostral by one or two scales. 

Two to three scales between supraorbital semicircles. A few 
supraocular scales distinctly enlarged, no well-defined supraocular 
disk. One to two elongate supraciliaries on each side followed by 
granules. Canthus distinct, seven canthal scales, the second longest. 
Eight to 1 1 loreal rows, scales small, subequal. 

Temporal and supratemporal scales granular. A very indistinct 
double row of enlarged granules at margin between temporal and 
supratemporal areas. Scales around interparietal enlarged, inter- 
parietal larger than or about equal to ear, separated from 
supraorbital semicircles by one to three scales on each side or in 
contact. 

Suboculars separated from supralabials by one row of scales, 
posteriorly grading into temporal granules, anteriorly diminishing 
gradually in size in front of eye. Seven to ten supralabials to center 
of eye. 

Mentals wider than deep, in straight line contact with eight to ten 
scales between infralabials, grading in size from the ihfralabials 
toward the center. Sublabials not clearly differentiated. Central 
gular scales small, swollen, even conical, grading laterally into much 
larger scales. 

Trunk. Two to three middorsal rows slightly enlarged, tricar- 
inate, imbricate, grading into small, keeled but juxtaposed flank 
scales, showing small granules between them. Ventral scales equal to 
or smaller than the middorsals, smooth, flat or swollen, sub- 
imbricate or juxtaposed, sometimes showing small granules. 

Dewlap. Large in male, extending onto first third of belly. Edge 
scales ca. equal to ventrals. Lateral scales in rows, several scales 
across, widely separated by naked skin. Barely indicated in female. 

Limbs and Digits. Largest arm and leg scales larger than 
ventrals and uni- to multicarinate. Supradigital scales multicarinate. 
Sixteen to 22 lamellae under phalanges ii and iii of fourth toe. 



6 BREVIORA No. 478 

Tail. Distinctly compressed to very strongly compressed, most 
strongly in the adult male, a low crest of unicarinate scales, the crest 
scales at least 2X those of the lateral rows. The other caudal scales 
uni- to multicarinate. No enlarged postanals in male. 

Size. The male type measures 98 mm snout to vent. A topotypic 
male measures 72 mm, while the largest topotypic female is 73 mm 
snout-vent length. The male from Urrao (CSJ 308) is 107 mm snout 
to vent; the unregenerated tail is 215 mm long. The female (CSJ 309) 
from the same locality has 75 mm snout-vent length. 

Color in Life. Normal Scott has provided notes from life for 
topotypic specimens. "Adult male: Dorsum with an obscure pattern 
of five dark brown blocks separated by lighter gray brown areas and 
broken by, on each side, a dorsolateral stripe of the same color. 
Dorsal crest paler and green. Tail and legs banded with dark gray 
brown and light gray green. On flanks a mixture of olive gray and 
orange flecks on a green ground. Side of head a mixture of orange 
and gray flecks, lips mottled and with blue flecks. Soles of feet olive 
brown. Throat with a series of orange and blue stripes. Chin mottled 
with orange, blue and gray, the blue predominant. Iris as in female 
but white ring tinged with green. Dewlap color complex: base with 
orange stripes on a blue gray ground, anterior third pale bluish rose, 
posterior portion white becoming pale blue toward belly. Juveniles 
colored much as females but green brighter on head, neck and sides. 
Adult female: Dorsum dark gray brown with gray black bar, lighter 
areas on neck, head and legs greenish gray. Sides mottled gray, olive 
green and black with a few light gray punctations. Side of head 
mostly greenish. Venter clear white, throat mottled green and 
white, underside of head patterned with white and gray, the chin 
with green and gray. Iris dark brown enclosing a narrow white 
ring." 

Juan Renjifo in a note to Stephen Ayala reports the color in life 
of the Amparrado maculigula (translated): "Color green (lichenate) 
with dark blotches on the back, the first above the shoulders. A 
blotch at the shoulder dark with a white spot in the center." 

Color in Preservative. Color now differs substantially from that 
reported for the fresh specimens. All blues, pinks and greens are 
gone. There are only light and dark grays but the complex pattern is 
retained. The dark crossbars of the middorsum contain light spots 
and the lower flanks tend to be vermiculate. The throat is boldly to 
weakly vermiculate, less strongly vermiculate in males than in 



1984 TWONEWSEMIAQUATIC ANOLES 

females, in which the heavily marked throat contrasts strikingly 
with the light and unmarked belly. The skin of the throat fan is gray. 

Habitat and Habits. The type locality, Quebrada San Lorenzo, 
is described as a stream about 10 m wide with occasional waterfalls, 
with a moderately steep gradient and huge boulders. A. maculigula 
was most common where there were 3 to 4 m diameter moss-covered 
boulders at the head of pools, when first seen most were on the 
upstream vertical face of the boulders in the spray zone and tried to 
escape under the overhanging edge of the boulders. In Quebrada 
San Lorenzo the smaller (lionotus group) species also occurred but 
was less common. (See further below.) No comparable details are 
available for the habitat of ICN 5917 and 5918, but the latter 
specimen is reported as on rocks in a stream. 

Comparisons. A. maculigula is clearly an alpha anole; a caudal 
vertebra dissected from a broken tail shows no trace of transverse 
processes. It is referred to the aequatorialis species group on the 
basis of very small head scales, large size and narrow digital 
dilations. Since its toe pads are "raised," i.e., overlap distally the 
proximal scales under the first phalanx, it is further referred to the 
eulaemus subgroup in contrast to species with pads "not raised," 
i.e., not overlapping but continuous with the scales of the first 
phalanx. (The latter are the aequatorialis subgroup.) 

Within the eulaemus subgroup, in which there are a confusing 
number of undescribed populations, A. maculigula is distinctive in 
its pustulose head scales, its small ventrals, usually smaller than the 
small dorsals, and in the relatively large size of the interparietal 
which is almost always larger than the ear, in the very compressed 
tail and in a distinctive coloration. It may well be unique among the 
known eulaemoid forms in its semiaquatic habits also. Certainly no 
other member of the group shows so compressed a tail or 
maculigula\ tendency to a lined pattern. 

The second and smaller species may very appropriately receive 
the Latin name that means "user of the same stream": 

Anolis rivalis 
(Figs. 4-7, 12, 14) 

Holotype: LACM 42124, adult male. 

Type locality: Belen, Rio Arquia, Antioquia, Colombia (6° 
15'N, 76° 39'W), Norman J. Scott coll., 17 April 1968. 



8 BREVIORA No. 478 

Paratypes: Antioquia: LACM 42128-133, 42135, 42137-139. 
MCZ 115720-722 from the type locality, Norman J. Scott coll.; 
LACM 42125-127, 42141, 45002-07, 51540, Finca Los Llanos, Rio 
Arquia, Philip Silverstone coll., 1968; ICN 5912, Camp Pegadorcito 
(Ingeominas), 45 minutes by helicopter from Medellin, on the height 
of the Cordillera Occidental between the towns of Frontino to the 
south and Dabeiba to the north, Rio Amparrad6, 805 m elevation, 
(about 6° 42'N, 76° 27'W,) Juan M. Renjifo and Vladimir Corredor 
coll., 13 September 1981; ICN 5913, Filo Amparrad6, the same area 
and collectors, 16 September 1981; ICN 5914, Camp Chontaduro, 
same area and collectors, 9-12 September 1981; ICN 5915, same 
area and collectors, 14 September 1981. Choco: LACM 72766, 
72772, MCZ 100353, Alto de Buey, P. Silverstone coll., 1968; ICN 
4053, Quebrada Mutata, 200 m, northern base of Alto de Buey, C. 
W. Myers, John Daly and Michael G. A. Hill coll., 18-24 October 
1978; SDSNH 31163, "Port Utria," = Puerto Utria (6° 02'N, 76° 
23'W), C. B. Perkins coll., 25 February 1938. 

Diagnosis. A member of the lionotus group of beta anoles, 
differing from the remainder of the group in the combination of 
small but not minute head scales (13 to 18 across snout between 
second canthals), one to two rows of scales between the supraorbital 
semicircles, interparietal in contact with the semicircles or separated 
by no more than two rows of scales, and by a zone of moderately 
large, indistinctly keeled, flat scales on the middorsum in 11 to 18 
rows. 

Description. Head. Scales small, uni- to multicarinate. Thirteen 
to 18 scales across snout between second canthals. Seven to nine 
scales border rostral posteriorly. Anterior nasal scale separated 
from rostral by one scale. Nine scales between supranasals. Scales in 
posterior portion of frontal depression smaller than those anteriorly 
placed. 

Supraorbital semicircles separated medially by one to three 
scales, in contact laterally with the largest scales of the supraocular 
disks which consist of a variable number of wrinkled or keeled 
scales, a few of which may be much larger than the others. One to 
three elongate supraciliaries continued posteriorly by a series of 
smaller scales. Canthus distinct, canthals 9. second largest. Six to 10 
loreal rows, lowermost largest. 

Temporals small and flat, not granular. A distinct double 
supratemporal row, dorsad of which nearly granular scales grade 



1984 



TWO NEW SEMIAQUATIC ANOLES 










Figure 4. /I. rivalis, new species. Type, LAGM 42124. Dorsal aspect of head. 




Figure 5. A. rivalis, new species. Type, LACM 42124. Lateral aspect of head. 



into the flat scales surrounding the large interparietal, usually larger 
than ear and in contact with the semicircles or separated by one to 
two scales. Scales posterior to interparietal not different in size from 
middorsals but grading into smaller nape scales which then grade 
posteriorly into the dorsals. Suboculars weakly keeled, narrowly in 
contact with supralabials or separated by one scale row, grading 



10 



BREVIORA 



No. 478 




Figure 6. A. rivalis. new species. Type, LACM 42124. Ventral view of chin. 



anteriorly into loreals, posteriorly more sharply distinct from the 
temporals. Six to nine supralabials to the center of the eye. 

Mental divided, wider than long, in contact with six or seven 
scales between infralabials. Gular scales smallest medially, quadrate, 
swollen. 

Trunk. Middorsal scales flat, hexagonal, wrinkled, in ca. 1 1 to 
17 rows, grading quite gradually into subgranular flank scales, 
larger in the center of the body than on the nape, becoming slightly 
smaller again in the sacral region and on the base of the tail. 
Ventrals smaller than dorsals, keeled, imbricate. 

Dewlap. Moderate, extending onto first third of belly, lateral 
scales weak, in densely packed rows, edge scales keeled, slightly 
larger than ventrals. 

Limbs and Digits. Scales on limbs strongly unicarinate except 
at knees and elbows where they are multicarinate. Supradigital 
scales multicarinate. Twelve to 18 scales under phalanges ii and iii of 
fourth toe. 



1984 



TWO NEW SEMIAQUATIC ANOLES 



11 




3 1Z 



12 



BREVIORA 



No. 478 





Figure 8. A. oxylophus, KU 34262. Frontal and middorsal scales. 





Figure 9. A. lionotus, KU 75951. Frontal and middorsal scales. 



1984 TWONEWSEMIAQUATIC ANOLES 13 





Figure 10. A. macrolepis, MCZ 133000. Frontal and middorsal scales. 




Figure 11. A. poecilopus, KU 113249. Frontal and middorsal scales. 



14 



BREVIORA 



No. 478 





Figure 12. A. rivalis, LACM 42129. Frontal and middorsal scales. 



Tail. Compressed without dorsal crest. Verticils indistinct. 
Enlarged postanals absent. 

Color as Preserved. Brown with darker flanks. Middorsum 
banded. Two narrow light lines on sides, one above shoulder, one 
starting at axilla. Spotting on lower flanks. Limbs banded. Belly 
and throat white or very weakly spotted. Dewlap white. Nape white 
to above level of ear where a darker margin sets it off against the 
dorsal light brown. 

Size. The type male is 62 mm in snout-vent length. MCZ 1 15722 
and LACM 42138, topotypic males, are each 64 mm snout to vent. 

Color in Life. Scott provides color notes on the topotypic 
specimens: "Adult male: Dorsum dark brown with thin green brown 
vertical bands on flanks. Several stripes from axilla to groin, the 
lower ones heavily suffused with red. Light areas on sides dusty 
rose. Venter yellowish white with heavy red suffusion on sides of 
abdomen, throat and underside of head. Iris dark chestnut enclosing 
a narrow yellow ring. Dewlap solid orange. Adult female: Dorsum 



1984 



TWO NEW SEMIAQUATIC ANOLES 



15 




Figure 13. A. rivalis from Quebrada Mutata, 200 m, northern base of Alto del 
Buey, Choc6, Colombia. Photo by C. W. Myers. 



as in male but with only a faint hint of red suffusion. Light areas of 
head white becoming green on snout. Venter yellowish white. 
Dewlap area with pale orange spot, throat anteriorly white. Iris as in 
male." 

Notes by C. W. Myers on the specimen obtained from Quebrada 
Mutata record the following colors: "Brown, changeable to 
brownish green, with lateral line and lower side of neck dirty white. 
Throat gray, dewlap light orange, venter greenish gray. Iris brown. 
Tongue light gray." 

Juan Manuel Renjifo has supplied color notes on ICN 5912 from 
the Rio Amparrad6 region: Greenish brown with bars of lighter 
greenish brown forming chevrons on the middle of the back. Labials 
and flanks rosy salmon. Venter yellowish cream. Gular sac white. 

One entry in Scott's field notes cites an unspecified rivalis from 
Belen as having an orange dewlap. There would appear to be a 
range of dewlap colors in this species. 

Habitat and Habits. At Quebrada San Lorenzo, Scott reports 
A. rivalis both to be less common and to occur on small rocks near 
the bottom of pools rather than on the huge moss covered boulders 
preferred by A. maculigula. They escaped by running from rock to 
rock over the water surface. Some were taken at night sleeping on 



16 BREVIORA No. 478 

vegetation overhanging the stream. He specifically mentions (letter 
of February 28, 1979): "Quebrada San Lorenzo was the only place 
where I got both aquatic species. Nearby Quebrada Barrero had less 
water and seemed to have only the "lionotus" [= rivalis] type." 

The single specimen of A. rivalis obtained by the Myers group in 
1978 was again found in a quebrada, but the notes do not cite any 
larger aquatic species. 

Renjifo in his field notes does not mention close or co-occurrence 
of the two species and confirms the semi-aquatic habits of rivalis in 
the Amparrado region. He cites ICN 5912 as (translated) "in forest 
on trunk above the stream, Vi meter above ground level;" ICN 5914 
"in stream on a stone;" ICN 5915 "sleeping on rocks in the stream 
alongside the current;" and ICN 5918 "in stream.... on rocks." 

Comparisons. A. rivalis is a member of the beta series and of the 
lionotus species group. 

The latter is a series of taxa (the other referred species: oxylophus 
Cope, 1875, lionotus Cope, 1861, poecilopus Cope, 1862, macro- 
lepis Boulenger 1911) that extend from Nicaragua to Ecuador, 
replacing one another with minimal or no overlap, so far as known. 
A sixth taxon belonging to this group occurs in northwestern 
Ecuador and southwestern Colombia. It will be described by 
Kenneth Miyata. All are characterized ecologically by "aquatic" 
habit and riparian habitat and (usually) by a zone of enlarged dorsal 
scales, more or less similar in size over an area of 14 to 20 rows, then 
grading laterally into the flank scales. They have also small keeled 
ventrals and a more or less well developed flank stripe. 

The species of this group ring the changes on just a few 
morphological characters: the size of the head scales, the number of 
scale rows between the supraorbital semicircles, the number of 
scales between the interparietal and the semicircles, the size of 
supratemporal and nape scales, and the size and also the keeling or 
lack of keeling of the middorsals. Each of these characters varies 
independently, and adjacent species tend to be sharply distinct in 
one or more features. 

No revision of the lionotus species group exists nor has South 
American macrolepis been recognized as a member of it until 
recently (Williams, 1976). 

Boulenger (191 1) in describing macrolepis did, indeed, associate it 
with poecilopus but also suggested a relationship to notopholis 
(humilis species group), to which it is only superficially similar. 



1984 



TWO NEW SEMIAQUATIC ANOl.ES 



17 




Figure 14. Map: Localities for semiaquatic anoles in Colombia and adjacent 
Panama. 



18 



BREVIORA 



No. 478 



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20 BREVIORA No. 478 

Taylor (1956) in his discussion of the lizards of Costa Rica called 
attention to the discrepancies between the type description of 
Uonotus Cope, 1861 (with a type locality in Panama) and that of 
oxylophus Cope, 1875 (without exact locality but presumably from 
Costa Rica), which was supposedly a synonym. However, lacking 
Panamanian specimens, he followed then and current usage in 
applying the name Uonotus to Costa Rican animals. My own 
examination of most of the material available in American 
museums shows significant differences between eastern Panamanian 
and western Panamanian and Costa Rican-Nicaraguan specimens. 
Campbell (1973) noted the character of true Uonotus that has most 
impressed me: dorsal scales about twice the size of the ventrals. In 
oxylophus (as in poecilopus) the dorsals and ventrals are about 
equal but in poecilopus the dorsals are strongly keeled, in oxylophus 
smooth or weakly keeled. Table 1 reports the characters dis- 
tinguishing members of this group. 

The new species is distinguished from all other members of the 
Uonotus group by a combination of characters: small but not minute 
head scales, few scales between the supraorbital semicircles, small to 
moderate supratemporal scales, nape scales distinctly smaller than 
the enlarged dorsals which are large, larger than the ventrals, 
wrinkled or weakly keeled and weakly imbricate. From poecilopus 
it is distinguished by its much larger middorsals, fewer rows between 
the supraorbital semicircles and its larger circum-interparietal 
scales. From macrolepis it is distinguished by its somewhat larger 
middorsals, the nape scales distinctly smaller than the large 
middorsals (rather than subequal) and by smaller head scales. 

Discussion. The sympatric occurrence of two semiaquatic 
anoles is not unique: within the Uonotus group there is the example 
of the local sympatry of poecilopus and Uonotus in the vicinity of 
the Panama Canal (Campbell, 1973). In this case the overlap zone is 
suspected to be narrow, but knowledge of the distributions is not 
adequate to demonstrate this. In the case of poecilopus and 
Uonotus, the two overlapping species are very different within the 
group in dorsal and head scale size. (In general, in the Uonotus 
group, although the species are primarily allopatric, the differences 
summarized in Table 1 are greater in adjacent than in distant taxa.) 

The two sympatric semiaquatic species here described are quite 
different in their affinities, belonging to different sections of the 
genus Anolis. They are adequately different in scale characters but 



1984 twonewsemiaouaticanoi.es 21 

not quite so strikingly as the most different species pairs within the 
lionotus group. The similarities in habits, in the pronounced flank 
stripes, and in the strongly compressed tails, are indeed more 
impressive than their differences. 

But these similarities are parallels only and are seen in other 
semiaquatic anoles not at all closely related, as Schwartz has 
demonstrated in describing a taxonomically quite isolated semi- 
aquatic anole from Hispaniola (Schwartz, 1978). The semiaquatic 
anoles, except for the allo-parapatric series that constitutes the 
lionotus group, are not at all a lineage, but only an ecomorph in the 
sense of Williams (1972, 1983). 

ACKNOWLEDGMENTS 

I am indebted to the curators who have provided me the 
opportunity to describe the new species: John Wright and Robert 
Bezy of the Los Angeles County Museum, Pedro Ruiz of the 
Instituto de Ciencias Naturales, Bogota, Colombia and Gregory 
Pregill of the San Diego Natural History Society. Norman Scott 
and Charles Myers have allowed me to quote from their field notes, 
and Pedro Ruiz has transmitted the field localities and observations 
of Juan Renjifo and Vladimir Corredor. Stephen Ayala has 
commented on the manuscript and provided the map which 
compares the distributions of Colombian aquatic anoles. Laszlo 
Meszoly has done the illustrations. The photograph of A. rivalis was 
provided by Charles Myers. 

LITERATURE CITED 

Boilenger, G. A. 1911. Description of new reptiles from the Andes of South 

America, preserved in the British Museum. Ann. Mag. Nat. Hist. ser. 8, 8: 

19-25. 
Campbell, H. W. 1973. Ecological observations on Anolis lionotus and A. 

poecilopus (Reptilia, Sauria) in Panama. Amer. Mus. Novitates, No. 2516, pp. 

1-29. 
Cope, E. D. 1861. Notes and descriptions of anoles. Proc. Acad. Nat. Sci. 

Phila., 1861: 209 215. 
1862. Contributions to Neotropical saurology. Proc. Acad. Nat. Sci. 

Phila.. 1862: 159-188. 
1875. On the Batrachia and Reptilia of Costa Rica. J. Acad. Nat. Sci., 



8: 93-157. 
Schwartz, A. 1978. A new species of aquatic anole (Sauria, Iguanidae) from 
Hispaniola. Ann. Carnegie Mus., 47: 261-279. 



22 BREVIORA No. 478 



Taylor, E. 1956. A review of the lizards of Costa Rica. Univ. Kansas Sci. 

Bull., 38: 3-222. 
Williams, E. E. 1972. The origin of faunas. Evolution of lizard congeners in a 

complex island fauna: a trial analysis. Evolutionary Biology, 6: 47-89. 
1976. South American anoles: the species groups. Pap. Avuls. Zool. S. 

Paulo, 29: 259-268. 
1983. Ecomorphs, faunas, island size and diverse end points in island 



radiations of Anolis pp. 326-370, 481 484. In Huey, R., E. Pianka, and T. 
Schoener (eds.). Lizard Ecology - Studies of a Model Organism, Harvard 
University Press, Cambridge, Mass. 



f- j' •''- 






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B R E V I iSPft A 

Museum of Comparative Zoology 



US ISSN 0006-9698 



Cambridge. Mass. 7 September 1984 Number 479 

AGONISTIC AND COURTSHIP DISPLAYS OF 
MALE ANOLIS SAGREI 

Michelle P. Scott 1 

Abstract. Male Anolis sagrei perform a dewlap fanning display and four different 
bobbing displays in agonistic and courtship encounters. Only one of the bobbing 
displays has a species-specific, stereotyped pattern. The total number of bobs and the 
number of bobs in each unit of this display and the use of the dewlap showed 
considerable variability both within and between individuals. The frequency 
distribution of the type of display used in aggressive encounters showed more inter- 
than intra-individual variability. Dewlap fanning displays were used at a relatively 
higher frequency before crests were raised, but the dewlap was used with the bobbing 
display at a relatively higher frequency by males after the erection of crests. 
Submissive animals displayed less frequently overall and were more apt to use a 
dewlap fanning display, but they were less apt to use the dewlap with a bobbing 
display than a dominant male. Male A. sagrei were less apt to use the species-specific 
stereotyped pattern in courtship than in agressive encounters. The "jiggle" bob 
display immediately preceeded half of the attempted matings. A dewlap fanning with 
erect posture was associated with the termination of a copulation. 

INTRODUCTION 

The brightly colored dewlaps and stereotyped bobbing displays of 
males of the iguanid lizard genus Anolis may communicate 
information regarding species, sex, reproductive state, dominance 
and territorial status, intentions, and level of arousal. Dewlap color 
alone sometimes does not appear to be an important element in 
species recognition and female choice for the solitary anole A. 
earolinensis (Greenberg and Noble, 1944; Crews, 1975a). However, 
in a complex Anolis fauna, color and dewlap proportions may 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. 



2 BREVIORA No. 479 

provide redundant information for species recognition (Williams 
and Rand, 1977). Jenssen's work (1970a) with Anolis nebulosus 
indicates that the male's species-specific bobbing display does 
indeed play a significant role in sexual selection and female choice. 

In examining anoline displays, recent researchers have focused on 
the degree of stereotypy (the amount and possible significance of 
variability within and among individuals) and the complexity of 
display repertoires of different species. Displays may vary either in 
temporal patterns or by the presence or absence of various 
components within the display. Species differ considerably in the 
amount of stereotypy of their displays, and the source of variability 
may be intra-individual or between individuals (Jenssen 1978). 
Anolis carolinensis (Crews, 1975b), A. aeneus (Stamps and Barlow, 
1973), and A. nebulosus (Jenssen, 1971) have very stereotyped 
displays. Each individual performs a consistent display, but there is 
variability between individuals. Different populations of A. nebu- 
losus have been examined and still further variability between 
populations was found. Other species, however, like A. limifrons 
(Jenssen and Hover, 1976) show greater intra-individual variability 
than inter-individual differences. At the extreme, almost all (97%) of 
the variability of A. townsendi displays is attributable to intra- 
individual variation (Jenssen and Rothblum, 1977). Anolis opal- 
inus, on the other hand, was found to be almost completely lacking 
stereotypy at any level, individual or population, in that there was 
no single discrete display pattern. However, 75% of the analyzed 
displays fit a theme which explained the highly variable bob number 
and display cadence (Jenssen, 1979). 

The complexity of the display repertoire of a species is another 
important feature. Researchers (e.g., Carpenter and Grubitz, 1961) 
first described only two basic types of displays, a courtship pattern 
of rapid head bobs and a species-unique "signature" pattern. The 
latter type was originally labeled an "assertion" display that, with 
modifiers, blends into a "challenge" display (Carpenter, 1967; 
Jenssen, 1970b, 1977; Crews, 1975b). Typically, if other display 
types were described, they consisted of the signature display plus 
modifiers (e.g., four-legged push-up posture preceding dewlap 
fanning), or they may be characterized by a different coordination 
of the dewlap extensions during the head bob sequence. Recently, 
researchers have found greater complexity in the display repertoire 
of some anoles (Hover and Jenssen, 1976; Jenssen and Rothblum, 



1984 DISPLAYS OF A NOLIS SAGREI 3 

1977). Anolis aeneus, A. limifrons, and A. townsendi, for example, 
have several clearly distinct display types which differ not just by the 
modifiers or the context in which they appear, but also by different 
bob patterns. 

This paper described the display repertoire of male Anolis sagrei 
in agonistic and in courtship situations. A description and analysis 
of the species-specific stereotyped display and its variable elements 
are given. An attempt is made to analyze these displays and their 
modifiers in terms of the social context in which they appear and the 
effect they have on conspecifics. The possible role of the species- 
specific signals used in sexual selection is discussed. 

MATERIALS AND METHODS 

Anolis sagrei has a wide distribution, occurring on Cuba, 
Jamaica, the Bahamas, Little Cayman, Cayman Brae, Swan Island, 
and coastal areas of Mexico and Belize, and has recently been 
introduced into Florida (Williams, 1976). It occupies a relatively 
open habitat. During the breeding season, males and females are 
strongly territorial. A male territory has been described by Evans 
(1936) as at least 50 m 2 and containing up to three females. In 
cultivated areas, it is typical for males to occupy a hedge row or 
fence at approximately 6 m intervals (Evans, 1936; Scott, personal 
observation). 

Twenty males (55-62 mm snout to vent length), 12 females and 8 
juveniles were collected near the Universtiy of South Florida, 
Tampa, in May 1979 when the mature males and females were 
sexually active. 

The lizards were held in experimental cages of two designs. In 
Design 1, one large male, a juvenile male and two to three females 
inhabited an aquarium 75 cm long X 32 cm deep X 45 cm high. Two 
of these aquaria so arranged were placed end to end with a 
removable opaque divider between them. Each contained a live 
plant and a prominent perching branch. The far side of each 
aquarium was covered by bark which was also favored as a perching 
site. Thus, when the opaque divider was removed, the two territorial 
males could see each other but were physically separated by a 
double pane of glass. In this design, 5 pairs of 10 different males 
were tested against each other three to five times. In Design 2, a wire 
cage 47 cm long X 51 cm deep X 39 cm high was divided in half 



4 BREVIORA No. 479 

longitudinally by a removable opaque partition. A single large male 
was housed in each half, which also contained a plant, a perch, and 
a bark wall. In these cages, the animals were in physical as well as 
visual contact with each other when the partition was removed. In 
this design, three pairs of six different males were effectively tested 
against each other only once since subsequent tests would have been 
declared "no contest" at the outset by the participants. These wire 
cages were kept in Sherer-Gillet environmental chambers with 
transparent doors. All animals experienced a constant environ- 
mental regime of 14L.10D photoperiod and a corresponding 
temperature cycle of 32:23° C. All animals received food (meal- 
worms and crickets) and water ad lib. Observation of the aquaria 
animals in Design 1 was made from behind a cardboard blind to 
ensure that the displays were directed at conspecifics and not the 
observer. The animals in the lighted environmental chambers of 
Design 2 were observed in a darkened room, making the observer 
relatively inconspicuous. The cages were arranged in each case so 
that no animal could see a conspecific except those in its 
experimental setup. When the partition was in place males were 
only occasionally observed displaying at the juveniles or females. 
Males were housed individually before they were placed in the 
experimental cages. 

All males were allowed at least one week to acclimate to a new 
cage before testing was begun. Agonistic tests were at least 15 
minutes long and continued 5 minutes after the situation appeared 
stable (that is, when there was no change in positions and display 
types used by the two animals). Courtship tests were terminated 
when the animals finished copulating. 

Male agonistic behavior was studied in both types of cage designs 
with the opaque dividers removed. Males in the aquaria of Design 1 
were repeatedly tested against each other whereas those in the wire 
cages of Design 2, which allowed physical contact, only met once 
because, once beaten, the subordinate animal did not display but 
retreated. Courtship behavior was observed only in the cages of 
Design 2; the partitions were left intact and a female was introduced 
into a wire cage which housed a single male. 

Parameters of male behavior were recorded with a 10 channel 
Esterline Angus operation recorder Model A. Displays were taped 
with a Sony video camera AVC 3210 with a 75 mm f 1 2.5 lens and 
analyzed with a Sony videorecorder AV 3650 with which the tapes 



1984 DISPLAYS OF A NOUS SA GREI 5 

could be slowed to 8% actual speed. The head and dewlap 
movements were transcribed onto a strip of acetate by running the 
acetate in front of the video screen at a constant known speed, using 
the gears of the event recorder and following the action with a felt 
tipped pen placed on the rostrum of the videotaped lizard. The time 
framework could be superimposed on the tracing making correc- 
tions for the video recorder speed and the gear ratio of the event 
recorder. Only animals from the wire cages of Design 2 were 
videotaped because the T.V. equipment was less conspicuous to the 
subjects. 

A total of 620 agonistic displays by 12 different males were 
observed in 18.8 hours. Two hundred twenty-nine displays of eight 
different courting males were observed in 12.4 hours. Thirty-one 
displays of three males in two entire agonistic events and three 
different males in entire courtship tests were taped and analyzed. 
Recording data on the video equipment and event recorder 
provided different benefits. When using the event recorder, the 
social context of the display and the response of the recipient could 
easily be included. In this way the meaning of the display could be 
defined by the response it provoked. With the video equipment it 
was possible to record data on the modifiers of the displays in fine 
enough detail to compare and discuss variability within and between 
individuals. The cadence of bobs could also be recorded to 
discriminate the type of bobbing display and to relate it to the social 
context. 

For purposes of discriminating dominant and submissive be- 
havior, the animals of all the tests in both cage designs were divided 
into two groups on the basis of their behavior at the end of the test. 
An animal was judged to have been acting dominant throughout the 
test if he was still facing his opponent at the end with an alert 
posture. Those anoles that ran from their opponent to the far side of 
the cage and had a posture low to the substrate were classed as 
submissive for the whole test. Since these classifications were 
somewhat subjective, borderline cases in which either animal 
showed a mixture of dominant and submissive behavior at the end 
of the test were not used in the following analysis. Within each 
group there was little variance in the frequency of each type of 
display performed (Table 1); therefore, for statistical purposes, data 
from all dominant males were pooled, as were those from all 
subordinate males. 



6 BREVIORA No. 479 

Table 1. Frequency ± variance of displays by type per minute of animals judged 
at the end of the test to be dominant or subordinate. 

Dominant animals Subordinant animals 

(N = 34) (N = 11) Difference 



Dewlap only 0.07 ± 0.01 0.07 ± 0.01 n.s. 

Bob only 0.43 ± 0.06 0.21 ± 0.04 t = 2.70, p < 0.01 

Bob + dewlap 0.16 ±0.02 0.01 ± 0.00 t = 3.49, p < 0.001 



Any encounter escalated (if it were going to) in a predictable 
fashion. Modifiers were added to displays and other behaviors 
appeared in the same order each time, as the states of arousal of the 
participants increased and their bodies became more tense. To 
facilitate analysis of behaviors and responses in agonistic en- 
counters, confrontations were divided into three stages which were 
differentiated by changes of body posture. In the first stage, a male 
had not yet erected either a nucal or dorsal crest or laterally 
compressed his body and was said to be in state A. In the second 
stage, a male had both crests (state B) and in the third, most aroused 
stage, he had laterally compressed his body as well (state C). 

Statistical tests used were t-test of proportions, arcsin trans- 
formed, to test the difference between the means of two samples and 
a one-way analysis of variance to examine the inter- and intra- 
individual variance in display stereotypy. Means are expressed ± the 
standard error. 

RESULTS AND DISCUSSION 

Displays and Modifiers 

A display is a sequence of behaviors, usually performed in its 
entirety, which communicates information about the sender. In the 
following context, it refers to the change in amplitude over time of a 
lizard's head and/ or dewlap and encompasses any consistantly 
associated stereotyped movements or postures. A modifier is a 
posture (static modifier) or movement (dynamic modifier) which is 
not always present with a particular stereotyped display but may be 
added (Jenssen, 1978). Anolis sagrei sometimes performed these 
modifying behaviors, particularly postures, separately from stereo- 
typed head bob displays. 

A brief description of the displays investigated follows. The 
frequency distributions of the display types, degree of stereotypy, 



1984 DISPLAYS OF A NOUS SA GREI 7 

and information conveyed are all discussed later in the context in 
which the display was used. 

Dewlap Fanning. The dewlap, which in Florida A. sagrei is 
bright orange with a yellow margin, is extended by the hyoid 
apparatus. The pattern and context of this action is extremely 
variable. It can occur as a single extension or in a fast or slow series 
of as many as 15 separate extensions. The series have no discernible 
pattern, and the display is used in all observed contexts and may be 
directed at a human observer as well as a male or female conspecific. 

Bobbing. The head can be bobbed from the neck, from mid- 
body, or as a four-legged push-up. Bobs from the neck or mid-body 
can occur singly or in a series of a single posture, or the display may 
begin as bobs from the neck and finish as bobs from the mid-body. 
These postures may demonstrate increasing states of arousal in that 
order. Bobs with a four-legged push-up posture are not performed 
singly. These four-legged push-up bobbing displays are typically 
8-10 bobs but may contain 18 to 20 and last as long as 30 seconds. 

There are three different sorts of bobbing displays. 

(A) Anolis sagrei can perform a species-specific stereotyped 
display which has a distinctive cadence to a series of bobs which 
may be called its "signature" display (Stamps and Barlow, 1973). It 
consists of four units: 2 quick initial bobs of increasing amplitude, a 
long bob, a short bob, followed by a series of 3 to 12 even-tempo 
bobs dampening in amplitude (Fig. 1). There may be variation in 
bob number of any unit of the pattern. If the dewlap is extended 
during the display, it always flashes on the third (long) bob and 
variably during the dampening series of bobs at the end. This 
signature display occurs in an assertion context (i.e., non-directed or 
low conflict situation), in both agonistic and courtship tests, and at 
every state of arousal. 

However, sometimes this bobbing display does not consist of this 
characteristic pattern. It may, however, have some similar elements 
to the signature pattern such as a long bob accompanied by a 
dewlap extension followed by a short bob. These displays also may 
or may not be accompanied by dewlap extensions and without them 
as a marker, the elements similar to the signature pattern often 
could not be identified. For example, in the display illustrated by 
Figure 2, the dewlap extension at 5 seconds marks what might be 
the long bob followed by the short bob characteristic of the 
signature pattern but it is unrecognizable by the pattern of bobs 
alone. These displays also may occur in any context. 



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10 



15 



lime (sec) 
Figure 3. The quick rhythmic bobbing display. 

(B) The second bobbing display is characterized by a quick 
(about 1-2/ sec) rhythmic bobbing with no inter-bob pause and 
without any dewlap extension (Fig. 3). The amplitude of these bobs 
was less than that of the bobs of the signature display, and usually it 
was performed with the head and body close to the substrate. In 
these tests, 49 of 53 times it was performed by an animal in a 
submissive situation. However, it was occasionally (4 times) 



1984 DISPLAYS OF ANOLIS SAGREI 9 

performed by a highly aggressive dominant male in a context when 
the opponent had a submissive posture. 

(C) The third type of bobbing display is jiggling action of the 
head. The amplitude of these bobs was very much less than that of 
other bobbing displays and the rate was much faster (8- 10/ sec). In 
these tests, all 34 times this display was used was in courtship 
situations and was performed by the male usually as he approached 
the female to take a neck grip prior to mating. It may be performed 
as a separate display or tacked on to the end of another. 

Modifiers. Many different behaviors may be used to modify a 
display. Some were consistently used in a particular context and 
state of arousal and often caused a consistent response in the 
recipient and therefore these modifiers may impart specific informa- 
tion on the intent and motivation level of the animal displaying. See 
Table 2 for a list of modifiers frequently used, their social context, 
and state of arousal when used. 

Male- Male Encounters 

The encounters between two males were staged such that they 
would both be expected to act territorially. A total of 12 different 
males were used in 26 tests. In Design 1, the two aquaria which each 
housed resident males were sufficiently large that the males were 
often 150 cm apart when they first saw each other, so that in 45% of 
the 25 tests they escalated aggression (see below) and were only 
prevented from combat by the glass partitions. When these same 
two males confronted each other in future tests, both continued to 
act aggressively. The situation in the wire cages of Design 2 was 
different. The animals were less than 50 cm apart when they first 
saw each other, and there was no physical partition. In every 
instance the rank order of the two was quickly apparent when one 
male fled without a contest; there was no escalation of aggression. 
The physical proximity seemed to be a deterrent to bluffing by the 
less aggressive male and did not allow for a slow escalation of 
aggressive behavior and evaluation of each other. 

The sequence of events in an escalating fight is typical of that 
described for other anoles (Crews, 1975b). Behaviors and modifiers 
to displays are added in a sequence that is predictable. In a aquaria 
of Design 1, males would display towards each other with first a 
nuchal crest and then a dorsal crest being erected. The whole body 



10 



BRF.VIORA 



No. 479 



Table 2. Modifiers used in displays of male A. sagrei and their context. The steps 
in escalation of an encounter from first display to conclusion (combat or neck grip) 
were consistently ordered (see text). The state of arousal refers to the stage of 
escalation. 



Modifier 



Display 



Context 



State of Arousal 



Dewlap extension Signature display 



Nuchal crest 
Dorsal crest 
Lateral compression 
Engorged throat 

Tail lift 
Tail lash 

Tongue protrusion 
Lip smack 
Slow approach 



Signature display or 
continued condition 

Signature display or 
continued condition 

Signature display or 
continued condition 

Usually a continued 
condition but can 
accompany signature 
display instead of 
dewlap extension 

In 4-legged push-ups 
of signature display 

Alone or with 
signature display 

Alone or with 
signature display 

Alone or with 
signature display 

Jiggle display 



Assertion, 
courtship & 
aggression 

Aggression 
Aggression 
Aggression 
Aggression 



Courtship & 
aggression 

Aggression 
Aggression 
Aggression 
Courtship 



All 

Moderate 

Mod. high 

Highest 

Moderate — highest 

Mod. high — highest 
Mod. high — highest 
Mod. high — highest 
Moderate — highest 
Highest 



can then be laterally compressed. (Table 3 illustrates that these 
successive posture changes indicate increasing states of arousal.) As 
the lizard's body becomes more tense, the bobbing action becomes 
more exaggerated, from nods to two-legged push-ups to four-legged 
push-ups. At this point increasing intensity of arousal is signaled by 
behaviors other than the bobbing displays. The males approach 
each other either slowly or with a rush. They will orient head to tail, 
lunge, and threaten by gaping their mouths. Because the animals 
were either physically separated or unwilling to engage in combat, 
no staged fight in either Design 1 or 2 proceeded past this point, but 
in a natural setting the fight will end with jaw grappling and one 
being physically pushed off the branch and chased off (Scott, 
personal observation). 



Dominant 




animals 


34 


Submissive 




animals 


11 



1984 DISPLAYS OF A NOL/S SAGREl II 

Table 3. The probability of an animal in an agonistic confrontation taking a 
more aggressive posture. A, B, and C are the states of arousal, and A — B is the 
probability of a male in the first state going to the second. An animal in state A has a 
body posture without nucal or dorsal crests or lateral compression of the body. An 
animal in state B has both crests and one in state C has crests and lateral 
compression. 

# tests A-B A - C B-C C-B C - A B - A 



0.91 0.09 0.74 

0.45 0.27 0.09 0.18 



Twenty-four displays by males clearly acting territorially were 
taped and analyzed; all had some recognizable components of the 
signature display. There was however a great deal of variability. 
Five of seven displays by one male, and four of five by another, were 
of the signature pattern. The remaining displays for these two 
animals were varied by having either one or three quick initial bobs 
instead of two. The third male whose displays were taped had only 
three of 12 showing the signature pattern. The number of initial 
bobs (2, 3, or 4), slow bobs (1 or 2), and quick bobs (0, 1, or 2) 
varied so that for some displays the signature pattern was barely 
recognizable. The mean number of bobs in these filmed displays was 
11.1 ± 0.4. The mean time elapsed was 17.4 ± 0.8 seconds. The 
coefficients of variation for the number of bobs in these variable 
units of the signature display in these agonistic encounters are: 24% 
for the initial bobs, 32% for the slow bob, 40% for the quick bob, 
and 23% for the total number of bobs in the display. Twenty of these 
24 taped displays were accompanied by dewlapping, whereas only 
23% of the bobbing displays included dewlap extensions in the total 
sample of all displays observed. Twenty-two of the taped bobbing 
displays were two-legged push-ups with crests erected; in 12 displays 
the body was laterally compressed as well, as the state of arousal 
during the confrontation increased. Each of the three animals gave 
one or two displays that began as four-legged push-ups, usually 
raising the tail on the up stroke of the bob (Fig. 4), and dampened to 
two-legged push-ups. Tail lifts were used as a modifier of two-legged 
push-ups in conjunction with nuchal and dorsal crests three times 
and once with lateral compression as well. One display which 
included crests and lateral compression also was accompanied by a 
tongue protrusion and a lip smack. 



12 



BREVIORA 



No. 479 




Figure 4. Aggressive display with a 4-legged push-up, nucal and dorsal crests, 
lateral compression of the body, and a tail lift. 



Information on the context of each display type was gained by 
pooling the data from all the staged encounters in both cage designs 
and examining the frequency distributions of display types (dewlap 
flashes, bobbing displays, and bobbing displays with dewlap 
extensions). Four bobs and dewlap flashes in a continuous sequence 
were arbitrarily chosen as the minimum to constitute a display. The 
mean proportion of dewlap fanning displays of all displays was 0.14 
±0.02. There was little intra-individual variation in the proportion 
of use of this display form from one test to another. In a one-way 
analysis of variance, 87% of the variation in the frequency of use of 
this display type was between individuals and 13% between tests of a 
single individual (F = 7.28, p < 0.01). This tendency was enhanced 
by the fact that each male in the cages of Design 1 was always 
matched against the same opponent and most animals responded in 
a characteristic fashion in each test. The frequency with which a 
bobbing display was accompanied by dewlap extension was 
similarly consistent. Eighty-three percent of the variation was 
between animals (F = 4.77, p < 0.01). The mean proportion of 
bobbing displays with dewlap modifier was 0.23 ± 0.03 (Fig. 5a). 

A male was judged to be either submissive or dominant on a test- 
by-test basis, but he probably performed as both a dominant and 
submissive individual during the test. Most animals that "lost" 
ultimately still responded with varying intensity to the challenge of 
the more aggressive one. In addition, the relative ranks of 
submission and dominance were not necessarily consistent even 



1984 



DISPLAYS OF ANOLIS SAGREI 



13 



dewlap bob + bob 
only dewlap only 



a. 


all males 


CO 


70 
60 


' N = 620 






o 

*■> 

o 


50 
40 
30 


• 








20 
10 


9 !«fi3l' 




b. dominant 




70 


males 


(Q 


60 




*•> 
O 

o 


50 
40 
30 




# 


20 
10 



N=486 




c. 



submissive 




70 


males 


"n 


60 




o 
o 


50 
40 
30 




cr 


20 
10 



N = 


= 6' 


\ 





Figure 5. Frequency distribution of display types performed by males in agonistic 
confrontations. All displays were used in a., but only those by clearly dominant or 
clearly submissive males were used in b. and c. 



14 BREVIORA No. 479 

when the same opponents were repeatedly matched in the separated 
aquaria. 

Submissive animals displayed (either bobbing or fanning) less 
than half as often (0.29/minute, N=ll) as dominant ones (0.63/ 
minute, N=34) (Table 1). Dewlap fanning formed a significantly 
higher proportion of the total displays by submissive animals than 
by dominant animals (x = 0.34 ± 0.08 vs. x = 0.08 ± 0.01; t = 2.29, 
p < 0.05) (Fig. 5b, c). A significantly lower proportion of the bob- 
bing displays of submissive animals was accompanied by dewlap 
extensions, x = 0.03 ± 0.01, than of dominant animals, x = 0.30 ± 
0.03 (t = 3.36, p < 0.01). Therefore, submissive animals use their 
dewlaps for fanning displays but not to modify bobbing displays. A 
significantly higher proportion of the bobbing displays of sub- 
missive animals were the rapid bob (Fig. 3), x = 0.25 ± 0.09 for 
submissive animals versus x = 0.01 ± 0.01 for dominant animals (t = 
2.99, p < 0.01). Dominant animals also took progressively more 
aggressive postures and reached a higher arousal state more 
frequently than submissive ones did (Table 3). 

The pacing of agonistic encounters varies with each fight, even 
when the combatants are known to each other. Of the pairs that 
always escalated to a full confrontation, usually the same individual 
was faster to be aroused. The mean latency time of aggressive 
animals for developing nuchal and dorsal crests was 4.6 ± 0.4 
minutes (N=38) and for lateral compression was 7.4 ± 0.5 minutes 
(N=26). There was a great deal of intra-individual variation in the 
kind of displays, if any, that were performed before both crests were 
erected and the body compressed. But the frequency distribution of 
display types performed in this low arousal state was different for 
animals that would be judged dominant or submissive at the end of 
the test, i.e., there were behavioral differences in the two groups 
from the beginning of the test (Fig. 6). 

A display by one male A. sagrei often prompted a display by the 
other in these tests. Forty percent of all displays seen occurred 
within 10 seconds after a display or single bob or dewlap flash by the 
opponent, but there was no particular tendency to use the same sort 
in answer as the one just seen. For example, a bobbing display was 
followed by dewlap fanning display or a bobbing display. Forty to 
50% of the fanning displays and the bobbing displays with and 
without dewlap modifier of both dominant and submissive animals 
was in response to the opponents display. There was a slightly 



1984 



DISPLAYS OF A NOUS SAGREI 









dewlap 
only 


bob + 
dewlap 


bob 
only 


a. dominant 
males 


ro 

»■« 
O 

•- 

o 


70 
60 
50 
40 
30 
20 
10 


N = 74 









b. submissive 




70 


males 


fO 


60 




o 
o 


50 
40 
30 






20 
10 



N = 32 



\1 



Figure 6. Frequency distribution of display types performed by dominant and 
submissive animals in agonistic confrontation before nucal or dorsal crests had been 
erected or the body had been laterally compressed. 

higher tendency for a dominant animal to respond with a bobbing 
display with a dewlap extension. Submissive animals performed 
almost no bobbing and dewlap displays, but those few that they did 
perform were all in response to the opponent's display (Table 4). 

Male- Female Encounters 

All male-female encounters were staged by introducing a female 
into the wire cage of Design 2 of a single resident male. Eight 
different males were used in 33 tests. The pattern of courtship 
behavior appears to be very similar to that described for other 
anoles (Noble and Bradly, 1933; Evans, 1938; Crews, 1977). Upon 



16 BREVIORA No. 479 

Table 4. Proportion and number of each type of display for submissive and 
dominant animals which were in response to a display by the opponent rather than 
spontaneously performed. 





Total 
displays 


Dewlap 
fanning 


Bobbing 
displays 


Bobbing 
displays 
+ dewlap 


Dominant 
animals 

Submissive 
animals 


40% (192) 
44% (28) 


35% (18) 
42% (8) 


37% (114) 
40% (17) 


49% (60) 
100% (3) 



introduction of a female, the territorial male performs the dewlap 
fanning display or the bobbing display with or without dewlap 
extensions either with the signature pattern or not. The bobbing 
displays seen in a courtship situation have none of the aggressive 
modifiers, such as crests and lateral compression, that characterize 
male-male encounters. The male then approaches the female, 
sometimes with a jiggling action with his head, takes a grip on her 
neck, and swings his tail beneath hers, juxtaposing his cloaca to hers 
and inserting a single hemipenis. More displays may follow mating. 
It has been reported for other anoles (Crews, 1977) that a receptive 
female will stand and arch her neck for the male to take a grip. In 
this study, no female was ever seen to show receptive behavior. In all 
but two of the 33 tests the male attempted to mate by taking or 
attempting to take a neck grip on the female. Eleven tests ended in 
copulation. In four of these, the matings were very short (<2 
minutes), and the male maintained his mating posture several 
minutes after the female escaped, indicating that intromission may 
not have occurred. Excluding those matings which may have been 
incomplete, the mean time for intromission was 6.7 ± 0.83 minutes. 
The anoline courtship display has been characterized as ending 
with a series of jiggly bobs (Crews, 1975b). In 18 of 33 courtships 
observed, A. sagrei males also performed a jiggle bob at the end of a 
bobbing display or alone as he approached the female. All eight 
males performed it in at least one test but only one male always 
courted using it. When this display was used, it usually immediately 
preceded an attempted neck grip (15 of 18 tests). This jiggle bob may 
be an important part of courtship and may be performed with 
greater regularity if the females showed signs of receptivity. 



1984 DISPLAYS OF ANOLIS SAGREI 17 

Of the seven video taped bobbing displays in courtship sequences 
of three males, only one was the signature pattern. The pattern of 
the rest could not be characterized. All four displays in the two 
courtships that ended in neck grips were four-legged push-ups with 
the tail lifted as the head bobbed up and the dewlap pulsed. This tail 
lift was a very common (28%) modifier in all courtship bobbing 
displays. 

The overall frequency of all types of displays in courtship 
sequences by 8 different males in 33 tests was 0.35/ minute which is 
about half that of a dominant male in an agonistic situation 
(0.66/ minute). As was the case of the agonistic tests, the data can be 
combined because variance of display frequency between per- 
formances in different tests was relatively low (0.05). The frequency 
of bobbing displays with dewlap flashes were similar in premating 
and postmating displays (0.32 vs. 0.21) which was similar to that in 
agonistic encounters (0.23). There was both inter- (57%) and intra- 
(43%) individual variation in the use of the dewlap (F = 1.3, p > 
0.20). The proportion of displays which were dewlap fanning 
displays were similar in courtship (0.15) and agonistic situations 
(0.14), but a higher proportion of the displays after mating (0.50) 
than before (0.10) were dewlap fanning (Fig. 7). Six of the seven 
complete matings, and none of the short matings, were followed by 
a dewlap fanning display, most of which had a distinctive erect 
posture (Fig. 8). But bobbing displays and further attempts to 
regain a neck grip followed one of the incomplete matings. Thus, 
bobbing displays may be predominantly a premating signal, and in 
this context fanning displays may be a signal of a successful 
copulation. 

CONCLUSIONS 

The male Anolis sagrei of this study show variability in both 
pattern and contextual use of displays. Anolis sagrei has only one 
species specific stereotypic pattern which may be used in every 
context. This signature display when used by males in the agonistic 
encounters was quite variable both in the total number of bobs and 
the number of bobs in each unit. In addition, the accompaniment of 
the dewlap extension was also variable; although, if it was used, it 
was always associated with one particular unit of the display, the 



BREVIORA 



No. 479 



dewlap bob + bob 

only dewlap only 



a. total 
displays 


(0 


70 
60 


N= 229 




o 

<*- 

o 


50 
40 
30 


• 






20 
10 


flM 




b. premating 
displays 



cr 



70 
60 


N= 201 




50 


- 




40 


• 




30 


- 




20 






10 


I^H ^^B 





c. postmating 


70 


displays "^ 


60 


o 
o 


50 
40 
30 


cr 


20 
10 



N= 28 




Figure 7. Frequency distribution of display types performed by males in courtship. 



1984 DISPLAYS OF A NOUS SAG RE/ 19 




Figure 8. Typical post-copulatory display of dewlap fanning and erect posture. 

long bob. This display is so variable that sometimes the signature 
pattern cannot be recognized. Video taped agonistic and courtship 
encounters showed that the species-specific signature pattern was 
used in almost all displays directed toward other territorial males, 
but was hardly or not at all recognizable in courtship displays. 

The only displays which are consistent in their form and use are 
not species-specific. The jiggle bob, although not always performed 
in courtships of unreceptive females, was never seen in any other 
context. The display of rapid rhythmic bobs, shown by submissive 
males, was not performed by courting males and only rarely by 
territorial males, and then modified by a very aggressive posture. 
The dewlap fanning display was seen in all contexts but was used 
consistently after copulation with a distinct erect posture and may 
be associated with successful copulation. It has not been described 
as such for other species, but probably occurs in A. carolinensis 
(Crews, personal communication) and may be used by other anoles. 

The color of the dewlap is also a species specific signal. But it may 
not be a symbol to aid species recognition as much as a means of 
emphasizing the bobbing display with the bright flash of color 
(Greenberg and Nobel, 1944; Williams and Rand, 1977). Dominant 
males in agonistic situations use the dewlap in bobbing displays 
significantly more than submissive males, but the dewlap is used for 
fanning displays at a significantly higher frequency by submissive 
males. Therefore, dewlap flashes alone may be less aggressive 
signals than when they accompany bobbing displays. 

If the dewlap per se were being used as a species-specific signal, 
one might hypothesize an increase in its use in courtship. However, 



20 BREVIORA No. 479 

fanning displays comprise a small proportion of premating displays, 
which is lower than its average use in other contexts, and the dewlap 
is used in bobbing displays at only slightly higher frequencies during 
courtship prior to mating than at other times. Therefore the two 
signals, the dewlap and the species-specific signature pattern, which 
one might expect to play a role in species recognition and female 
choice, are not particularly associated with courship, at least not 
when the animals are at relatively close range as they were in these 
tests. 

Anolis sagrei originates from Cuba, an island with a very complex 
Anolis fauna (22-28 species, Williams, 1969). The stereotyped 
display pattern is more variable than one might expect with so many 
congeneric species needing reproductive isolation (Ruibal, 1967). 
Information to enable species recognition may be redundantly 
coded by many physical and behavioral characteristics (Williams 
and Rand, 1977). This seems to be the case with A. opalinus on 
Jamaica, which has no discrete display pattern even though it is 
sympatric with its two closest relatives, A. garmani and A. grahami. 
All three species have similar dewlap color and display structure but 
differ greatly in body size, shape, and color (Jenssen, 1979). 

The signature display and increased use of the dewlap with the 
display are associated with territorial status in agonistic encounters. 
But neither shows an increase in use as the encounter escalates. The 
state of arousal and the willingness to escalate may be communi- 
cated by the modifiers of the displays, especially body posture and 
shape, and probably by the timing of the events. The more 
aggressive male of a pair was usually the first to erect his crests, 
engorge his throat, and laterally compress his body. Small shifts in 
posture or position in tense moments are probably also used as 
aggressive signals. Characterization of the behavioral repertoire of 
A. sagrei has been elusive (Ruibal, 1967) because of this subtlety. 

ACKNOWLEDGMENTS 

I would like to thank E. E. Williams, D. Crews, W. Gartska, T. A. 
Jenssen, and A. S. Rand for their critical comments on earlier drafts 
of this paper; J. F. A. Traniello for his technical help and 
encouragement; B. Scott for the art work; and J. S. Godley for 
organizing the lizard hunt. This study was supported in part by 
NIMH Research Scientists Development Award 00135 to D. Crews. 



1 984 DISPLAYS OF A NOUS SA GREI 2 1 

LITERATURE CITED 

Carpenter, C. C. 1967. Aggression and social structure in iguanid lizards, pp. 

87-105. In W. Milstead (ed.) lizard Ecology: a Symposium, Columbia Univ. 

Missouri Press. 
Carpenter, C. C, and G. Gribitz. 1961. Time-motion study of a lizard. 

Ecology 42: 199-200. 
Crews, D. P. 1975a. Effects of different components of male courtship behavior 

on environmentally induced ovarian recrudescence and mating preferences in 

the lizard, Anolis carolinensis. Anim. Behav., 23: 349-356. 
1975b. Inter- and intraindividual variation in display patterns in the 

lizard, Anolis carolinensis. Herpetologica. 31 : 37-47. 
1977. The annotated anole: studies on the control of lizard reproduction. 



Am. Sci., 65: 428-436. 
Evans, L. T. 1936. A study of a social hierarchy in the lizard, Anolis carolinensis. 

J. Genet. Psychol., 48: 88-111. 
1938. Courtship behavior and sexual selection of Anolis. J. Comp. 

Psychol., 26: 475-497. 
Greenberg, B., and G. K. Noble. 1944. Social behavior of the American 

chameleon (Anolis carolinensis Voight). Physiol. Zool., 17: 392-439. 
Greenberg, N. 1977. A neurethological study of the display behavior in the 

lizard, Anolis carolinensis (Reptiiia, Lacertilia, Iguanidae). Am. Zool., 17: 

191-201. 
Hover, E., and T. Jenssen. 1976. Descriptive analysis and social correlates of 

agonistic displays of Anolis limifrons (Sauria: Iguanidae). Behav., 58: 173-191. 
Jenssen, T. A. 1970a. Female response to filmed displays of Anolis nebulosus 

(Sauria: Iguanidae). Anim. Behav., 18: 640-647. 
1970b. The ethoecology of Anolis nebulosus (Sauria: Iguanidae). J. 

Herp., 4: 1 38. 
1971. Display analysis of Anolis nebulosus (Sauria: Iguanidae). Copeia, 

1971: 197-209. 
1977. Evolution of anoline lizard display behavior. Amer. Zool., 17: 

203-215 
1978. Display diversity in anoline lizards and problems of interpretation, 

pp. 269-285. In N. Greenberg and P. D. MacLean (eds.) Behavior and 

Neurology of Lizards, NIMH. 
1979. Display behavior of male Anolis opalinus (Sauria, Iguanidae): a 



case of weak display stereotypy. Anim. Behav., 27: 173-184. 
Jenssen, T. A., and E. Hover. 1976. Display analysis of the signature display of 

Anolis limifrons (Sauria: Iguanidae). Behav., 57: 227-240. 
Jenssen, T. A., and L. M. Rothblum. 1977. Display repertoire analysis of Anolis 

townsendi (Sauria: Iguanidae) from Cocos Island. Copeia, 1977: 103-109. 
Noble, G. K., and H. T. Bradley. 1933. The mating behavior of lizards: its 

bearing on the theory of sexual selection. Ann. N. Y. Acad. Sci., 35: 25-100. 
Rand, A. S., and E. E. Williams. 1970. An estimate of redundancy and 

information content of anole dewlaps. Am. Natur., 104: 99 103. 



22 BREVIORA No. 479 

Ruibal, R. 1967. Evolution and behavior in West Indian anoles, pp. 1 16-140. In 

W. W. Milstead (ed.). Lizard Ecology: a Symposium, Columbia Univ. Missouri 

Press. 
Stamps, J., and G. Barlow. 1973. Variation and stereotypy in the displays of 

Anolis aeneus (Sauria: Iguanidae). Behav., 48: 67-94. 
Williams, E. E. 1969. The ecology of colonization as seen in the zoogeography of 

anoline lizards on small islands. Quart. Rev. Biol., 44: 345-389. 
1976. West Indian Anoles: a taxonomic and evolutionary summary. I. 

Introduction and species list. Breviora Mus. Comp. Zool. No. 440, pp. 1-21. 
Williams, E. E., and A. S. Rand. 1977. Species recognition, dewlap function 

and faunal size. Am. Zool., 17: 265-274. 



MUS. COMP. ZOOt 
LIBRARY 



JUN 1 7 1985 

B R ffSV I R A 

Museum of Comparative Zoology 



US ISSN 0006-9698 



Cambridge, Mass. 21 Jine 1985 Number 480 

THREE NEW LIZARDS OF THE 

GENUS EMOIA (SCINCIDAE) 

FROM SOUTHERN NEW GUINEA 

Walter C. Brown 1 and Fred Parker 2 

Abstract. Three new species of skinks in the genus Emoia arc described from 
provinces south of the central mountain range. New Guinea. 

INTRODUCTION 

The genus Emoia is widespread through much of the Indo- 
Australian Archipelago and the islands of the Pacific, and ranges 
into the Philippines in the north and northeastern Australia and 
New Hebrides in the south. The greatest diversity of species occurs 
in New Guinea and surrounding smaller islands. The three new 
species described in this paper are from south of the central moun- 
tain range in New Guinea. 

One species is based on a series of specimens collected primarily 
by the junior author in the Fly River and Bamu River drainages. 
Western Province, Papua New Guinea. The second ranges from 
Western Province westward through southern Irian Jaya. Both spe- 
cies are in the E. bauciini-submetallica group as defined by Brown, 
1953. The third species which belongs to the E. physicae group 
(Brown, 1953) has a wide range in several provinces of southern 
Papua New Guinea. The last two, in addition to the series collected 
by the junior author, are also represented in earlier collections. 



'Department of Herpctology. California Academy of Sciences, San Francisco. 
California 941 18. and, Menlo College. Menlo Park. California 94025. 
-7 1 7 Ross River Road. Kirwan. Queensland. Australia. 



2 BREVIORA No. 480 

These had been misidentified as juveniles of E. p. physicae, E. p. 
tropidoiepis, or E. baudini. 

Emoia aurulenta new species 
(Fig. 1) 

Holotype: MCZ 142327, adult male, collected at Migalsimbip in 
the upper Fly River drainage at an elevation of about 1,200 m. 
Western Province, Papua New Guinea, Fred Parker Coll., 16 August 
1972. 

Paratypes: The following localities in Western Province, Papua 
New Guinea, Migalsimbip: SAM 11637; AM 40778; MCZ 
142322-26, 142328-30, 152263-71, 152273 75, CAS 154186, 
AMNH 111718; UPNG 6477-78, 6480, 6483; Derongo: MCZ 
131895-900; AMNH 103961; Menemsore: MCZ 131901; Emeti: 
MCZ 152265-67, 152291, AMNH 111715; Tingkem: MCZ 131894. 

Diagnosis. This species differs from the other New Guinean spe- 
cies of the Emoia baudini section in the following combination of 
characters: (1) small size, 37+ to 49.5 mm snout-vent length at 
maturity; (2) pattern of gold-orange markings on the sides of the 
neck and the lateral surfaces of the body in life (these fade to dirty, 
silvery-white in preservative); (3) low number of midbody scale 
rows; 26 to 32 (mean 28+), rarely greater than 30; (4) number of 
subdigital lamellae, 39 to 48 beneath the fourth toe; and (5) number 
of paravertebral scale rows between the parietals and base of tail 45 
to 52. 

Description. A small Emoia, snout-vent length of 41.3 to 49.5 mm 
for 16 males and 37.4 to 47.5 mm for eight females; habitus moder- 
ately slender; snout moderately tapered, bluntly rounded, its length 
35 to 40% of head length and 52 to 60% of head breadth; head 
breadth 64 to 69% of head length and 15 to 18% of snout-vent 
length; eye moderately large, its diameter 7 1 to 85% of snout length 
and 40 to 45% of head breadth; ear diameter about one third to one 
half of eye diameter; rostral broader than high, forming long, nearly 
straight suture with frontonasal; supranasals widely separated, nar- 
rowly triangular, in contact with anterior loreal; prefrontals moder- 
ately to widely separated; frontal longer than broad, rounded 
posteriorly, shorter than fused fronto-interparietal shield, in contact 
with first and second supraoculars; four large supraoculars; 
parietals large, in contact posteriorly; one pair of nuchals; anterior 



1985 



NEW SPECIES OF EMOIA 




Figure I. An adult E. aurulenla. 



loreal higher and shorter than posterior, in contact with second, or 
first and second supralabials; six to seven upper labials, fifth (very 
rarely sixth) enlarged and beneath eye; six to seven lower labials; 
dorsal scales smooth, vertebral rows not distinctly enlarged; 26 to 32 
(usually 28 to 30) midbody scale rows; 45 to 52 (mean 48.3) paraver- 
tebral rows between parietals and base of tail, ventrals about same 
size as dorsals; preanals somewhat enlarged; limbs well developed, 
length of extended hind limb 90 to 1 10% of axilla-groin distance and 
47 to 54% of snout-vent length; 39 to 48 (mean 42.6) smooth, 
rounded lamellae beneath fourth toe; 10 to 14 lamellae under first 
toe; rank of adpressed toes from longest to shortest four, three, two 
to five, one; tail slender and much longer than body. 

Measurements (in mm) of Holotype. Snout-vent length 49.0; 
axilla-groin distance 24.8; hind limb length 24.1; head length 1 1.9; 
head breadth 8.1; snout length 4.35; eye diameter 3.4; ear diameter 
1.5; tail length 76.4. 

Color in Preservative. The basic ground color on the dorsum 
ranges from light olive greenish-brown to brown marked by lighter 
and darker blotches, usually involving all dorsal scale rows (or occa- 
sionally not including the paravertebral rows). The light and dark 



4 BREVIORA No. 480 

blotches may occupy alternate scales or involve two or more scales. 
The upper lateral surfaces are dark brown, bordered dorsally by a 
row of scattered, single or clustered, pale scales. There are also 
scattered pale scales between and on the limbs. The neck is marked 
by pale and dark blotches, and the posterior labials by dark bars. 
The venter is dirty white to grayish, lightest on the chin and in the 
limb regions. 

In life, the top of the head is bronze with a lighter line from above 
the eye to the nuchals or for a few specimens extending further 
posterior onto the neck. The dorsum is gray or brown with alternate 
black scales or paravertebral rows of black spots in a checkerboard 
pattern. The upper flanks are black or mottled black and brown. 
The lower flanks and usually the side of the neck are marked by 
golden yellow spots. The venter is yellowish to cream. 

Comparison. Emoia aurulenta is related to Papuan species of the 
E. baudini-submetallica group, but is readily distinguished in life by 
such features of the color pattern as the checker-board pattern on 
the dorsum and the prominent golden yellow spots on the sides. It 
also is characterized by a lower number of midbody scale rows than 
other known species of this section. Also, the number of subdigital 
lamellae is greater than for most other species of this group, only 
overlapping slightly with E. s. popei and two other undescribed 
species (Table 1). 

Etymology. The name aurulenta refers to the golden yellow spots 
on the sides. 

Note on Reproduction. No data are available for this species. 

Note on Habitat. This is a diurnal species which is active on the 
rain forest floor beneath the unbroken canopy. 

Range. Emoia aurulenta is known only from the Fly and Bamu 
River drainages in the Western Province, Papua New Guinea. 

Another undescribed species represented by samples from several 
populations south of the central mountain range between Western 
Province in Papua New Guinea and the Jamur Lake area in western 
Irian Jaya can not be identified with any previously described spe- 
cies of Emoia. Four specimens from various localities in Western 
Province, Papua New Guinea were collected by Fred Parker in 1969 
and 1971. A series of specimens in the Leiden Museum collected 
during a 1955 survey and the 1959 expedition along the Digul 
River and several of its tributaries in southeast Irian Jaya and sev- 
eral collected earlier at more westerly locations (identified as E. 



1985 



NEW SPECIES OE EMOIA 



baudini) also belong to this species. These were collected in part by 
the British Ornithological Expedition in the Mimika River area in 
1913 and in part by the Royal Netherlands Geographical Society 
Expedition in the Jamur Lake area in 1959. 

Emoia aenea new species 
(Fig. 2) 

Lygosoma baudini, (part) Boulenger, 1914, Trans. Zool. Soc. Lon- 
don, 20: 259. 

Holotype: MCZ 131949, adult male, collected at Menemsore, 
Western Province, Papua New Guinea, Fred Parker coll., 30 March 
1969. 

Paratypes: The following localities in Western Province, Papua 
New Guinea, Matkomrae: MCZ 144393, Emeti: MCZ 144386, 
Kiunga: MCZ 131948; the following localities in Irian Jaya, Mimika 
River area: BMNH 1913.11.1.81-82, 1913. 10.31. 164F; Gariau, 
Jamur Lake area: RMHN 21278; lower Digul River, Tanah Merah: 
RMHN 21180-85, 21273-74, 21276-77; 21279; Tanah Tinggih: 
RMHN 21275, 21280-82; Kouh: RMHN 21186 89; CAS 156680; 
Mariang: RMHN 21190-94. 




Figure 2. An adult E. aenea. 



BREVIORA 



No. 480 



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1985 NEW SPECIES OF EMOIA 7 

Diagnosis. This species can be differentiated from other species of 
the E. submetallica complex on the basis of the following combina- 
tion of characters: (1)31 to 36 midbody scale rows; (2) 49 to 54 
paravertebral scale rows between the parietals and the base of the 
tail; (3) 39 to 46 rounded lamellae under the fourth toe; (4) sixth 
(rarely fifth) upper labial enlarged and beneath the eye; (5) snout- 
vent length at maturity 53.5 to 71.0 mm; (6) such features of the 
color pattern as dorsum medium brown (in life bronzy anteriorly), 
sometimes with darker blotches in longitudinal series mostly lateral 
to the paravertebral scale rows; upper lateral surfaces (two or three 
rows of scales) darker brown to blackish, lower lateral surfaces 
grayish slate; venter whitish to ivory, occasionally more grayish on 
head and abdominal regions, pale spot on neck absent. 

Description. An Emoia of intermediate size, snout-vent length 
53.5 to 71.8 mm for 1 1 males and 59.6 to 69.5 mm for three females; 
habitus moderately robust, limbs well developed; snout moderately 
tapered, bluntly rounded, its length 54 to 62% of head breadth and 
34 to 39% of head length; head breadth 58 to 69%) (rarely less than 
60%) of head length and 14 to 17% of snout-vent length; eye rela- 
tively large, its diameter 67 to 90% of snout length and 36 to 50% of 
head breadth; ear diameter one quarter to nearly one half of eye 
diameter; rostral broader than high, forming long, nearly straight 
suture with frontonasal; supranasals narrow, elongate, in contact 
with anterior loreal; prefrontals moderately to widely separated; 
frontal longer than wide, about same length as fused fronto- 
interparietal shield, broadly rounded posteriorly, in contact with 
first and second supraoculars; four large supraoculars; seven or 
eight supraciliaries; parietals large, in broad contact posteriorly; one 
pair of nuchals; anterior loreal shorter and higher than posterior, in 
contact with first and second, second, or second and third upper 
labials; usually six to eight upper labials, sixth (occasionally fifth) 
enlarged and beneath eye; seven or eight lower labials; dorsal scales 
smooth, paravertebral scales not or scarcely enlarged; midbody 
scale rows 31 to 36; paravertebral scale rows between parietals and 
base of tail 49 to 54; preanals not or slightly enlarged; limbs well 
developed, length of extended hind limb 91 to 120% of axilla-groin 
distance and 45 to 54% of snout-vent length; 39 to 46 rounded 
lamellae under fourth toe; 10 to 14 lamellae under first toe; rank of 
adpressed toes from longest to shortest four, three, two, five, one; 
tail longer than body. 



8 BREVIORA No. 480 

Measurement (in mm) of Holotype. Snout-vent length 71.0; 
axilla-groin distance 34.7; hind limb length 31.7; head length 17.8; 
head breadth 10.0; snout length 6.2; eye diameter 5.7; ear diameter 
1.5; tail length 104±. 

Color in Preservative. The dorsal ground color ranges from tan to 
a vaguely grayish-brown or a light brown. It may be nearly uniform 
or be marked by a longitudinal row of dark brown blotches or 
sometimes dark brown, narrow, marginal lines on the scale rows 
lying lateral to the paravertebral rows. The top of the head, except 
sometimes the margins of the scales is relatively uniform. The upper 
lateral surface is marked by a narrow, darker brown band varying 
from one or two to three or four scale rows in breadth, narrowing 
on the neck and head and scarcely evident on the snout. This band is 
sometimes bordered by scattered, pale scales along its dorsal margin 
and there are numerous pale scales on the lower lateral surfaces, but 
the dark band itself it usually unmarked. The upper part of the 
upper labials are dusky, and there are some dark marks on some of 
the lower labials. The venter is whitish ivory, unmarked, and the 
undersurface of the tail has scattered, small, blackish or grayish 
spots. 

In life the head and neck are bronzy, the belly white. The Kiunga 
specimen has a patch of pink just anterior to the vent, the Menem- 
sore specimen a patch of orange. The dorsal scales are smooth and 
iridescent. 

Etymology. The name aenea refers to the bronzy coloration on 
the top of the head and anterior body. 

Comparisons. This species is close to E. submetallica obscura and 
E. submetallica popei in scale counts. In color E. aenea is more 
similar to E. s. obscura than to E. s. popei, lacking the small, whitish 
blotch on the side of the neck which is typical of the latter. E. aenea 
also appears to be slightly larger in size and the number of subdigital 
lamellae may be slightly greater than for either of the two subspecies 
of submetallica (Table 1 ), although it is closest to E. s. popei in this 
character. Also the sixth upper labial is usually the one which is 
enlarged and beneath the eye for E. s. popei and E. aenea, whereas 
it is the fifth for E. s. obscura. 

Note on Reproduction. RMHN 21281, a gravid female measuring 
62.2 mm snout-vent length, has two eggs in the oviducts. 



1985 NEW SPECIES OF EMOIA 9 

Habitat Note. Parker's specimens were found in primary forest in 
areas of high rainfall. All localities are at low elevation between sea 
level and 100 m. 

Range. Known from Western Province, Papua New Guinea, and 
Mimika River, Digul River and Jamur Lake areas in southern Irian 
Jaya. 

Emoia physicina new specices 
(Fig. 3) 

Holotype. MCZ 152287, adult male, collected near Emeti, 
Western Province, Papua New Guinea at an altitude of about 100 
m, Fred Parker coll., October 1971. 

Paratypes. Same locality as holotype: MCZ 142567-72, 152283-86, 
152288, 152290, AMNH 111713-14; other localities in Western 
Province, Menemsore: MCZ 131945-46, 152280; Kiunga: MCZ 
152281-82; Derongo: AMNH 111712, MCZ 131938-39, 131942; 
Matkomrae: MCZ 131943-44; localities in other provinces of 
Papua New Guinea, Chimbu Province, Karimui: MCZ 97308-09, 
99193-96, 109579-82, 109584-96, CAS 1 17731, 1 17750, 1 18769-70, 
118807; Soliabeda: CAS 117733 34, MCZ 109509, 109589-96, 
109598-99; Dege: MCZ 90750; Bomai: MCZ 90478-501; Eastern 
Highlands Province; AMNH 98570a-b; Gulf Province, Pio River: 
AMNH 102233. CAS 118871-72; MCZ 109601-03; Koni: CAS 
117726-27; MCZ 109514, 109529, 109532; Uraru: CAS 117753-54, 
MCZ 109572-75, 109577- 78; Oroi: MCZ 109833; Weiana: MCZ 
109521-23; localities in Irian Jaya, Assike on the Digul River: 
RMHN 5081a-b. 

Diagnosis. This species can be distinguished by the following 
combination of characters: (1) relatively weak keels on the dorsal 
scales, especially on the posterior half of the body; (2) small size, 
39.0 to 50.7 mm snout-vent length; (3) relatively low midbody scale 
count 30 to 38; (4) fifth upper labial enlarged and beneath the eye; 
(5) number of vertebral scale rows between the parietals and base of 
tail 45 to 54, and (6) number of fourth toe lamellae 34 to 43. 

Description. A relatively small Emoia, snout-vent length 39.0 to 
50.7 mm for twelve mature males and 43.3 to 50.0 mm for fourteen 
mature females; habitus moderately slender with well-developed 
limbs; snout moderately tapered, broadly rounded at tip, its length 



10 



BREVIORA 



No. 480 









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Figure 3. An adult /.'. physicina. 

51 to 60% of head breadth and 35 to 39% of head length; head 
breadth 60 to 72%- of head length and 15 to 18% of snout-vent 
length; eye relatively large, its diameter 72 to 85% of snout length 
and 39 to 46% of head breadth; ear moderate, its diameter about 
two fifths to three fifths of eye diameter, usually with two lobules 
anteriorly; rostral broader than high, forming long, convex or trun- 
cate suture with frontonasal; prefrontals moderately to widely 
separated; frontal slightly longer than broad, about as long as 
fronto-interparietal shield, in contact with two anterior supraocu- 
lars; four large supraoculars; frontoparietals fused with interparietal 
into one shield; parietals in contact; one pair of nuchals; anterior 
loreal shorter and slightly higher than posterior loreal, in contact 
with first or first and second upper labials; seven or eight upper 
labials, fifth (very rarely sixth) enlarged and beneath eye; six or 
seven lower labials; dorsal scales, at least posteriorly, with three 
weak to moderate keels; 30 to 38 scale rows at midbody; 45 to 54 
(rarely greater than 53) paravertebral rows between parietals and 
base of tail; limbs well developed, length of extended hind limb 91 to 
105%- of axilla-groin distance and 44 to 54% of snout-vent length; 34 
to 43 rounded lamellae beneath fourth toe (only one specimen has 



1985 NEW SPECIES OF EMOIA 11 

more than 40); 8 to 10 beneath first toe; rank of adpressed toes from 
longest to shortest four, three, five, two, one; tail much longer than 
body. 

Measurements (in mm) of Holotype. Snout-vent length 48.1; 
axilla-groin distance 21.9; hind limb length 22.6; head length 12.9; 
head breadth 8.3; snout length 4.5; eye diameter 3.4; ear diameter 
1.6; tail length 74.8. 

Color in Preservative. The dorsum (six to eight scale rows) is 
medium brown, nearly uniform or with rows of darker brown spots 
marginal to the paravertebral rows. The lateral surfaces, at least five 
to six upper scale rows, are usually much darker brown, nearly 
uniform or with scattered single or small groups of pale or whitish 
scales and often one or two light scales on neck. The upper labials 
are dusky, at least on the dorsal half, and the lower labials may or 
may not have dusky blotches. The venter is grayish slate, more dusky 
tan or ivory in the limb region. 

For living specimens, the dorsum is grayish-brown to brown, 
usually with two rows of paravertebral dark spots. Lateral surfaces 
are blackish with scattered light spots on the lower flanks. Some 
specimens have the upper and lower surfaces of the snout suffused 
with red. 

Etymology. The name physicina refers to the fact that the species 
has been confused with juveniles of other keel-scaled species such as 
E. phy sitae. 

Comparisons. Emoia physicina can be distinguished from all 
other known species of the E. physicae section except E. callistica 
(1) on the basis of its small size, (2) the very weak keels which 
separate it from all but E. kuekentha/i, (3) in having the fifth instead 
of sixth upper labial enlarged and beneath the eye. It differs from E. 
callistica in lower number of midbody scale rows and subdigital 
lamellae, in the distinct nuchals, and in the very weak rather than 
strong keels on the dorsal scales. 

Note on Reproduction. Gravid females CAS 110360, 49.3 mm 
snout-vent length and CAS 1 18770, 50.0 mm snout-vent length have 
two eggs in the oviducts. One hatchling (MCZ 90750) measures 24 
mm snout-vent length. 

Habitat Note. This species occupies the rain forest floor in areas 
shaded by the canopy. Specimens do bask in sunflecked areas, how- 
ever. It is strictly diurnal. 






l2 ,^ 



BREVIORA No. 480 



Range. E. physicina is known from the following provinces in 
southern Papua New Guinea: Eastern Highlands, Gulf, Chimbu, 
and Western. 

ACKNOWLEDGMENTS 

We wish to thank Pere Alberch and Ernest E. Williams, Museum 
of Comparative Zoology (MCZ), Allen E. Greer, Australian 
Museum (AM), Richard G. Zweifel, American Museum of Natural 
History (AMNH), John Pernetta, University of Papua New Guinea 
(UPNG), R. C. Drews and A. E. Leviton, California Academy of 
Sciences, and Terry D. Schawner, South Australian Museum 
(SAM) for the loan of material used in this study. The senior author 
also thanks the trustees of the Australian Museum and the Science 
and Industry Endowment Fund, Commonwealth Scientific and 
Industrial Research Organization of Australia, for their financial 
assistance while studying in the museums of Australia. Robert 
Drewes and Allen Greer have read the manuscript and been most 
helpful with their suggestions. 

LITERATURE CITED 

Brown. W. C. 1953. Results of the Archbold Expeditions No. 69. A review of 
New Guinea li/ards allied to Emoia baudini and Emoia physical'. Amer. Mus. 
Novitates. No. 1627. pp. I 25. 



'' JS ;2r-«iP<. 

b r e;ti o r a 

Museum ofjfibniparative Zoology 

US ISSN 0006-9698 

Cambridge, Mass. 21 June 1985 Number 481 



A NEW ANOLIS OF THE LIONOTUS GROUP FROM 

NORTHWESTERN ECUADOR AND SOUTHWESTERN 

COLOMBIA (SAURIA: IGUANIDAE) 

Kenneth Miyata 1 

Abstract. Anolis lynchi, new species, is described from several lowland rain 
forest localities in northwestern Ecuador and southwestern Colombia. It is allied to 
the semiaquatic anoles of the lionolus species group, and appears to be most closely 
related to Anolis poecilopus Cope of Panama and northwestern Colombia. The 
lionolus species group and the known distribution of its South American representa- 
tives is discussed. 

INTRODUCTION 

Much of the complexity of the South American Anolis fauna is 
contained within the narrow confines of the wet Pacific coastal 
region of Colombia and Ecuador (see Table 2 in Williams, 1976). 
Although these anole communities may be richer than similar 
Amazonian communities (up to perhaps twelve sympatric species 
vs. six in Amazonia), a major feature of the region seems to be more 
restricted ranges of the individual species and narrower microhabi- 
tat preferences. Localities close together geographically may have 
strikingly different faunas, and within a single fauna there may be 
distinct forest and forest edge components. 

Until recently there were few representative series of most South 
American anoles. Moderate to large series of some species from the 
Andes and their western and eastern flanks have been collected 



'Deceased; formerly Museum of Comparative Zoology, Harvard University, Cam- 
bridge, Massachusetts 02138. 



2 BREVIORA No. 481 

during the past decade by workers interested primarily in frogs, who 
find sleeping anoles at night. These have sometimes been the ones 
most poorly represented in museum collections; in some cases it 
seems as if a different anole fauna can be found at night. Field 
parties from the University of Kansas led by Dr. John D. Lynch 
have night-collected a series of anoles from the vicinity of Santo 
Domingo de los Colorados, Provincia del Pichincha, Ecuador. 
Some of these prove to represent a new species, which I here name: 

Anolis lynchi, new species 
(Figs. 1-4) 

Holotype: MCZ 124406, an adult male from Santo Domingo de 
los Colorados, 600 m elevation, Provincia del Pichincha, Ecuador, 
R. W. Henderson coll., 31 July 1968. 

Paratypes: ECUADOR: Provincia del Pichincha: KU 178953, 
Santo Domingo de los Colorados, 580 m, elevation J. D. Lynch 
coll., 12 June 1977; KU 178954-178958, 2 km E, I km S Santo 
Domingo de los Colorados, 600 m elevation, T. Berger and J. D. 
Lynch coll., 13 June 1977, and KU 178959 from the same locality, J. 
D. Lynch coll., 1 1 July 1977; MCZ 1 57 1 56, 2 km E, 1 km N Santo 
Domingo de los Colorados, 620 m elevation, J. D. Lynch coll., 31 
July 1977; Provincia de Esmeraldas: USNM 21 1222, 1-2 km W El 
Placer, 390 410 m elevation, J. A. Peters coll., 1 December 1958. 

Refered specimens: COLOMBIA: Departmento del Cauca: 
AMNH 107864-70, 109598-602, Quebrada Guangui, 0.5 km above 
Rio Patia, upper Saija drainage, 100 200 m elevation. 

Diagnosis. Anolis lynchi can be distinguished from all other 
known species of South American Anolis by the high number of 
scales across the snout ( 1 8-29 at level of second canthals), the small, 
subequal scales in the supraocular area, distinct pale flank stripes, 
and the greatly enlarged bilobed hemipenes of adult males. 
' Description. Head. Head scales small, flat to slightly granular. 
(Counts for holotype are in parentheses.) Eighteen to 29 (28) scales 
across snout at level of second canthals. Seven to 10 (8) scales 
border rostral posteriorly. Circumnasal scale separated from rostral 
by one to two (2) scales. Nine to 12 (10) scales between supranasals. 
Supraorbital semicircles distinct, raised, separated medially by 
three to six (4) scales. Supraocular scales small, subequal, slightly 



1985 



ANOLIS LYNCH I 




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BREVIORA 



No. 481 




Figure 2. Anolis lynchi, new species. Lateral view of head of holotype (MCZ 
124406). 



keeled to granular, 11 to 19 (18) across widest point. Supraciliaries 
three to five (4). Canthus distinct in adults, less so in juveniles; five 
to six (5) canthals, the second the largest. Loreal rows seven to ten 
(10), subequal. Nasal area slightly swollen in adults. 

Temporals and intertemporals granular. Interparietal slightly 
smaller than ear opening, separated from supraorbital semicircles 
by four to six (4) scales. Interparietal much smaller than ear opening 
in some specimens. Scales anterior to interparietal enlarged or sub- 
equal (enlarged) with respect to temporals; scales posterior to inter- 
parietal slightly flattened or granular (flattened), larger than, but 
grading into, dorsal scales. No enlarged supratemporal rows. Dis- 
tinct thin fold extending from supraorbital semicircles laterally to 
mid-orbit, where it extends posteriorly above ear opening and fades 
out in axillary region. 

Suboculars slightly keeled, separated from supralabials by one 
row of scales. Seven to 10(10) supralabials to center of orbit. 

Mental groove extending approximately three quarters across, 
each part much wider than deep. Six to eight (7) scales in contact 
with mental posteriorly. Eight to II (11) infralabials. Chin scales 
granular, except for some which are slightly keeled. 



1985 



A NOUS LYNCH I 




Figure 3. Anolis lynchi, new species. Dorsal view of head of holotype (MCZ 
124406). 



BREVIORA 



No. 481 




Kigure 4. Anoli.s lynchi, new species. Ventral view of head of holotypc (MCZ 
124406). 



Dewlap. Dewlap of adult males medium-size, extending from 
under the midpoint of the orbit posteriorly to a point just under the 
axilla. Scales form 13 to 16 longitudinal rows when dewlap 
extended, but arranged irregularly and not parallel. Dewlap scales 
narrowly compressed laterally and elongate, especially at the edge. 
Dewlap of adult females small. 

Trunk. Middorsal scales flattened or slightly keeled (flat); two 
medial rows slightly larger than adjoining rows. Lateral scales gran- 
ular, grading into larger, subimbricate ventral scales. Ventral scales 



1985 A NOUS LYNCH I 7 

slightly to moderately keeled; keeling more prominent midventrally 
than anteriorly or posteriorly. 

Limbs and Digits. Scales on limbs smooth to lightly keeled; 
generally smoother on upper arm and thigh than on lower arm or 
calf. Scales on forearm subequal; those on hind limbs are larger 
anteriorly. Keeling on limb scales unicarinate. Supradigital scales 
multicarinate, usually with two longitudinal keels. Fifteen to 18 (17) 
lamellae under phalanges ii and iii of fourth tow. Distal phalanx 
raised slightly above adjacent phalanx. 

Tail. Base of tail greatly swollen in adult males, slightly thicker 
than trunk in pelvic region, tapering abruptly about one quarter 
SVL behind the vent. Tail slightly compressed laterally, height 
about 20 percent more than width. Caudal scales unicarinate, more 
strongly keeled ventrally than laterally or dorsally. Males with a 
distinctly keeled middorsal scale row beginning where the swollen 
tail base ends. Postanals not enlarged. 

Hemipenes. The hemipenes of adult males are greatly enlarged, 
accounting for the extreme swelling at the base of the tail. The 
everted hemipenes are bifurcate, the split extending back about one 
third of the distance from the distal end to the base. The distal end 
has small calyces that become larger proximally. The shaft has large 
folds, becoming smoother on the base. The sulcus spermaticus has 
several longitudinal folds. 

Measurements. Adult males 54.2 to 61.8 mm snout-vent length 
(SVL), adult females 49.5 to 59.3 mm SVL. 

Color as Preserved. The dorsum ranges from cinnamon-brown 
to deep fuscous. Males show indistinct lighter brown longitudinal 
stripes dorsolaterally; females have a distinct gray middorsal stripe. 
Some males show a broken and indistinct series of small, elongate 
dark brown marks middorsally. Both sexes show traces of indistinct 
pale and dark brown lateral stripes that are very thin just above a 
prominent muddy gray flank stripe. The gray is heavily flecked with 
brown, but the flank stripes are very distinct in most specimens. 
Below this flank stripe is a dark brown stripe heavily flecked with 
gray. The gray flank stripe extends to the orbit, and the sides of the 
head are gray with brown flecks and spots. The venter is an almost 
immaculate light gray to cream, with some gray flecking laterally. 
The undersides of the limbs and tail are lightly mottled with brown. 

Color in Life. The following description of Ecuadorian speci- 



8 BKEVIORA No. 481 

mens from Santo Domingo de los Colorados is compiled from the 
field notes and Kodachrome transparencies of living animals pro- 
vided by John D. Lynch. 

The dorsum is brown to reddish brown, with an obscure series of 
dark brown middorsal blotches in males and a pale gray middorsal 
stripe in females. The flank stripe is dull cream and is bordered 
above by a thin, yellowish cream stripe. The flanks are mottled 
brown, with an olive wash in some specimens. The sides of the head 
are off-white with dark brown marks under the orbits. The chin, 
venter, and underside of the limbs are cream; the underside of the 
tail of females and the underside of the tail of males posterior to the 
swollen tail base is yellowish. The dewlap is orange with white 
scales. The iris is bright coppery brown. The tip of the tongue is 
gray. 

Color notes of the Colombian specimens were provided by 
Charles W. Myers. They were similar to the Ecuadorian specimens, 
except Myers noted grayish chins, a dull gray reticulum under the 
limbs, and the dewlaps were dull orange (Myers' italics). Myers also 
noted that the throat lining was unpigmented. 

Etymology. The species is named after John D. Lynch, whose 
indefatigable excursions into the Ecuadorian night are responsible 
for many of the known specimens. 

Variation. The Colombian specimens referred to A no/is lynchi 
differ somewhat from the Ecuadorian sample. They have fewer 
scales across the snout at the second canthals ( 18-24 vs. 25-29) and 
fewer scales across the supraocular region ( I 1 - 1 5 vs. 15-18). Both 
counts reflect slightly larger head scales in the northern population. 
There also appears to be a slight difference in color of living ani- 
mals. The Ecuadorian specimens had immaculate cream venters 
while those of the Colombian specimens were cream with a dull 
gray reticulum under the limbs and a grayish throat. The dewlaps of 
adult males from both areas were basically orange, but those of the 
northern population were characterized as a dull orange while those 
of the southern populations were a fairly bright orange. 

Despite the differences seen between the Ecuadorian and Colom- 
bian populations, 1 regard them as conspecific. The differences may 
well be clinal, and there is no evidence for sharp discontinuities in 
character states that might reflect breaks in gene flow. However, 
since there is a possibility that the Colombian populations may be 
distinct, I have not designated these specimens as paratypes. 



1985 A NOUS LYNCH/ 9 

Comparisons. Anolis lynchi most closely resembles A. poecilo- 
pus of Panama and northwestern Colombia. The two are closely 
related (see discussion below), but can be distinguished by the char- 
acters summarized in Table 1. 

The reduced supraocular scales of A. lynchi are paralleled in two 
other groups of South American anoles. The latifrons group 
includes several giant anoles with adult SVLs of 125 mm and more. 
The aequatorialis group is a poorly known complex of medium to 
larger size anoles (adult SVLs generally over 80 mm). Species in 
both groups lack transverse processes on the posterior caudal verte- 
brae and are alpha anoles in the terminology of Etheridge (1960). 
Since Anolis lynchi has these transverse processes, it falls into the 
beta section of the genus. Anolis lynchi can be easily distinguished 
from any known member of the latifrons or aequatorialis groups by 
its smaller adult size (less than 65 mm SVL vs. 80 mm or more), 
much smaller supraocular scales (11-19 across), smaller scales 
across the snout (at least 18 at level of second canthals), and the 
greatly enlarged, bifurcate hemipenes of adult males. 

Anolis granu/iceps, a small beta anole of uncertain relationship 
from western Colombia and northwestern Ecuador, shows a similar 
reduction in the size of the supraocular scales, but the central scales 
in the supraocular area are slightly enlarged and keeled, while those 
of lynchi are subequal and usually granular. The hemipenes of 
granu/iceps are small and not bifurcate, and the small male dewlap 
is golden orange. 

Natural History. All of the Ecuadorian specimens of A. lynchi 
for which there are field notes were collected at night as they slept 
on low vegetation adjacent to small streams in closed-canopy rain 
forest. They were sleeping head-up on leaves and small ferns along- 
side, or overhanging, the creek beds, usually within 1 m of the 
ground. The Colombian specimens were collected in a forested 
ravine on low vegetation near a stream during the day. It appears 
that A. lynchi is associated with streamside lowland rain forest 
habitats, although admittedly collectors may have concentrated on 
streamside habitats, or the lizards may simply have been easier to 
see in these areas. Charles W. Myers, who collected the Colombian 
sample, believes that it is not an aquatic anole in the same sense as 
A. poecilopus. At his collecting site this niche was apparently occu- 
pied by A. macro/epis, another member of the lionotus group. 
However, the association between A. lynchi and streamside forest 



10 BREVIORA No. 481 

habitats seems real, at least in the southern part of its distribution. 
Despite extensive collecting in the vicinity of Santo Domingo de los 
Colorados, the only available specimens were found in the imme- 
diate vicinity of forest streams. Other anoles found within 200 m of 
one such creek (2 km E, 1 km N Santo Domingo de los Colorados) 
include A. chforis, fraseri, gracilipes, peraccae, princeps, and an 
undescribed species related to vittigerus. 

Discussion. Williams (1976) recognized two South American 
anoles, A. poecilopus Cope and A. macrolepis Boulenger, in his 
lionotus species group. He has subsequently described a third, A. 
rivalis Williams, 1984. The group was characterized in Williams' 
1976 key by the presence of a zone of enlarged flat or slightly keeled 
middorsal scales. In A. macrolepis and A. rivalis the enlarged mid- 
dorsals are considerably larger than the lateral scales and are 
arranged in from 11 to 17 longitudinal series. In A. poecilopus the 
enlarged middorsals are only slightly larger than the laterals, into 
which they grade. In A. lynchi the zone of enlarged middorsals is 
scarcely evident. There are only two slightly larger and flatter para- 
vertebral scale rows, and the difference in size and shape is slight. 
The lack of an obvious zone of enlarged middorsal scales, and the 
extreme reduction in the size of scales on the supraocular discs, at 
first obscured the relationships of A. lynchi and suggested an asso- 
ciation with the aequatorialis group (sensu Williams, 1976). How- 
ever, the presence of transverse processes on the posterior caudal 
vertebrae made this impossible. 

The large bilobed hemipenes of A. lynchi provide the clearest 
morphological evidence linking A. lynchi with the lionotus species 
group. Many anoles show a tendency for the distal end of the 
hemipenis to be bilobed, but few show a strongly forked hemipenis 
with the bifurcation extending as much as a third the length of the 
everted organ. This condition characterizes several semiaquatic 
anoles in Central America and northwestern South America allied 
to A. lionotus Cope. In addition to a semiaquatic streamside habi- 
tat and bilobed hemipenes, these lizards share a similar body pat- 
tern with distinct lateral stripes. The group consists of six currently 
recognized species: A. lionotus of central Panama, A. oxylophus of 
Costa Rica, Nicaragua and western Panama, A. poecilopus of east- 
ern Panama and northwestern Colombia, A. rivalis and A. 
macrolepis of western Colombia, and A. lynchi. Each is distinct in 



1985 A NOUS LYNCH I II 

external morphology, and occupies largely exclusive ranges. Anolis 
lynchi and A. macrolepis have both been collected at Quebrada 
Guangui in Cauca, Colombia and A. poecilopus and A. lionotus 
occur in sympatry in central Panama (Campbell, 1973), but the 
zones of overlap are probably narrow. 

Anolis lynchi is the southernmost representative of the lionotus 
group. The distribution of the South American species of the group 
is shown in Figure 3. The range of A. poecilopus, which on the basis 
of external similarity seems to be the closest relative of A. lynchi, is 
separated from that of A. lynchi by a substantial gap of approxi- 
mately 1,000 km. The intervening area is inhabitated by A. 
macrolepis and A. rivalis, which are not morphologically close to 
either species. 

ACKNOWLEDGMENTS 

John D. Lynch collected most of the specimens of the new species 
and provided information about the localities, field notes, and 
Kodachrome transparencies. Charles W. Myers of the American 
Museum of Natural History (AMNH) loaned Colombian speci- 
mens and field notes. Ernest E. Williams shared his wealth of 
knowledge on South American Anolis and commented on the 
manuscript. Stephen Ayala commented on the manuscript and 
answered questions regarding Colombian specimens. Other speci- 
mens were loaned by William E. Duellman, Museum of Natural 
History of the University of Kansas (KU). W. Ronald Heyer and 
Ronald 1. Crombie, National Museum of Natural History (USNM), 
and Arnold G. Kluge, University of Michigan Museum of Zoology 
(UMMZ). My own unsuccessful efforts to collect Anolis lynchi in 
Ecuador were partially funded by Earthwatch and the Center for 
Field Research of Belmont, Massachusetts and by the Museu de 
Zoologia of the Universidade de Sao Paulo. I thank Jeannie 
Sellmer and Laszlo Meszoly for the illustrations. 



12 



BREVIORA 



No. 481 



OVER 1000 METERS 
100 KILOMETERS 




Figure 5. Distribution of the Anulis lionotus species group in South America 
and lower Central America. Triangles: A. lynchi, n. sp. Solid squares: A. macrolepis. 
Open squares: A. rivalis. Circles: A. poecilopus. Localities for A. macrolepis and A. 
rivalis were provided by Stephen Ayala and Ernest E. Williams. 



1985 A NOUS LYNCH I 13 



LITERATURE CITED 

Cam FBI i i ., H. W. 1973. Ecological observations on Anolis lionotus and Anolis 

poecilopus (Rcptilia. Sauria) in Panama. Amer. Mus. Nat. Hist. Novitatcs. 

2516: I 29. 
ETHiKiiKil-. R. E. I960. The relationships of the anoles (Reptilia: Sauria: lguani- 

dae). An interpretation based on skeletal morphology. Ann Arbor, University 

Microfilms. xi\ + 236 pp. 
Williams. E. E. 1976. South American anoles: the species groups. Pap. Avuls. 

Zool.. S. Paulo. 29: 259 268. 
1984. New or problematic Anolis from Colombia. III. Two new semia- 

quatic anoles from Antioquia and Choco. Colombia. Breviora Mus. Comp. 

Zool.. No. 478. pp. 1 22. 



Vt >S. COMP. ZOOL 

B R E~XJ R A 

of Co2B^»arative Zoology 



i ] c ! 

US ISSN 0006-9698 



Cambridge, Mass. 21 June 1985 Number 482 

NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. IV. 

ANOLIS ANTIOQVIAE, NEW SPECIES OF THE 

ANOLIS EULAEMUS SUBGROUP FROM 

WESTERN COLOMBIA. 

Ernest E. Williams 1 

Abstract. A new species of the Anulis eulaemus subgroup of the A. aequatorialis 
species group from the western part of the Departmento Antioquia, Colombia is 
closer to A. fitchi Williams and Duellman of southern Colombia and northern 
Ecuador than to neighboring A, ventrimaculatus Boulenger and A. eulaemus Bou- 
lenger of the Departments of Valle and Risavalda. 

Three adult female Anolis from the Cordillera Occidental of 
Antioquia, Colombia represent a distinctive new species of the 
eulaemus subgroup of the Anolis aequatorialis species group. Other 
members of this subgroup include A. eulaemus Boulenger, A. ven- 
trimaculatus Boulenger, A. gemmosus O'Shaughnessy, A. maculig- 
ula Williams and A. fitchi Williams and Duellman, all ranging 
farther to the south in Colombia and Ecuador. Several novelties in 
the genus Anolis have recently been discovered in Antioquia, so it 
seems appropriate to emphasize the interest of the new material by 
naming this lizard: 



'Museum of Comparative Zoology, Harvard University, Cambridge. Massachusetts 
02138. 



BREVIORA 



No. 482 




\ \ 



i i 



\ 





\ 


c 


, 1 


T 


\ \ 




\ \ 


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\ \ 




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3 



1985 



ANGUS ANTIOQUIAE 




Figure 2. Anolis uniioquiae, 1 ACM 72763. Lateral view of head. 



A no/is antioquiae, new species 
(Figs. 1-4) 

Type: INDERENA 0277, adult female. 

Type locality: Along a road paralleling Quebrada Chaparral, 
Rio San Juan drainage, 10 km E of Andes (town), western Antio- 
quia, Colombia, 2200 2300 m. Philip A. Silverstone, Carlos Arturo 
Escobar and Luis Carlos Acevedo Cuartas coll., July 1971. 

Paratypes. Antioquia: LACM 72763, adult female, same local- 
ity as type. CSJ 310, adult female, Urrao. 

Description. Head. Head scales minute, anteriorly weakly uni- 
to multicarinate, posteriorly pustulose. Sixteen to 19 scales across 
snout between second canthals. Six to seven scales border rostral 
posteriorly. Anterior nasal scale separated from rostral by one scale 
or in contact. 

Supraorbital semicircles separated medially by four to five scales. 
The medial supraocular scales enlarged, wrinkled or keeled, but 
supraocular disk not or very weakly defined. Three short supercil- 
iaries on each side, followed by granules. Canthus projecting later- 
ally, sharp-edged, slightly overhanging loreal region. Second and 
third or third and fourth or fourth and fifth scales longest. Ten to 
twelve loreal rows. 



BREVIORA 



No. 482 




Figure 3. Anolis anlioquiae. LACM 72763. Dorsal view of head. 



1985 



ANOUS ANTIOQUIAE 




Figure 4. Anolis anlioquiac, LACM 72763. Ventral view, of head. 



6 brfviora No. 482 

Temporals and supratemporals granular. An intertemporal dou- 
ble row of enlarged scales hardly indicated. No interparietal. Scales 
in the interparietal area granular posteriorly, grading into parietal 
area, bounded by oblique lateral ridges which converge on but do 
not meet a blunt occipital knob, larger scales alongside the 
supraorbital semicircles. Ear moderate but less than half eye 
aperture. 

Suboculars weakly keeled, narrowly and irregularly in contact 
with the supralabials. Six to seven supralabials to below the center 
of the eye. 

Mental completely or incompletely divided, each half as wide as 
deep, indented medially by six to seven small swollen smooth scales 
between the large smooth sublabials, only the first of which on each 
side is in contact with the infralabials (on one side of INDERENA 
0277 the second sublabial also is in contact with the infralabials). 
Central gular scales smooth or keeled, imbricate, projecting, sub- 
equal, becoming rather abruptly larger, laterally adjacent to the 
infralabials. 

Trunk. A central zone of small keeled juxtaposed scales grading 
into smaller projecting and bluntly or sharply pointed flank scales 
that are juxtaposed or separated by minute granules. Ventrals 
larger, smooth, juxtaposed or subimbricate, not in transverse rows. 

Dewlap. Moderate (female), extending posteriorly to the level 
of the axilla. Scales close packed, more or less projecting, smaller 
than ventrals. 

Limbs and digits. Scales on limbs uni- to multicarinate, the 
largest (on anterior thigh) as large or larger than ventrals, on digits 
multicarinate. Digital dilations narrow, 21 to 22 lamellae under 
phalanges ii and iii of fourth toe. 

Tail. Compressed, no dorsal crest. Tail scales small, larger 
below, all keeled except those under the base of tail smooth. Two 
middorsal rows enlarged. 

Size. The snout-vent length of the INDERENA type is 72 mm, 
of CSJ 310 77 mm, and of LACM 72763 74 mm. The tail is broken 
in all three specimens. 

Color (as preserved). Light purplish with brown markings 
expanding at intervals on the dorsum (Type and LACM 72763) or 
dark purplish with a dorsal zone brown with a black line on each 
side (CSJ 310). Throat and belly very weakly spotted on a light grey 



1985 



ANOLIS ANTIOQUIAE 



Table I . Comparison of Anolis amiuquiae and A. fitchi. 



antioquiae 



fitchi 



head 
scales 

number between 
second canthals 

postrostrals 

superciliaries 



canthals 

loreal rows 
interparietal 

suboculars 



supralabials to 
center of eye 



middorsals and 
flank scales 



ventrals 



lamellae under 
4th toe 



minute, uni- and 
multi-carinate 

16-19 

6-7 

the 3 anterior short followed 
by granules 

sharply keeled, slightly over- 
hanging loreals 

10-12 

none 

narrowly and irregularly in 
contact with supralabials 



6-7 

indented medially by 6-7 
small scales between large 
suhlahials 

a dorsal zone of keeled juxta- 
posed scales grading into 
smaller conical flank scales 
juxtaposed or separated by 
minute granules 

larger than dorsals, smooth, 
subimbricate or juxtaposed, 
not in transverse rows 



21-22 



small, multicarinate, tubercu- 
late or wrinkled 

11-18 

6 9 

one anterior elongate followed 
by one or two shorter ones 
and granules 

blunt, not overhanging loreals 

8-11 

smaller than ear, separated 
from semicircles by 3-6 scales 

separated from supralabials by 
one row or narrowly in 
contact 

8-11 

in transverse contact with 5-8 
scales between infralabials. 
Suhlahials not differentiated. 

2 4 middorsal rows slightly en- 
larged, keeled, swollen, sub- 
imbricate, lateral granules con- 
ical, juxtaposed 

larger than dorsals, smooth, 
imbricate or subimbricate, 
tending to be in transverse 
rows 

21-24 



8 BREVIORA No. 482 

ground (Type) or light purplish laterally smudged medially (CSJ 
310). The limbs are very weakly banded, the tail darkish above, 
lighter below. Dewlap white, blotched and spotted with intense 
black. 

Color in life. P. A. Silverstone has provided color notes for the 
Type and LACM 72763: "Dorsum green with black median stripe 
with paired light green spots. Venter brown with white spots. Dew- 
lap red orange with black blotches and yellow border." 

Ecology. Silverstone reports the two specimens collected by his 
party "on ferns at the edge of forest next to pasture." 

Comparisons. Anolis antioquiae is closest to recently described 
Anolis fitchi Williams and Duellman. In both the female has a 
blotched dewlap. In both there is a noticeable indication of a small 
round parietal spine or convexity (Fig. 2) just anterior to the 
enlarged median name scales. This small prominence is not evident 
in other species of the group. Of the other recognized species of the 
eulaemus subgroup, two — A. ventrimaculatus and A. gemmosus — 
entirely lack a dewlap in females. In the one female referred to A. 
eulaemus the dewlap is much reduced, apparently with uniformly 
dark skin. All three examined specimens of A. antioquiae lack an 
interparietal; this scale is always present in A. fitchi and in A. 
eulaemus, but sometimes absent in A. gemmosus and often so in A. 
ventrimaculatus. 

A. antioquiae is unique in one respect: the very sharp canthal 
ridge projecting above the loreal region, although an apparently 
trivial character, has not been seen by me in any other species of the 
group or indeed in any Anolis. Within the group the three subequal 
short superciliaries are unusual. The median angular indentation of 
the mental by postmental scales contrasts with the straight trans- 
verse border seen at the mental gular contact in A. fitchi. The 
conical flank scales often surrounded by granules are again very 
different from other members of the group. 

Although this species is apparently allopatric to — somewhat to 
the north of — A. eulaemus and A. ventrimaculatus, and although 
only females are known (and the female reported as that of eulae- 
mus is questionably so referred), there is no question of conspecific- 
ity or even of very close relationship. Even with A. fitchi, very 
distant in Napo in Ecuador and Putamayo in Colombia, despite 
more resemblances, there is no demonstration of close affinity 
except for position with the same subgroup of a species group. 



1 985 A SOI. IS A NTIOQL I A E 9 

ACKNOWLEDGMENTS 

I am grateful to Jorge Hernandez of INDERENA, John Wright 
and Robert Bezy of the Los Angeles County Museum (LACM), 
and Marco A. Serna of the Colegio San Jose, Medellin, Colombia 
(CSJ) for the loan of specimens. Laszlo Meszoly made the 
drawings. 

LITERATURE CITED 

Williams, E. E.. and W. E. Di ii i man. 1984. Anolis fitchi, a new species of the 
Anoli.s acquaiorialis group from Ecuador and Colombia, pp. 257 266. //; Seigel 
R. A. ci al. (eds.). Vertebrate Ecology and Systematics A Tribute to Henry 
Fitch. Lawrence. Kansas. 



., us CO«*f° U 



B R E JU ^S R A 



ur»»»' 



.iiseniii of Comparative Zoology 



US ISSN 0006-9698 



Cambridge, Mass. 21 June 1985 Number 483 

NOTES ON PRISTIDACTYLUS (SQUAMATA: IGUANIDAE) 

Richard Etheridge 1 and Ernest E. Williams 2 

Abstract. Pristidactylus is diagnosed as a genus of austral South American 
anoloid iguanid lizards allied to Enyalius and possibly paraphyletic with respect to 
Leiosaurus, Aperopristis, and Diplolaemus. West of the Andes Pristidactylus torqua- 
tus occurs in forested coastal areas and the inland Cordilleras of central Chile; P. 
valeriae (new comb.) and P. alvaroi (new comb.) are restricted to disjunct patches of 
relict forests to the north in Santiago Province. East of the Andes four species occur 
in Argentina. P. casuhatiensis lives in high, wet, rocky meadows atop the Sierra de la 
Ventana in southern Buenos Aires Province, and P. achalensis occurs in a similar 
habitat atop the Sierra de Cordoba in Cordoba Province. P. scapulatus is known 
from arid, rocky habitats at disjunct localities in the Cordilleras and precordilleras 
from western Catamarca Province southward to Mendoza Province, and on isolated 
basaltic mesetas at lower altitudes from Mendoza southward to Chubut Province. In 
cordilleran and precordilleran populations adult females virtually lose the bold 
juvenile pattern whereas those from the basaltics do not. Although the latter are not 
recognized nomenclatorially, the name P. araucanus is available for them. P. fascia- 
tus, type species of the genus, lives in a narrow belt of sandy steppe from San Juan 
Province southward to Rio Negro and extreme southern Buenos Aires Provinces. It 
has frequently been incorrectly referred to as P. araucanus. Leiosaurus bardensis is a 
synonym of P. fasciatus. A key to the species of Pristidactylus is provided. 

INTRODUCTION 

Pristidactylus is a genus of austral South American iguanid 
lizards with a disjunct distribution in Chile and Argentina, and an 
exceedingly complex taxonomic history. Fitzinger (1843: 58) de- 
scribed Pristidactylus as a subgenus of Leiosaurus Dumeril and 
Bibron, 1837, and designated Leiosaurus fasciatus Dumeril and 



'Department of Biology, San Diego State University, San Diego, California 92182. 
2 Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. 



2 BREVIORA No. 483 

Bibron, 1837 as its type species. Boulenger (1885: 127) employed 
Pristidactylus, but only for the species fasciatus. In this century, up 
until 1976, fasciatus and all of the other species now referred to 
Pristidactylus were variously placed in Leiosaurus, Urostrophus 
Dumeril and Bibron, 1837 and Cupriguanus Gallardo, 1964. In 1976 
Etheridge {in Paull, Williams, and Hall, 1976: 10) pointed out that if 
Barrio (1969) was correct in placing Cupriguanus araucanus Gal- 
lardo, 1964 in the synonomy of Leiosaurus fasciatus Dumeril and 
Bibron, 1837 and if the latter were congeneric with Cupriguanus 
achalensis Gallardo, 1964, type species of Cupriguanus, then Cupri- 
guanus is a junior synonym of Pristidactylus Fitzinger, 1843. In this 
work seven species are referred to Pristidactylus: achalensis Gal- 
lardo, 1964; alvaroi Donoso-Barros, 1975; casuhatiensis Gallardo, 
1968; fasciatus Dumeril and Bibron, 1837; scapulatus Burmeister, 
1861; torquatus Philippi, 1861; and valeriae Donoso-Barros, 1966. 
Pristidactylus in this modern sense has never had a proper diag- 
nosis, nor have the species within it been adequately distinguished 
from one another. We plan a future, more formal review of Pristi- 
dactylus and related taxa; we here propose only to provide a diag- 
nosis and discussion of the genus as now understood, enough 
description of the named forms to permit their recognition, and 
lastly a key which is arranged to display the relationships of the 
species as we now understand them. These notes are based on exam- 
inations of 221 specimens of Pristidactylus in European, North and 
South American museums (see Acknowledgments), and field work 
in Argentina by Etheridge in late January, February, and March 
during the austral summer of 1983. 

Pristidactylus Fitzinger 1843 

1843 Leiosaurus (Pristidactylus) Fitzinger, Syst. Rept., Wien, 1: 16. — Type species 
(by indication): Leiosaurus fasciatus Dumeril and Bibron 1837. 

1845 Leiosaurus (Pentadactylus) Gray, Cat. Spec. Liz. Brit. Mus., London, 224. — 
Type species (by monotypy and original designation) Leiosaurus fasciatus 
D'Orbigny in Dumeril and Bibron 1837. 

1885 Pristidactylus — Boulenger, Cat. Liz. Brit. Mus., London, 2: 127. 

1964 Cupriguanus Gallardo, Neotropica, B. Aires, 10(33): 127. — Type species (sub- 
sequent designation by Gallardo, 1967: Cupriguanus achalensis Gallardo 
1964.) 

1976 Pristidactylus — Etheridge in Paull, Williams, and Hall, Breviora, Mus. Comp. 
Zool., No. 441, p. 10. 



1985 PRISTIDACTYl.US 3 

Diagnosis. Pristidactylus is a member of the "anoloid" group of 
iguanid genera, specified by the acquisition of endolymphatic sacs 
that extend back between the parietal and occipital bones into the 
dorsal neck musculature (Etheridge and de Queiroz, in prepara- 
tion). Within anoloids Polychrus is considered the sister group of 
the remaining genera, the latter having lost femoral pores and 
acquired a spinulate Oberhauchen with flat cell borders. Within this 
latter group Pristidactylus together with Diplolaemus, Leiosaurus, 
Aperopristis and Enyalius retain four pairs of sternal ribs and have 
acquired a unique structure of the distal two or three subdigital 
scales: each has a median longitudinal groove. Although grooved 
distal subdigitals are obscure or lacking in forms with distinctly 
multicarinate subdigital scales (P. fasciatus, A. catamarcensis, A. 
paronae, E. leechii, and E. brasiliensis), their presence is considered 
a derived characteristic of the group, their absence the result of loss. 
This subgroup containing Pristidactylus forms the sister taxon to a 
group formed by the remaining anoloid genera: Urostrophus, Ani- 
solepis, Aptycholaemus, Chamaeleolis, Phenacosaurus, Chamaeli- 
norops, and Anolis, all of which have three rather than four pairs of 
sternal ribs. 

Relationships among the genera that form the subgroup contain- 
ing Pristidactylus remain unresolved, and a preliminary analysis 
indicates that Pristidactylus itself may be paraphyletic. Neverthe- 
less, Pristidactylus can be diagnosed within this group. It differs 
from Aperopristis, Diplolaemus, and Leiosaurus in having a black 
bar within the antehumeral fold and marked sexual dichromatism; 
it further differs from Aperopristis and Leiosaurus in having fully 
autotomic caudal vertebrae and a different color pattern, and from 
Diplolaemus in possessing pterygoid teeth. Pristidactylus differs 
from Enyalius in having the caudal scales smooth proximally and 
keeled distally (keeled throughout in Enyalius); slightly convex 
rather than conical mesoptychials; wide, smooth, scarcely imbricate, 
lamellarlike supradigitals rather than sharply unicarinate, rhomboi- 
dal and imbricate supradigitals; the prefrontal-frontonasal part of 
the skull roof rounded rather than flat; a more robust body (some- 
what compressed in Enyalius); shorter limbs (adpressed hind limb 
reaches to between the antehumeral fold and orbit in Pristidactylus, 



4 BREVIORA No. 483 

to the orbit or beyond in Enyalius); and a shorter tail (tail/ snout- 
vent 0.73-1.60 in Pristidactylus, 1.60-3.30 in Enyalius). 

Juveniles of Pristidactylus and Diplolaemus have a light grey or 
yellowish background with bold, dark crossbands over the head, 
body, tail, and limbs, with a characteristic horseshoe-shaped mark- 
ing over the back of the head, and the crossbands of the back 
usually constricted laterally. In both sexes with increasing size the 
dark markings typically break up or acquire irregular borders. 
Large adult male Pristidactylus become nearly unicolor except for a 
conspicuous, black antehumeral bar, and have at most only a faint 
trace of the juvenile pattern. The juvenile pattern, though altered, 
remains bold in adult females, except for the northern populations 
of P. scapulatus. The juvenile markings show various patterns of 
ontogenetic change in Diplolaemus, but never disappear entirely, 
and are identical in large individuals of both sexes. The distinctive 
butterfly or fleur-de-lis markings of Leiosaurus and Aperopristis are 
present at hatching and change little in either sex. Thus, marked 
sexual dichromatism is a characteristic of Pristidactylus, shared 
with Enyalius, but distinguishing it from Aperopristis, Diplolaemus 
and Leiosaurus. 

Boulenger in 1885 transferred Leiosaurus torquatus, and in 1889 
Leiosaurus scapulatus, to Urostrophus Dumeril and Bibron, 1837; 
type species U. vautieri Dumeril and Bibron, 1837. Gallardo (1964) 
recognized that a number of differences exist between Urostrophus 
vautieri and Leiosaurus scapulatus. He proposed the genus Cupri- 
guanus to include scapulatus and two additional species from 
Argentina described in the same work, araucanus from Laguna 
Blanca, Neuquen Province, and achalensis from the Pampa de 
Achala in Cordoba Province. He further suggested that Leiosaurus 
torquatus probably should be transferred to Cupriguanus. Gallardo 
pointed out that Cupriguanus differs from Leiosaurus (in which he 
included Diplolaemus and Aperopristis) in having compressed and 
pectinate infradigitals at the bases of the first, second, and third toes 
of the hind foot. The pectinate scales on the inner border of the first, 
second, and third toes, conspicuous in P. fasciatus and emphasized 
by Gallardo (1964) as characteristic of Cupriguanus, are not uni- 
formly developed in all species of Pristidactylus, and are least prom- 
inent in P. casuhatiensis. Also, these scales are conspicuously 
swollen, though not pectinate, in Diplolaemus, so that this feature is 



1985 PR/STIDACTYI.LS 5 

not as clear-cut as Gallardo thought. Gallardo also pointed out that 
Prist idactylus differs from Urostrophus in having a shorter tail, 
more labial scales and more scales between the supralabials and 
suboculars, no projecting scales around the eye, the trunk and tail 
not compressed, and the fourth toe shorter. Gallardo's (1979) use of 
the binomial Pristidactylus vautieri for Urostrophus vautieri was 
unjustified and probably unintended. As indicated earlier, Urostro- 
phus is a member of the sister taxon of the group containing Pristi- 
dactylus, distinguished from the latter by the loss of a pair of sternal 
ribs and the absence of grooved distal subdigital scales. Thus, 
although not all of Gallardo's characters for Cupriguanus (= Pristi- 
dactylus) are useful, Pristidactylus can be distinguished from Uro- 
strophus as well as from other anoloid genera as indicated in the 
diagnosis. 

Cei (1973a) compared Cupriguanus with Leiosaurus, Aperopris- 
tis, and Diplolaemus. His concept of Cupriguanus was based upon 
the Argentinian species, achalensis, scapulatus, casuhatiensis, and 
"fasciatus" (—araucanus). Leiosaurus bardensis (— Pristidactylus 
fasciatus) was included with belli and catamarcensis in the genus 
Leiosaurus, and Aperopristis was considered monotypic with the 
species paronae. Diplolaemus included the species darwinii, bibro- 
nii, and leopardinus. He pointed out correctly that Diplolaemus 
differs from the others in lacking pterygoid teeth. The subocular was 
said to be elongate in Diplolaemus and Cupriguanus, represented by 
an arc of small, subequal scales in Leiosaurus and Aperopristis; 
however, we find this condition variable in both Diplolaemus and 
Pristidactylus. The subdigital scales were said to be smooth in 
Diplolaemus and Cupriguanus, unicarinate or multicarinate in Lei- 
osaurus, and multicarinate in Aperopristis. The subdigital scales 
are smooth in Diplolaemus and multicarinate in Aperopristis; how- 
ever, in Pristidactylus they are smooth, indistinctly unicarinate or 
multicarinate. Cei's description of the subdigital scales of A. paro- 
nae and L. catamarcensis as having three to five distinct keels is 
correct, but his statement that the subdigital scales of belli have one 
to three keels is not entirely in accord with our observations. In 64 
specimens of L. belli horn Chubut, Mendoza, Neuquen, Rio Negro, 
and Santa Cruz Provinces we find the subdigital scales either 
entirely smooth, as stated in the type description (Dumeril and 
Bibron, 1837), or with one, rarely two, indistinct keels, as mentioned 



6 BREVIORA No. 483 

by Koslowsky (1898). The distal caudal scales were said to be 
smooth in Diplolaemus, but we find they are keeled in D. darwinii, 
as first noticed by Stejneger (1909). 
A precis of the recognized species of Pristidactylus follows: 

Pristidactylus fasciatus (Dumeril and Bibron 1837) 

1837 Leiosaurus fasciatus Dumeril and Bibron, Erpet. Gen., Paris, 4: 244. — Type 

locality: not specified, specimen shipped from Buenos Aires. Restricted type 

locality (D'Orbigny, 1847): Carmen de Patagones, Buenos Aires Province, 

Argentina. 
1843 Leiosaurus ( Pristidactylus) fasciatus — Fitzinger, Syst. Rept., Wien, 1: 58. 
1845 Leiosaurus ( Ptenodactylus) fasciatus — Gray, Cat. Spec. Liz. Coll. Brit. Mus., 

London, 224. 
1885 Pristidactylus fasciatus — Boulenger, Cat. Liz. Brit. Mus., London, 2: 127. 
1896 Pristidactylus fasciatus — Koslowsky, Revta Mus. La Plata, 7: 447. 
1933 Leiosaurus fasciatus — Burt and Burt, Trans. Acad. Sci. St. Louis, 27(1): 30. 
1968 Leiosaurus bardensis Gallardo, Neotropica, Buenos Aires, 14(43): 5, unnumb. 

Fig. — Type locality: Puelen, Cochico, Altos de Chochico, La Pampa Province, 

Argentina. 
1973 Cupriguanus bardensis — Cei and Roig, Deserta, Mendoza, 4: 71. 
1978a Pristidactylus bardensis — Cei and Castro, Publ. Occ. Inst. Biol. Anim. Univ. 

Nac. Cuyo, Ser. Sci., 5: 3. 

Diagnosis. Pristidactylus fasciatus is unique within the genus in 
having multicarinate subdigital scales and the tail less than 50% of 
the total length. 

Taxonomy. The type description of fasciatus was based upon a 
juvenile specimen (42 mm snout-vent) collected on the wall of the 
fort at Carmen de Patagones in extreme southern Buenos Aires 
Province, just north of the mouth of the Rio Negro. Dumeril and 
Bibron (1837: 244) attribute the name to D'Orbigny, who collected 
the animal and for a time observed it alive. D'Orbigny (1847), in an 
expanded description with notes on the live animal, described it as a 
"charming species with all of the body a lively yellow, the top of the 
head brown, two large, transverse bands on the back of the head, six 
on the back all interrupted on the flanks. The upper part of the tail 
is likewise ringed with black, just as are the limbs." 

Dumeril and Bibron's (1837) description of fasciatus is for the 
most part applicable to juvenile individuals of all species now 
referred to Pristidactylus, except that the subdigital scales are said 
to be multicarinate, and the internal border of the foot has a dentic- 
ulation, both variable in the genus, and the tail is shorter than in 



1985 PRISTIDACTYLUS 7 

other forms. That the tail was mutilated was noted in the type 
description, but the figure provided by D'Orbigny (1847) shows an 
animal with a complete tail, and when measured on the illustration 
the tail appears to be between 0.8 and 0.9 times the snout-vent 
length. 

Boulenger (1885) listed Pristidactylus fasciatus on the authority 
of Dumeril and Bibron (1837), but inexplicably stated that the tail is 
nearly twice as long as the head and body. Koslowsky (1896) 
reported specimens under that name from Rio Negro and Neuquen, 
the latter having subdigital keels completely lacking, and later Kos- 
lowsky (1898), under the name Leiosaurus fasciatus, reported a 
juvenile from Neuquen with subdigital scales tricarinate at the 
extremities but otherwise smooth. In the latter the tail/ snout-vent 
ratio was 1.18. It seems likely that Koslowsky's specimens from 
Neuquen are referable to the form described from Laguna Blanca by 
Gallardo (1964) as Cupriguanus araucanus rather than to P. fascia- 
tus. In the former, the tail is longer than the snout-vent length and 
the subdigital scales are smooth except for the grooved distal sub- 
digitals. Koslowsky's Rio Negro specimen was not described. Kos- 
lowsky's (1898) use of the binomial Leiosaurus fasciatus was fol- 
lowed by Burt and Burt (1933) and Lieberman (1939). 

Barrio (1969) compared the types of Pristidactylus fasciatus and 
Cupriguanus araucanus Gallardo, 1964. He concluded that they 
were synonyms because in both the juvenile pattern was one of 
distinctive dark crossbands on the head, body, limbs, and tail; there 
were pectinate scales on the infradigitals of toes 1, 2, and 3; and 
measurements taken from D'Orbigny's (1847) figure yield a snout- 
vent/ tail ratio of 0.89, only slightly larger than that of C. araucanus. 
However, the distinctive juvenile markings are characteristic and 
very similar in all species of Pristidactylus and Diplolaemus, most 
other species of Pristidactylus have pectinate subdigitals on the foot, 
and according to Gallardo's (1964) figures and our own calculations 
the snout-vent/ tail length ratio in specimens from Laguna Blanca, 
the type locality of araucanus, is greater than 1 .0. Barrio noted that 
Gallardo had specifically rejected the synonomy of araucanus with 
fasciatus because of the shorter tail in the latter, and that Gallardo 
must have obtained his measurements from the illustration of 
D'Orbigny, Although Barrio directly compared the holotypes of 
both species he apparently failed to note the presence of distinctly 
multicarinate subdigitals in fasciatus, as contrasted with the smooth 



8 BRFVIORA No. 483 

subdigitals of araucanus. The synonomy of araucanus with fas ciat us 
was accepted by Cei (1973a, 1973b, 1975, 1979), Cei and Castro 
(1978a, 1978b), and Cei and Roig (1973). 

Gallardo (1968) described Leiosaurus bardensis from Cochico, La 
Pampa Province and reported additional specimens from Ishigua- 
lasto, San Juan Province, some distance north of the type localities 
of fasciatus and araucanus. He compared the species in detail with 
Diplolaemus darwinii and D. bibronii, but not with any of the 
species now referred to Pristidactylus. The species was recognized as 
Cupriguanus bardensis by Cei and Roig (1973), and as Pristidacty- 
lus bardensis by Cei and Castro (1978a, 1978b). Its distinctive fea- 
tures, i.e., multicarinate subdigital scales, pectinate subdigitals on 
the foot, and a tail shorter than head and body, are just those 
characteristics which distinguish P. fasciatus from other species of 
Pristidactylus. We have examined the types of bardensis and fascia- 
tus and a dozen other specimens from Rio Negro, Mendoza, and 
San Juan Provinces, and we find they agree in all essential details. 

Distribution and Habitat. Pristidactylus fasciatus has been taken 
at widely scattered localities between Ishigualasto (±30°10'S, 
57°50'W) at about 1,700 m altitude in the north and the type locality 
near sea level in the south. All records are within the Southern 
Chaco Province, or Monte (Cei, 1979). Cei and Castro (1978a) 
stated that in the Bermejo Basin of San Juan Province it occurs in 
areas of sand dunes. In Mendoza Province the species lives on flat, 
sandy steppe, at altitudes between 1,000 and 1,750 m, and Gallardo 
(1968) stated that at Chochico in La Pampa Province (type locality 
of bardensis) it lives in loose sand, in small burrows of Microcavia 
sp. constructed in mounds of sand that accumulate around the bases 
of Prosopis alpataco. Other species of Argentinian Pristidactylus 
appear always to be associated with rocky areas, and at the same 
latitudes occur at higher altitudes. South of La Pampa Province 
there are no specific locality records except for the type locality, but 
Koslowsky (1898) reported the species from Rio Negro Province, 
and we have seen a specimen in the Zoologiske Museum, Universi- 
tetes K0benhaven also from "Rio Negro." 

Pristidactylus scapulatus (Burmeister) 

1861 Leiosaurus scapulatus Burmeister, Reise La Plata-Staat., Halle, 2: 523. Type 
locality: Sierra de Uspallata, near Uspallata, about 5,000 ft. altitude, Men- 
doza Province, Argentina. 



1985 PRISTIDACTYLUS 9 

1861 Leiosaurus multipunctatus Burmeister, Reise La Plata-Staat., Halle, 2: 524. 

Type locality: Sierra de Uspallata, near Paramillo, about 8,000 ft. altitude, 

Mendoza Province, Argentina. 
1889 Urostrophus scapulatus Boulenger, Proc. Zool. Soc. London, 144; PI. 15, 

Fig. 2. 
1941 Leiosaurus scapulatus— Mutter, Zeitschr. f. Naturwiss, 94: 184. 
1964 Cupriguanus araucanus — Gallardo, Neotropica, B. Aires, 10(37): 129; Fig. 2, 3 

— Type locality: Laguna Blanca, Neuquen Province, Argentina. 
1964 Cupriguanus scapulatus — Gallardo, Neotropica, B. Aires, 10(37): 128. 
1976 Pristidactylus scapulatus- Etheridge, in Paull, Williams, and Hall, Breviora, 

Mus. Comp. Zool., No. 441, p. 10. 

Diagnosis. Pristidactylus scapulatus differs from P.fasciatus and 
P. casuhatiensis in having perfectly smooth subdigital scales, from 
P. achalensis in having the crowns of the posterior marginal teeth 
slightly tapered with small anterior and posterior cusps, and from 
the Chilean forms {torquatus, alvaroi, and valeriae) in having one or 
two scales interposed between the nasal and lateral postrostral 
scales. 

Taxonomy. The descriptions of Leiosaurus scapulatus and L. 
multipunctatus Burmeister, 1861 were accompanied by the descrip- 
tion of a third species, Leiosaurus marmoratus, from several locali- 
ties west of Catamarca. Boulenger (1889) stated that scapulatus 
represented the adult male, multipunctatus the adult female, and 
marmoratus the juveniles of a single species, which he referred to as 
Urostrophus scapulatus. Muller (1928) pointed out that the types of 
marmoratus belonged to the genus Phrynosaura Werner in Burger. 
1907 (subsequently transferred to Ctenoblepharis by Cei [1974] and 
then to Liolaemus [Cei, 1980]). Muller (1941) later confirmed Bou- 
lenger's synonomy of multipunctatus with scapulatus after compar- 
ing the types at the Museum in Halle. 

As presently recognized, Pristidactylus scapulatus is a species of 
the high Cordilleras and precordilleras of western Argentina in the 
provinces of Catamarca, San Juan and Mendoza (Cei, 1979). How- 
ever, we now believe that specimens from the basaltic tablelands of 
southern Mendoza, western Neuquen and southern Chubut pro- 
vinces, previously referred to P. araucanus and later to P.fasciatus (see 
above discussion under fasciatus), may be conspecific with P. scapu- 
latus. In characters of scalation and proportions, and in the color 
pattern exhibited by adult males, specimens from the cordilleras and 
precordilleras cannot be distinguished from those from the basaltic 
tablelands. In these populations as in all other forms of Pristidacty- 
lus, the bold juvenile pattern becomes progressively more obscured 



10 BREVIORA No. 483 

in males, becoming faint or fading entirely, except for the black 
antihumeral bars, in large adults. 

Adult females from the basaltic tablelands do, however, differ 
from those of the Cordilleras and precordilleras in the extent to 
which they retain the juvenile pattern. Females 98 mm snout-vent 
length and larger, from northwestern San Juan Province, and from 
the Sierra de Uspallata and the vicinity of Paso de Choique in 
western Mendoza Province usually have numerous scattered dark 
brown spots on both upper and lower surfaces, but retain only a 
faint indication of the juvenile crossbands (see Boulenger, 1889, Fig. 
2). Those from the basaltic meseta around Volcan Payun in south- 
western Mendoza Province, from the vicinity of Laguna Blanca in 
western Neuquen Province, and from Meseta Canquel in southern 
Chubut Province, also usually have scattered dark spots, but retain 
with little fragmentation or loss of contrast the bold pattern of 
juveniles. Especially prominent is the dark, horseshoe-shaped mark 
across the back of the head. Retention of a bold pattern in adult 
females is characteristic of all other species of Pristidactylus, thus 
the loss of pattern in adult females in the Cordilleras and precordille- 
ras is unique within the genus. 

A question that we have not resolved to our own satisfaction is 
how to treat this apparent step-clinal variation in the ontogenetic 
development of the female color pattern taxonomically. Our six 
populations are widely separated from one another, except for Paso 
del Choique and Payun, where the step cline is abrupt. The pattern 
of small, dark spots on a washed-out greyish background character- 
istic of the Cordilleran females, like that of the males, matches very 
well the mostly granitic rocks of their habitat, whereas the bold 
female pattern is more cryptic on the dark brown, eroded substrate 
of the basaltic mesetas. We may be dealing with relict populations 
that are now no longer in contact, and the situation appears com- 
parable to a chain of islands extending from north to south, so that 
any taxonomic decision must perforce be arbitrary. 

Serological comparisons of specimens from Uspallata, Choique, 
Payun, and Batra (near Payun) show a high degree (>84%) of homo- 
geneity (Cei and Castro, 1975). Future karyological, electrophoretic 
or immunological studies may shed light on the problem. For now, 
however, we follow a conservative nomenclature, formally recogniz- 
ing only scapulatus, although in the key below we utilize adult 



1985 PRISTIDACTYLUS 11 

female color pattern and locality to separate the cordilleran and 
precordilleran populations from those of the basaltics. If the latter 
are eventually recognized taxonomically the name P. araucanus is 
available (see discussion under P. fasciatus). 

Distribution and Habitat. The most northerly record is an 
unspecified locality in the high Cordilleras of Catamarca Province 
(Koslowsky, 1895). In northwestern San Juan Province at an alti- 
tude of 3,800 m in the Reserva Provincial de San Guillermo 
(29°18'S, 69°15'W) they are closely associated with granitic out- 
crops and live in burrows under stones or at the bases of small 
bushes, and appear to be active in the early morning and late after- 
noon. They occur in similar habitats but at lower altitudes in the 
Sierra de Uspallata (32°30-41'S, 69°00-06'W; 2,000-3,000 m) and 
the vicinity of Paso de Choique (36°25-27'S, 69°25-45'W; 1,800- 
2,000 m) in western Mendoza Province. Fewer than 50 km eastward 
of Paso de Choique they occur on the basaltic meseta dominated by 
Volcan Payiin (36°26'S, 69°16-25'W; 1,800-2,000 m), where they 
are also closely associated with stony areas and are active in the 
morning and late afternoon. Likewise they occur in similar habitats 
but at lower altitudes to the south, in the vicinity of Laguna Blanca 
(39°03-45'S, 70°23-37'W; 1,200 m) in western Neuquen Province, 
and the Meseta Canquel (44° 10-33'S, 68° 16-20'W; 900 m) in south- 
ern Chubut Province. A single specimen is known from Puerto 
Madryn on the coast in northeastern Chubut. 

Pristidactylus achalensis (Gallardo) 

1964 Cupriguanus achalensis Gallardo, Neotropica, Buenos Aires, 10(33): 132; Fig. 

4. — Type locality: Posta de Pampa de Achala, Cordoba Province, Argentina. 
1976 Pristidactylus achalensis — Etheridge in Paull, Williams, and Hall, Breviora, 

Mus. Comp. Zool., No. 441, p. 10. 

Diagnosis. Pristidactylus achalensis is unique within the genus in 
having the crowns of the posterior marginal teeth rather strongly 
compressed linguo-labially, flared in an anterior-posterior direction, 
and deeply cusped, and in having an adult male color pattern of 
uniform bright green with an immaculate yellowish belly. 

Distribution and Habitat. The species is limited to a small region 
in western Cordoba Province, from the Pampa de San Luis 
(31°20'S, 64°46'W) in the north, southward at altitudes of 2,000 to 



12 BREVIORA No. 483 

2,500 m to Cerro Champaqui (3 1°59'S, 64°56'W). It is abundant on 
the Pampa de Achala, where the habitat is a high pampa with many 
low rock outcrops that emerge from a wet meadow covered with low 
herbaceous vegetation. Numerous small sinkholes full of rain water 
dot the landscape. The bright green adult males are conspicuous 
throughout the day. Unlike other members of the genus this species 
is omnivorous, feeding on a variety of insects, including butterflies, 
and flowers (di Tada et al., 1977a and 1977b). Its diet is reflected in 
the species' unique tooth form. Gallardo (1967) reported that 130 
individuals were obtained by two collectors in a period of three 
hours on Cerro Champaqui in February. 

Pristidactylus casuhatiensis (Gallardo) 

1968 Cupriguanus casuhatiensis Gallardo, Neotropica, B. Aires, 14(43): 2; unnumb. 

fig. — Type locality: Sierra de la Ventana, Buenos Aires Province, Argentina. 
1976 Pristidactylus casuhatiensis — Etheridge in Paull, Williams, and Hall, Brevi- 

ora, Mus. Comp. Zool., Cambridge, No. 441, P. 10. 

Diagnosis. Pristidactylus casuhatiensis is unique within the genus 
in having one or two indistinct keels on most of the subdigital scales, 
the posterior marginal tooth crowns swollen, and in having an adult 
male color pattern consisting of a green background with a fine, 
dark reticulum, especially prominent on the sides; their venter is 
immaculate yellow. The adult females have a brown background 
color with obscure transverse bars. 

Distribution and Habitat. The species is known only from Cerro 
Tres Picos (38° 27'S, 62° 1 2'VV) and Cerro de la Ventana (38°08'S, 
61°47'W) in the Sierra de la Ventana, at altitudes of about 1,000 m 
in southern Buenos Aires Province. The habitat is one of a high 
pampa with flat rocks, herbaceous vegetation and small sinkholes 
full of rainwater. P. casuhatiensis feeds on beetles, spiders, and 
terrestrial snails (Plagiodontes patagonicus) (Gallardo, 1970), the 
latter perhaps correlated with its distinctive tooth form. According 
to Gallardo (1970), the populations from Cerro Tres Picos and 
Cerro de la Ventana exhibit slight differences in their characteristics. 

Pristidactylus lorquatus (Philippi) 

1861 Leiosaurus lorquatus Philippi in Philippi and Landeck, Arch. Naturgesch., 
Berlin, 27(1): 295. Type locality: in the neighborhood of Concepcion, Chile. 



1985 PRISTIDACTYLUS 13 

1861 Leiosaurus valdivianus Philippi in Philippi and Landeck, Arch. Naturgesch., 
Berlin, 27(1): 298.— Type locality: in the Province of Valdivia, Chile. 

1885 Urostrophus torquatus~Bou\enger, Cat. Liz. Brit. Mus., London, 2: 124. 

1970 Cupriguanus torquatus — Donoso-Barros, Bol. Mus. Nac. Hist. Nat. Chile, 
Santiago, 49(24): 86. 

1979 Pristidactylus torquatus — Montecinos Espinoza and Formas, Herp. Review, 
10(4): 121. 

Diagnosis. Pristidactylus torquatus differs from the Argentinian 
species in having the nasal scale in broad contact with the lateral 
postrostral, and from the Chilean species P. alvaroi and valeriae in 
having more than 140 scales around midbody, and generally smaller 
and more numerous scales overall. Some individuals have an 
inconspicuous and incomplete row of slightly enlarged middorsal 
scales, but they are not continuous on the lumbar region as is 
alvaroi. 

Distribution and Habitat. Pristidactylus torquatus occurs in for- 
ested areas of the coastal and inland Cordilleras of central Chile from 
Curico Province (35° 10'S) southward to southern Llanquihue Prov- 
ince (41°50'S), and on Isla de Chiloe (Donoso-Barros, 1966; For- 
mas, 1979; Montecinos Espinosa and Formas 1979). The species is 
arboreal, but also forages in dense brush (Donoso-Barros, 1966). 

Pristidactylus alvaroi (Donoso-Barros) new comb. 

1975 Cupriguanus alvaroi Donoso-Barros, Bol. Soc. Biol. Concepcion, 47(1974): 
221; Figs. 3-5. - Type locality: Cerro El Roble, Santiago Province, Chile. 

Diagnosis. Pristidactylus alvaroi differs from other members of 
the genus in having a markedly compressed tail, a middorsal row of 
enlarged scales continuous on the back, at least on the lumbosacral 
region, and a different color pattern. 

Distribution and Habitat. The species is known only from the 
type locality in Chile, where it occurs in relict clusters of Nothofagus 
forest at the base of Cerro El Roble, Santiago Province (32°58'S, 
71°01'W). It is said to be abundant (Donoso-Barros, 1975). 

Pristidactylus valeriae (Donoso-Barros) new comb. 

1966 Urostrophus valeriae Donoso-Barros, Rept. Chile, Santiago, 369: PI. 83. - 

Type locality: Alhue, Chile. 
1975 Cupriguanus valeriae Donoso-Barros, Bol. Soc. Biol. Concepcion, 47( 1974): 

223. 



14 BREVIORA No. 483 

Diagnosis. Pristidactylus valeriae differs from Argentinian spe- 
cies and from torquatus in having fewer, larger scales all over, with 
fewer than 140 scales around midbody. It differs from alvaroi in 
lacking a continuous row of enlarged middorsal scales on the back, 
and in color pattern, and in not having a strongly compressed tail. 

Distribution and Habitat. The species is known in Chile from 
Nothofagus forests of the Cordillera de la Costa near Alhue 
(34°09'S, 71°24'W), and from Cerro El Roble (32°58'S, 71°01'W), 
Santiago Province (Donoso-Barros, 1966). According to Donoso- 
Barros (1966) the species is not as arboreal as P. torquatus, and is 
often found in dense thickets and relict formations of Chusquea 
quila. 

Key to the Species of Pristidactylus 

la. Scales larger and less numerous, i.e., nasal scale in broad contact with lateral 
postrostral; nasal in narrow contact with anterior supralabial, or narrowly 
separated from it by contact of lateral postrostral and anteromost loreolabial; 
largest supraoculars equal to or larger than scales of the supraorbital semicir- 
cles at narrowest width of frontal region; 6 to 8 scales in a horizontal line across 
widest part of supraorbital region between superciliaries and supraorbital semi- 
circle; 8 to 16 scales bordering supralabials above Chilean species 2 

lb. Scales smaller and more numerous, i.e., one or two scales interposed between 
nasal and lateral postrostral; nasal scale well separated from anterior suprala- 
bial by one or two scales. Largest supraoculars smaller than scales of the 
supraorbital semicircles at narrowest width of frontal region; 8 to 10 scales in a 
horizontal line across widest portion of supraorbital region between supercilia- 
ries and supraorbital semicircles; 15 to 23 scales bordering supralabials above. . . 
Argentinian species 4 

2a. Scales smaller and more numerous, i.e., scales around midbody more than 140; 
scales bordering supralabials above more than 13; 13 to 22 scales across tem- 
poral region in a straight line from postoculars to anterior margin of ear 

torquatus 

2b. Scales larger and less numerous, i.e., scales around midbody fewer than 140; 
scales bordering above supralabials fewer than 13; 10 to 14 scales across tem- 
poral region in a straight line between postocular and anterior margin of ear ... 3 

3a. Tail more strongly compressed; a continuous row of enlarged middorsal scales 
on the lumbar region; dorsum greyish, venter yellow, iris red; tail conspicuously 
banded in adults alvaroi 

3b. Tail slightly compressed; a middorsal scale row absent on the lumbar region; 
dorsum bluish, venter reddish, iris blue; tail in adult not conspicuously banded; 
(antehumeral black mark absent in m&\zfide Donoso-Barros 1966). . . valeriae 

4a. Subdigital scales distinctly multicarinate; tail less than 48% total length 

fasciatus 



1985 PRISTIDACTYLUS 15 

4b. Subdigital scales smooth or with one or two weak keels; tail more than 48% 
total length 5 

5a. Subdigital scales with one or two weak keels; crowns of posterior marginal 
teeth somewhat swollen, the anterior and posterior cusps absent or only faintly 
indicated; adult male dorsal pattern a fine, dark reticulum, especially evident on 
the sides casuhatiensis 

5b. Subdigital scales perfectly smooth; crowns of posterior marginal teeth some- 
what compressed, anterior and posterior cusps moderate or small but distinctly 
present; adult male dorsum nearly uniform, a fine reticulation absent 6 

6a. Crowns of posterior marginal teeth flared, more strongly compressed, with 
larger anterior and posterior cusps; adult male uniform blue-green, green or 
yellow-green above achalensis 

6b. Crowns of posterior marginal teeth tapered, slightly compressed, with small 
anterior and posterior cusps; adult male grey or yellowish-grey above, with or 
without small, scattered dark spots scapulatus 7 

7a. Adult female color pattern obscure, typically broken into scattered dark spots, 
with dorsal cross bars faintly or not all indicated; no distinct horseshoe-shaped 
mark on the back of the head Precordilleras and Cordilleras 

7b. Adult female color pattern consisting of bold dark and light markings, a con- 
spicuous dark horseshoe-shaped mark across the back of the head; throat and 

belly usually with scattered dark spots 

Payiin, Laguna Blanca, and Canquel 

ACKNOWLEDGMENTS 

We are grateful to the following individuals for permission to 
examine specimens in their care: A. G. Kluge, The University of 
Michigan Museum of Zoology; H. W. Greene, Museum of Verte- 
brate Zoology, Berkeley; R. Sage, private collection; G. Peters and 
R. Giinther, Zoologisches Museum der Humboldt-Universitat, Ber- 
lin; J. Cranwell, Museo Argentino de Ciencias Naturales, Buenos 
Aires; P. Alberch, Museum of Comparative Zoology, Harvard Uni- 
versity; S. W. Braestrup, Zoologiske Museum, Universitetes K0ben- 
haven; T. Cekalovic, Instituto de Zoologia, Universidad de Concep- 
tion; K. Klemmer, Natur-Museum Senckenberg, Frankfurt-am- 
Main; W. Ladiges and H. Koepcke, Zoologisches Museum, 
Hamburg; J. D. Williams, Museo de La Plata; W. E. Duellman, 
Museum of Natural History, University of Kansas; A. G. C. Gran- 
dison and E. N. Arnold, British Museum (Natural History); L. Cas- 
tro, Instituto de Biologia, Universidad Nacional de Cuyo, Mendoza; 
R. Braun, Instituto Argentino de Investigaciones de las Zonas Ad- 
das, Mendoza; F. Achaval, Universidad de Uruguay; W. Hellmich 
and U. Gruber, Zoologisches Sammlung des Bayerischen Staates, 



16 BREVIORA No. 483 

Mtinchen; J. Guibe, Museum National d'Histoire Naturelle, Paris; 
C. McCoy, Carnegie Museum of Natural History; A. Leviton, Cali- 
fornia Academy of Sciences; H. La Gilia, Museo Historia Natural 
de San Rafael; R. Laurent, Fundacion Miguel Lillo, Tucuman; P. 
E. Vanzolini, Museu de Zoologia da Universidade de Sao Paulo; U. 
Parenti, Museo Civico de Storia Naturale, Torino; J. Eiselt and F. 
Tiedeman, Naturhistorisches Museum, Wien; and G. Zug, National 
Museum of Natural History, Washington. 

Funds for travel in the United States and field work in Argentina 
were provided by a grant from the National Geographic Society 
Committee on Research and Exploration. For providing laboratory 
facilities and access to field vehicles we wish to thank Eng. Rolando 
Braun, Director of the Instituto Argentino de Investigaciones de las 
Zonas Aridas, Mendoza, and Dr. Oswald H. Sala, Director of the 
Centro Nacional Patagonico in Puerto Madryn. We also thank Don 
Eleodoro A. Sanches, Jefe Division de la Fauna of San Juan Prov- 
ince for permission to work in the Reserva Provincial de San 
Guillermo. 

We especially wish to express our gratitude to Dr. Jose M. Cei for 
his tireless efforts to make our field work in Argentina a success, 
and to Fernando Videla and Jorge D. Williams for their assistance 
in the field. 



LITERATURE CITED 

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xiii + 497 pp. 
. 1889. On some specimens of lizards in the Zoological Museum of Halle 

(Saale). Proc. Zool. Soc. London, 1889(11): 143 145. 
Burger, O. Estudios sobre reptiles chilenos. An. Univ. Chile, Santiago, 121(2): 

147-155. 
Burmeister, H. 1861. Reise durch die La Plata Staaten mit besonderer Ruck- 

sicht auf die physische Beschaffenheit und den Culturzustand der Argentinische 

Republik. Ausgefurt in den Jahren 1857, 1858, 1859 und 1860. Halle, 2: iv + 538 

pp. 
Burt, C.A., and M.D. Burt. 1933. A preliminary checklist of the lizards of 

South America. Trans. Acad. Sci. St. Louis, 28(1 2): 1-104. 
Cei, J. M. 1973a. Comentarios sobre algunos generos de iguanidos: Diplolaemus, 

Leiosaurus. Aperopristis y Cupriguanus. Physis, Buenos Aires, Sec. C. 32(85): 

269-276. 



1985 PRISTIDACTYLUS 17 



1973b. Distribucion geografica y caracteres poblacionales de Cuprigua- 



nus fasciatus (D'Orbigny) (Sauria, Iguanidae). Physis, Buenos Aires, Sec. C, 
32(85): 255-262. 

— . 1974. Two new species of Ctenoblepharis (Reptilia, Iguanidae) from the 
arid environment of the central Argentina, Mendoza Province. J. Herp., 8(1): 
71-75. 

— . 1975. The present taxo-serological works in the field of herpetology at 
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. 1979. The Patagonian herpetofauna, pp. 309 339. In W. E. Duellman, 

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1980. Remarks on the taxonomic status and specific characters of Lio- 



laemus tnarmoratus (Burmeister). J. Herp., 14(2): 192-193. 
Cei, J. M., and L. P. Castro. 1975. A serological contribution to the taxonomic 

status of Cupriguanus, a South American genus of iguanid lizards. Serol. Mus. 

Bull., New Brunswik, 51: 5-6. 
. 1978a. Datos preliminares sobre las componentes de la herpetofauna de 

la Provincia de San Juan. Pubis Occ. Inst. Biol. Anim., Univ. Nac. de Cuyo, 

Mendoza, Ser. Cien., 5: 1-4. 

1978b. Atlas de los vertebrados inferiores de la region de Cuyo. Pubis 



Occ. Inst. Biol. Anim., Univ. Nac. de Cuyo, Mendoza, Ser. Cien., 2: 1-38. 

Cei, J. M., and G. V. RoiG. 1973. Fauna y ecosistemas del Oeste arido argentino. 
I. Reptiles de la Provincia de Mendoza. Deserta, Mendoza, 4: 69-91. 

di Tada, I., R. Martori, B. Kutner, and A. Ocana. 1977. Reproductive cycle 
and feeding behavior of Cupriguanus achalensis (Sauria:Iguanidae). Herp. Rev., 
8(3), suppl. p. 6. 

di Tada, I., R. Martori, B. Kutner, A. Ocana, and M. Beatriz. 1977. Herpe- 
tofauna endemica de la Pampa de Achala (Cordoba, Argentina) I Reun. Ibero- 
amer. Zool. Vert. La Rabida, 1977: 493-512. 

Donoso-Barros, R. 1966. Reptiles de Chile., Santiago, xclvi + 458 pp. 

. 1970. Catologo herpetologico chileno. Bol. Mus. Nac. Hust. Nat. Santi- 
ago, 31: 49 124. 

. 1975. Nuevos reptiles y anfibios de Chile. Bol. Soc. Biol. Concepcion, 



48(1974): 217-229. 

D'Orbigny, A. 1847. Voyage dans l'Amerique meridionale. . .pendant les annees 
1862. . . 1833. Paris, 5(1): 1-12; Pis. 1-15. 

Dumeril, A. M. C, and G. Bibron. 1837. Erpetologie generale ou Histoire 
Naturalle complete des reptiles. Paris, 4: 1-572. 

Fitzinger, L. 1843. Systema Reptilium. Fasciculus primus, Amblyglossae. Wien, 
vi + 106 pp. 

Formas, J. R. 1979. La herpetofauna de los bosques temperados de Sud Amer- 
ica, pp. 341 369. In W. E. Duellman (ed.), The South American herpetofauna: 
its origin, evolution and dispersal. Mus. Nat. Hist. Univ. Kansas, Monog., 7: 
1-485. 

Gallardo, J. M. 1964. Los generos "Urostrophus" D. et B. y "Cupriguanus" 
gen. nov. (Sauria, Iguanidae) y sus especies. Neotropica, Buenos Aires, 10(33): 
125-136. 



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. 1967. A proposito del genero Cupriguanus Gallardo (Sauria, Iguani- 

dae). Neotropica, Buenos Aires, 13(41): 56. 

. 1968. Dos nuevas especies de Iguanidae (Sauria) de la Argentina. Neo- 
tropica, Buenos Aires, 14(43): 1-8. 

. 1970. Interesante hallazgo en Sierra de la Ventana. Camping, Buenos 

Aires, 12: 67-68. 

. 1979. Composition, distribution y origen de la herpetofauna Chaquena, 



pp. 299-307. In W. E. Duellman (ed.). The South American herpetofauna: its 

origin, evolution and dispersal. Mus. Nat. Hist. Univ. Kansas., Monog., 7: 

1-485. 
Gray, J. E. 1845. Catalogue of the specimens of lizards in the British Museum. 

London, xxviii + 289 pp. 
K.SOLOWSK.Y, J. 1895. Batrachios y reptiles de Rioja y Catamarca (Republica 

Argentina) recogidos durante los meses de Febrero a Mayo de 1895. Rvsta Mus. 

La Plata, 6: 357-370. 
. 1896. Sobre algunos reptiles de Patagonia y otras regiones argentinas. 

Rvsta Mus. La Plata, 7: 447-457. 
. 1898. Enumeration sistematica y distribution geografica de los reptiles 



argentinos. Rvsta Mus. La Plata, 8: 161-200. 
Liebermann, J. 1939. Catalogo sistematico y zoogeografico de los lacertillios 

argentinos. Physis, 16(48): 61-82. 
Montecinos Espinosa, R. M., and J. R. Formas. 1979. Distribution of the 

Chilean lizard Pristiciactylus torquatus. Herp. Rev., 10(4): 121-122. 
Moller, L. 1928. Herpetyologische Mitteilungen, Zool. Anz. 77: 61-84. 
. 1941. Uber die in der Sammlungdes Zoologischen Institutes der Univer- 

sitat Halle a. Saale aufbewahrten Amphibien und Retilientypen. Zeitschr. f. 

Naturwiss, 94: 182-205. 
Paull, D., E. E. Williams, and W. P. Hall. 1976. Lizard karyotypes from the 

Galapagos Islands: chromosomes in phylogeny and evolution. Breviora, Mus. 

Comp. Zool., No. 441, pp. 1-31. 
Philippi, R. A., and L. Landeck. 1861. Neue Wirbelthiere von Chile. Arch. 

Naturgesch., Berlin, 27(1): 289-301. 
Stejneger, L. 1909. Batrachians and Reptiles, Part II, pp. 211-224. In W. B. 

Scott (ed.), 1905-1911; Reports of the Princeton University Expeditions to 

Patagonia, 1896-1899, Vol. Ill, 1, Zoology, Princeton, xiii + 374 pp. 



B R E u »iKI R A 

Museum of Comparative Zoology 



US ISSN 0006-9698 



Cambridge, Mass. 21 June 1985 Number 484 

MALE AGGRESSIVE BEHAVIOR IN A PAIR 
OF SYMPATRIC SIBLING SPECIES 

Jonathan B. Losos 1 

Abstract, lntraspecific encounters were staged between adult male Anolis mar- 
canoi and A. cyhoies in order to describe their aggressive behavior. The major 
component of stationary behavior is the dewlap display, accompanied by various 
static and dynamic modifiers. Advancing behaviors include moving toward an oppo- 
nent, threatening attack, and actual biting. The agressive behavior of A. marcanoi is 
progressive and hierarchically structured. By contrast, the behavior of A. cyhoies is 
less elaborate and ritualized than that of its sibling and lacks progressive structure. 

INTRODUCTION 

Defined simply as "pairs or even larger groups of related species 
which are so similar that they are considered as belonging to one 
species until a more satisfactory analysis clears up this mistake," 
(Mayr, 1942) sibling species have been discovered at an increasing 
pace as the level of systematic analysis has become finer, passing 
from examinations of morphological characters to investigations of 
molecular properties. 

The different patterns of distribution of sibling species raise inter- 
esting ecological questions. On one hand, sibling species may be 
completely allopatric, which raises questions more about the evolu- 
tionary forces operating to produce such species than about the 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. Present Address: Museum of Vertebrate Zoology and Department of 
Zoology. University of California. Berkeley, California 94720. 



2 BREVIORA No. 484 

ecological relationships between them. On the other hand, parapat- 
ric and sympatric sibling species distributions bear directly on eco- 
logical theory. In theory, two species occupying the same niche 
cannot long coexist unless differences sufficient to lessen competi- 
tive pressures evolve. The parapatric ranges of many sibling species 
pairs or groups attest to the difficulty such species have in coexisting 
(e.g., the members of the Rana pipiens complex in North America 
[Pace, 1974] and the Anolis chlorocyanus [Williams, 1965] and A. 
ricordii [Schwartz, 1974] groups in Hispaniola). One would expect 
that sympatric sibling species would exhibit differences in mor- 
phology, physiology, or behavior that would permit coexistence. 
Indeed, resource partitioning of some kind is observed in almost all 
known cases of sympatric sibling species (Mayr, 1942; Schoener, 
1977). 

In this regard, the curious case of the Hispaniolan sibling species 
Anolis cybotes and A. marcanoi warrants attention. Anolis cybotes, 
the larger of the two, is found throughout the island, while A. 
marcanoi is restricted to a small area in the southwest portion of the 
Dominican Republic (Williams, 1975). The two are classic sibling 
species, consistently separable morphologically only by the color of 
the dewlap of males and the throat of females: Anolis marcanoi has 
a red dewlap or throat, while that of A. cybotes is usually either 
white or yellow, depending on locality (Williams, 1975). Electro- 
phoretic studies by Webster (1975), however, clearly reveal that the 
two are distinct species. Table 1 provides a review of differences 
between the species. 

Ecologically, the species are just as similar. Unlike the cases of 
sibling species already discussed, no apparent differences in physi- 
ology or microhabitat preference have been discovered. They are 
patchily sympatric throughout the range of A. marcanoi and have 
been noted on adjacent fenceposts in some areas (Hertz, 1980), 
while in other locations only one species is present. Hertz suggests 
that A. marcanoi may be better adapted for hotter, more open 
microhabitats, but no consistent differences have been noted in the 
field (Williams, 1975; Hertz, 1980; personal observation). 

At issue is the manner in which these species interact. Are the 
differences between the species in dewlap color sufficient to allow 
species recognition, or are there also behavioral differences of 
importance? If species recognition is highly accurate, do the similar- 



1985 



ANOUS SIBLING SPECIES BEHAVIOR 



Table I. Differences between Anolis marcanoi and Anolis cvhotes* 



A. cvhote. 



A. marcanoi 



range 
dewlap color 

scale characters 

maximum snout- 
vent length 

heat tolerance: 
Experimental 
Voluntary 
Maximum 

Critical Thermal 
Maximum 



widespread throughout 
Hispaniola 

normally white to yellow, 
except some populations on 
the extreme end of the 
southwestern peninsula in 
Haiti, which are red 

middorsal and midsacral 
scales sometimes enlarged 

81 mm 



34.3° C 



Peravia Province, south- 
west Dominican Republic 

red 



middorsal and midsacral 
scales rarely enlarged 

65 mm 



38.4° C 



35.7° C 



40.5° C 



*After Williams. 1975 and Hertz, 1980. 

ities between the species lead to resource competition and interspe- 
cific aggression? The goal of this study is to provide a thorough 
description of intraspecific male aggressive behavior in the two spe- 
cies. In themselves, reports on the intraspecific male aggressive 
behavior of both species are important because detailed characteri- 
zations of the components and progression of male aggressive 
behavior in Anolis are not common (the only reports of comparable 
detail are Greenberg and Noble, 1944, and Greenberg, 1977, on A. 
carolinensis and Jenssen 1979a and 1979b on A. opalinus). Furth- 
ermore, in order to analyze interspecific interactions, it is first neces- 
sary to understand the dynamics of intraspecific behavior. This 
knowledge then may be used as a control with which interspecific 
aggressive behavior can be compared. 

MATERIALS AND METHODS 

In January 1983, adult male A. marcanoi were collected past the 
first ford on the El Recodo road, approximately 6 to 10 km north of 



4 BRFVIORA No. 484 

Bani, Peravia Province, Dominican Republic. Adult A. cybotes 
were collected at that locality and in Santo Domingo, Dominican 
Republic. The lizards were housed in terraria in the Biological 
Laboratories, Harvard University, and provided with perches, small 
potted plants, and a soil substrate, and offered several crickets one 
to two times a week and water three to four times a week. Room 
temperature was maintained at approximately 26 to 30°C. Individu- 
als were separated from other males in the same terrarium by card- 
board partitions. 

Encounters between two conspecific males were staged between 23 
March and 16 December 1983. Lizards were placed in the parti- 
tioned halves of either a 41 X 22 X 20 cm (the first 14 A. marcanoi 
encounters) or a 74 X 32 X 29 glass terrarium (the last 12 A. marca- 
noi and all 19/1. cybotes encounters) covered with a mesh lid and 
provided with perches in each half of the terrarium. Lizards were 
left undisturbed in the terrarium for a period ranging from 17 hours 
to four days, but usually lasting two days. Water was only provided 
when an encounter had to be postponed, forcing the period to 
exceed two days. Lizards were allowed to acclimate in order to gain 
familiarity and enhance territorial defense. 

Approximately 15 to 30 minutes prior to an encounter, a 250 W 
infrared light was switched on, warming the terrarium to 25 to 32°C 
and markedly increasing the activity level of the lizards. Tempera- 
ture in the room in which the encounters were staged was approxi- 
mately 20° C. 

Encounters were initiated by lifting the partition. All lights in the 
room were extinguished, except two 60 W desk lamps pointed 
directly into the terrarium which lit the display arena and made it 
difficult for the lizards to see observers in the room. Encounters 
were videotaped on a Panasonic NV-8050 time lapse recorder with a 
MTI-65 video camera and a Canon 16-100 mm TV zoom lens and 
then analyzed at normal and slow-motion speeds on a Panasonic 
WV-5350 video monitor. Encounters normally lasted 45 minutes, 
but were terminated when one lizard tried repeatedly to escape from 
the other, and were extended when the lizards were still intensely 
interacting at the end of the alloted time. 

Twenty-one A. marcanoi and ten A. cybotes were used in these 
encounters. Lizards were paired randomly, with the constraints that 
they had not been housed in the same terrarium, that they had no 



1985 ,4 AW/VS SIBLING SPECIES BEHAVIOR 5 

prior experience with each other (two exceptions), and that there 
was no more than a 5 mm difference in snout-vent-length. 

RESULTS 

The aggressive behavior of the lizards can be divided into two 
categories: stationary and advancing behavior. In stationary behav- 
ior, the lizard remains in one area and primarily moves its head, 
bobbing with or without extension of the dewlap. Several modifiers 
of this display, either morphological changes or movements of body 
parts, are correlated with aggressive intensity, particularly in A. 
marcanoi. The level of intensity of A. marcanoi could be judged by 
the correlated appearance of behavioral and morphological re- 
sponses. As displays progressed, lizards generally displayed a 
sequence of actions and display modifiers, adopted a more threaten- 
ing posture with body and head raised and apparent size maxim- 
ized, and became more active. By contrast, in A. cybotes there was 
little correlation between the appearance of particular aspects of 
aggressive behavior. Thus, level of intensity was more difficult to 
gauge. 

In advancing behavior, the lizard moves toward its opponent. In 
the early, less intense stages, the lizard makes various intention 
movements, while later stages involve lunging and biting. 

The male aggressive behavior of the two species differed in two 
major respects. First, though elements of both stationary and 
advancing behavior were displayed by A. cybotes, many of the 
components of these behaviors were absent or much less elaborate 
than those exhibited by A. marcanoi. Second, the ordered appear- 
ance of progressively more aggressive behaviors, so integral a part 
of male-male interactions of A. marcanoi, was absent in the behav- 
ior of A. cybotes. In most cases, particularly among stationary be- 
haviors, any particular behavior did not necessarily follow any 
other particular behavior, nor did particular behaviors connote dif- 
ferent levels of aggression. 

Stationary Behavior 

The primary stationary display of the lizards was the dewlap 
display, composed mainly of head-bobbing and dewlap extension. 



6 BRFVIORA No. 484 

Dewlap Display 

A no/is marcanoi 

There were generally three levels of intensity of the display exhi- 
bited by A. marcanoi. Intensity is best correlated with the degree to 
which the lizard elevated its body above the substrate. 

The low level dewlap display, often the first display the lizard 
would make in an encounter, involved little body elevation. The 
body occasionally remained completely in contact with the sub- 
strate, but usually the anterior portion of the body was slightly lifted 
by extending the front limbs, holding them out to the side in the 
typical reptilian stance (Fig. la). The result was that the forequar- 
ters were raised several mm off the substrate. The snout was also 
angled upward at 30 to 45 degrees to the ground. The head usually 
was kept in the vertical plane of the body. During the process of 
raising the body and snout, the dewlap usually was extended com- 
pletely with several rapid head bobs. On a number of occasions, the 
forebody distinctly raised first, and then the dewlap extended, but 
the two usually occurred simultaneously. 

This display occurred primarily at the start of an encounter, 
shortly after the partition had been raised, though sometimes it was 
omitted altogether. Occasionally, when an encounter had proven 
indecisive and the lizards had calmed, they would revert to this level. 

The intermediate dewlap display involved an increase in body 
elevation. Generally, it took one of two forms: either the forelimbs 
were completely extended with the snout pointed at a very high 
angle (75 to 90 degrees), giving the whole body a very steeply sloped 
configuration, or all four legs pushed up, with the hind legs out to 
the side, elevating the whole body off the substrate (Fig. lb). The 
tail was then usually also lifted, either held rigid and completely 
elevated, or slightly arched with the posterior half dropping to the 
substrate. The tail was also on occasion held higher than the body or 
was even curled back above it. The dewlap was extended completely 
in this display; the snout's higher angle and the higher elevation of 
the forequarters (at least a little higher in the second variation than 
in the low level dewlap display) allowed the dewlap to be lifted 
completely off the substrate. 

In the highest level of intensity display, clearly distinct from the 
first two, the entire body was elevated high off the substrate (Fig. 
lc). In the full elevation display, the forelimbs were held under the 



1985 



A NOUS S1BI ING SPECIES BEHAVIOR 




(b) 





Figure I. Male Anolis marvanoi performing (a) low-level, (b) intermediate, and 
(c) full elevation displays. Display modifiers shown here, such as crest erection and 
head spot darkening, were noted in all three display levels, through they were more 
common in the more aggressive levels. 



8 BREVIORA No. 484 

body and almost completely straightened, while the hindlimbs were 
pushing up greatly either sprawled out to the side or also under the 
body. The tail was usually rigidly held straight back or even held 
higher than the body. The snout was very high, and the dewlap was 
well off the substrate. 

Several particular aspects of the display varied independently of 
these levels, though in some cases relationships were noted. The 
most variable, and perhaps most significant, component of the dis- 
play involved the dynamics of dewlap extension. In general, the 
degree of extension varied as a function both of time elapsed and 
intensity of the encounter. The dewlap was usually completely 
extended and maintained or pulsed (retracted and reextended) to a 
barely perceptible extent at the beginning of encounters and when a 
high level of intensity was observed. As encounters progressed into 
series of alternating exchanged displays, dewlap extension tended to 
decrease both in extent and duration. Often, the dewlap would only 
be pulsed out briefly at the end of a series of bobs; the amount of 
dewlap extended varied similarly, sometimes only revealing a slight 
bit of the dewlap, and sometimes not at all. If intensity increased, 
for example as a result of an advance by one of the lizards, dewlap 
extension would increase in subsequent bobbing. 

Anolis cy botes 

Anolis cybotes'on\y has one level of dewlap display, comparable 
to the low level dewlap display of A. marcanoi. A lizard displayed 
with its posterior half in contact with the substrate, while its anterior 
was raised a variable amount by extension of the forelegs (Fig. 2), 
either out to the side or directly under the body. The snout also was 
angled upward to a variable degree. On rare occasions, usually when 
it was on the ground, a lizard displayed with all four legs out to the 
side, pushing its body up off the ground. 

The dewlap was usually extended, at least in the initial bout of 
displaying, with concurrent head-bobbing. The amplitude, number, 
and even presence of bobs varied. On occasion, usually later in an 
encounter, the dewlap was extended and retracted without any head 
movement at all. As with A. marcanoi, sometimes the dewlap was 
extended and then retracted, while other times it was maintained at 
full extension. 



1985 ANOUS SIBLING SPECIES BEHAVIOR 




Figure 2. Typical display posture of male Anolis cyboies. The forequarters are 
elevated, but the hindquarters are firmly planted on the substrate. The nuchal crest is 
occasionally erected more fully, and the dorsal crest is sometimes apparent. The 
dewlap often is extended more fully. 

Head Bobbing 

Anolis marcanoi 

Though the bobs of the head usually were part of the dewlap 
extension process, similar to the fanbob of A. aeneus (Stamps and 
Barlow, 1973), in later stages of the encounter, especially when the 
lizards were alternating displays, the bobs were increasingly empha- 
sized and independent of dewlap extension. A series of 2 to 10 bobs, 
either of the normal amplitude or greatly exaggerated with several 
times that amplitude, were performed without dewlap extension, or 
as a distinct precursor to extension. At other times, the dewlap was 
pulsed out with exaggerated jerking bobs, similar to the jerkbob of 
A. aeneus. The lizards sometimes bobbed several times at full dew- 
lap extension, with a slight pulse of the dewlap during each bob and 
slight retractions in between. This usually occurred when the display 
intensity was high. In several cases, a subordinate lizard raised and 
lowered its head extremely slowly with large amplitude and no dew- 
lap extension. This was usually repeated several times. 

Jenssen (1983) found that A. cybotes performed only one stereo- 
typed head bobbing pattern, in contrast to the greater repertoires of 
other Anolis (Jenssen, 1977, 1978). A detailed investigation was not 
conducted, but from an analysis of the diplays of several lizards, it 
appears that A. marcanoi has at least two distinct display types. 
However, neither the extent of variation nor the degree of stereo- 
typy of the head-bobbing patterns were determined. It is possible 



10 



BREVIORA 



No. 484 



also that A. marcanoi has several other head-bobbing patterns that 
were not discovered. No relationship between level of dewlap dis- 
play and head-bobbing pattern was found. 

In the first pattern, equivalent to the signature display (Stamps 
and Barlow, 1973; Jenssen, 1978), the head was bobbed once or 
twice with great amplitude, often with a pause in between bobs, and 
then rapidly bobbed another 7 to 17 times with the snout remaining 
elevated (Fig. 3). The dewlap began to expand between bobs six to 
nine, at which time the bobbing would begin to slow. Several times 
this display was prefaced with a lowering and raising of the head, 
akin to, though slower than, the "head-dipping" 1 of A. cybotes 
(Jenssen, 1983). Jenssen (personal communication) pointed out the 
close similarity between this display pattern and the sole one 
reported for A. cybotes. Only the number of initial large-amplitude 
bobs — one or two in a A. marcanoi, three in A. cybotes — 
distinguishes the two, attesting to the close relationship of the two 
species. This pattern was observed most often early in encounters, 
though it was also seen sometimes at high levels of intensity. This 
description must be considered tentative, however, because it is 
based on only four displays of three lizards. 

A second display, probably functioning as a challenge display 
(Jenssen, 1978), was often seen late in encounters, especially when 
lizards were fairly close to each other and alternating displays. It 
was characterized by a large amplitude dip of the head, a pause, and 
then three or four rapid low amplitude bobs with the head not quite 




Figure 3. Sample display-action pattern graph of the first head-bobbing pattern 
ot Anolis marcanoi. The line represents approximate elevation through time. The 
shaded region represents dewlap extension through time. This pattern was sometimes 
preceded by a dip of the head. 



1985 A NOUS SIBLING SPECIES BEHAViOR I 1 

dropping back to its original level (Fig. 4). Sometimes, there was a 
pause after the bob, with the head often not returning to its original 
level. Usually the dewlap was brought out only at the end of the 
bobbing sequence, if at all. This description is based on an analysis 
of nine displays by four lizards. It did seem to be fairly stereotyped, 
however, and was observed in the displays of many of the other 
lizards. 

Intensity was reflected in the length and magnitude of dewlap 
display and head-bobbing. More intense encounters tended to have 
longer displays with greater number of bobs and pulses and greater 
dewlap extension. The pace of the displays was also faster in more 
intense displays. If these intense encounters did not immediately 
progress into more direct aggressive action, however, displays 
tended to become shorter and less animated, often devolving into 
the alternating bouts of medium speed bobbing mentioned above. 

Anolis cybotes 

Because Jenssen has already extensively examined the stereo- 
typed head-bobbing patterns of A. cybotes, they were not investi- 
gated here. Two distinct methods of bobbing were noted in A. 
cybotes. In one, by far the more common in A. marcanoi, the whole 
head was bobbed, passively moving the dewlap along with it, but 
without greatly changing its amplitude. This method of bobbing 
produces the head-bobbing display action patterns so extensively 
studied in Anolis. 




Figure 4. Sample of the second head-bobbing pattern of Anolis marcanoi. 
Though not indicated here, the dewlap was occasionally extended at the end of the 
display. 



12 BRF.VIORA No. 484 

By contrast, dewlap bobbing — rarely exhibited by A. marcanoi, 
but commonly performed by A. cybotes — primarily involved mov- 
ing the dewlap up-and-down by raising and lowering the posterior 
portion of the hyoid, causing a large amplitude change in the dew- 
lap, but moving the head only slightly. This bobbing type was usu- 
ally preceded by the first type and appeared when the lizard was 
displaying intensely. Only the first type was involved in dewlap 
extension. 

The quick inverted head bob or "head dip" noted in wild A. 
cybotes by Jenssen was observed on a number of occasions. His 
observation that head dips were often performed independently of 
dewlap displays was confirmed, though no long series of head dips, 
which Jenssen also reported, were noted in these experiments. 

Sialic Modifiers 

Several morphological responses — Jenssen's static modifiers 
(Jenssen and Hover, 1976; Jenssen, 1977, 1978) — were also indica- 
tive of level of intensity, particularly in A. marcanoi. 

Anolis marcanoi 

Both the nuchal and dorsal crests were often erected during dis- 
plays (Fig. lb, and lc). Often, they were erected, retracted, and 
erected again several times in an encounter. The nuchal crest always 
appeared before and disappeared after or simultaneously with the 
dorsal crest. The crests often at first were erected fully, but then 
retracted to only a fraction of their full size as the encounter 
continued. 

The presence of fully erected crests indicated a high intensity 
state, but their absence did not imply the converse. When the dis- 
tance separating the lizards was not great, actions more aggressive 
than the full elevation display were generally accompanied by full 
crest erection, as were most full elevation and some intermediate 
elevation displays. Crests were noted less frequently when the 
lizards were displaying from across the large terrarium. Lizards that 
appeared clearly subordinate — indicated by display level, activity, 
posture, and, retrospectively, by the outcome of the encounter (see 
below) — were much slower to raise their crests and maintained them 
generally for a shorter period. 



1985 ANOUS SIBl INC. SPECIES BEHAVIOR 13 

Similarly, the appearance of a dark spot on the side of a lizard's 
head correlated broadly with intensity. Between the eye and the 
tympanic opening, a black circular spot slightly larger than the 
tympanic opening would appear when a lizard was aroused (Fig. 
1c). A similar well-defined dark patch with a similar function has 
been reported for A. carolinensis (Greenberg and Noble, 1944; 
Greenberg, 1977), but in that species the patch is rectangular and 
directly postorbital without intervening undarkened scales. The 
appearance of the spot in both species seems clearly a function of 
arousal; lizards handled during transfer to and from the experimen- 
tal arena almost invariably exhibited it and often tried to bite. Many 
other Anolis display an irregular darkening in that region when 
aroused (G. C. Mayer, personal communication; personal observa- 
tion). Dominant lizards were much more likely to exhibit the spot 
than subordinate ones, which rarely displayed it except at the high- 
est levels of intensity, and only then when putting up resistance 
instead of fleeing. As with the nuchal and dorsal crests, black spots 
were usually apparent, especially on dominant lizards, at high levels 
of intensity, and sometimes at lower levels, ranging from low level 
dewlap displays to full elevation displays. Darkening of the spot 
almost invariably was preceded by erection — though not necessarily 
full erection — of the crests. As with crest erection, the spot appeared 
much more frequently when the distance separating the lizards was 
not great. 

Body orientation and appearance also were important compo- 
nents of the display. Lizards, especially dominant ones, increased 
their apparent body size several ways. The apparent size of the head 
was increased by erecting the nuchal crest, engorging the head, and 
lowering the hyoid apparatus of the throat. Such enlargement is a 
common aggressive response in iguanids (Greenberg and Noble, 
1944; Carpenter, 1967). Presumably, dewlap extension has the same 
effect of making the lizard appear larger. When retracted, the dew- 
lap remained apparent, ranging in size from a slight rim to a fairly 
large crescent, making the head region appear larger. The body was 
also made apparently larger by expansion of the dorsal crest and 
lateral compresion of the body. Subordinate lizards, as well as dom- 
inant lizards displaying after winning an encounter (indicated by 
quiescence or flight of their opponents) rarely attempted to increase 
their apparent size. 



14 BRFVIORA No. 484 

Lizards also attempted to assume a position that would expose 
the greatest part of their bodies to the other lizard, increasing their 
apparent size. When both lizards were displaying, the most common 
position was a parallel alignment, each lizard exposing its broadside 
to the other. As with many other Anolis (e.g.. Carpenter, 1965, 
1967), when not aligned parallel, one or both lizards (usually the one 
with less apparent broadside exposure to the other) would turn its 
head, so that the dewlap when extended would be parallel to the 
other's head and thus appear as large as possible. Maximizing 
apparent size was most obvious when one lizard was on a perch 
while the other was nearby on the ground. In these instances, dis- 
playing in an upright position on the branch would not present as 
large an image as possible to the lizard on the ground. Invariably, 
the lizard on the branch would display on the side of the branch. 
The closer the lizard on the ground was to the branch, the greater 
would be the angle from the vertical of the lizard on the branch. 
When the lizard on the ground was directly underneath the branch 
of the other lizard, the latter would arrange itself horizontally, at a 
90° angle from the upright. By contrast, a lizard displaying to 
another lizard also on a perch would always display directly upright 
on the perch. 

Anolis cybotes 

As with A. marcanoi, several morphological responses were noted 
as part of the male-male interactive behavior of A. cybotes. In the 
latter species, however, lack of a particular static modifier — or of all 
static modifiers — did not necessarily correlate with low levels of 
intensity. 

Like A. marcanoi, A. cybotes possesses both a nuchal and a 
dorsal crest. The crests appear to be smaller in A. cybotes and are 
fully erected much less frequently. Often, the dorsal crest was visible 
only as a low ridge along the lizard's back. Crest erection appears to 
serve the same function in both species, increasing apparent size and 
indicating heightened level of intensity. In the eight encounters in 
which dominance could be determined (with A. cybotes, dominant 
lizards could only be determined post facto; dominance could not be 
determined by the presence of hierarchically higher displays and 
modifiers as it could with A. marcanoi), five of the dominant lizards 
displayed crest erection. In two of the three other instances, no other 






1985 4 /Vtf/./S SIBLING SPECIES BEHAVIOR 15 

preliminaries to a direct attack occurred. Only two of the eight 
subordinate lizards displayed crest erection, both in encounters in 
which the dominant lizard also displayed crest erection. 

Anolis cybotes does not have a well-defined circular head spot 
like that of A. marcanoi. It can, however, change skin hue (as can A. 
marcanoi); often, dominant lizards became lighter. An ill-defined 
darkening of the skin in the postorbital region of the head, where 
the dark spot in A. marcanoi is located, was noted on several occa- 
sions, usually when the lizard was aroused by another male or was 
handled. Only in one lizard, however, was anything approaching a 
circular spot apparent. 

As with A. marcanoi, A. cybotes attempted to increase its appar- 
ent size in various ways. This was observed more often in dominant 
lizards, though in some cases both lizards attempted to increase 
their size. Apparent size was increased in several ways, all also 
exhibited by A. marcanoi. Most common, other than dewlap exten- 
sion, was enlargement of the head. Lizards also increased their 
apparent size by lateral compression and throat enlargement. 

Dynamic Modifiers 

Several display modifiers involved moving body parts and thus 
constituted dynamic modifiers (Hover and Jenssen, 1976; Jenssen 
1977, 1978). Instead of bobbing with the head, on several occasions 
lizards raised and lowered the body by pushing up with the legs, 
primarily the forelegs. Push-ups have been widely reported among 
Anolis (e.g., Greenberg, 1977; Jenssen, 1979a), and iguanids in gen- 
eral (Carpenter, 1967), but no correlation with intensity was disco- 
vered in A. marcanoi or A. cybotes. 

Anolis marcanoi 

The full elevation display was the most active level of dewlap 
display. The rear of the body was moved independently in several 
ways. Most common were push-ups with the rear legs, usually in 
series of two to six causing the posterior to oscillate greatly, similar 
to "rearing" observed in A. opalinus (Jenssen, 1979a). The precur- 
sor of these pelvic push-ups could be seen in several intermediate 
dewlap displays in which the rear legs were raise and lowered 
slightly as the dewlap was thrust out during head-bobbing. In the 
full elevation display, the lizards also sometimes jumped backwards 



16 BRFVIORA No. 484 

several times in rapid succession, forcing the posterior of the body 
up-and-down in a manner similar to that caused by hind-leg push- 
ups. Similar behavior in the same context has been noted in male A. 
lineatopus (Rand, 1967). The full elevation display was usually only 
performed in response to a dewlap display or other action by the 
other lizard in an encounter. Posterior movements were never per- 
formed without such provocation. 

Head orientation was also related to intensity level. Throughout 
the dewlap displays, and especially at low intensity levels, lizards 
changed their head orientation frequently and usually rather slowly. 
The snout was often pointed at the other lizard, but this action in 
itself did not seem highly significant. On the other hand, a lizard 
rapidly turning its snout toward the other, either at the end of a 
display by the former or during the display of the latter, appeared to 
indicate a high level of intensity. Generally, pointing occurred dur- 
ing intermediate or full elevation displays or more intense behavior 
and was more often performed by the dominant lizard in the en- 
counter. When a dominant lizard rapidly pointed its snout at a 
displaying subordinate lizard, the latter often ceased its display, 
retracted its dewlap, and dropped to a less elevated posture. 

Anolis cybotes 

There was no indication in A. cybotes of anything resembling the 
pelvic push-ups of A. marcanoi. The posterior was always station- 
ary and usually firmly on the substrate, though occasionally the 
entire body was lifted off the substrate. 

Pointing with the snout at an opponent was commonly seen in A. 
cybotes, though neither as frequently nor in as defined a manner as 
in A. marcanoi. Several times, a lizard rapidly turned, its snout and 
even its whole body directed at its opponent, briefly paused, and 
then leapt at or toward it. The derivation of pointing in A. marcanoi 
may thus be revealed; perhaps pointing in A. marcanoi has become 
ritualized, detached from its original threat of imminent attack. 
Pointing was never observed by a lizard which would subsequently 
lose an encounter. 

A common action more frequently displayed by A. cybotes than 
by A. marcanoi was tilting or cocking the head downward toward 
an opponent. This often occurred when the lizard was on a branch, 
looking down upon an opponent on the ground and often was 



1985 ASOI.IS SIBLING SPECIES BEHAVIOR 17 

associated with some movement or action by the other lizard. 
Though cocking of the head may simply allow better vision of the 
other lizard's actions, it probably has a communicative function, 
perhaps as an intention movement, indicating awareness of the 
other lizard's actions and readiness to respond. Anolis also com- 
monly performs this behavior prior to attacking potential prey items 
(G. C. Mayer, personal communication). Dominant lizards more 
commonly performed this behavior. 

Advancing Behavior 

Past the stationary display, the steps of increasing intensity of 
aggression become more stereotypical, involving movement toward 
an opponent and threat of or actual attack, though often inter- 
rupted by more bouts of unilateral, alternating, or simultaneous 
dewlap displays. In most cases, the progression to increasing levels 
of intensity was only stopped by fleeing or quiescence of the subor- 
dinate lizard, particularly in A. marcanoi, though several times both 
lizards ceased behaving and assumed less intense postures without 
either clearly "winning" the encounter. 

Anolis marcanoi 

Figure 5 presents the flow of advancing behaviors of A . marcanoi. 

Rapid pointing of the snout toward the other lizard was the low- 
est level of advancing behavior as well as a display modifier, though 
it was still usually tied to the interspersed dewlap displays. 

Taking one or several steps toward the other lizard is the first 
overt threat of attack. At close proximity, stepping toward the other 
lizard threatened imminent attack and often caused the other lizard 
to cease its display and/ or assume a more subordinate position, 
retreat one or several steps, or seek escape from the encounter. 

Also at closer proximity, intent, or at least threat, to attack was 
indicated by the assumption of a "poised" position. The body was 
held off the ground with the legs out to the side and appearing ready 
to spring. The head, enlarged by crest and throat expansion, with 
the dewlap not usually greatly extended, was tilted somewhat for- 
ward and toward the other lizard. Rarely was this position assumed 
without a subsequent attack. At this time, the lizards were usually in 
the faceoff position (Carpenter, 1962, 1978), their bodies parallel 
with heads pointed in opposite directions. When displaying on a 



18 



BREVIORA 



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1985 ANOUSSm.mC} SPECIES BEHAVIOR 19 

perch, this poised position often took the form of a crouch, clearly 
threatening a jump toward the other lizard. Echelle et al. (1971), in a 
study of aggressive behavior in nine Costa Rican Anolis, noted the 
assumption of a similar position in males that subsequently attacked 
their opponents. 

Usually following the poised position was an advance toward the 
rear of the other lizard, culminating in an open-mouthed lunge 
toward the other lizard's hindquarters. Biting was never observed at 
this stage. Often the tongue was bunched at the front of the open 
mouth. Though tongue-bunching is characteristic of aggressive 
interactions in many Anolis species (e.g., Greenberg, 1977), it does 
not appear to have consistent importance in A. marcanoi. Attacking 
individuals always lunged with mouth open and often kept the 
mouth open while displaying after lunging. In addition, two lizards 
opened their mouths in threat displays without immediately lunging 
afterwards. 

The attacked lizard responded either by jumping back and flee- 
ing, or pivoting, usually with a jump backwards, away from the 
attacking lizard, with its head turned over the shoulder toward the 
attacking lizard. The two lizards, now in semi-circular positions 
with heads at opposite ends, displayed at each other, bodies com- 
pletely elevated off the ground and snouts angled up, with mouths, 
at least initially, wide open. Occasionally, the attacking lizard con- 
tinued moving toward the other's rear so that both circled around, 
maintaining the same relative position. In several instances, the 
attacked lizard, after displaying, lunged for the other's rear, though 
in no instance was it successful in driving the former off. In all but 
two cases, after a sequence of displays, lunges, and circling motions, 
the attacked lizard fled and attempted to escape the terrarium by 
running to a distant wall and clawing wildly at it. 

In two instances, the two lizards locked jaws for less than two 
seconds, shaking each other violently, until they released, jumping 
backwards, with the attacked lizard fleeing. In one of these in- 
stances, the attacking lizard chased the other lizard several steps 
before ignoring it. 

In all instances in which the losing lizard fled, the winning lizard 
continued displaying for up to several minutes. Slowly, the lizard's 
displays and postures descended the stages of intensity, though they 
would temporarily reascend if the other lizard came nearby in its 



20 BRF.VIORA No. 484 

attempts to escape the terrarium. Ultimately, the winning lizard 
would assume a non-aggresive posture, stretched out on a perch or 
the ground with its head on the substrate or barely elevated and its 
rear legs stretched back behind it. The overall effect was to increase 
its length while decreasing its apparent height. This is the normal 
resting position of the lizards when undisturbed. 

A no/is cybotes 

Advancing behavior was progressively ordered to an extent, but 
not completely (Fig. 6). In the most structured sequences, a lizard 
performed some display behaviors and then moved towards its 
opponent, either jumping off a branch or running half or more of 
the distance separating the lizards. This was usually done at a rapid 
pace. The advancing lizard then adopted a stalking, "poised" posi- 
tion, somewhat like that exhibited by A. marcanoi, but with all four 
legs out to the side, seemingly prepared to spring, with the long axis 
of the body pointed directly at the other lizard. This was not always 
seen, and sometimes a lizard hardly paused between moving toward 
and actually attacking the other lizard. 

The act of biting an opponent was very different in the two spe- 
cies. Encounters involving A. marcanoi only resulted in biting when 
both lizards continued to behave intensely. The sequence of faceoff 
position-lunging-biting was seen only once in the A. cybotes 
encounters. Instead, most incidents of biting involved very little 
ritualized behavior. One lizard leapt or ran at the other and bit it 
behind the head, attempting to pin it to the substrate. Attempts to 
escape by the attacked lizard often resulted in that lizard carrying 
the other around on its back. The attacked lizard often was not 
displaying vigorously and in several cases was trying to escape from 
the attacking lizard. Ano/is marcanoi rarely exhibited chasing 
behavior, but this was common in A. cybotes. In the latter species, 
on occasion, lizards were chased all over the aquarium until they 
could be cornered and bitten. 

Most surprising were two instances in which a lizard, with no 
preliminaries, ran directly and rapidly at the other lizard, which also 
had exhibited no aggressive behavior, and bit it behind the head. In 
both cases, the attacking lizard persistently bit and chased the other 
lizard until the encounter was ended. This was reminiscent of the 
"vicious biting attack" termed "abnormal" by Greenberg and Noble 



1985 



4 NOUS Sim INC. SPECIES BF HAVIOK 



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22 BREVIORA No. 484 

in A. Carolinensis (1944). This behavior was observed by these 
investigators in 18 or 800 males, as well as in some females and 
surgically-altered males. 

Submissive Behavior 

A no/is mareanoi 

As indicated previously, subordinate individuals usually did not 
posture as intensely as dominant lizards. Dewlap displays were gen- 
erally less elevated, shorter in duration, less frequent, and often 
without crests or dark spots. Subordinate lizards also did not as 
often attempt to enlarge their apparent head or body size. 

More submissive behavior generally took the form of staying very 
low to the ground, with the head elevated little and the snout angled 
up only slightly, if at all. Carried to the extreme, completely submis- 
sive lizards laid completely flat on the ground, appearing as incon- 
spicuous as possible. 

At higher levels of intensity, particularly in response to advancing 
behavior, submissiveness was usually indicated by fleeing the en- 
counter and attempting to escape the aquarium. 

Anolis ey botes 

One of the most surprising aspects of A. cybotes" behavior is the 
lack of submissive behavior in the lizards that were judged, by the 
eventual outcome of an encounter, to be subordinate. Unlike A. 
mareanoi, these lizards did not behave less vigorously or aggres- 
sively, nor did they necessarily display fewer behaviors connoting 
higher levels of intensity than the dominant lizard in the encounter. 
In several cases, the subordinate lizard initiated aggression by mov- 
ing toward the other lizard. This was never seen in the A. mareanoi 
encounters. 

More generally, lizards did not adopt a submissive posture as A. 
mareanoi did. They sometimes moved away or displayed less, but 
they did not flatten themselves or otherwise appear inconspicuous. 
Often, a lizard appeared to be trying to escape the aquarium, though 
neither persistently nor frantically enough to warrant ending an 
encounter. When this proved futile, the lizard turned back and 
behaved aggressively again toward the other lizard. Such behavior 
was rarely exhibited by A. mareanoi. 



1985 4 JV0/./6' SIBLING SPECIES BEHAVIOR 23 

Index of Aggression 

In order to quantify levels of intensity in interactions, an index of 
aggression, similar to that developed for Puerto Rican anoles by 
Ortiz and Jenssen (1982) was constructed. The index assigns progres- 
sively higher point values for actions indicating greater intensity 
(Table 2). This index, then, can be used to compare quantitatively 
the level of aggressive intensity of A. marcanoi in inter- and intra- 
specific encounters (Losos. in press). Points for any behavior are 
scored only once per encounter even though many behaviors are 
repeated periodically. The score of a lizard in an encounter is the 
sum of points awarded for behaviors which it exhibited. 

The behavior of A. marcanoi was progressive, with the less 
aggressive behaviors preceding the more intense ones. Because this 
was .not the case for A. cybotes, no index of aggression could be 
developed for that species. Occasionally, the actions of A . marcanoi 
during the dewlap display did not necessarily follow a set hierarchy 
of increasing intensity either. The most notable exception occurred 
when a lizard, usually at a distance from its opponent, sometimes 
moved toward the other before pointing its snout rapidly and 
directly at it. 



Table 2. Index of Aggression used to score encounters between adult male Anolis 
marcanoi. 

Points Action or Display Modifier 

1 low level and intermediate dewlap display 

2 full elevation dewlap display 

2 crests erected 

3 head spot apparent 

3 lifting the pelvic region and or moving backwards during 

dewlap display 

3 pointing snout rapidly at opponent at conclusion of own 

display or during opponent's display 

4 moving toward opponent (including jumping onto or off of 

perch) 

6 adopting poised position 

7 lunging toward opponent 

8 locking jaws 

8 opponent flees 



24 BREVIORA No. 484 

Several notes on the construction of this index are necessary. Only 
two levels of dewlap display are scored although three were recog- 
nized. The intermediate dewlap display stage was considered only 
slightly more intense than the low level dewlap display, and conse- 
quently the two were grouped together. Though the nuchal crest is 
almost always erected before and retracted after the dorsal crest, 
they are not considered independent behaviors denoting different 
levels of aggressive intensity. The scores of the lizards agree well 
with the qualitative evaluations of the 26 encounters. The encoun- 
ters can be split into four categories based on their outcome: 1. 
neither lizard supplants the other; 2. displaying and moving toward, 
one flees; 3. lunging, one flees; and 4. jaw-locking, one flees. 

The index well represents the level of intensity of aggression in an 
encounter (Table 3). Even within particular categories, encounters 
that qualitatively appeared to be more intense scored higher on the 
index of aggression. The one discrepancy involves several of the 
higher scoring encounters in the first category which scored higher 
than several of the lower scores in the second category. In the 
former encounters, the dominant lizard was fairly aggressive, adopt- 
ing the poised position, but then subsiding without forcing the other 
lizard to flee. In the latter encounters, on the other hand, the subor- 
dinate lizard fled immediately after the dominant one jumped down 
from a perch toward it. These encounters probably would have 
scored appreciably higher if the subordinate lizard had not so 
promptly fled; the points awarded for forcing the other lizard to flee 
were not sufficient to compensate for those lost from the abrupt 
termination of the encounter. Notwithstanding this one shortcom- 
ing, the index appears to satisfactorily evaluate levels of intensity in 
aggressive encounters. 

A second confirmation, with four exceptions, of the index's accu- 
racy comes from comparison of the scores of the lizards judged 
dominant by their behavior, level of activity, and posture with the 
scores of those judged subordinate. In no case did the subordinate 
lizard achieve a higher score than the dominant lizard, while in four 
instances both lizards received the same score. 

DISCUSSION 

The male aggressive behavior of A. marcanoi appears very similar 
to that of most other Anolis studied, both overall and in particular 



1985 ANGUS SIBLING SPECIES BEHAVIOR 25 

Tabic 3. Results of encounters between male Anolis mananoi. 





Lizards Paired 








(I.D. Number. 


Dominant Lizard 






Higher Scoring 


Score Subordinate 




Encounter Outcome 


Lizard First) 


Lizard Score 


Total Score 


neither flees 


9 18 










4 12 


1/1 


2 




5-9 


5/1 


6 




12-2 


6/1 


7 




14-16 


8/3 


II 




2 13 


6/6 


12 




1-7 


8/8 


16 




12 6 


1 8 1 1 


29 




14 3 


18/11 


29 




1-10 


22; 15 


37 




19 5 


23 14 


37 


displaying, one flees 


3 9 


15/6 


21 




1-5 


21/4 


25 




3-9 


26/6 


32 




2-16 


23/11 


34 




13 17 


23/12 


35 




13-7 


29 14 


43 




5-4 


32/18 


50 


lunging, one flees 


11-7 


35/17 


52 




3-5 


39/14 


53 




14-7 


39/15 


54 




7-13 


33/31 


64 




13-12 


36/28 


64 


jaw-locking. 


16-18 


38/30 


68 


one flees 


19-15 


41/29 


70 




9-2 


47/26 


73 



detail. The progressive nature of increasingly aggressive behavior 
has long been noted (Greenberg and Noble, 1944; Carpenter, 1967; 
Gorman, 1968; Jenssen, 1977, 1978; Hover and Jenssen, 1976; Ortiz 
and Jenssen, 1982). As in the study of five Puerto Rican Anolis by 
Ortiz and Jenssen, the behaviors and modifiers observed generally 
appeared in a progressive fashion, with some characteristically 
appearing early in encounters and others appearing progressively 
later as intensity increased. In most cases there was a clear linear path 
from one stage to another, though with one major exception, mov- 
ing forward, which often preceded snout pointing. As noted above, 



26 BkivioRA No. 484 

the ordering of some behaviors and display modifiers depended on 
distance separating the lizards. At a given distance, behaviors and 
modifiers generally did not appear prematurely, though many ear- 
lier behaviors and modifiers recurred at higher levels of intensity. A 
comparison of the index of aggression derived for A . marcanoi with 
that derived by Ortiz and Jenssen for Puerto Rican Anolis shows 
great similarity, though the latter is more detailed than the index 
presented here. 

Particular behaviors also resemble those reported for related spe- 
cies. Stamps and Barlow ( 1973) reported several stages in the dew- 
lap display of A. aeneus: fan, fanbob, bob, and jerkbob. Not only 
wereall four stages observed in A. marcanoi, but they appeared, 
very approximately, in equal proportions to those of A. aeneus. 
Crest erection, assumption of the faceoff position, lunging, and jaw- 
locking have been observed in a number of species. Maximizing 
apparent size by body configuration and orientation is also preval- 
ent among Anolis (Jenssen, 1977) and iguanids in general (Carpen- 
ter, 1967). 

However, there are some differences in the behavior of A marca- 
noi as well. Pointing with the snout has been reported for only one 
other species, A. carolinensis (and then only at very close range 
[Cooper, 1977] ). Anolis carolinensis is also the only species reported 
to exhibit a well-defined postorbital dark spot. Behaviors noted 
occasionally in A. marcanoi, such as open mouth threats and mouth- 
smacking, are of greater importance in other species such as A. 
opalinus (Jenssen, 1979a) and A. aeneus (Stamps and Barlow, 
1973), while actions never observed in A. marcanoi, such as tongue 
protrusion and foreleg lifting, have been observed in many other 
species (e.g.. Rand, 1967; Jenssen, 1979a; Jenssen and Rothblum, 
1977; Ortiz and Jenssen, 1982). It appears that as well as having its 
own stereotyped display-action patterns, each species of Anolis has 
its own behavioral repertoire, sharing some behaviors with other 
species while possessing a few unique behaviors of its own. 

With regard to A. cybotes, two related questions merit address- 
ing. Why is the aggressive behavior of A. cybotes so much simpler 
than that of A. marcanoi, and, indeed, of all Anolis examined? And 
why is there such little order in the appearance of aggressive behav- 
iors, unlike the highly ordered and progresive pathways of increas- 
ing aggression in A. marcanoi and other Anolis? 



1985 I NOUS SIBI l\(i SIM C IES BEHAVIOR 27 

Before these questions can be answered, several points of caution 
must be raised. First, it is possible that the experimental arena was 
not large enough to elicit the full range and ordering of natural 
aggressive behavior. Much larger experimental arenas have been 
used by other investigators (e.g., Jenssen, 1970, 1975; Jenssen and 
Hover, 1976; Stamps and Barlow, 1973), which was not possible in 
this case. If this is true, however, it is curious that the same problem 
did not occur in A. marcanoi, which is so similar to A. cybotes. 
Second, the lizards had been in captivity for more than ten months 
when the last of these experiments were performed. Though most 
appeared to be in reasonably good health, the long stretch of captive 
care may have taken its toll on their behavior. The lizards did, 
however, react quickly and in seemingly normal ways during the 
encounters; there was just no pattern from lizard-to-lizard or 
encounter-to-encounter. Third, the encounters involving A. cybotes 
were staged during the winter, a refractory period for males of the 
species in which their aggressive behavior is diminished in the wild 
(Jenssen, personal communication). Several A. marcanoi tested dur- 
ing the winter (which presumably is the refractory period for this 
species as well), after the same length of time in captivity, exhibited 
typical behavior. 

Jenssen (1983) argues that the single stereotyped head-bobbing 
display pattern of A. cybotes is a primitive trait with respect to all 
Ano/is. The rest of A. cybotes' aggressive behavior — dewlap dis- 
plays, display modifiers, and advancing behavior — are also simpler 
and less ritualized than that of its sibling. A. marcanoi. That this 
simplicity could not be derived from the more complex behavior of 
A. marcanoi, however, is not clear. 

Even if these behaviors are retained primitive ones, the ultimate 
question, why they have not been expanded and elaborated as in 
other species, still needs an answer. Jenssen (1983, personal com- 
munication) suggests that the determinants of social spacing in A. 
cybotes — a large lizard with large territories, few neighbors, low 
turnover, and low density — would mitigate against many territorial 
challenges. Lizards would know their neighbors and rarely chal- 
lenge them. Intrusion would thus be a matter to be seriously dealt 
with because interlopers are likely to be lizards without a territory 
seeking to displace a resident, and because, due to low turnover, 
available territories would be hard to come by. Consequently, an 



28 BKI VIORA No. 484 

intrusion would rapidly result in a fight and elaborate communica- 
tory rituals would be unnecessary. This hypothesis can explain the 
simplified aggressive behavior of A. cybotes compared to that of A. 
distichus (with which it was contrasted by Jenssen), but why such 
radical differences with its sibling species, A. marcanoi, and other 
truck-ground ecomorphs (sensu Williams, 1972), such as A. crista- 
tellus, A. cooki, A. gundlachi (Ortiz and Jenssen, 1982), and A. 
lineatupus (Rand, 1967)? All that can be said is that nothing is 
known about the spacing and turnover of A. marcanoi populations, 
so direct comparisons with its sibling species cannot be drawn. 

One important point can be made about the lack of distinct sub- 
missive behavior in A. cybotes. In A. marcanoi, submissive behavior 
appeared to have a pacifying effect on the dominant lizard. Only 
rarely and to a limited extent did a dominant lizard continue to act 
aggressively toward a clearly submissive opponent. By contrast, 
submissive behavior elicited no such complacency in A. cybotes. 
Most incidents of biting occurred upon lizards that were not dis- 
playing high levels of aggression. Consequently, submissive behav- 
ior did not seem to gain any advantage to the lizard displaying it. 
Rather, perhaps the best defense if escape is impossible is to put up 
an aggressive front, threatening retaliation in case of attack. 

Whether these findings and speculations relate to natural behav- 
ior needs to be verified in the field. Studies are needed on the social 
structure and spacing of populations of both A. marcanoi and A. 
cybotes in nature. Futhermore, it is important to find out whether 
A. cybotes does, indeed, need a much greater space between indi- 
viduals than A. marcanoi for naturally progressive behavior to 
appear. 

Other than expanding the general pool of knowledge of Anolis 
behavior, the data here reported hopefully will be of value in inves- 
tigating the ecological and behavioral interactions of these sibling 
species. Knowledge of the intraspecific behavior of males of both 
species will serve as a control with which to compare the results of 
experiments investigating interspecific male aggressive behavior and 
the mechanisms of species recognition. 

ACKNOWLEDGMENTS 

I am deeply indebted to Sibel Akyol, Pere Alberch, Ernest E. 
Williams, and, particularly, Greg Mayer for their continuous 



1985 ANOUS SIBLING SPECIES BEHAVIOR 29 

encouragement, advice, and support. I am also deeply grateful to 
Sixto and Ivon Inchaustegui for their assistance when I was in the 
Dominican Republic. Thanks also to Emily Gale, Carlos Garcia, 
Harry Greene, Paul Hertz, Thomas Jenssen, James Knight, Jose 
Rosado, the members of the Herpetology Department, Museum of 
Comparative Zoology, and the staff of the Biological Laboratories, 
Harvard University, for their help, and to Crimson Camera for the 
loan of a video camera. Laszlo Meszoly and Gene Christman kindly 
drew the illustrations. 

LITERATURE CITED 

Carpenter, C. 1962. Patterns of behavior in two Oklahoma lizards. Amer. Mid. 
Nat.. 67: 32-51. 

1965. Display of the Cocos Island anole. Herpetologica. 21: 256-260. 

_. 1967. Aggression and social structure in iguanid lizards, pp. 87-105. In 

W. W. Milstead (ed.). Lizard Ecology: A Symposium. Columbia. Mo., Univer- 
sity of Missouri Press, ix + 300 pp. 

1978. Ritualistic social behaviors in lizards, pp. 253-267. In N. Greenberg 



and P. D. MacLean (eds.) Behavior and Neurology of Lizards: An Inter- 
disciplinary Colloquium. Rockville. Md., NIMH, vii + 352 pp. 

Cooper, W. E. 1977. Information analysis of agonistic behavioral sequences in 
male iguanid lizards, Anolis carolinensis. Copeia, 1977: 721-735. 

Echelle, A. A.. A. F. EciiELLE, and H. S. Fitch. 1971. A comparative analysis 
of aggressive display in nine species of Costa Rican Anolis. Herpetologica. 27: 
271-288. 

Gorman, G. C. 1968. The relationships of Anolis of the roquet species group 
(Sauria: Iguanidae) — 111. Comparative study of display behavior. Breviora Mus. 
Comp. Zool., No. 284, pp. 1-31. 

Greenberg, B., and G. K. Noble. 1944. Social behavior of the American chame- 
leon (Anolis carolinensis). Physiol. Zool. 17: 392-439. 

Greenberg, N. 1977. A neuroethological study of display behavior in the lizard 
Anolis carolinensis. Amer. Zool.. 17: 191-201. 

Hertz, P.E. 1980. Comparative physiological ecology of the sibling species Ano- 
lis cvbotes and A. marcanoi. J. Herp.. 14: 92-95. 

Hover, E. L., and T. A. Jenssen. 1976. Descriptive analysis and social correlates 
of agonistic displays of Anolis limi/rons. Behavior, 58: 173-191. 

Jenssen. T. A. 1970. The ethoecology of Anolis nebulosus (Sauria, Iguanidae). J. 
Herp.. 4: I 38. 

1975. Display repertoire of a male Phenacosaurus heterodermus (Sauria: 

Iguanidae). Herpetologica, 31: 48-55. 

1977. Evolution of anoline lizard display behavior. Amer. Zool.. 17: 

203-215. 

1978. Display diversity of anoline lizards and problems of interpretation, 

pp. 269-285. In N. Greenberg and P. D. MacLean (eds.). Behavior and Neurol- 



30 BREVIORA No. 484 

ogy of Lizards: An Interdisciplinary Colloquium. Rockville, Md., NIMH, vi + 

352 pp. 

1979a. Display modifiers of Anolis opalinus. Herpetologica, 35: 21 30. 

1979b. Display behavior of male Anolis opalinus (Sauria, Iguanidae): A 

case ol weak display stereotypy. Anim. Bchav., 27 173-184. 

1983. Display behavior of two Haitian lizards. Anolis cybotes and Anolis 



distic/uis, 552 569. //; A. G. J. Rhodin and K. Miyata (eds.). Advances in 

Herpetoloffy and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 

Cambridge. Mus. Comp. Zool.. x + 725 pp. 
Ji nssi n, T. A., and E. L. HoVKK. 1976. Display analysis of the signature display 

of Anolis limi/rons. Behavior. 57: 227 240. 
Jenssen, T. A., and L. RoniHi.tM. 1977. Display repertoire analysis of Anolis 

townsendi (Sauria, Iguanidae). Copeia, 1977: 103-109. 
Mavr. E. 1942. Systemalics and the Origin of Species. New York, Columbia 

University Press, x + 334 pp. 
Ortiz, P. R.. and T. A. Jenssen. 1982. Interspecific aggression bewteen lizard 

competitors, Anolis cook i and Anolis cristatellus. Z. Tierpsychol., 60: 227 238. 
Pace, A. E. 1974. Systematic and biological studies of the leopard frogs (Rana 

pipiens complex) of the United States. Miscellaneous Publications of the 

Museum of Zoology. Univ. of Michigan, 148: I 140. 
Rand. A. S. 1967. Ecology and social organization in the iguanid lizard Anolis 

lineatopus. Proc. U.S. Nat. Mus., 122: I 79. 
Sciioener, T. W. 1977. Competition and the niche, pp. 35-136. //; C. Gans and 

D. W. Tinkle (eds.) Biology ol the Reptilia, Vol. 7. London, Academic Press, viii 

+ 720 pp. 
Schwartz, A. 1974. An analysis of variation in the Hispaniolan giant anole 

Anolis ricordii Dumeril and Bibron. Bull. Mus. of Comp. Zool., 146: 89 146. 
Stamps, J. A., and G. W. Barlow. 1973. Variation and stereotypy in the displays 

of Anolis aeneus. Behavior, 47: 67 94. 
Webster, T. P. 1975. An electrophoretic comparison of the Hispaniolan lizards 

Anolis ( yhoies and A. marcanoi. Breviora Mus. Comp. Zool., No. 431, pp. 1-8. 
Williams, E. E. 1965. The species of Hispaniolan green anoles (Sauria: Iguani- 
dae). Breviora Mus. Comp. Zool., No. 227, pp. 1-16. 
. 1972. The origin of faunas. Evolution of lizard congeners in a complex 

island fauna: a trial analysis. Evol. Biol., 6: 47 89. 

1975. Anolis marcanoi new species: sibling to Anolis cybotes: descrip- 



tion and field evidence. Breviora Mus. Comp. Zool.. No. 430, pp. 1-9. 



r 



BREVIORA 

Museum of Comparative Zoology 

US ISSN 0006-9698 

Cambridge, Mass. 29 August 1986 Number 485 



THE ANATOMY AND RELATIONSHIPS OF 

STEREOPHALLODON AND BALDWINONUS 

(REPTILIA, PELYCOSAURIA) 

Donald Brinkman 1 and David A. Eberth 2 

Abstract. As representatives of pelycosaurs from the earliest terrestrial deposits 
of the Texas and New Mexico Permo-Pennsylvanian sequence, Stereophallodon and 
Baldwinonus are important in our understanding of pelycosaur evolution. However, 
their variable placement in the phylogenetic schemes of Romer and Price (1940), 
Langston (1965), and Reisz (1980) has underscored the fact that their morphology is 
poorly known and that their relationships are openly questioned. Reanalysis of 
previously described specimens and heretofore undescribed specimens has allowed 
for a reevaluation of both the morphology of Stereophallodon and Baldwinonus and 
the phylogenetic relationships of these taxa with other pelycosaurs. 

Stereophallodon is interpreted as the sister-taxon of Ophiacodon on the basis of 
five derived features: the shape of the cultriform process, the ventrally directed 
basipterygoid process, the anteroposteriorly oriented basipterygoid articular sur- 
faces, the presence of a ridge that is triangular in cross section extending dorsally from 
the caniniform-tooth buttress, and the presence of a notch on the postero-medial 
edge of the quadrate. Stereophallodon is primitive with respect to Ophiacodon in 
retaining the following characters: dorsal vertebrae that have flattened ventral sur- 
faces and subparallel sides, a femur with an adductor crest running along the mid- 
ventral surface of the bone, and a long neck on the astragalus. Apomorphic features 
of Stereophallodon are the presence of two greatly enlarged caniniform teeth sup- 
ported by a caniniform-tooth buttress and the triangular cross section of the prox- 
imal end of the fibula. 



'Tyrrell Museum of Palaeontology, Box 7500, Drumheller, Alberta, Canada, 
T0J 0Y0 

2 Tyrrell Museum of Palaeontology, Box 7500, Drumheller, Alberta, Canada, 
T0J 0Y0 



SEP 1 

H, 
UN1VER- 



2 breviora No. 485 

Baldwinonus is represented by the weathered remains of a single individual from 
New Mexico. It is considered related to Stereophallodon on the basis of shared 
derived features of the maxilla. 

INTRODUCTION 

In a previous paper (Brinkman and Eberth, 1983), the interrela- 
tionships of a number of genera of pelycosaurs were considered. The 
genera included representatives of most of the families recognized 
by Romer and Price (1940), and thus the analysis served as a test of 
their long accepted phylogeny. The relationships of the Ophiaco- 
dontidae, Edaphosauridae, and Sphenacodontidae that they pro- 
posed were supported by this analysis, but the relationships of the 
Varanopseidae and the Caseidae were not. Rather than considering 
Varanops and Aerosaurus as primitive sphenacodonts and Casea as 
a primitive edaphosaurian, it was concluded that these genera are 
members of a single clade that is primitive in a number of features 
with respect to the clade that includes Dimetrodon, Edaphosaurus, 
and Ophiacodon. 

Brinkman and Eberth restricted their analysis to better preserved 
taxa in order to establish ". . .a framework in which detailed taxo- 
nomic and morphological revisions of individual genera and fami- 
lies can be interpreted." (Brinkman and Eberth, 1983: 3). Since that 
review, two problematic taxa, Stereophallodon and Baldwinonus, 
have been studied by the authors and their relationships with other 
pelycosaurs have been reevaluated. In this paper, we describe the 
morphology of these two taxa and present our phylogenetic 
conclusions. 

Stereophallodon was first described by Romer (1937: 90) and was 
characterized by the presence of "ophiacodont" vertebrae and two 
greatly enlarged caniniform teeth. In a later description (Romer and 
Price, 1940), it was stated that Stereophallodon differed from 
Ophiacodon in the presence of two large caniniform teeth, the 
greater development of the prefrontal pocket, abrupt decrease in 
length of the postcaniniform teeth, and in showing a tendency 
towards a greater ventral flattening of the centra. In the same paper, 
Romer and Price (1940) named a new pelycosaur, Baldwinonus 
trux, and described it as having ophiacodont-like quadrate, 
vertebral and tooth morphology, but sharing with Stereophallodon 
the specialized characters of enlarged caniniform teeth, close-set 



1985 STEREOPHALLODON AND BALDWINONUS 3 

zygapophyses and flattened ventral centra. Noting similar carnivo- 
rous specializations in the diminutive Eothyris, considered by 
Romer and Price to be an especially primitive ophiacodont, and 
Stereorhachis, a pelycosaur from Autun, France, with Ophiacodon- 
like postcrania, Romer and Price erected the taxon Eothyrididae for 
the inclusion of Eothyris, Baldwinonus, Stereophallodon, and 
Stereorhachis and characterized the family as follows: 

Ophiacodont pelycosaurs, primitive in most known 
regards but paralleling the higher sphenacodonts in the 
development of much enlarged canines and showing a 
tendency towards elongation of the vertebral column 
(Romer and Price, 1940: 246-247). 

The occurrence of Stereophallodon in the earliest terrestrial 
deposits of the Texas Permo-Pennsylvanian sequence (Pueblo 
Formation) led Romer and Price to suggest that the eothyridids 
represented an early radiation of carnivorous pelycosaurs that, 
ecologically, were supplanted by sphenacodonts and were, in 
themselves, a phyletic dead end. Contrary to this opinion, Watson 
(1954, 1957) suspected a close phylogenetic relationship between 
Eothyris and the caseids, Casea and Cotylorhynchus. According to 
Romer and Price (1940) the caseids were derived from an obscure 
group, the Nitosauridae, the taxa of which appear to be based on 
fragmentary remains of questionable association (Reisz, 1980). 
Watson's insight gained more firm support following Vaughn's 1958 
description of Colobomycter pholeter, a small pelycosaur from the 
Fort Sill fissure fills of Oklahoma that displays strong affinities to 
both Eothyris (short face, anteroposteriorly thickened postorbital 
bar, large supratemporal, large parietal foramen, large naris and 
similar maxillary dentition) and caseids in general (rugose skull, 
short face, large naris, large supratemporal, large parietal foramen 
and the presence of an infraorbital foramen). Langston (1965) 
described a new pelycosaur from the Cutler Formation of north- 
central New Mexico (Oedaleops) that likewise showed a number of 
features shared in common by the eothyridids Eothyris and 
Colobomycter and the caseids. His reevaluation of the eothyridid 
relationships within the Pelycosauria led him to conclude (as had 
Vaughn) that the Eothyrididae (including Stereophallodon, Stereo- 



4 BREVIORA No. 485 

rhachis, Baldwinonus, Eothyris, Oedaleops, Colobomycter, as well 
as three genera no longer accepted as members of that family, the 
very poorly known Delorhynchus, Bayloria and Tetraceratops) 
were antecedent to the caseids and that, in contradiction to Romer 
and Price, the nitosaurids represented a phyletic dead end group of 
ophiacodonts convergent with caseids in only a few characters of 
proportion. Most recently, Reisz (1980), in a cladistic analysis of 
pelycosaur families, concluded that the relationship between eothy- 
ridids {Eothyris and Oedaleops) and caseids was well founded, 
citing the shared derived characters of an overhanging rostrum, an 
elongate narial opening, and a maxillary contribution to the ventral 
border of the orbit. 

While the Eothyris, Oedaleops, Colobomycter / Caseidae relation- 
ship appears now to be well established, the position of Stereophal- 
lodon, Stereorhachis, and Baldwinonus are less certain. Langston 
(1965) chose to retain these taxa within the Eothyrididae although 
he cited the enlarged caniniform teeth as the only reason for doing 
so and remained uncertain of the validity of the grouping. Reisz 
went one step further and removed Stereorhachis and Stereophal- 
lodon from the Eothyrididae and placed them in the Ophiacodon- 
tidae, (1980: 566, 568). He questioned the reality of Baldwinonus, 
identifying the maxilla and neural spines of the type specimen as a 
sphenacodont and the vertebrae as pertaining to an ophiacodont. In 
our previous paper, (Brinkman and Eberth, 1983) we have 
questioned Reisz's phylogeny and have proposed a different 
arrangement. For this reason and the fact that the relationships of 
Baldwinonus, Stereophallodon, and Stereorhachis, and the validity 
of Baldwinonus remain uncertain, and because a number of new 
specimens of Stereophallodon have become available, we have 
undertaken a review of the specimens of these taxa available to us 
for examination. Stereorhachis, known from the Stephanian near 
Autun, France, was not available for study and must unfortunately 
remain excluded from the following analysis. In considering the 
position of Baldwinonus and Stereophallodon the phylogenetic 
conclusions of our earlier study were tested and our understanding 
of the morphological diversity present in pelycosaurs has been 
increased. 



1985 



STEREOP HALLO DON AND BALDWINONUS 



MATERIALS AND METHODS 

The methods used in interpreting polarities of the character-states 
are those outlined by Brinkman and Eberth (1983: 3). In that 
analysis, the polarity of character-states was interpreted using 
outgroup comparison. All other reptiles were considered the closest 
sister-group to the pelycosaurs; diadectomorphs were considered 
the closest sister-group to the pelycosaurs and all other reptiles; and 
anthracosaur amphibians were considered the sister-group of the 
clade composed of the foregoing taxa. This framework is used as a 
basis for interpreting polarities in this study as well. 

The following specimens were examined during the course of this 
work: 

AMNH 4780: (type and only known specimen of Baldwinonus 
trux) fragmentary remains of a single individual 
including maxilla, quadrate, twelve partial verte- 
brae, and partial ribs; 

MCZ 1535: (type of Stereophallodon ciscoensis) fragmentary 

remains of a single individual including the pre- 
maxilla, maxilla, quadrate, articular, left half of 
occiput, and five centra; 

MCZ 1944: fragmentary remains of a single large individual 

identified as Stereophallodon on the basis of the 
maxilla and articular; 

MCZ 6618: fragmentary remains of a single large individual, 

identified as Stereophallodon on the basis of the 
quadrate and articular; 

AMNH 4768: associated vertebrae, sacral ribs and premaxilla of 
a single large individual, identified as Stereophal- 
lodon on the basis of the premaxilla and the 
similarity of the vertebrae with those preserved in 
MCZ 6618. 

A number of isolated elements present in the collections from 
Prideaux Pocket, a bonebed in the Pueblo Formation, can be 
identified as belonging to Stereophallodon. These include: MCZ 
6358, three premaxillae; MCZ 6354, five maxillary fragments; MCZ 
6352, two prefrontals; MCZ 6353, one postfrontal; MCZ 6348, five 



6 breviora No. 485 

basioccipitals; MCZ 6349, three basi-parasphenoids; MCZ 6350, 
four quadrates; MCZ 6359, eight dentary fragments; MCZ 6352, 
three articulars; MCZ 6371, one axis centrum; MCZ 6357, partial 
centrum of a cervical vertebra; MCZ 6355, twenty mid-dorsal 
centra; MCZ 6356, twenty-two centra from various positions along 
the vertebral column; MCZ 6768, three ilia; MCZ 7083, part of the 
distal end of a femur; MCZ 6766, proximal end of femur; MCZ 
6765, two proximal ends of fibulae; and MCZ 6767, four proximal 
ends of tibiae. 

The following acronyms are used in this paper; AMNH, 
American Museum of Natural History; MCZ, Museum of Com- 
parative Zoology; UCMP, University of California, Berkeley, 
Museum of Paleontology. 

DESCRIPTION 

Stereophallodon 

An almost complete premaxilla is preserved in AMNH 4768 (Fig. 
1A-B). This has a posteriorly sloping dorsal ramus and a tooth 
bearing base. A foramen is present on the posterior surface of the 
premaxilla at the base of the dorsal ramus. Sockets for five teeth are 
present in the premaxilla. The diameter of the anterior two sockets 
is about twice that of the more posterior sockets. The premaxilla of 
Stereophallodon is like that of Ophiacodon (Fig. ID) and different 
from that of the sphenacodontines (Fig. IE) in the structure of the 
dorsal ramus. In Stereophallodon and Ophiacodon the posterior 
edge of the dorsal ramus meets the tooth-bearing base of the bone 
just above the maxillary suture. In the sphenacodontines, the 
posterior edge of the dorsal ramus meets the base of the premaxilla 
anterior to the maxillary suture. 

The maxilla (Fig. 1C) is represented by a number of specimens 
which show a caniniform-tooth buttress and adjacent areas. The 
sutural surface for the palatine extends forward onto the buttress to 
a point above the exposed posterior edge of the second caniniform 
tooth. The maxilla is broken at the dorsal apex of the buttress. The 
horizontal cross section at this point shows that a rodlike brace 
extended dorsad from the buttress along the inner surface of the 
maxilla as in Ophiacodon. In the sphenacodontines no such ridge is 
present; the caniniform-tooth buttress is a broad, gently contoured 
wedge of bone that merges smoothly and evenly with the medial 



1985 



STEREOPHALLODON AND BALDWINONUS 





Figure 1. The right premaxilla of Stereophallodon in A) lateral and B) medial 
views; C) the right maxilla of Stereophallodon in medial view; D) the right 
premaxilla of Ophiacodon in lateral view, and E) the right premaxilla of 
Sphenacodon in lateral view. Arrows on the premaxillae indicate the posteroventral 
edge of the dorsal ramus of the premaxilla. Stereophallodon premaxilla drawing 
based on AMNH 4768, reconstructed portion based on the articular surface for the 
premaxilla preserved on the maxilla in MCZ 1944; shaded portion of the maxilla 
drawing based on MCZ 1535, with the outline drawn on the basis of MCZ 1944; 
Ophiacodon drawing based on UCMP 40035; and Sphenacodon drawing based on 
UCMP 83047. 

Abbreviations: BUT, caniniform-tooth buttress; BUT, DOR, EX, dorsal extension 
of the caniniform-tooth buttress; F, foramen; M S S, maxillary sutural surface; N S S, 
nasal sutural surface; PAL S S, palatine sutural surface; V S S, vomer sutural surface. 



surface of the maxilla. Moreover, a precaniniform-tooth step is not 
present in Stereophallodon. Rather, the ventral edge of the maxilla 
slopes gently dorsally and bears well-developed precaniniform teeth. 

Two large caniniform teeth are present on the maxilla, each 
having a basal diameter of about three times that of any of the more 
posterior teeth. At least three teeth are present anterior to the 
caniniform teeth. The postcaniniform teeth, as described by Romer 
and Price (1940), are extremly short, measuring about one sixth the 
length of the caniniform teeth, and are more broadly exposed on the 
internal surface of the maxilla than on the external surface. 

Fragments of the dermal skull elements present in the type 
specimen include the prefrontal, postfrontal, and frontal. Isolate d 



8 



BREVIORA 



No. 485 



prefrontals are present in the Prideaux Pocket locality. These 
elements show that the frontal of Stereophallodon is like that of 
Ophiacodon in having a small lateral lappet. The prefrontal has a 
lateral pocket anterior to the orbit, although this does not seem 
significantly larger than a similar pocket seen in the prefrontal of 
larger specimens of Ophiacodon. 

Pterygoids are preserved in MCZ 1944 (Fig. 2 A). The right and 
left pterygoids are preserved together in a single block with the 
transverse flange of the right pterygoid visible in dorsal view and the 
quadrate ramus of the left pterygoid visible in medial view. The tip 
of the transverse flange of the right pterygoid is preserved separately 
(Fig. 2C). As in Ophiacodon (Fig. 2B), a tympanic flange extends 





Figure 2. The right pterygoid of A) Stereophallodon and B) Ophiacodon in 
medial view and C) lateral view of the transverse flange of the pterygoid of Stereo- 
phallodon. Drawing of Stereophallodon a composite based on right and left 
pterygoid preserved in MCZ 1944. Ophiacodon drawn from MCZ specimen. 

Abbreviations: BSPT, basipterygoid articular surface; TRANS, transverse flange 
of the pterygoid; TYMP FLANGE, tympanic flange; ?Q, ?Quadrate. 



1985 



STEREOPHALLODON AND BALDWINONUS 



medially from the ventral edge of the quadrate ramus of the 
pterygoid. The transverse flange of the pterygoid is like that of 
Ophiacodon in that its lateral edge is thinner relative to its height 
than that of Dimetrodon. This difference is consistent in both large 
and small specimens of Dimetrodon and Ophiacodon and thus is 
independent of size. As in Ophiacodon, a single row of many small 
teeth is present on the ventral edge of the transverse flange. Five 
teeth are preserved, and more were probably present on the missing 
portion of the transverse process (dashed area, Fig. 2A). 

Quadrates are present in MCZ 1535 (Fig. 3A-C) and MCZ 6618. 
Four isolated quadrates are present in the material from Prideaux 
Pocket. The articular surface is differentiated into two condyles 
with a saddle-shaped surface extending between them. The lateral 
condyle has articular surface extending from its medial to its lateral 
surface. The medial condyle has articular surface developed only on 
its lateral and ventral surfaces. A stapedial pit is present on the 
medial edge of the bone just above the medial condyle. This is a 
shallow depression containing a round tuberosity. In posterior view 
the dorsal edge of the quadrate is notched. This notch is similar in 
position to the shelf on the posteromedial surface of the quadrate as 
in Ophiacodon and is considered a less well-developed stage of the 
same character. The shape of the articular surface in the two genera 
is different in that the lateral condyle is broader mediolaterally in 
Stereophallodon than in Ophiacodon. 

The left half of the occiput is preserved in MCZ 1535 (Fig. 4). 
Although not visible in this view, the fused supraoccipital and 






Figure 3. The right quadrate of Stereophallodon in A) posterior, B) medial, and 
C) articular views. Drawing based on MCZ 1535. 



Abbreviations: LAT COND, lateral condyle; MED COND, medial condyle; 
NOTCH, notch in dorsal edge of quadrate; S, stapedial pit. 



10 BREVIORA No. 485 

TAB 



PRO 




Figure 4. Left half of the occiput of Stereophallodon, posterior view. Drawing 
based on MCZ 1535. 

Abbreviations: F M, dorsal edge of foramen magnum; PP, postparietal; PRO, 
prootic; TAB, tabular. 

prootic end laterally in an oval area of unfinished bone as in 
Ophiacodon. Romer and Price (1940) identified a piece of bone on 
the lateral corner of the supraoccipital as a displaced postparietal. 
However, a separate element is present medial to this in a position 
normally occupied by the postparietal in all other pelycosaurs and is 
thus more reasonably interpreted as that element. The more lateral 
bone is therefore interpreted as a fragment of the tabular. 

A nearly complete basi-parasphenoid is present in MCZ 1944 
(Fig. 5), and, based on this specimen, a number of fragmentary 
basi-parasphenoids can be identified among the Prideaux Pocket 
material. As in Ophiacodon, the basipterygoid processes are 
directed ventrally and the long axis of their articular surfaces are 
oriented anteroposteriorly. The cultriform process is like that of 
Ophiacodon in being nearly as wide as it is high. This is unlike the 
condition in Dimetrodon and Edaphosaurus where the cultriform 
process is more bladelike in cross section. The basi-parasphenoid 
wings are broadly separated. A small shelf covers the anterior 
portion of the groove between the wings. 

Five isolated basioccipitals from Prideaux Pocket can be referred 
to Stereophallodon (Fig. 6) on the basis of their large size. No other 
pelycosaur from that locality is as large. These are like the 
basioccipitals of Ophiacodon in that the occipital condyles are 



1985 



STEREOPHALLODON AND BALDWINONUS 



11 




Figure 5. Basi-parasphenoid of Stereophallodon in A) dorsal and B) ventral 
views. Drawing based on MCZ 1944. 

Abbreviations: BS S. basisphenoid shelf; BST TUB, basipterygoid tubercula; 
BS W, basi-parasphenoid wings; CUL PR, cultriform process; D S, dorsum sella; 
P F, pituitary fossa. 



VAG F 





OPS 




Figure 6. Basioccipital of Stereophallodon in A) posterior, B) right lateral, C) 
dorsal, and D) ventral view. Drawing based on MCZ 6348. 

Abbreviations: BS S, basi-parasphenoid sutural surface; OP S, opisthotic sutural 
surface; VAG F, medial edge of vagus foramen; XII, foramen for twelfth cranial 
nerve. 



flattened inferiorly. Also, in both genera, the ventral surface is 
marked by a series of ridges and grooves between the basioccipital 
tubercula. In Dimetrodon and Edaphosaurus, no ridges are 
present on the ventral surface of the basioccipital between the 
basioccipital tubercula. 



12 



BREVIORA 



No. 485 



Articulars are present in MCZ 1535 (Fig. 7), MCZ 1944, and 
MCZ 6618. Three articulars are known from the Prideaux Pocket 
locality. The most complete of these is MCZ 6618, in which the 
angular and prearticular remain in sutural contact with the 
articular. The shape of the cotyles correspond to the condyles of the 
quadrate: the lateral cotyle is a cup-shaped depression, and the 
medial cotyle faces medially. No retroarticular process is present on 
the articular. Instead, a ridge extends from the ventral edge of the 
angular to the posterior tip of the articular. A distinct process for 




PROC 






PROC 



Figure 7. Articular of Stereophallodon in A) dorsal, and B) medial views, and the 
posterior end of the jaw of C) Ophiacodon and D) Dimetrodon. Stereophallodon 
drawing based on MCZ 1535, Ophiacodon and Dimetrodon from Romer and Price 
(1940). 



Abbreviations: A, articular; ANG, angular; C, coronoid; LAT COT, lateral 
cotyle; MED COT, medial cotyle; PRA, prearticular; PROC, medial process on 
angular; PTM, pterygoid process of articular; RP, retroarticular process; SANG, 
surangular. 



1985 STEREOPHALLODON AND BALDWINONUS 13 

muscle attachment is present on the angular just anterior to the 
articular. As in Ophiacodon and Varanops, this is a medially 
directed process that is triangular in dorsal view. In Dimetrodon 
and Edaphosaurus the process is formed by the articular and it 
extends ventromediad. The articular has a ridge on its ventral 
surface that marks the limit of contact with the angular. As in 
Ophiacodon, Edaphosaurus, and Dimetrodon, the angular extends 
below the adductor fossa as a ventrally extended platelike element. 

The anterior end of the dentary is preserved in specimen MCZ 
1944, and a number of dentary fragments are known from Prideaux 
Pocket. In contrast to the slender dentary of Ophiacodon, the 
dentary of Stereophallodon is robust and deep. The anterior three 
teeth are enlarged, although they are smaller than the caniniform 
teeth of the maxilla. Behind these, the dentary narrows medio- 
laterally, presumably to allow the upper caniniform teeth to pass 
lateral to the dentary. 

Twenty associated vertebrae are present in AMNH 4768. These 
include two sacral, four caudal, four lumbar, and ten dorsal 
vertebrae. Some of these are preserved in articulation. An atlas 
centrum and axis are present in MCZ 1944. The axis and four centra 
from the cervical or anterior dorsal region are present in MCZ 1535, 
and a number of isolated centra are known from Prideaux Pocket. 
An atlas intercentrum from this locality is referred to Stereophal- 
lodon on the basis of its large size. Thus most morphological areas 
along the vertebral column are represented, although the total 
number of presacral vertebrae and the number of vertebrae present 
in each area is unknown. 

The atlas centrum is directly comparable to the fused atlas cen- 
trum and axis intercentrum of an adult Ophiacodon in outline, 
position of notochordal pits, and width of the ventral surface. Thus, 
although no sutures are visible, it is assumed that the axis intercen- 
trum is fused to the atlas centrum in this specimen. 

The axis is keeled. Anteriorly, a well-developed lip is present to 
receive the intercentrum (Fig. 8 A). The transverse processes are 
short and slop ventrolaterally. 

The four more posterior cervical and anterior dorsal centra 
preserved in MCZ 1535 show a transition from a keeled ventral 
surface on the cervical centra to a flattened ventral surface 
possessing a pair of widely separated ventral ridges on the anterior 



14 



BREVIORA 



No. 485 





Figure 8. A J) vertebrae of Stereophallodon. A) axis, B) anterior cervical verte- 
bra; C) mid-cervical vertebra; D) anterior dorsal vertebra; E) dorsal vertebra; F) 
lumbar vertebra; G) sacral vertebra in lateral and ventral views. H) dorsal vertebra in 
proximal view; I) lumbar vertebra; J) and sacral vertebra in distal views. K-N) cross 
sections of dorsal vertebrae of K) Stereophallodon; L) Ophiacodon; M) Edaphosau- 
rus; and N) Dimetrodon. Drawings A-D based on MCZ 1535; E-I based on AMNH 
4768; and K-N from Romer and Price (1940). 

Abbreviations: CAP ART SURF, capitular articular surface; IC, intercentrum; 
KEEL, mid-ventral keel; LIP, anterior lip on axis; POST ZYG, postzygapophysis; 
PRE ZYG, prezygapophysis; RIB, lumbar rib in place; SAC RIB, sacral rib; TRANS 
PR, transverse process. 



dorsal centra. The most anterior of the cervicals is broadly rounded 
in cross section; its ventral surface possesses a midline keel and two, 
more lateral, longitudinal ridges (Fig. 8B). The intercentrum is 
preserved in place. Without it, a well-developed lip would be 
present. A large articular surface for the head of the rib is present on 
each side of the centrum. This extends onto the intercentrum. The 



1985 STEREOPHALLODON AND BALDWINONUS 15 

second vertebra in this series (Fig. 8C) is approximately wedge- 
shaped in cross section, but has a pair of closely placed ridges 
ventrally rather than a single mid-ventral keel. The capitular 
articular surfaces are less well developed than in the more anterior 
vertebra. A lip to receive the intercentrum is not present. The 
following two vertebrae, the best of which is illustrated in Figure 8D, 
are more rectangular in cross section. The pair of ridges on the 
ventral surface of the centrum are widely separated (Fig. 8D). The 
articular surface for the capitular head of the rib is very small in 
comparison to the more anterior vertebrae. 

Ten dorsal vertebrae are present in AMNH 4768. These are 
subrectangular in end view and in cross section (Fig. 8H). The 
ventral surfaces of each centrum is flattened and does not possess a 
keel (Fig. 8E). The ventral half of the lateral surfaces of the centra 
are subparallel to one another dorsoventrally. A small articular 
surface for the capitular head of the rib is located near the dorsal 
edge of each centrum. The transverse processes are short and are 
separated from the capitular articular surfaces by a small notch. The 
zygapophyses are set close to the midline and are strongly sloped. 
The neural spines are short, their height being about one and a half 
times the height of the centrum. The spines are expanded 
anteroposteriorly at their tips. 

Four lumbar vertebrae are present in AMNH 4768 (Fig. 8F). 
These differ from the dorsal vertebrae in being wider and having a 
trefoil-shaped cross section. In end view (Fig. 81) the centrum is 
round, rather than sub-rectangular as in the dorsal vertebrae. The 
articular surfaces for the ribs on the lumbars are larger than those 
on the dorsal vertebrae, and in some specimens ribs remain in place, 
fused to the centrum. The zygapophyses are more nearly horizontal 
than they are in the dorsal vertebrae. The neural spines are similar 
to those of the dorsal vertebrae in their proportions and structure. 

The sacral vertebrae can be identified by the exaggeration in the 
size of the rib articular surfaces. The ventral half of the centrum is 
compressed mediolaterally, forming a rounded ventral ridge that 
extends between the ends of the centrum (Fig. 8G). The zyga- 
pophyses are similar to those of the dorsal vertebrae in being large, 
sub-horizontally oriented surfaces. The neural spine is narrower 
anteroposteriorly than in the more anterior vertebrae, and slopes 
posteriorly. The centrum is oval in end view (Fig. 8J). 



16 BREVIORA No. 485 

The caudal vertebrae are round in cross section. The articular 
surfaces for the ribs are large, the capitular articular surfaces being 
slightly smaller than the tubercular surfaces. Both the proximal and 
distal edges of the centrum are deeply bevelled for the intercentra. 

The anterior cervical vertebrae of Stereophallodon are like those 
of Ophiacodon, Edaphosaurus, and the sphenacodontines: the cen- 
tra are keeled and the anterior end is strongly bevelled for the inter- 
centrum. The dorsal vertebrae of Stereophallodon bear little 
resemblance to those of Ophiacodon, Edaphosaurus, or the sphena- 
codontines. In Ophiacodon, the mid-dorsal vertebrae are wedge- 
shaped in cross section (Fig. 8L) and the posterior dorsal vertebrae 
are circular in end view and in cross section. Vertebrae that are 
subrectangular in cross section like those of the dorsal vertebrae of 
Stereophallodon (Fig. 8K) are not present. The dorsal vertebrae of 
Edaphosaurus differ from those of Stereophallodon in that in cross 
section the centra are rounded rather than flattened ventrally (Fig. 
8M). The sphenacodontines have dorsal vertebrae that are strongly 
keeled (Fig. 8N). The lumbar vertebrae of Stereophallodon are 
unlike those Ophiacodon, Edaphosaurus, or the sphenacodontines 
in having a centrum that is trefoil-shaped in cross section. 

The proximal end of the ulna is present in MCZ 6618 (Fig. 9). The 
element is broad, medio-laterally flattened, and its posterior edge is 
strongly convex. The trochlear notch is shallow and has a distinct 
articular surface for the proximal end of the radius. The ulna of 
Stereophallodon is like that of Ophiacodon in being broad at the 
level of the trochlear notch. In Edaphosaurus and Dimetrodon the 
ulna is narrower in this area. 

The acetabular region of the pelvis is present in MCZ 6618 (Fig. 
10). Three partial ilia are present in the collection from the Prideaux 
Pocket. The ilium lacks an anterior process and has only a short 
posterior process. As in Ophiacodon and Ruthiromia, a shelf 
extends medially above the area of articulation with the sacral rib. 
This shelf is located at the dorsal edge of the iliac blade. 

The distal end of the femur is present in MCZ 6618 (Fig. 11). As 
in primitive reptiles generally, the dorsal surface of the distal end of 
the femur has a well-developed intercondylar groove. The finished 
bone covering this groove extends around the distal end of the 
femur towards the fibula. The shaft of the femur is circular in cross 
section and the adductor crest is a low ridge extending along i*s 



1985 



STEREOPHALLODON AND BALDWINONUS 



17 





Figure 9. Proximal end of ulna of A) Stereophallodon and ulna of B) Ophiaco- 
don, and C) Dimetrodon. Drawing of Stereophallodon based on MCZ 6618. Draw- 
ings of Ophiacodon and Dimetrodon from Romer and Price (1940). 




Figure 10. Partial pelvis of Stereophallodon in lateral and medial views. Drawing 
based on MCZ 6618. 

Abbreviations: IL, ilium; PUB, pubis; SAC, articular surface for sacral rib; 
SUPRA AC BUT, supra-acetabular buttress. 



mid-ventral surface. This is different from the condition in Ophiaco- 
don, where the ridge is a sharp crest located along the fibulad edge 



18 



BREVIORA 



No. 485 






Figure 1 1 . Right femur of A) Stereophallodon, B) Ophiacodon and C) Dime- 
trodon. Stereophallodon drawn from MCZ 6618. Ophiacodon and Dimetrodon 
from Romer and Price (1940). 

Abbreviations: ADD CR, adductor crest. 



of the bone. In Ruthiromia, Casea, and Edaphosaurus, the adductor 
crest is located in a mid-ventral position, but this is a sharp crest 
rather than a low rugosity, and gives the femur a tear-drop-shaped 
cross section. 

The proximal end of the left tibia and the shaft and the distal end 
of the right tibia are preserved in MCZ 6618. Three proximal ends 
of tibiae are present in the material from Prideaux Pocket. This 
element (Fig. 12) is much more fully ossified than any tibia known 
in Ophiacodon, so detailed comparison is not possible with that 
genus. The element differs from the tibiae of Dimetrodon and 
Edaphosaurus in that the outline of the lateral condyle of the 
articular surface is circular, rather than oval in articular view. The 
shaft has well-developed ridges on both the medial and lateral 
surfaces. Well-developed ridges are present on the lateral edges of 
all other pelycosaur tibiae examined. A slight ridge is present on the 
internal edge of the tibia in Ophiacodon, although it is not as 
strongly developed. Ruthiromia is similar to Stereophallodon in 
having a well-developed ridge on the internal surface of the tibia. 

The proximal and distal ends of a fibula are present in MCZ 6618. 
Two proximal ends of fibulae are present in the material from 



1985 



STEREOPHALLODON AND BALDWINONUS 



19 




MED CON 



Figure 12. Left tibia of Stereophallodon in articular and anterior views. Anterior 
view is a composite drawing based on right and left elements present in MCZ 6618. 

Abbreviations: CNEM, cnemial tuberosity; LAT CON, lateral condyle, MED 
CON, medial condyle. 



Prideaux Pocket (Fig. 13). The proximal end of the fibula is 
triangular in proximal view, with the base of the triangle forming 
the articular surface for the femur. A muscle scar is present on the 
lateral tip of the triangle. The proximal end of the fibula is unlike 



20 



BREVIORA 



No. 485 








Figure 13. A-C) the proximal end of the right fibula of Stereophallodon, in A) 
lateral view, B) medial view, and C) proximal view; D) left fibula of Stereophallodon, 
actual length unknown; E) Ophiaeodon, and F) Dimetrodon. Stereophallodon draw- 
ing based on MCZ 6618, Ophiaeodon and Dimetrodon from Romer and Price 
(1940). 

Abbreviations: FEM, articular surface for femur; TAR, articular surface for 
tarsus; TUB, tuber of uncertain function. 



those of other pelycosaurs in its triangular shape. In Ophiaeodon, 
Dimetrodon, and Edaphosaurus the proximal end of the fibula is 
rectangular in end view and has a proximo-distally oriented 
tuberosity on its lateral edge. The distal end of the fibula of Stereo- 
phallodon is wide and dorso-ventrally compressed. The astragalar 
and calcaneal articular surfaces are set at a low angle to each other 
and are differentiated by a slight narrowing of the distal end of the 
bone. Ophiaeodon, Edaphosaurus, and Ruthiromia are like Stereo- 
phallodon in having broad distal ends of the fibulae. In sphenaco- 
dontines, the distal end of the fibula is narrower mediolaterally. 



1985 



STEREOPHALLODON AND BALDWINONUS 



21 



A nearly complete left astragalus and the distal half of the right 
astragalus are present in MCZ 6618 (Fig. 14). As in primitive 
reptiles generally, this element is L-shaped with a tibial articular 
surface located on the lateral branch of the L, and the dorsal branch 
contributes to the support of the fibula. A perforating notch is 
present on the lateral face of the astragalus. The proportions of the 
astragalus are like those of the element in Edaphosaurus in that the 
neck is elongate, constituting nearly half of the proximodistal length 
of the bone. This is unlike the condition in Ophiacodon, where the 
neck is less than 20% of the length of the element. 







Figure 14. The astragalus of Stereophallodon in A) dorsal and B) ventral views, 
and the astragalus of C) Ophiacodon and D) Dimetrodon. Stereophallodon drawing 
based on MCZ 6618. The shaded area is the preserved portion of the right astragalus; 
the dotted outline is based on the preserved portion of the left astragalus. Ophiaco- 
don and Dimetrodon from Romer and Price (1940). 



Abbreviations: 
surface for tibia. 



PER FOR, medial edge of perforating foramen, TIB, articular 



22 



BREVIORA 



No. 485 



Baldwinonus 

A fragmentary right maxilla is preserved and displays 25 teeth or 
alveoli including two that are greatly enlarged (Fig. 15). In lateral 
view the ventral edge of the maxilla is nearly straight posterior to 
the caniniform teeth and shows a steep incline anterior to them. No 
edentulous step is present as in sphenacodontines. The bases of five 
precaniniform teeth are present. These increase in size posteriorly. 
All were apparently erupted at the time of death. The base of one 
caniniform tooth is present and is suboval in cross section with a 
shallow dorsoventral groove along its anterior and posterior 
surfaces. The base of this tooth is nine times greater in cross- 
sectional area than those of the adjacent teeth as originally observed 
by Romer and Price, 1940. Eighteen postcaniniform teeth or alveoli 
are present. Those teeth that are fully preserved show compression 
and recurvature only at the tips. Slight uncrenulated cutting edges 
are present along these tips. 

In medial view, the alveolar ridge is swollen above the caniniform 
teeth forming a caniniform-tooth buttress. This swelling accommo- 
dates the sockets for the caniniform teeth. A horizontal cross section 
of the dorsal-most extent of the buttress is triangular in outline and 
thus indicates that the buttress extended dorsad against the medial 
surface of the maxillary wall as a vertical rod or ridge. Such a 



BUT 



PMSS 




Figure 15. A) the right maxilla of Baldwinonus in internal view. Arrows mark the 
position of B) cross-section through the maxilla. Drawing of AMNH 4780. 



Abbreviations: BUT, caniniform buttress; MX PAL V S, maxillary/ palatine ves- 
sel scar; PAL S S, palatine sutural surface; PM S S, premaxilla sutural surface. 



1985 STEREOPHALLODON AND BALDWINONUS 23 

structure is seen in Ophiacodon but not in the sphenacodontines 
where the caniniform-tooth buttress extends dorsally as a broad 
smoothly contoured wedge. The presence of a rodlike extension is 
also indicated by the down turning of the maxillary/ palatine vessel 
scar as it passes forward along the medial surface of the buttress. An 
identical condition exists in Ophiacodon where the maxillary 
palatine vessel is downturned presumably to avoid contact with the 
edges of the rodlike extension. In sphenacodontines, where the 
buttress swelling smoothly extends into adjacent surfaces over a 
broad area at its dorsal terminus, the vessel scar extends forward 
with no downturning. 

Posterior to the buttress the aveolar ridge contracts, forming, in 
cross section, a V-shaped structure (Fig. 15B). In sphenacodontines, 
the alveolar ridge remains rectangular in cross section. 

The medial half of a left quadrate is preserved (Fig. 16). The 
articular surface is a saddle-shaped joint. The articular surface of 
the medial condyle is developed only on its lateral-facing surface. 
The lateral condyle is not fully preserved. The stapedial pit contains 
a strong tuberosity, presumably marking the site of contact with the 
stapes. A notch is present on the dorsal edge of the bone. In all these 
features, the quadrate of Baldwinonus is like that of Stereophallo- 
don. 

Thirteen partial vertebrae are present. Each can be placed in one 
of three groups defined on the basis of the morphology of the 
centrum and transverse process, and the relative position of these 





Figure 16. The left quadrate of Baldwinonus in posterior, medial, and articular 
views. Drawing based on AMNH 4780. 

Abbreviations: LAT CON, lateral condyle; MED CON, medial condyle; 
NOTCH, notch in dorsal edge of quadrate; S, stapedial pit. 



24 



BREVIORA 



No. 485 



groups can be established on the basis of comparison with 
Stereophallodon. 

The first group comprises four vertebrae; each possesses a sharp, 
ventral keel, a wedge-shaped cross section, and a bevelled antero- 
ventral edge. This group is represented by the specimen illustrated in 
Figure 17 A. When preserved, the transverse process slopes postero- 
laterally and is about equal in length to the width of the centrum. It 
has a large oval articular surface for the rib. The neural spine, 
preserved on an unillustrated specimen measures about twice the 
height of the centrum. The slope of the transverse process indicates 
that these vertebrae are located in the cervical region of the vertebral 
column. 

The second group comprises three vertebrae, two of which are 
illustrated, Figure 17B and C. These vertebrae are longer than those 
of the first group and possess a centrum whose ventral surface 
displays two, close-set parallel ridges. A transverse process, pre- 
served on one of these, is a short laterally directed structure with a 
ventral buttress extending to the anterior end of the centrum. These 
vertebrae correspond to the anterior dorsal vertebrae of Stereo- 
phallodon. 









Figure 17. Presacral vertebrae of Baldwinonus. A) anterior cervical vertebra, B) 
mid-cervical vertebra, C) posterior-cervical vertebra, and D) dorsal vertebra. Draw- 
ings based on AMNH 4780. 



Abbreviations: CAP ART SURF, capitular articular surface. 



1985 STEREOPHALLODON AND BALDWINONUS 25 

Six vertebrae constitute the third group and are best exemplified 
by the illustrated specimen, Figure 17D. These vertebrae possess 
smooth, flat ventral surfaces. The lateral surfaces of the ventral half 
of the centra are approximately parallel dorsoventrally, and thus the 
centra are subrectangular in cross section. These vertebrae cor- 
respond to the dorsal vertebrae of Stereophallodon. 

DISCUSSION 

Ophiacodon and Stereophallodon 

In the review of pelycosaurs presented by Brinkman and Eberth, 
(1983) two major clades were recognized. The first includes Casea, 
Ruthiromia, Varanops, and Aerosaurus. This clade is referred to 
below as the Casea-Varanops clade. It was defined on the basis of 
three derived character-states: 1) the presence of a secondary ridge 
leading from the distal end of the deltopectoral crest to a more 
medial position on the proximal end of the humerus; 2) the presence 
of a contact between the maxilla and quadratojugal with these 
bones raised to form a ridge along the contact and; 3) the presence 
of basipterygoid processes directed laterally and articular surfaces 
elongate mediolaterally. The second includes Ophiacodon, Edapho- 
saurus, Dimetrodon, and Sphenacodon. This clade is referred to 
below as the Ophiacodon- Dimetrodon clade. It is defined on the 
basis of four shared derived characters: 1) a concave ventral cheek 
margin; 2) a posteriorly sloping anterior margin of the premaxilla; 
3) a dorsal process of the stapes that articulates with the ventral 
surface of the paroccipital process of the opisthotic and; 4) an 
extended ventral plate of the angular in the region of the articular. 

The Stereophallodon material at hand includes only one of the 
skeletal regions used in defining the Casea-Varanops clade (the 
basipterygoid processes of the basi-parasphenoid). In Stereophallo- 
don the processes extend ventrad and possess smooth antero- 
posteriorly elongate articulating surfaces. Such a morphology is 
absent in members of the Casea-Varanops clade and had been 
considered a unique derived character of Ophiacodon (Brinkman 
and Eberth, 1983). 

The present Stereophallodon material includes two of the four 
skeletal regions used in defining the Ophiacodon- Dimetrodon clade 
(the premaxilla and the angular). The premaxilla is like those of 
members of the Ophiacodon- Dimetrodon clade in that it does not 



26 BREVIORA No. 485 

project anteriad as is the case primitively in pelycosaurs. The 
angular shares with those of members of the Ophiacodon- Dimetro- 
don clade the presence of an extended ventral plate in the region of 
the articular. Thus these character-states indicate that Stereophallo- 
don is a member of the Ophiacodon- Dimetrodon clade. 

Within the Ophiacodon- Dimetrodon clade, Stereophallodon is 
most parsimoniously placed as the closest sister-taxon of Ophiaco- 
don (Fig. 18) and forms with Ophiacodon a monophyletic taxon 
defined by five characters. Two of these, the ventral orientation of 
the basipterygoid processes and the antero-posteriorly oriented 
articular surfaces have been discussed above and are present 
elsewhere in pelycosaurs only in Ophiacodon. 

Three additional features not discussed by Brinkman and Eberth 
(1983) can be interpreted as derived and shared by only Ophiacodon 
and Stereophallodon. In interpreting the polarity of these charac- 
ters, the methodology used in Brinkman and Eberth (1983) was 
employed. The outgroups used are, 1) members of the Casea- 
Varanops clade, which is the closest sister-taxon of the Ophiacodon- 
Dimetrodon clade and, 2) the taxa that were used as outgroups in 
interpreting the polarities of character-states within pelycosaurs. 
The polarity of character-states was interpreted according to the 
distributions shown in Brinkman and Eberth (1983: Fig. 2). 



Casea Varanops Aerosaurus Ophiacodon Edaphosaurus Dimetrodon Sphenacodon 




Figure 18. Cladogram showing the interrelationships of seven genera of pelyco- 
saurs. From Brinkman and Eberth (1983). 



1985 STEREOPHALLODON AND BALDWINONUS 27 

One of these features is the presence of a ridge that is triangular in 
cross section extending dorsally from the caniniform-tooth buttress. 
This is not present in Edaphosaurus or the sphenacodontines. The 
condition is not known in the members of the Casea-Varanops 
clade, but in other reptiles used as outgroups, including Paleothyris, 
Captorhinus, and Diadectes, no ridge is present. Thus the absence 
of such a ridge is interpreted as primitive for pelycosaurs, and its 
presence is interpreted as a derived feature. Since it is only known in 
Stereophallodon and Ophiacodon, it supports the hypothesis that 
these genera are members of the same clade. 

The second feature is the presence of a notch on the postero- 
medial edge of the quadrate. Although differing in detail, the 
similarity in position of this and the shelf on the medial surface of 
the quadrate of Ophiacodon indicates that they are different 
variants on a single character-state. No other pelycosaur shows a 
notch or shelf in this position. None is present in captorhinids or 
diadectomorphs. The absence of such a notch, therefore, is 
interpreted as primitive for pelycosaurs, and the condition in Stereo- 
phallodon and Ophiacodon is interpreted as a derived character- 
state shared by those two genera. 

The third character-state is the presence of a cultriform process 
that is wide in cross section. In Dimetrodon, Edaphosaurus, 
Varanops, Casea, and Diadectes, the cultriform process is bladelike 
(that is, much higher than it is wide) in cross section. In Stereophal- 
lodon and Ophiacodon, the cultriform process is wider than it is 
high and has a broad U-shaped cross section. The presence of the 
bladelike cultriform process in both the members of the Casea- 
Varanops clade and in Diadectes indicates that it is primitive for 
pelycosaurs. The wide cultriform process is therefore interpreted as 
a derived character state that is shared by Stereophallodon and 
Ophiacodon. 

An alternative hypothesis of relationship that would place Stereo- 
phallodon as a member of the Dimetrodon-Sphenacodon clade is 
weakly supported by two characters: the presence of an enlarged 
caniniform-teeth buttress and two greatly enlarged caniniform 
teeth. Although the Dimetrodon-Sphenacodon clade was not 
discussed by Brinkman and Eberth (1983), a number of shared 
derived characters have been cited by Reisz (1980) and Reisz (in 
press) as defining the clade. These are the presence of 1) an 



28 breviora No. 485 

edentulous step anterior to the caniniform teeth; 2) enlarged 
caniniform teeth; 3) a swollen caniniform tooth buttress; 4) a 
reflected lamina of the angular; 5) a tall septomaxilla; 6) keeled 
dorsal vertebrae; 7) a ventrolateral^ oriented paroccipital process; 
8) a short lacrimal; 9) a tall skull and; 10) a strongly convex 
maxillary margin. The absence in Stereophallodon of an edentulous 
step anterior to the caniniform teeth, a reflected lamina, keeled 
dorsal vertebrae, and ventrolateral^ oriented paroccipital processes 
indicates that Stereophallodon cannot be a member of this clade. 
Furthermore the absence in Stereophallodon of characters that 
define the Edaphosaurus-Dimetrodon-Sphenacodon clade (basi- 
pterygoid articular surfaces at right angles; absence of a shelf 
between the basipterygoid tubera, a well-developed lateral lappet of 
the frontal, a downturned prearticular, a pterygoideus process 
formed by the articular, a quadrate ramus of the pterygoid without 
a tympanic flange, a well-developed lateral process of the supra- 
occipital, lateral excavations on the neural arches, transverse 
processes of vertebrae lacking a ventral web of bone) indicates that 
Stereophallodon is not the sister taxon of the Dimetrodon- 
Sphenacodon clade. Thus Stereophallodon is interpreted as the 
closest sister-taxon of Ophiacodon that has paralleled members of 
the Dimetrodon-Sphenacodon clade in the development of enlarged 
caniniform teeth supported by a large caniniform-tooth buttress. 
This hypothesis of relationship is in agreement with the recom- 
mendation of Reisz (1980) that Stereophallodon be separated from 
the Eothyrididae and placed in the Ophiacodontidae. 

In some characters, Stereophallodon and Ophiacodon are differ- 
ent. Outgroup comparison was used to determine whether the 
character-states that occur in Stereophallodon are primitive for the 
Ophiacodon- Stereophallodon clade or apomorphic for Stereophal- 
lodon. The outgroups used in this analysis are the Edaphosaurus- 
Dimetrodon clade (the sister-taxon of the Ophiacodon- Stereophal- 
lodon clade), the Casea-Varanops clade (the sister-taxon of the 
Ophiacodon- Dimetodon clade), other reptiles (the sister-taxon to 
the pelycosaurs) and diadectomorphs (the sister-taxon to all of 
these). The polarity of character-states was interpreted according to 
the distributions shown in Brinkman and Eberth (1983: Fig. 2). 

One of the differences between Stereophallodon and Ophiacodon 
is in the structure of the centra. In Stereophallodon the dorsal 



1985 STEREOPHALLODON AND BALDWINONUS 29 

vertebrae are rectangular in end view, and have flat ventral surfaces 
and subparallel sides. In Ophiacodon vertebrae of the cervical and 
dorsal region are wedge-shaped in cross section and vertebrae of the 
lumbar region are circular in end view and in cross section. The 
dorsal vertebrae of Casea are like those of Stereophallodon. Those 
of Ruthiromia differ in detail, but are similar to Stereophallodon in 
that the posterior dorsal vertebrae tend to have a flat ventral 
surface. Also, the trefoil shaped cross section seen in the lumbar 
vertebrae of Stereophallodon can be considered a structural ante- 
cedent to the "pinched-in" condition of the lumbar vertebrae of 
Ruthiromia (see Eberth and Brinkman, 1983). Since Casea and 
Ruthiromia are primitive members of the Casea- Varanops clade, 
and Stereophallodon is primitive within the Ophiacodon- Dimetro- 
don clade, the morphology of the centra of the dorsal vertebrae seen 
in Stereophallodon is interpreted as primitive with respect to Ophia- 
codon. Inherent in this conclusion is the interpretation that this 
centrum morphology is primitive for pelycosaurs. 

A second difference between Stereophallodon and Ophiacodon is 
seen in the structure of the femur. The femur of Ophiacodon 
possesses a sharp adductor crest that runs along its postero-ventral 
edge. Stereophallodon differs from Ophiacodon and is like other 
pelycosaurs in that the adductor crest is located in a mid-ventral 
position. The wide distribution of this latter character-state indicates 
that it is a primitive feature in pelycosaurs. Thus in this feature, 
Stereophallodon is more primitive than Ophiacodon. 

A third difference between Stereophallodon and Ophiacodon is 
seen in the structure of the astragalus. In Stereophallodon the 
astragalus has a long neck, in Ophiacodon it is short. The astragalus 
of Ophiacodon has been thought to be primitive, but establishing 
polarities has been difficult (Eberth and Brinkman, 1983). A long 
neck is present in Ruthiromia (the primitive member of the 
Varanops clade), Edaphosaurus (the primitive member of the 
Edaphosaurus-Sphenacodon-Dimetrodon clade), and Stereophallo- 
don. Thus the Stereophallodon character-state is interpreted as the 
primitive condition for pelycosaurs. 

A fourth difference between Stereophallodon and Ophiacodon is 
seen in the structure of the quadrate. The quadrate of Stereophallo- 
don is distinctive in the shape of the articular surface. Edaphosaurus, 
Ophiacodon, the sphenacodontids, Casea, and the varanopseids 



30 breviora No. 485 

each have a different quadrate articular morphology. Thus it is not 
possible to determine which is the primitive pattern using outgroup 
comparison. 

A fifth difference between Stereophallodon and Ophiacodon is 
seen in the structure of the fibula. The fibula of Stereophallodon is 
distinctive in the triangular cross-sectional shape of its proximal 
end. In Ophiacodon, the fibula has a rectangular proximal end. This 
is also the case in Edaphosaurus, sphenacodontines, and the 
varanopseids. Thus the character-state seen in Ophiacodon is 
interpreted as the primitive character-state and that seen in Stereo- 
phallodon is considered apomorphic for that genus. 

Finally, Stereophallodon differs from Ophiacodon in the presence 
of a caniniform-tooth buttress and the large size of the caniniform 
teeth. As discussed above, these are interpreted as derived features 
acquired in parallel with the derived members of the Dimetrodon- 
Sphenacodon clade. 

In summary, Stereophallodon is interpreted as the closest sister- 
taxon of Ophiacodon. Its position in the Ophiacodon- Edaphosaurus- 
Dimetrodon clade is supported by seven shared derived character- 
states. Two of these are shared with all other members of the clade; 
five are derived features shared only with Ophiacodon. The two 
features shared with all other members of the clade are the presence 
of a platelike extension of the angular beneath the articular and the 
posterior slope of the dorsal ramus of the premaxilla. The features 
that unite Stereophallodon with Ophiacodon are the shape of the 
cultriform process of the basi-parasphenoid, the ventrally directed 
basipterygoid process, the anteroposteriorly oriented basipterygoid 
articular surfaces, the presence of a ridge that is triangular in cross 
section extending dorsally from the caniniform-tooth buttress, and 
the presence of a notch on the postero-medial edge of the quadrate. 
Stereophallodon shares five features with Ophiacodon that are 
primitive relative to Edaphosaurus and the sphenacodontines. 
These are the presence of a triangular process located medial and 
anterior to the articular, the presence of a tympanic flange on the 
ventral edge of the quadrate ramus of the pterygoid, the absence of 
an anterior process on the ilium, the presence of a shelf covering the 
anterior end of the groove between the basisphenoid wings, and the 
absence of a retroarticular process. Stereophallodon is primitive 
with respect to Ophiacodon in retaining the following characters: 



1985 STEREOPHALLODON AND BALDWINONUS 31 

dorsal vertebrae that have flattened ventral surfaces and subparallel 
sides, a femur with an adductor crest running along the mid-ventral 
surface of the bone, and a long neck on the astragalus. Stereophallo- 
don is derived relative to Ophiacodon in the presence of two greatly 
enlarged caniniform teeth supported by a caniniform-tooth buttress, 
and a fibula that has a triangular cross section at its proximal end. 

Baldwinonus 

Before considering the systematic position of Baldwinonus it is 
necessary to consider whether or not more than one individual is 
present in the type specimen. This was questioned by Reisz (1980), 
who identified the centra as ophiacodont elements, and the maxilla 
and neural spines as sphenacodont. Thus Reisz concluded that at 
least two individuals were present. 

In identifying the maxilla as that of a sphenacodont, rather than a 
sphenacodontine — the only group of sphenacodonts known from 
New Mexico — Reisz recognized that it differs significantly from the 
maxilla of other better known sphenacodontines. Also, in identify- 
ing the vertebrae as ophiacodont vertebrae Reisz recognized that 
they were not from Ophiacodon, the only ophiacodontid present in 
New Mexico. In the features in which the vertebrae and maxilla of 
Baldwinonus are different from the sphenacodontines and Ophiaco- 
don, they are like Stereophallodon. The quadrate of Baldwinonus, 
not considered by Reisz, is also like Stereophallodon and unlike the 
quadrate of Ophiacodon or the sphenacodontines. While studying 
the type specimen, it was possible to piece together many of the 
vertebral fragments. Some of the resulting vertebrae are in articula- 
tion, and in one case a neural spine, identified as a sphenacodont 
neural spine by Reisz, could be attached to a centrum. Thus, it is 
concluded that the type specimen of Baldwinonus is most probably 
a single individual that is morphologically similar to Stereophal- 
lodon. 

In most features in which both Baldwinonus and Stereophallodon 
are known, Baldwinonus is simply a smaller version of Stereophallo- 
don. A difference in the structure of the maxilla is present: in Stereo- 
phallodon, the palatine sutural surface extends onto the caniniform- 
tooth buttress; in Baldwinonus, the palatine sutural surface ends 
just posterior to the caniniform-tooth buttress. On the basis of this 
difference, Baldwinonus is considered a valid genus. 



32 breviora No. 485 

Two derived character-states that unite Stereophallodon and 
Ophiacodon are also present in Baldwinonus; therefore, the three 
genera can be combined in a monophyletic group. These are the 
presence of the rodlike dorsal extension of the caniniform-tooth 
buttress on the internal surface of the maxilla and the presence of a 
notch in the postero-medial surface of the quadrate. 

That Baldwinonus and Stereophallodon may be more closely 
related to each other than either is to Ophiacodon is indicated by the 
shared derived character-state of greatly enlarged caniniform teeth 
supported by a buttress on the internal surface of the maxilla. Other 
character-states of Stereophallodon that were interpreted as being 
derived, specifically the structure of the proximal end of the fibula, 
are not known in Baldwinonus. 

Character-states shared by Baldwinonus and Stereophallodon 
that can be considered primitive for pelycosaurs in general are seen 
in the structure of the vertebrae. The vertebrae of Baldwinonus are 
like those of Stereophallodon in that the dorsal vertebrae have flat 
ventral surfaces and subparallel sides. Thus the discussion of the 
polarity of the vertebral features of Stereophallodon given above 
also applies to Baldwinonus. 

The structure of the maxilla of Baldwinonus is more fully known 
than that of Stereophallodon. Like the maxilla fragments of Stereo- 
phallodon, it is only superficially similar to the sphenacodontines. 
The differences include the absence of the following derived 
sphenacodontine characters: a maxillary step, a reduced tooth 
count, compressed teeth with fully developed cutting edges, an 
evenly tapering caniniform-tooth buttress, and a rectangular post- 
buttress alveolar ridge. 

In summary, from the characters discussed here, it is concluded 
that Baldwinonus is closely related to Stereophallodon. These genera 
are interpreted as primitive members of a clade that also includes 
Ophiacodon, and therefore replace Ophiacodon as the most 
primitive known member of the Ophiacodon- Dimetrodon clade 
(Fig. 19). Character-states shared by Stereophallodon, Baldwinonus, 
and primitive members of the Casea-Varanops clade can be 
considered primitive for pelycosaurs. One such feature is the 
structure of the centra of the dorsal vertebrae. This is of significance 
in providing evidence that pelycosaurs, as traditionally defined, had 
a monophyletic origin. 



1985 STEREOPHALLODON AND BALDWINONUS 33 



Casea Ruthpromia Aerosaurus Voranops Ophiacodon Stereophallodon Baldwinonus Edaphosaurus Sphenacodon Dimetrodon 




Figure 19. Cladogram showing the interrelationships of Stereophallodon, Bald- 
winonus and the pelycosaurian genera considered by Brinkman and Eberth (1983) 
and Eberth and Brinkman (1983). 



ACKNOWLEDGMENTS 

The authors are greatly indebted to the scientific and curatorial 
staff of the American Museum of Natural History for permission 
to study specimens in their care. The manuscript was read by Phil 
Currie. His critical comments were greatly appreciated. The 
specimen drawings, except for Figures 1C-E, 5, and 7A-B, were 
made by Linda Krause of the Tyrrell Museum of Palaeontology. 
Figures 1C and 7A-B were made by Donna Sloan of the Tyrrell 
Museum. The manuscript was typed by Judy McMillan. The 
comments of two anonymous reviewers also greatly improved the 
manuscript. The responsibility for any errors in methodology or 
conclusion rest solely with the authors. 



LITERATURE CITED 

Brinkman, D. B., and D. A. Eberth. 1983. The interrelationships of pelycosaurs. 

Breviora No. 473, 35 pp. 
Eberth, D. A., and D. B. Brinkman. 1983. Ruthiromia elcobriensis, a new 

pelycosaur from El Cobre Canyon, New Mexico. Breviora, No. 474, 26. pp. 
Langston, W., Jr. 1965. Oedaleops campi (Reptilia: Pelycosauria). A new genus 

and species from the Lower Permian of New Mexico and the family 

Eothyrididae. Bull. Texas Mem. Mus., 9: 1-46. 



34 breviora No. 485 

Reisz, R. R. 1980. The Pelycosauria: a review of phylogenetic relationships, pp. 

553-592. In A. L. Panchen (ed.), The Terrestrial Environment and the Origin of 

Land Vertebrates, London and New York, Academic Press. 

. Pelycosauria. Handbuch der Palaeoherpetologie. Fischer, Stuttgart. 

Romer, A. S. 1937. New genera and species of pelycosaurian reptiles. Proc. New 

Engl. Zool. Club, 16: 89-96. 
Romer, A. S., and L. I. Price. 1940. Review of the Pelycosauria. Geol. Soc. Am. 

Spec. Pap. No. 28, 538 pp. 
Vaughn, P. P. 1958. On a new pelycosaur from the Lower Permian of 

Oklahoma, and the origin of the family Caseidae. J. Paleont., 32: 981-991. 
Watson, D. M. S. 1954. On Bolosaurus and the origin and classification of 

reptiles. Bull. Mus. Comp. Zool., Ill: 299-249. 
. 1957. On Millerosaurus and the early history of the sauropsid reptiles. 

Phil. Trans. Roy. Soc. Lond., B, 240: 325-400. 



BREVIORA 

Museum of Comparative Zoology 

US ISSN 0006 9698 

Cambridge, Mass. 29 August 1986 Number 486 



THELODVS MACINTOSHI STETSON 1928, 

THE LARGEST KNOWN THELODONT 

(AGNATHA: THELODONTI) 

Susan Turner 1 

Abstract. Material attributed to the thelodont Thelodus macintoshi Stetson, 
from the Silurian of New Brunswick, Canada, is shown to be heterogeneous. It 
comprises not only the type thelodont, T. parvidens Agassiz 1839, but also a species 
of Loganella similar to L. ludlowiensis or L. martinssoni (Gross 1967) and an 
acanthodian resembling Gomphonchus. Scales of T. bicostatus Hoppe and T. trilo- 
batus Hoppe are found in association with those of T. parvidens for the first time. 
Measurements of Stetson's type material confirm that T. parvidens is the largest 
known thelodont, with a total length of about one meter. The vertebrate fauna 
including these thelodonts could be as old as late Llandovery (within the Long Reach 
Formation) or as young as early Pridoli (in the Jones Creek Formation). The New 
Brunswick thelodonts are similar in preservation to those from the Lower Silurian 
fish beds of Scotland, but their phylogenetic affinities are closer to those from the 
Silurian of England and the Baltic region. 

INTRODUCTION 

In 1928 Harold Stetson described articulated thelodont material, 
preserved in calcareous concretions, from a site in the Silurian rocks 
near Nerepis, Kings Co., New Brunswick, eastern Canada. This was 
the first definite thelodont material found in North America. The 
site, at Cunningham's Brook (Cunningham's Creek on the map of 
MacKenzie 1964), has also yielded the heterostracan Cyathaspis 
acadica (Matthew), (see Denison 1964), spines of an acanthodian 
related to Climatius (Denison 1956, p. 385), and a possible anaspid, 

V&^^ 

'Queensland Museum, Fortitude Valley, Queensland 4006, Australia 



. 



TV 



2 breviora No. 486 

Ctenopleuron nerepisense Matthew 1907, which remains indetermi- 
nate (e.g., Woodward 1920). Associated invertebrates include cera- 
tiocarids, conodonts and a xiphosuran (Denison 1956; Blieck 1982). 

MATERIALS AND METHODS 

The nodules containing this thelodont material were collected 
from siltstones by Stetson and W. E. Schevill of the Museum of 
Comparative Zoology during a Harvard expedition in 1927, follow- 
ing up the lead of William Macintosh of the Natural History 
Society of New Brunswick. The type material is housed at the 
Museum of Comparative Zoology, Harvard University. 

Subsequently, during the 1950s and 1960s, Robert Denison (then 
of the Field Museum of Natural History, Chicago) collected more 
material from two separate horizons. Denison referred all the thelo- 
donts he collected to Thelodus macintoshi Stetson. In 1956 he stated 
that the specimens came from the Long Reach Formation, follow- 
ing the lead of Matthew (1888) who stated that the fish beds were 
in the Mascarene Group, Division 2 which was subsequently called 
the Long Reach Formation (e.g., MacKenzie 1951). In 1964 Deni- 
son corrected his statement by referring the Cyathaspis acadica 
material to the Jones Creek Formation, on the basis of MacKenzie 's 
1951 field appraisal. This interpretation was followed by Blieck 
(1982). 

Evidence, however, from the 1964 maps of the region by Mac- 
Kenzie suggests that all the fish beds are within the older Long Reach 
Formation. The position of the two formations has been confused 
because in the 1950s, and on the 1964 Saint John and Hampstead 
maps, MacKenzie placed the Jones Creek Formation below the 
Long Reach Formation. In fact, the reverse order is the case (Berry 
and Boucot 1970; McCutcheon 1981). One recent source places the 
fish beds definitely within the Long Reach Formation (Smith 1966). 
In the notes of the 1964 Hampstead map there is even a reference to 
a new locality in the Long Reach Formation for "a primitive fish" in 
a brook, northeast of Armstrong Corner. This specimen was not 
identified and I have no further information about it. It may be that 
the cyathaspids did not come from the same horizon as the thelo- 
donts and other elements of the fauna. 

In the early 1970s, Denison sent me, on request, a sample of loose 
scales from one of the nodules from the New Brunswick site. I 



1986 THELODUS MACINTOSHI 3 

attempted to make thin sections, but the histological structure of the 
scales was disrupted by post-mortem algal or fungal attack, or 
recrystallization (see PI. 1). In general shape, however, the scales 
seemed identical to those of T. parvidens. I also examined a speci- 
men of T. macintoshi in the British Museum (Natural History), 
collected from New Brunswick by W. Graham-Smith in 1937. On 
the basis of these investigations, I suggested that T. macintoshi was 
virtually identical to T. parvidens, at least in its scale morphology 
(Turner 1973, 1976). [I also stated that the Long Reach Formation 
was the stratotype, following Denison 1956; see also Blieck 1982]. 
From a perusal of Stetson's figures in 1968 I decided to write a 
paper on the closure of Iapetus, the early Palaeozoic ocean (Turner 
1970). For if T. parvidens and T. macintoshi were the same species, 
then New Brunswick and the Anglo-Welsh region must have been 
closely aligned in the late Silurian. 

In his original work Stetson (1928, Figs. 1-3) described and fig- 
ured scales he thought looked like typical T. parvidens scales but 
which bore long thin extensions on the posterolateral rims of the 
crowns (Figs. 2, 3). This character prompted him to distinguish T. 
macintoshi as a separate species. In spring 1983 I examined both 
Stetson's and Denison's collections. Specimens PF are housed at the 
Field Museum of Natural History, Chicago (FMNH), MCZ at the 
Museum of Comparative Zoology, and BMNH at the British 
Museum of Natural History. 

MATERIALS EXAMINED 

MCZ 2035 HOLOTYPE Thelodus macintoshi Stetson 1928 is 
T. parvidens Agassiz with T. trilobatus scale. Area at 
least 30 + X 250 mm. Holotype includes thin 
sections. 

MCZ 2037 Paratype T. macintoshi is T. parvidens. 

MCZ 2036 Paratype T. macintoshi is Loganella cf. L. ludlo- 
wiensis (Gross 1967). 

MCZ 13007 in part to 13015, includes the specimens from which 
some thin sections were cut, are referred to T. parvid- 
ens Agassiz. 

MCZ 13007 in part; one nodule has Gomphonchus type scales. 

MCZ 13014 includes T. bicostatus (Hoppe 1931) scales. 

MCZ 13012 is at least 240 mm long. 



4 BREVIORA No. 486 

PF 1805 T. macintoshi is T. parvidens, area 300 X 170 mm 

(Denison coll.). 
PF 1809 T. macintoshi is T. parvidens horizon A, area at least 

300 X 170 mm (Denison coll.). 
PF 1804 T. macintoshi is T. parvidens, horizon B (Denison 

coll). 
PF 1808 loose thelodont scales in coprolite includes T. par- 

videns, T. bicostatus and T. trilobatus, horizon B. 
PF 1803 is a coprolite containing Gomphonc hus type scales. 

PF 1802 is a coprolite with thelodont scales, T. parvidens, 

horizon B. 
PF 3475 jumbled T. parvidens scales in coprolite, (Denison 

coll. 1961). 
PF 3476 T. macintoshi - a good articulated T. parvidens, 

incomplete 150 X 90 mm in area (Denison coll. 

1961). 

PF 3477 and PF 1801 are acanthodians and PF 1698 is an acantho- 
dian ?spine. At least three uncatalogued specimens, including 126-52 
and 1969 Denison coll. FMNH, are coprolites from horizon B con- 
taining well-preserved T. parvidens scales. 

BMNH P.52444. T. macintoshi (Graham-Smith coll.) is T. parvid- 
ens with T. trilobatus? scales. 



Plate 1. Histology of thelodont scales from coprolitic nodules from Cunning- 
ham's Brook, Nerepis, New Brunswick (slides prepared at Field Museum, Chicago). 

A) Vertical section through large scales of T. parvidens showing growth lines in 
orthodentine and invading hyphae or boring algae; anterior to left, slide 4187; 

B) Small scale of T. parvidens in sagittal section interposed between two larger scales 
of T. parvidens, slide 4187; C) Sagittal section through a scale of T. parvidens with 
a much enlarged anterior base (in the manner of T. laevis Pander (see Gross 1967; 
Karatajute-Talimaa 1978), and the pulp canal; anterior to left, slide 4192; D) Young 
scale of Thelodus trilobatus with large open pulp cavity, in sagittal section; anterior 
to right, slide 4192; E) Scale of Thelodus bicostatus in transverse vertical section 
showing growth lines, slide 4187. F) Scale of T. bicostatus or T. trilobatus showing 
growth lines and invading hyphae or algae, slide 4187; G) Horizontal section 
through crown of T. parvidens scale, slide 4192. All to same size, approx. X60. 



1986 



THELOD US MA CIN TOSH I 




BREVIORA 



No. 486 




Figure 1. Thelodus macintoshi Stetson 1928= Thelodus parvidens Agassiz 1838. 
Holotype, MCZ 2035. Close dotting: small T. parvidens scales in part, towards the 
sides of the heart-shaped nodule (probably the region of the pectoral fins). Larger 
scales in center; good articulated patches shown. Asterisk mark: one scale of T. 
trilobatus. Parallel bars: cracks in the nodule with slickensides. Clear rectangles: 
specimen labels. Arrow points to anterior. 



kESULTS 

Material. Examination of Stetson's specimens and his thin sec- 
tions (to which the 1928 plates do not do justice), and of Denison's 
material, reveals that the material attributed to T. macintoshi is 
heterogeneous: it comprises at least two genera of thelodonts, and 
one specimen is actually an acanthodian. T. macintoshi (s.s.) is 
almost certainly synonymous with T. parvidens. I propose that T. 
macintoshi Stetson should be formally considered a junior synonym 
of T. parvidens, but I shall refer to T. macintoshi in an informal 
sense to distinguish the New Brunswick material from other 
material. 



1986 



THELODUS MACINTOSH I 




Figure 2. Isolated scales from nodule PF 9603 from horizon B of Denison, 
Cunningham's Brook, Nerepis, New Brunswick. A) T. parvidens, lateral view show- 
ing neck riblets, PF 9603. 1 ; B) T. parvidens, ventral view, PF 9603.2; C) T. bicos- 
tatus, dorsal view of broken crown, PF 9603.3; D) T. trilobatus, ventral and dorsal 
views, PF 9603.4; E) T. trilobatus, lateral and dorsal views, PF 9603.5; F) T. 
trilobatus, dorsal view of smooth crown, PF 9603.6; G) T. trilobatus, lateral and 
ventral views, PF 9603.7. 



BREVIORA No. 486 




Figure 3. Scales on paratype of T. macintoshi, MCZ 2036. A) Loganella sp., 
two scales seen in cross section; B) Loganella sp., body scale in crown view showing 
unbroken posterolateral spinelets; CD) Loganella sp., body scales in crown view 
with small anterior basal process and posterolateral neck spinelets. 

The scales of the T. macintoshi specimens are large, up to 1.5 mm 
long, and in an advanced stage of growth with well-developed bases 
(see Fig. 2A, PI. 1A-C); they are undoubtedly scales of a mature 
animal. They differ from European examples of T. parvidens 
(including the type specimen) in the clear expression of the numer- 
ous riblets on the neck region (PI. 2). However, as most European 
scales are waterworn to some degree it seems possible that the full 
extent of neck ribbing in the type species has never been clearly seen 
in the isolated scales. Gross did show one scale with 12 riblets on 
one side of the neck (1967, PI. 1, Fig. 3 A) but their number is usually 
much smaller. The slight differences in rib ornament on the neck of 
scales may reflect local variation within a single population of The- 
lodus parvidens. 

Furthermore, some of the specimens, including coprolitic masses 
and thin sections, show that scales of T. bicostatus type are part of 
the T. parvidens squamation (see Fig. 2). Despite the fact that the 
scales of T. bicostatus are very distinctive, both Gross (1967) and 
Turner (1973, 1976) thought they might be special scales of T. par- 
videns, while retaining the specific name bicostatus. One scale on 
the type specimen (MCZ 2035) appears to belong to T. trilobatus 
(see Fig. 1). A patch of small striated scales among the larger rhom- 
boid T. parvidens scales on specimen BMNH P52444 could also 
belong to T. trilobatus. Scales of this species are also seen in copro- 
litic masses (Fig. 2D-G). Thin sections of scales from the nodules 



1986 



THELODUS MAC IN TOSH I 




Plate 2. Scales of Thelodus parvidens from coprolitic nodule from Cunningham's 
Brook, Nerepis, New Brunswick. A) Detail of neck in lateral view showing riblets 
on lower neck, PF 9603.8, approx. X60; B) Scale in lateral view, PF 9603.8, 
approx. X45; C) Scale in posterobasal view showing pulp cavity, PF 9603.10, 
approx. X45. 



confirm the presence of T. parvidens in association with T. bicosta- 
tus and T. trilobatus (see PI. 1). Thus the T. macintoshi material 
confirms the synonymy of T. parvidens, T. trilobatus and T. bicos- 
tatus postulated initially by Gross (1967). 

The most interesting discovery in this re-examination of Stetson's 
type material is the presence of at least one specimen of a loganiid. 
The paratype of T. macintoshi (MCZ 2036) is not a thelodontidid, 
but a specimen of Loganella sp. cf L. ludlowiensis (Gross 1967) or 
perhaps cf L. martinssoni (Gross 1967). [The genus name Logania 
formerly attributed to this and other loganiid species is preoccupied 
{Logania Distant — Lepidoptera; Whitley 1976). Here I propose to 
use the replacement name Loganella.] The figures in Stetson's paper 
depicting scales with rim spines actually represent sagittal sections 
through Loganella scales (see Fig. 3). The scales in the nodule are 
sometimes broken through, exposing the slit-like pulp canal and 



10 BREVIORA No. 486 

navicular shape of typical loganiid body scales (cf Gross 1967 e.g., 
Figs. 11 I-K). 

At least two specimens from Nerepis, one in each of the collec- 
tions at MCZ and the Field Museum, are not thelodonts but 
acanthodians. The scales are those of an ischnacanthid cf 
Gomphonchus. 

It is not really difficult to see why Stetson did not recognise the 
different thelodonts in his fauna. Before 1928 only a few cross- 
sections of Thelodus scales, except those of T. parvidens, had been 
illustrated. Knowledge of the histological structure of scales did not 
become refined until the comprehensive study by Gross (1967), in 
which he separated the genus Logania from Thelodus. The MCZ 
houses the thin sections Stetson used; those cut from MCZ 2036 
show the teardrop-shaped outlines of Loganella scales cut in horiz- 
ontal section, with a slit-like pulp cavity. Nine slides of nodule 
pieces containing scales of T. macintoshi are housed at the Field 
Museum. 

In the main, these scales are well-preserved and the sections show 
clearly that the scales belong to T. parvidens: some sections of the 
scales are mere "ghosts," the internal structure of the dentine having 
been recrystallised. One slide, PF slide 4187/52-102b, contains 
cross-sections of two scales of T. bicostatus (Hoppe 1931) (see PI. 
IE). Many of the scales of T. parvidens, including the bicostatus and 
trilobatus forms, exhibit clear incremental lines in the dentine; there 
are from nine to 1 1 in large scales and two to three in small scales 
(see PI. 1). The incremental lines in thelodont scales are a measure of 
scale growth (Gross 1967). Whether the lines represent annual or 
seasonal increments may never be ascertained; if the animals were 
living in warm near-equatorial waters, as has been suggested for 
Siluro-Devonian vertebrates on other occasions (e.g., Halstead and 
Turner 1973), then it seems possible that the lines represent seasonal 
fluctuations in the availability of calcium and/ or phosphate. Some 
scales are also penetrated by what look like fungal hyphae, similar 
to those described by Goujet and Locquin (1977) and Karatajute- 
Talimaa (1978) (see PI. 1 A, F). 

Size of T. parvidens. Several of the nodules (e.g., those contain- 
ing the holotype, see Fig. 1) contain articulated squamation which 
indicates that the thelodonts were very large. Stetson was correct in 
estimating that T. macintoshi was longer than Tur inia pagei (Powrie 



1986 THELODUS MAC1NTOSHI 11 

1870), the largest known thelodont at that time. He considered that 
some eight to ten inches would need to be added to the length of the 
holotype Turinia, making an estimate of 22 inches (around 550 mm) 
for T. macintosh!. Study of the type specimen shows that scales 
towards the center are very large, around 1.5 mm square, and 
closely packed in neat diagonal rows, whereas towards the antero- 
lateral margins the scales are progressively smaller, around 0.5 mm 
square and along the lateral margins the very small scales are much 
disrupted (Fig. 1). I suggest that, by comparison with the type spec- 
imen of Turinia pagei (see Turner 1982), the type specimen exhibits 
part of the cephalothorax extending to the region of the proximal 
tips of the pectoral fins. The specimen measures about 300 mm long 
by 250 mm wide. Given that the cephalothorax was probably about 
one quarter to one third of total body length, this New Brunswick 
thelodont might have had a length between 900 and 1200 mm. Other 
nodules in the collections examined show areas of squamation at 
least as large as that in the type specimen. With a length around one 
meter, these specimens of T. parvidens are the largest thelodonts 
found to date. This more than confirms Stetson's prediction about 
the size of the animals. 

Age of the Nerepis fauna. Matthew (1888) considered the fish- 
bearing beds to be of Niagaran age because of the associated fauna, 
which includes Ceratiocaris pusillis and the xiphosuran Bunodella 
horrida (Denison 1956). Bailey and Mclnnes (1888) reported that 
Matthew regarded the beds to be about the same age as those con- 
taining Palaeaspis in the United States, that is, the Medina and 
Clinton Groups, and equivalent to Divisions 2 and 3 of the Anticosti 
Group and Groups B and B 1 of Arisaig. Westoll (1958a) placed the 
beds within the Wenlock, equivalent to the Lockport of the United 
States. 

Both the Long Reach and Jones Creek Formation beds lie within 
the Mascarene Group, referred to as the Upper Silurian by the 
Canadian Geological Survey (MacKenzie 1964). When MacKenzie's 
maps were published in 1964 it was still not certain whether the 
Jones Creek Formation lay above, or below, the Long Reach For- 
mation. The latter has been dated as Upper Llandovery/ Lower 
Wenlock (C6), because of its Costistricklandia-Eocoelia community 
(Berry and Boucot 1970). MacKenzie (1951) had placed the Long 
Reach Formation above the Jones Creek Formation, but Berry and 



12 BREVIORA No. 486 

Boucot, following a statement by MacKenzie, considered that it 
must rest below and that the Jones Creek Formation was of Lud- 
low/ Pridoli age (see also Blieck 1982). McCutcheon ( 1981) has reit- 
erated this interpretation, which would imply a large time gap 
between the two formations. I am not able to ascertain from the 
literature if this hiatus is discernible in the field. On their correlation 
chart Berry and Boucot (1970) show the Long Reach Formation 
extending from ?Upper Llandovery to Ludlow, but based on the 
brachiopod fauna, they prefer a Late Llandovery/ Early Wenlock 
age. The Jones Creek Formation is possibly as old as Upper Ludlow 
(Ludfordian), for, as Berry and Boucot point out, the beds contain a 
Salopina community which is thought to signify Ludlow shallow 
water conditions in the Welsh Borderland (Lawson 1975). 

Berry and Boucot compared the Jones Creek Formation with the 
Pembroke Formation of Maine. Devonian fish material has been 
discovered in the Eastport Formation of this region by Denison 
(Field Museum collections) but is not described yet. As the strati- 
graphy and structure of the Nerepis area seem complex, perhaps a 
more thorough search for fish remains in equivalent Siluro- 
Devonian beds nearby in Canada and in the United States would 
help clarify the succession. 

The association of cyathaspids, acanthodians and T. parvidens 
(including trilobatus and bicostatus varieties) and a Loganella sp. cf. 
L. ludlowiensis in New Brunswick could be as old as early Wenlock 
and no younger than early Downton. A similar assemblage of these 
thelodont species and acanthodian scales has been found in sedi- 
ments as old as Lower Wenlock in the Welsh Borderland (Turner 
1973) and also in the Upper Llandovery of Norway (Turner 1984), 
and it is also found throughout the Upper Ludlow and early Down- 
ton of Europe (Turner 1973; Karatajute-Talimaa 1978; Marss 
1982b). Marss (1982b) suggests that the pre-late Ludlow loganiid 
in the Welsh Borderland is in fact L. martinssoni. The possibility 
that the New Brunswick loganiid may belong to this latter species 
cannot be ruled out. Cyathaspidids are known from supposed 
Upper Llandovery and Wenlock sequences in the Canadian Arctic 
but details of these early forms are not yet published (Denison 1964; 
Thorsteinsson 1967; Dineley and Loeffler 1976). In their review of 
cyathaspids Dineley and Loeffler (1976) compared Cyathaspis acad- 
ica with C. banksi, known from the Ludlow and early Downton of 



1986 THELODUS MACINTOSHI 13 

the Welsh Borderland. It seems likely, however, following the rea- 
soning of Elliott (1978) and of Dineley and Loeffler, that cyathas- 
pids occur earlier in Canada than in Europe. More work on the 
Canadian cyathaspids might help clarify their relationships and 
biostratigraphic significance. 

DISCUSSION 

Implications. As I predicted (Turner 1970), T. maeintoshi is 
very closely related if not identical to the type thelodont T. parvid- 
ens. Also the type material contains T. bicostatus and T. trilobatus. 
These three scale forms, T. parvidens, T. bicostatus and T. triloba- 
tus, are elements of the Baltic-Anglo-East Canadian Silurian fauna 
(the T. parvidens assemblage of Turner 1973). This assemblage may 
represent a facies fauna indicative of the onset of "red bed" 
conditions. 

If the thelodonts do come from the Long Reach Formation and 
the age of the fish beds is confirmed as early Wenlock, or even late 
Llandovery, this will add weight to the predicted occurrence of T. 
parvidens throughout the southern Laurasian region in the early 
Silurian. The appearance of T. parvidens before the late Wenlock is 
disputed by Karatajute-Talimaa (1978), who has not found it in any 
of the older localities which she has investigated. However, the fact 
that T. parvidens did appear in the late Llandovery is confirmed by 
its discovery in the Norwegian succession. If this is the oldest occur- 
rence then there must have been the possibility of dispersal of T. 
parvidens between the west Baltic, the Welsh Borderland and New 
Brunswick. This would imply a shallow water connection between 
at least two of these three regions in mid-Silurian times. 

The T. parvidens assemblage could also represent a cline, with the 
T. maeintoshi form to the "west" of the range in New Brunswick, 
and forms such as T. costatus (Pander 1856), T. sculp tilis Gross 
1967, and T. admirabilis Marss 1982 to the "east" in the southern 
Baltic — even, perhaps, including T. marginatus Karatajute-Talimaa 
1978. T. trilobatus, T. bicostatus, and T. pugniformis Gross 1967 
would be included in T. parvidens as varieties because all possess a 
similar histological structure. These thelodonts would be placed 
within Blieck's Ichthyofacies IIA (Blieck 1982). Blieck (1982, Fig. 5) 
envisaged the New Brunswick site as located within the Appalachian 
channel with a possible link to the Welsh Borderland and elsewhere 



14 BREVIORA No. 486 

in western Europe via a narrow shallow water passage. New Bruns- 
wick was almost certainly in connection with the Anglo-Welsh 
cuvette, Norway and probably elsewhere in the Baltic hinterland at 
some point during late Llandovery to early Downton times (see also 
Turner and Tarling 1982). 

Scales of T. parvidens have also been found in the Ludlow Moy- 
dart Formation of Nova Scotia ((/)rvig in Boucot et al. 1974) and 
those of "Thelodus sp." in the Upper Silurian (Lower Devonian?) 
Oriskany Sandstone of Nictaux Falls, Nova Scotia (Eastman 1907, 
Gardiner 1966). Thus there are indications that the T. parvidens 
fauna was quite widespread in eastern Canada by late Silurian 
times. 

Environment. Denison (1956); Robertson (1957); and White 
(1958) considered that the fauna at Nerepis indicated a marginal 
marine or non-marine environment. From the state of preservation 
it would seem that the animals were living in or near a quiet back- 
water, possibly asupratidal pool or a lagoon. The style of preserva- 
tion in the New Brunswick fish beds is virtually identical to that in 
the Lower Silurian fish beds of southern Scotland (see e.g., Ritchie 
1968). At Lesmahagow and Hagshaw thelodonts, anaspids, ceratio- 
carids, eurypterids and rare xiphosurans are found, often contained 
in nodules (considered to be coprolitic) within black shales. Some 
near-complete thelodonts are preserved in large nodules, and 
numerous scales occur in the center of nodules, probably the 
remains of consumed thelodonts. At Cunningham's Brook these 
coprolitic masses seem to be confined to one horizon (Denison's 
horizon B). There is little or no evidence of transport seen on any of 
the specimens; thelodont scales are not noticeably waterworn and a 
range of scale size is seen in the disarticulated scale masses in 
nodules as well as in articulated squamation (see Figs. 1, 2). 

Denison ( 1 964) discussed the state of preservation of the cyathas- 
pids and found that one specimen, also found in a coprolite, was a 
juvenile on the evidence of its thin plates and scales. This also sug- 
gests that the fish were living and dying in a low-energy 
environment. 

The large size of the thelodonts in the Nerepis fauna suggests that 
this was an ideal environment for these creatures which until now 
have invariably been thought of as small insignificant agnathans. 



1986 THELODUS MACINTOSH! 15 

The thelodonts in the Lower Silurian of Scotland, belonging to 
Loganella scotica Traquair 1898, which seem to have lived in a 
similar environment, also exhibit a range of size from a few centime- 
ters (juveniles) to quite large individuals (up to an estimated 400 
mm). If similarity of faunal components and style of preservation 
are taken into account then the Nerepis site and Scottish lower fish 
beds could be of equivalent age, which would be late Llandovery to 
early Wenlock (e.g., Rolfe 1973). 

ACKNOWLEDGMENTS 

I would like to thank Mrs. Ruth Romer, Dr. and Mrs. R. H. 
Denison, Dr. W. E. Schevill and M. C. Schaff for their help and 
hospitality in Cambridge, Mass. Dr. Denison made helpful com- 
ments upon the manuscript. Dr. and Mrs. John Bolt, J. Clay 
Bruner, and Dr. and Mrs. W. Turnbull gave me their help and 
hospitality while I was in Chicago. The opportunity to examine the 
Field Museum material was provided by the award of a grant from 
the Thomas Dee Visiting Scientist Fellowship. Dr. Godfrey S. Now- 
lan helped me to obtain information on the geology of New Bruns- 
wick. Dr. Roger Miles allowed me to examine and cast the BMNH 
specimen then under his care. Part of this work was done under the 
auspices of an Honorary Research Fellowship of the Queensland 
Museum, for which I thank Dr. Alan Bartholomai and the Board of 
Trustees. 

LITERATURE CITED 

Agassiz, J. R. L. 1839. Fishes, p. 605. In R. I. Murchison. The Silu- 
rian System, founded on geological researches in the counties of Salop, Here- 
ford, Radnor, Montgomery, Caermarthen, Brecon, Pembroke, Monmouth, 
Gloucester, Worcester, and Stafford; with descriptions of the coal-fields and 
overlying formations. London: John Murray, xxxii + 768 pp. 

Bailey, L. W., and W. McInnes. 1888. Report on explorations and surveys in 
portions of northern New Brunswick and adjacent areas in Quebec, and in 
Maine. United States Annual Report, Geol. Natural Hist. Surv. Can. Ill, part II, 
Report M: 1-52. 

Berry, W. B. N., and A. J. Boucot. 1970. Correlation of the North American 
Silurian rocks. Geol. Soc. Am. Spec. Pap. No. 102: 1-289. 

Blieck, A. 1982. Les grandes lignes de la biogeographie des Heterostraces du 
Silurien superieur-Devonien inferieur dans le domaine nord-atlantique. 
Palaeogeography, Palaeoclimatology, Palaeoecology, 38: 283-316. 



16 BREVIORA No. 486 

Boucot, A. J., J. F. Dewey, D. L. Dineley, R. Fletcher, W. K. Fyson, J. G. 

Griffin, C. F. Hickox, W. S. McKerrow, and A. M. Ziegler. 1974. The 

Geology of the Arisaig area, Antigonish County, Nova Scotia. Geol. Soc. Am. 

Spec. Pap. No. 139: 1-191. 
Denison, R. H. 1956. A review of the habitat of the earliest vertebrates. Fieldi- 

ana. Geology, 11: 359-457. 
1964. The Cyathaspididae, a family of Silurian and Devonian jawless 

vertebrates. Fieldiana, Geology, 13: 311-473. 
Dineley, D. L., and E. J. Loeffler. 1976. Ostracoderm faunas of the Delorme 

and associated Siluro-Devonian formations, Northwest Territories, Canada. 

Spec. Pap. Palaeont. No. 18: 1-218. 
Eastman, C. R. 1907. Devonic fishes of the New York formations. Memoir of the 

N.Y. State Mus. Nat. Hist., Albany, 10: 1 235. 

Elliott, D. K. 1978. New Pteraspididae from the Canadian Arctic, p. 27. PADS 

Abstracts. International Symposium on the Devonian System 1978. The 

Palaeontological Assoc, Univ. Bristol: 61 pp. 
Gardiner, B. G. 1966. Catalogue of Canadian fossil fishes. Roy. Ont. Mus. Life 

Sciences, Contrib. No. 68: 1-154. 
Goujet, D., and M. V. Locquin. 1979. Decouverte de spores fongiques dans les 

ecailles de poissons et d'agnathes paleozoiques: Mycobystrovia lepidophaga 

gen. et sp. nov. 104e Congres National des Societes Savantes, Bordeaux, 

sciences, Fascicule 1: 87-99. 
Gross, W. 1967. Uber Thelodontier-Schuppen. Palaeontographica, 127A: 1-64. 
Halstead, L. B. and S. Turner. 1973. Silurian and Devonian ostracoderms, pp. 

67-79. In A. Hallam (ed.). Atlas of Palaeobiogeography. Amsterdam: Elsevier, 

531 pp. 
Hoppe, K. H. 1931. Die Coelolepiden und Acanthodier des Obersilurs der Insel 

Oesel. Palaeontographica 76: 35-94. 
Karatajute-Talimaa, V. N. 1978. Silurian and Devonian Thelodonts of the 

USSR and Spitsbergen. Vilnius: Mosklas, 1-334 [in Russian]. 
Lawson, J. D. 1975. Ludlow Benthonic Assemblages. Palaeontology 18 (3): 

509-525. 
Mackenzie, G. S. 1951. Geol. Surv. Can. Pap. 1951-15 (out of print, not seen). 
1964. Geology, Saint John, New Brunswick. Geol. Surv. Can. Map 

1113A, 1 inch: 1 mile, 1:63360. 
1964. Geology, Hampstead, New Brunswick. Geol. Surv. Can. Map 



I114A, 1 inch: 1 mile, 1:63360. 
Marss, T. 1982a. Thelodus admirabilis n. sp. (Agnatha) from the Upper Silurian 

of the East Baltic. Eesti NSV Teaduste Akadeemia Toimetised 31, K. Geol. no. 

31: 112 116. 
. 1982b. Vertebrate zones in the East Baltic Silurian, pp. 97-105. In D. 

Kaljo and E. Klaaman (eds.), Ecostratigraphy of the East Baltic Silurian. 

Tallinn: Academy of Sciences Estonian SSR, Inst. Geol. 
Matthew, G. F. 1888. On some remarkable organisms of the Silurian and 

Devonian rocks in Southern New Brunswick. Proc. Trans. Roy. Soc. Can., 

Ottawa, Section 6(1)6, Section 4: 49-62. 



1986 THELODUS MACINTOSH! 17 



1907. A new Genus and a New Species of Silurian Fish. Trans. Roy. Soc. 



Can. (3), 1, section 4: 7-9. 
McCutcheon, S. R. 1981. Revised stratigraphy of the Long Reach area, south- 
ern New Brunswick: evidence for major, northwestward-directed Acadian 

thrusting. Can. J. Earth Sciences, 18 (3): 646-656. 
Pander, C. H. 1856. Monographie der fossilen Fische des silurischen Systems der 

russisch-baltischen Gouvernements. St. Petersburg: x + 1-91. 
Powrie, J. 1 870. On the earliest known vestiges of vertebrate life; being a descrip- 
tion of the fish remains of the Old Red Sandstone of Forfarshire. Trans. Geol. 

Soc. Edinburgh, 1: 284-301. 
Ritchie, A. 1968. New evidence on Jamoytius kerwoodi White, an important 

ostracoderm from the Silurian of Lanarkshire, Scotland. Palaeontology, 11, 1: 

21-39. 
Robertson, J. D. 1957. The habitat of early vertebrates. Biol. Rev. Camb. 32: 

156-87. 
Rolfe, W. D. I. 1973. Excursions 14 and 15, Lesmahagow and Hagshaw Hills, 

pp. 105-126. In B. J. Bluck (ed.). Excursion Guide to the Geology of the 

Glasgow Districts. Glasgow Geol. Soc. Glasgow: 181 pp. 
Smith, J. C. 1966. Geology of Southwestern New Brunswick, pp. 1-12. In W. H. 

Poole (ed.), Geology of parts of Atlantic provinces. Guidebook, Geol. Assoc. 

Can. Miner. Assoc, Halifax. 
Stetson, H. C. 1928. A New American Thelodus. Amer. J. Science, 16: 221-31. 
Thorsteinsson, R. 1967. Preliminary note on Silurian and Devonian ostraco- 

derms from Cornwallis and Somerset Islands, Canadian Arctic Archipelago, pp. 

45-47. In Evolution des Vertebres. Problems actuel de Paleontologie, 163, 

Paris: C. N. R. S. Masson et Cie. 
Traquir, R. H. 1898. Report on Fossil Fishes. Summary of Progress for 1897, 

Geol. Surv. G. B., 72-76. 
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Nature, London, 227, No. 5253: 90. 
1970b. Fish help to trace continental movements. Spectrum, Bull. Cent. 

Off. Inf., No. 79: 8-10. 
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Soc, 129: 557-84. 
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Catalogus I: Animalia, Pars 122: 1 14 pp. 
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tests of Lower Palaeozoic continental reconstructions. Palaeogeography, 
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Westoll, T. S. 1958. Stratigraphical Framework, pp. 235-239. In T. S. Westoll 

(ed.), Studies on Fossil Vertebrates. Univ. London: Athlone Press: 263 pp. 
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sess.: 25-34. 



REVIORA 



Museum of Comparative Zoology 

US ISSN 0006 9698 

Cambridge, Mass. 29 August 1986 Number 487 

THE IDENTIFICATION OF LARVAL PARASUDIS 

(TELEOSTEI, CHLOROPHTHALMIDAE); WITH 

NOTES ON THE ANATOMY AND RELATIONSHIPS 

OF AULOPIFORM FISHES 

Karsten E. Hartel and Melanie L. J. Stiassny 1 

Abstract. Larvae and transforming individuals of Parasudis are identified and 
described for the first time. The phylogenetic integrity of the family Chlorophthalmi- 
dae is investigated and restricted to the sistergenera, Parasudis and Chlorophthal- 
mus. Two derived morphological characters support the monophyly of an extended 
Ipnopidae which now includes the genus Bathysauropsis. The "rostral cartilages" in 
aulopiform fishes are reviewed and the homology of these and associated structures 
throughout the lower Neoteleostei is questioned. 

INTRODUCTION 

Parasudis truculenta (Goode and Bean, 1895) is a relatively com- 
mon offshore fish found from off northern New England to equato- 
rial Brazil. Adult specimens have been taken only in bottom trawls 
at depths of ca.180 to 480 m, but dietary studies suggest that Para- 
sudis moves off the bottom to feed in mid water (Mead 1966). The 
species is hermaphroditic, and ripe specimens have been collected 
from February to May (Mead 1960). Neither the eggs nor the larvae 
of this species have yet been described (Okiyama 1984). 

The purpose of this paper is to describe larval and transforming 
Parasudis, and to comment on aspects of anatomy and relation- 
ships. There is considerable confusion in current literature regarding 



'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. Order of authorship is alphabetical. 

U 



SEP 1 



2 BREVIORA No. 487 

the systematics and classification of Rosen's (1973) Aulopiformes 
(see Johnson 1982; Okiyama 1984; Rosen 1985; Stiassny 1986). The 
resolution of this problem is beyond the scope of this paper and for 
the purpose of this study we adhere to Rosen's classification of the 
Order (1973:509). 

MATERIALS AND METHODS 

Specimens are from the Woods Hole Oceanographic Institution 
(WHOI) collections housed at the Museum of Comparative Zool- 
ogy, Harvard University (MCZ); the Atlantic Research Centre, New 
Brunswick (ARC); the DANA collections of the Zoological 
Museum, University of Copenhagen (ZMUC); the National Mu- 
seum of Natural History, Washington, D.C. (USNM); the British 
Museum of Natural History (BMNH) and the University of Miami 
(UMML). The specimens used in this study are listed in appendix 
under material examined. 

Counts and measurements follow Hubbs and Lagler (1964). For 
osteological examination, selected specimens were cleared and 
stained following the procedure of Dingerkus and Uhler (1977); 
these specimens are indicated "c.s." under material examined. 
Anatomical drawings were made with the aid of a Zeiss SV-8 stereo- 
microscope with a camera lucida attachment. Osteological and syn- 
desmological nomenclature follows Stiassny (1986). 

DIAGNOSIS 

Parasudis larvae are characterized by the distinctive bulbous 
shape of the snout, the pattern of snout, opercular and caudal pig- 
mentation, as well as by a particular basihyal form and dentition. 
The larvae are further distinguished from those of other chloro- 
phthalmoid taxa by having 38 to 39 vertebrae and/ or myomeres 
(Okiyama 1984:208). 

DESCRIPTION 

Larvae 
(Figs. 1A, BandC) 

Preflexion or flexion larvae are not present among the material 
examined, and the following description is based on 85 postflexion 



1986 LARVAL PARASUDIS RELATIONSHIPS 3 

specimens ranging from 10.6 to 80 mm standard length (SL). Spec- 
imens of 15.4 and 30.6 mm SL are illustrated in Figs. 1A-C. The 
body is elongate, oval to round in cross section at maximum depth; 
maximum depth 6.4 to 9.4 times in SL; maximum body width sub- 
equal or equal to maximum body depth (7.8 to 1 1.7 in SL). Head 
length (HL) 2.6 to 3.9 times in SL, probably negatively allometric. 
Bulbous snout rounded in anterodorsal profile, 2.2 to 3.0 times in 
HL. Eyes large, 3.5 to 5.0 times in HL. The eyes often appear 
stalked or partially stalked (Fig. IB). The presence (or absence) of 
stalked eyes may be a result of damage incurred during collection; 
however, stalked and non-stalked specimens do appear in the same 
samples. The eyes of both larval and transforming individuals lack 
the "keyhole" shaped aphakic space characteristic of adult 
chlorophthalmids. 

All larvae examined appear to have a full complement of fin 
elements, although, the fragile fin rays are often damaged. The 
origin of the dorsal fin (10 rays) is slightly posterior to the base of 
the pectoral fin (17 rays). The narrow-based adipose fin is posi- 
tioned above the second or third anal ray (8-9 anal rays). The pecto- 
rals are at mid-body level with their bases almost under the thin 
opercular flap. Pelvic fin (9 rays) origin is below the anterior half of 
the dorsal fin. The caudal fin is forked even in the smallest speci- 
mens. Procurrent caudal rays are present and increase in number 
with growth. The anus is located well caudad of the pelvic fin base 
but closer to the insertion of the pelvic than to the insertion of the 
anal fin. 

The dentition remains relatively constant throughout the larval 
period. Both the premaxillae and the dentaries bear single rows of 
evenly spaced caniniform teeth (Fig. 2). Anteriorly, these larval 
teeth are recurved, but are markedly retrose in the posterior aspect 
of both upper and lower jaws. Two to three small recurved 
caniniform teeth are implanted along the anteriolateral aspect of the 
dentary (Fig. 2). 

In most specimens a few small caniniform teeth are inserted along 
the body of the palatine cartilages, and in the ventral hypobranchial 
apparatus a markedly elongate basihyal cartilage bears small 
recurved fang-like teeth on its anterior margin (Fig. 3A). 



BREVIORA 



No. 487 






Figure 1. Parasudis truculenta. (A) Lateral view of larva 15.4 mm SL (MCZ 
62400). (B) Dorsal view of head of same specimen. (C) Lateral view of larva 30.6 mm 
SL (MCZ 62399). (D) Lateral view of transforming juvenile 85 mm SL (MCZ 62401). 
Note that the dotted lines on the body in (A) and (C) represent myosepta while the 
solid lines in (D) represent the pigmented edges of scale pockets. Drawn by S. 
Landry. 



1986 



LARVAL PARASUD1S RELATIONSHIPS 



PMX 




Figure 2. Isolated buccal jaws of larval Parasudis (MCZ 62397, 38 mm SL). 
Abbreviations: Dent, dentary; Mx, maxilla; Pmx, premaxilla. 



Pigmentation 

Body color in preserved specimens is a uniform opaque white 
although very infrequently a light tan or brown wash is present. Live 
coloration is unknown, but based on observations of live pretrans- 
formation Chlorophthalmus (Hartel personal observation), living 
Parasudis larvae are presumed to be almost translucent in seawater. 

Opercular pigmentation covers an area almost as large as that of 
the eye and at times may be quite dark. Opercular pigmentation is 
present in the smallest specimen examined. 

Initially, a small group of melanophores is situated on the caudal 
peduncle above the midline. With growth (>30 mm SL) the pig- 
mented area becomes roughly triangular in outline with the pos- 
terior lower corner extending below the midline (Fig. 1 C). A melan- 
ophore distinct from the triangular patch and situated in the upper 
corner of the caudal base is found in specimens greater than 22 mm 



BREVIORA 



No. 487 




Figure 3. Parasudis. (A) Ventral gill arch elements and hyoid apparatus of 
larva 30 mm SL (MCZ 62398). (B) First epibranchial and associated structures of 
adult specimen (MCZ 40561). Chlorophthalmus. (C) Ventral gill arch elements of 
larva 22 mm SL (MCZ 62403). (D) First epibranchial and associated structures of 
adult specimen (MCZ 40509). 



1986 LARVAL PARASUDIS RELATIONSHIPS 

SL, and a second is usually developed at the lower corner of the 
caudal base by 30 mm SL. 

Pigmentation in the snout region first appears as a single melano- 
phore above the anterior end of the maxilla at 14 mm SL. Addi- 
tional melanophores are added posteriorly in a line just above the 
maxilla and by 50 mm SL, the row reaches a point almost under the 
anterior margin of the orbit. 

Lateral body pigmentation begins as one or two mid-body 
melanophores at 22 mm SL. The melanophores increase laterally in 
two groups. Four are present under the dorsal, and three are under 
the adipose fin by 24 mm SL. At 40 mm SL the melanophores, 
which are always found directly over a myomere, form an almost 
continuous band. There are one, two or sometimes three melano- 
phores per myomere. 

Deep pigment is present in the dorsal midline as a small dark spot 
just posterior to the dorsal fin. Ventrally a similar small midline spot 
is found at mid-body. An additional line of deep pigment is present 
along the midventral caudal peduncle. All of these deeper markings 
become less noticeable with growth as the muscle tissues become 
denser. Internal pigment is often visible through the bodywall below 
the base of the pectoral fin (Fig. 1 A). This darkened area is densely 
pigmented mesentery supporting the viscera and areas of visceral 
peritoneum wrapping partially around the developing anterior 
viscera. The pigmented mesentary extends caudad almost to the 
level of the anal fin origin (Fig. 4). 

Transforming specimens 
(Fig. ID) 

This description is based upon two specimens, 75.5 mm SL (MCZ 
57922) and 85 mm SL (MCZ 62401), illustrated in Fig. ID. The 
larger specimen is only 5 mm larger than the largest available larval 
specimen (ARC 6879), yet marked morphological changes are evi- 
dent. 

General body proportions remain constant, but the form of 
the head is altered by the loss (via reabsorption?) of the character- 
istic bulbous larval snout. The smaller individual retains a few of the 
larval retrose jaw teeth and has a strong basihyal dentition. In the 
larger transforming fish the jaws are edentulate, apparently having 
lost the larval teeth and not yet having developed the juvenile/ adult 



BREVIORA 



No. 487 




P.m 



Figure 4. Internal pigmentation of larval Parasudis (based on two specimens; 
MCZ 63055, 40 mm SL and MCZ 63056, 31 mm SL). Drawn by S. Landry. 

Abbreviations: B.c, body cavity; P.m, pigmented mesentery; V.p, visceral 
peritoneum. 

dentition. A few small palatine teeth are present, and small teeth are 
embedded in the dermis overlying the basihyal element. As the 
smaller of the two transforming specimens is badly damaged and 
contorted, the remaining description is based solely upon the 85 mm 
SL specimen. 

The diagnostic larval pigmentation pattern is faint but the rem- 
nants of snout, opercular, lateral and caudal pigmentation are still 
discernible. Patches of scales are present and, as is indicated in Fig. 
ID, these have a distinctive pattern of implantation. Each scale 
pocket is located on a continuous flap of skin which is strongly 
pigmented along its distal margin. The resultant "herringbone" 
pattern is clearly visible in adult specimens (Mead 1966: Fig. 47) 
and where scales have been lost, corresponds to approximately two 
pigment lines per scale. 

Transformation apparently occurs rapidly. We have seen a fully 
transformed juvenile (MCZ 41840) the same length as the larger 
transforming larva. The fully transformed fish has the definitive 
adult dentition (see Mead 1966), is scaled, and has lost almost all 
traces of larval pigmentation. During transformation the snout 
becomes increasingly flattened and "shrunken," lending the fish its 
characteristic "duckbilled" appearance. Apparently, a loss (reab- 
sorption?) of the somewhat gelatinous larval body tissues results in a 






1986 LARVAL PARASUDIS RELATIONSHIPS 9 

more attenuated body form in the recently transformed juvenile. 
Halliday (1968: Fig. 1) illustrates a 99.0 mm SL transformed juve- 
nile that shows initial body deepening. 

The internal pigmentation of the anterior viscera is present in 
transforming specimens. Apparently during transformation, pig- 
ment migrates from the visceral peritoneum onto the parietal peri- 
toneum, and the pigmentation of the caudad extension of the 
mesentery is greatly reduced. In fully transformed juveniles the pa- 
rietal peritoneum is strongly pigmented throughout the abdominal 
cavity but little pigment is associated with the visceral peritoneum. 

We have examined a number of transforming Chlorophthalmus 
specimens from both the North Atlantic Ocean and the Coral Sea. 
In each of these specimens, tranformation occurs at a considerably 
smaller size (ca. 35 to 40 mm SL) than that observed in our Para- 
sudis specimens. 

DISTRIBUTION 

The western Atlantic specimens we have examined come from 
areas off the Brazilian coast to areas east of the southern tip of Nova 
Scotia (Fig. 5). Included in our study is one eastern Atlantic 
specimen (MCZ 62402). Preliminary morphometric analysis of this 
larva and its eastern Atlantic locality indicate that it may represent 
Parasudis fraser-brunneri (Poll 1953). However, due to a lack of 
additional material for detailed study and the close morphological 
resemblance of P. fraser-brunneri to P. truculenta (see Mead 
1966:184), we are unable to confirm the specific identity of the 
eastern Atlantic specimen. 

All of the larval Parasudis were taken in midwater trawls. Speci- 
mens from the WHOI opening and closing nets (MOC 10 and 20) 
indicate that the larvae are most commonly taken above 150 m (five 
collections: 70 to m; to 100 m; 70 to 140m; to 203 m; 70 to 150 
m). Information from non-closing nets indicate similar distribu- 
tions. However, other collections taken with non-closing nets indi- 
cate that the larvae may approach the surface rather closely, since 
Parasudis larvae are found in collections taken above 50 m. Except 
for a few larvae from the relatively shallow Straits of Florida, all 
specimens were taken in trawls over ocean bottoms from 1937 to 
4934 m deep. 



10 



BREVIORA 



No. 487 





HO 60 


40* 


0" ( 


) 








O 






*C 






40 




r iO ( 


£- 






) 


-r?^ 






wq *y 


o 






o 


o o ^y 






6° o 














,0 
















7 r" 


« =£><£■ 






AJ 










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<£> 


AT) 










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Figure 5. Distribution of larval Parasudis examined during this study. Open 
circles: P.truculenta. Solid circles: transforming larvae. Triangle: P.?fraser-brunneri 
larva (MCZ 62402). Symbols may represent more than one sample. 



GENERIC PLACEMENT 

As currently constituted the family Chlorophthalmidae (=Chlo- 
rophthalminae of Sulak 1977 and Nelson 1984) comprises three 
genera of benthic fishes (Okiyama 1984): Chlorophthalmus Bona- 
parte, 1840 (18+ species), Parasudis Regan, 1911 (two species) and 
Bathysauropsis Regan, 1911 (three species). Representatives of the 
former two genera inhabit the North Atlantic; Bathysauropsis is 
known only from the deep waters (2000 to 2500 m) of the South 
Atlantic and South Pacific (Mead 1966). 

Based on adult morphology, Mead (1966) presented a key to 
chlorophthalmid genera; we note that Parasudis is further distin- 
guished from the other two genera by having a single gill raker along 
the anterior border of the first epibranchial (Fig. 3B). In both Chlo- 
rophthalmus (Fig. 3D) and Bathysauropsis, the first epibranchial 
bears at least two or more lath-like denticulate gill rakers along its 
anterior margin. 

Predorsal number distinguishes the three genera. Adult and larval 
Parasudis have a single predorsal bone (Figs. 6A, B). Chlorophy- 
thalmus have two (Figs. 6C, D), adult Bathysauropsis have three. 



1986 



LARVAL PARASUDIS RELATIONSHIPS 



11 




Figure 6. Parasudis. (A) Posterior region of the neurocranium, vertebral 
column and predorsal bones of adult specimen (MCZ 40561 ). (B) Larva 30 mm SL 
(MCZ 62398). Chlorophthalmus. (C) Posterior region of the neurocranium, vertebral 
column and predorsal bones of adult specimen (MCZ 41444). (D) Larva 22 mm SL 
(MCZ 62403). Arrows indicate predorsal bones. 



12 BREVIORA No. 487 

Johnson (1982) discussed predorsal (=supraneural) number among 
aulopiforms and noted a trend towards predorsal reduction 
throughout the assemblage. He cited notosudids (=scopelosaurids) 
and Omosudis as other taxa with a single predorsal. Sulak (1977) 
stated that ipnopine genera bear one of two predorsals. 

The anal position and lack of anal pigmentation during trans- 
formation of Parasudis contrasts with that of Chlorophthalmus 
where the anus is situated closer to the pelvic base (see Mead 1966), 
and is highly pigmented during transformation. Other pigment 
characters of the anterior visceral cavity separate Parasudis and 
Chlorophthalmus. In Parasudis the larval pigmentation is confined 
to the visceral peritoneum and associated mesentery, while in Chlo- 
rophthalmus it is a discrete patch closely associated with the parietal 
peritoneum which lines the body cavity. In addition, larval Parasu- 
dis differs from larval Chlorophthalmus by the presence in the 
former of a caudad extension of pigmented mesentry (Fig. 4). Deep 
visceral pigmentation in other aulopiforms is discussed by Johnson 
(1974). 

COMPARISON WITH OTHER AULOPIFORM TAXA 

To assess the phylogenetic significance of certain features de- 
scribed in Parasudis we compare them here with those found in 
Chlorophthalmus and other aulopiform taxa. Clearly, the following 
anatomical comparisons are far from exhaustive, and we have 
selected only those features which appear to be phylogenetically 
informative at the present level of analysis. 

Rosen (1971) found basihyal teeth of a similar, although larger, 
form and distribution in larval Chlorophthalmus (see also our Fig. 
3C), as well as in the ipnopid larvae he described (see Okiyama 1981 
for other ipnopid larvae). Sulak (1977) described comparable basi- 
hyal dentition in larval Bathypterois, and Bathytyphlops. Rosen 
(1971) noted that a similar basihyal dentition is unknown in any 
adult fish (although the pattern is approximated in Glossanodon). 
Sulak (1977:80) stated that the ". . .condition appears to be unique 
to the juvenile stages of basal myctophiform fishes." However, until 
the full limit of the distribution of this type of larval dentition are 
known, particularly in larval aulopids and neoscopelids, we can 
draw little phylogenetic inference from its occurrence in Parasudis 
larvae. 



1986 LARVAL PARASUDIS RELATIONSHIPS 13 

Basihyal morphology easily distinguishes Chlorophthalmus from 
Parasudis, which in Chlorophthalmus is short and strongly spatu- 
late (Figs. 3A, C), and in Parasudis is elongate. These differences, 
although less markedly than in larvae, are present in the ossified 
adult elements and help to distinguish the taxa (compare Figs. 7B, 
C). Basihyal morphologies similar to those of larval Chlorophthal- 
mus are found in the larvae of Bathypterois (Sulak 1977; personal 
observation) and other ipnopids (Rosen 1971; Okiyama 1981), as 
well as in adult Aulopus, Bathysaurus and Scopelosaurus (Bertelsen 
et ah, 1976; personal observation). Thus, based upon its limited 
distribution, we interpret the broad and elongate basihyal of larval 
Parasudis as an autapomorphic feature. 

Interestingly, adult ipnopids share a derived condition of the basi- 
hyal element, which is relatively tiny [or entirely absent in some 
bathypteroids (Mead 1966)] and obliquely aligned (Fig. 7A). A sim- 
ilar condition of the basihyal is present in adult Bathysauropsis, 
where unfortunately the larvab condition is unknown (Okiyama 
1984).2 

Certain stomiatoids have a similar basihyal arrangement, how- 
ever, we agree with Weitzman (1974) that the presence of a moder- 
ately sized and horizontally aligned basihyal is primitive for the 
Stomiiformes as a whole. Among alepisauroids (sensu Rosen 1973), 
a small (toothed) basihyal is found in certain scopelarchid taxa 
(Johnson 1974) and in Bathysaurus, but a small vertically aligned 
basihyal appears to be restricted in its distribution among aulopi- 
forms to adult Bathysauropsis and the Ipnopidae. 3 The common 
occurrence of this basihyal morphology suggests an immediate 
common ancestry between them. 

Corroboration of the phyletic integrity of a grouping comprising 
Bathysauropsis and the ipnopids is found in the degree of develop- 
ment of the branchiostegal membranes and gular fold. Mead 
(1966:115) described the region in bathypteroids in the following 



2 We have been able to examine only two specimens of Bathysauropsis gracilis but 
have kindly been given access to notes and radiographs made by K. Sulak after his 
examination of specimens of B. gigas and B. mayanus. 

following Johnson (1982) and Okiyama (1984) the Ipnopidae is taken here to 
include the Bathypteroidae of Mead ( 1 966) and is equivalent to the Bathypteroidae of 
Marshall and Staiger (1975) and the sub-family Ipnopinae of Sulak (1977) and 
Nelson (1984). 



14 



BREVIORA 



No. 487 




Figure 7. Basihyal and associated structures in adult (A) Bathypterois (MCZ 
57624). (B) Chlorophthalmus (MCZ 41444). (C) Parasudis (MCZ 40561). 



Abbreviations: Bbl-2, basibranchials 12; Bhy, basihyal; Cbl, ceratobranchi- 



al 1. 



terms which apply equally to Bathysauropsis and the ipnopids: 
"Branchiostegal membranes broad and separate from one another 
and from the isthmus, overlapping anteriorly, and crossed near the 
tip of the jaw by a thick but narrow gular fold ..." This morphology 
appears to be unique among aulopiforms and although a poorly 
developed gular fold is present in some stomiiform fishes (e.g., 
Gonostoma and Polymetme) it is not as elaborate as that found in 
Bathysauropsis and the ipnopids. 

Chlorophthalmus shares with Parasudis the pigmented scale 
pocket skin flap character. In Parasudis the dorsal and ventral 
scale rows converge rostrad, "herringbone" fashion, upon the lateral 
line (Mead 1966; Fig. 47), while in Chlorophthalmus, dorsal and 
ventral scale rows diverge from the midline resulting in a slightly 
"zig-zag" pattern (Mead 1966; Fig. 46). This highly distinctive feature 
of squamation is unique to Parasudis and Chlorophthalmus, and 
is here interpreted as a synapomorphy uniting the two genera. 

In the eyes of adult Chlorophthalmus (Theisen 1965; Fig. 3), as 
well as in adult Parasudis, a "keyhole" shaped aphakic space is 
present. Although well-developed aphakic spaces are not uncom- 
mon to the eyes of many benthic marine fishes (Munk 1966; Mar- 
shall 1971), a "keyhole" shaped space is restricted to adults of these 



1986 LARVAL PARASUDIS RELATIONSHIPS 15 

two genera. A similar space is absent in the eyes of Bathysauropsis 
and it is here interpreted as a synapomorphy of Parasudis and 
Chlorophthalmus. 



RELATIONSHIPS 

In reviewing the literature pertaining to Parasudis and its sup- 
posed relatives, it is clear that many problems remain (Johnson 
1982; Rosen 1985; Stiassny 1986). However, at the intrafamilial 
level, aside from the suggestion that Parasudis may share a close 
phylogenetic relationship with paralepidid alepisauroids (Mead 
1966), most authors who have considered the chlorophthalmids 
(e.g., Gosline et al. 1966; Marshall and Staiger 1975; Sulak 1977; 
Nelson 1984; Stiassny 1986) imply that Chlorophthalmus and Para- 
sudis are closely related, and our own investigation confirms a sister- 
group relationship between these two taxa. The genus Bathysaurop- 
sis is poorly known anatomically, and consequently, its phylogenetic 
placement is less clear. Mead (1966) suggested that further investiga- 
tion of this genus may indicate a need for familial reallocation, a 
finding that is corroborated by this study. We propose that the 
notion of an "intermediate" or "transitional" position of Bathysau- 
ropsis, forming a "link" between Parasudis and Chlorophthalmus 
on the one hand and the ipnopids on the other, and thus serving as 
". . .the primary basis of the present incorporation of the ipnopine 
genera into the Chlorophthalmidae" (Sulak 1977:64) be replaced by 
a phylogenetic scheme in which Bathysauropsis is removed from the 
Chlorophthalmidae and incorporated within the Ipnopidae (Fig. 8). 
This relocation of Bathysauropsis is based on our observations of 
the presence of a small, obliquely aligned basihyal and a well- 
developed gular fold (Fig. 8: characters one and two). The position 
of Aulopus in relation to these taxa is here unresolved; larval 
aulopid material was unavailable for this study. Elsewhere Stiassny 
(1986) argued that Chlorophthalmus, Parasudis and Aulopus (but 
not Bathysauropsis and the other ipnopid genera) share with cte- 
nosquamate fishes an advanced type of palatine/ maxillary associ- 
ation and morphology (see also Rosen 1985 for a consideration of 
aulopid relationships). For this reason Aulopus is included in our 
cladogram, albeit in an unresolved position (Fig. 8). 



16 



BREVIORA 

WULOPOIDEI" 



No. 487 
CTENOSQUAMATA 



O^ cf <** C^ <* ^ 




Figure 8. Cladogram of eurpterygian relationships, incorporating the results of 
the present study. Characters are the presence of: 1) A small obliquely aligned basi- 
hyal bone; 2) A thick well-developed gular fold; 3) A "keyhole" shaped aphakic space; 
4) A peculiar scale pocket morphology and pigmentation; A) See Rosen (1985) for 
synapomorphies uniting the Aulopoidei with the Ctenosquamata; B) See Stiassny 
(1986) for synapomorphies uniting Aulopus, Chlorophthalmus and Parasudis with 
the Ctenosquamata; C) See Lauder and Liem ( 1983) for synapomorphies uniting the 
Ctenosquamata. Note: Following Rosen (1973), "other aulopoids" are the Bathy- 
sauridae and Notosudidae. "Other Ipnopidae" are Bathypterois, Bathytyphlops, 
Bathymicrops and Ipnops. 

Significance of the Rostral Cartilage 

Given this admittedly tentative scheme of relationships for the 
chlorophthalmids, it is extremely interesting to note the highly var- 
ied condition of the so-called "rostral cartilage" in these and in other 
aulopiform taxa. The "rostral cartilage" of both larval and adult 
Parasudis is a single median structure which is bound by a well- 
developed ethmo-rostroid ligament to both the ethmoid and pre- 
maxillae (Figs. 9 A, B). The attachment of the cartilage to the ethmoid 
is strong in the larval fish and becomes weaker with growth. In 



1986 LARVAL PARASUDIS RELATIONSHIPS 17 

Parasudis, the rostral cartilage stains normally with Alcian Blue and 
is therefore presumably fully chondrified hyaline cartilage. In con- 
trast, the "cartilage" in Chlorophthalmus (Figs. 9C, D; Rosen 1985; 
Fig. 40C) is paired, and the "cartilages" are associated with the 
symphyseal processes of the premaxillae. An ethmo-rostroid liga- 
ment is lacking, but the whole region is invested with loose connec- 
tive tissue fibers. These rostral structures do not stain normally with 
Alcian Blue; staining is weak and diffuse. Similarly, in Aulopus 
(Rosen 1985; Fig. 41C; Stiassny 1986; Fig. 6) the single median 
structure does not stain as hyaline cartilage and is highly fibrous. 
Theisen (1965) illustrated paired rostral structures in Ipnops similar 
to those of Chlorophthalmus. Rosen (1985) also illustrated paired 
"rostral cartilages" of an apparently remarkably similar type and 
arrangement in the stomiiform, Maurolicus. Other stomiiform taxa 
examined in this study either lack a "rostral cartilage" entirely (e.g., 
Photichthys), or bear a single median structure ligamentously 
bound to the ethmoid region (e.g., Gonostoma and Diplophos). We 
have been unable to locate "rostral cartilages" in either Bathypterois 
or Bathysauropsis. In both of these genera the premaxillae are 
bound to the ethmoid region by a well-developed ethmo-rostroid 
ligament. We interpret the similarities between the rostral morphol- 
ogies of Chlorophthalmus and Ipnops (and Maurolicus) to be 
homoplasious. To assess them otherwise, that is, to argue for the 
phyletic alignment of Chlorophthalmus with Ipnops (and Mauroli- 
cus) on the basis of this one "rostral cartilage" character would 
require losses and reversals in a series of other characters (see Fink 
and Weitzman 1982; Marshall and Staiger 1975). 

Bertelsen et al. (1976) described and illustrated the rostral mor- 
phology of certain notosudids where a large (often ossified) "rostral 
cartilage" is bound to the ethmoid and the premaxillae by a well- 
developed ethmo-rostroid ligament similar to that of the stomiiform, 
Gonostoma (Stiassny 1986), or to that illustrated here for Parasudis. 

Rostral morphologies are also confusing among other aulopiform 
lineages; for example, among synodontids, Synodus (Fig. 10A) 
bears a large medial and fully chondrified "rostral cartilage" 
strongly bound to the ethmoid. Saurida (Fig. 10B), on the other 
hand, lacks the structure as do both Harpadon and Bathysaurus. 
The paralepidid Sudis bears small paired hyaline cartilages, which 
in adults are associated with the symphyseal processes of the pre- 



18 



BREVIORA 



No. 487 



-R.CART" PMX 




Figure 9. Parasudis. (A) Ethmovomer and upper jaws of adult specimen in 
dorsal view (MCZ 40561). (B) Ethmovomer of larva 30 mm SL (MCZ 62398) in 
dorsal view. Chlorophthalmus. (C) Ethmovomer and upper jaws of adult specimen 
(MCZ 40564) in dorsal view. (D) Ethmovomer of larva 22 mm SL (MCZ 62403) in 
dorsal view. 






1986 



LARVAL PARASUDIS RELATIONSHIPS 



19 



maxillae. In larvae these structures appear more strongly bound to 
the ethmoid. Finally, in Alepisaurus a condition much like that 
described in Salmo (Fink and Weitzman 1982; Rosen 1985) per- 
tains. Paired hyaline cartilages are strongly bound to the inner faces 
of the premaxillary symphyseal processes. 

Even from this cursory review it is clear the homologies of rostral 
morphologies and other associated structures throughout the lower 
Neoteleostei are questionable. By the level of the Acanthomorpha, 
the situation has stabilized such that there appears little doubt of the 
homology of the single median chondrified rostral cartilage strongly 
bound to the premaxillary ascending processes by a well-developed 
maxillo-rostroid ligament (Stiassny 1986). Until the distribution 
and homologies of the non-acanthomorph rostral structures are bet- 
ter understood, perhaps the term rostral cartilage should be re- 
stricted to that structure in the Acanthomorpha. When referring to 
the various rostral structures in non-acanthomorph neoteleosts 
before their homologies are resolved, the term "rostral cartilage" or 
"rostral structure" should be employed within quotation marks. 

ACKNOWLEDGMENTS 

We would like to thank the following individuals and institutions 
for the loan of material and /or helpful information and comments: 




PMX 




Figure 10. Ethmovomer and upper jaw of adult specimens of (A) Synodus 
(MCZ 47490). (B) Saurida (MCZ 561 1 1). 

Abbreviations: Fr, frontal; Me, mesethmoid; Me.p, mesethmoid process; Mx, 
maxilla; Pal, palatine; Pmx, premaxilla; "R.cart", "rostral cartilage". Ligaments: II, 
ethmo-rostroid; IV, median palato-maxillary; XI, ethmo-maxillary; XII, palato- 
premaxillary. 



20 BREVIORA No. 487 

E. Bertelsen (ZMUC), R. H. Gibbs and R. P. Vari (USNM), R. K. 
Johnson (FMNH), D. F. Markle (formerly ARC), K. J. Sulak 
(ARC), J. Webb (Boston University), and A. Wheeler (BMNH). 
For their critical review of the manuscript we are grateful to J. E. 
Craddock (WHOI), G. D. Johnson (USNM), and K. F. Liem 
(MCZ). The late Sally Richardson offered invaluable comments on 
this paper and we dedicate our first work on larval fishes in her 
memory. We are especially grateful to Sally Landry for her skillful 
preparation of Figures 1 and 4. 

This research was carried out during the tenure of NSF BSR 
84-07449 (MLJS). NSF DEB 77-23726 (K. F. Liem) provided sup- 
port for the transfer, sorting and curation of the WHOI specimens 
used in the study. 

LITERATURE CITED 

Bertelsen, E., G. Krefft and N. B. Marshall. 1976. The fishes of the family 
Notosudidae. Dana Rept., 86: 3-114. 

Dingerkus, G. and L. D. Uhler. 1977. Enzyme clearing of Alcian Blue stained 
whole small vertebrates for demonstration of cartilage. Stain Tech., 52(4): 
229-232. 

Fink, W. L. and S. H. Weitzman. 1982. Relationships of the stomiiform fishes 
(Teleostei), with a description of Diplophos. Bull. Mus. Comp. Zool., 150(2): 
31-93. 

Gosline, W. A., N. B. Marshall and G. W. Mead. 1966. Order Iniomi. In 
Fishes of the Western North Atlantic. Mem. Sears Fdn. mar. Res., 1: 1-18. 

Halliday, R. G. 1968. Occurences of Parasudis trunculentus (Goode and Bean) 
1895 (Iniomi: Chlorophthalmidae) off La Have Bank, Nova Scotia. J. Fish. Res. 
Bd. Canada, 25(2): 421-422. 

Hubbs, C. L. and K. F. Lagler. 1964. Fishes of the Great Lakes Region. Ann 
Arbor: Univ. Michigan Press, 213 pp. 

Johnson, R. K. 1974. A revision of the alepisauroid family Scopelarchidae (Pis- 
ces: Myctophiformes). Fieldiana (Zool.) 66: 1-249. 

1982. A revision of the alepisauroid families Evermannellidae and Scope- 
larchidae: Systematics, morphology, interrelationships and zoogeography. Field- 
iana (Zool.) N. S. 12: 1-252. 

Lauder, G. V. and K. F. Liem. 1983. The evolution and relationships of the 
actinopterygian fishes. Bull. Mus. Comp. Zool., 150: 95-197. 

Marshall, N. B. 1971. Explorations in the lives of fishes. Cambridge: Harvard 
Univ. Press, 204 pp. 

Marshall, N. B. and J. C. Staiger. 1975. Aspects of the structure, relationships 
and biology of the deep-sea fish Ipnops murrayi (Family Bathypteroidae). Bull. 
Mar. Sci., 25(1): 101-111. 



1986 LARVAL PARASUDIS RELATIONSHIPS 21 

Mead. G. W. I960. Hermaphroditism in archibenthic and pelagic fishes of the 

order Iniomi. Deep-Sea Research, 6: 234-235. 
1966. Family Chlorophthalmidae. In Fishes of the Western North Atlan- 
tic, Mem. Sears Fdn. mar. Res., 1: 162-189. 
Munk, O. 1966. Ocular anatomy of some deep-sea teleosts. Dana Rept., 70:1-62. 
Nelson, J. S. 1984. Fishes of the World. 2nd Ed. New York. Wiley and Sons, 

ix + 523 pp. 
Okiyama, M. 1981. A larval Ipnops and its possible metamorphosis. Jap. J. Ich- 

thyol., 28(3): 247-253. 
1984. Myctophiformes:Development. In H. G. Moser, (ed. -in-chief), 

Ontogeny and Systematics of Fishes. Spec. Pubis. Am. Soc. Ichthyol. Herptol., 

No.l: 206-218. 
Rosen, D. E. 1971. The Nacristiidae, a Ctenothrissiform family based on juvenile 

and larval scopelomorph fishes. Am. Mus. Novit., 2452: 1-22. 
1973. Interrelationships of higher euteleostean fishes. In Greenwood, P. 

H., R. S. Miles, and C. Patterson (eds.), Interrelationships of Fishes. Zool. J. 

Linn. Soc. 53 Suppl. 1: 397-513. 
1985. An essay on euteleostean classification. Am. Mus. Novit., 22877: 



1-45. 

Stiassny, M. L. J. 1986. The limits and relationships of the acanthomorph tele- 
osts. J. Zool., Lond., (B)l: 1-50. 

Sulak, K. J. 1977. The systematics and biology of Bathypterois (Pisces, Chlo- 
rophthalmidae) with a revised classification of benthic myctophiform fishes. 
Galathea Rept., 14:49-108. 

Theisen, B. 1965. On the cranial morphology of Ipnops murrayi Gunther, 1878 
with special reference to the relations between the eyes and the skull. Galathea 
Rept., 8:7-17. 

Weitzman, S. H. 1974. Osteology and evolutionary relationships of the Sternop- 
tychidae, with a new classification of stomiatoid families. Bull. Am. Mus. Nat. 
Hist., 153(3): 329-478. 

APPENDIX 

MA TERIAL EXAMINED 

Parasudis material 

The single figure given in parenthesis after the catalogue number 
is SL; if more than one specimen is registered the number of speci- 
mens is given and followed by their size range. 

P. truculent a 

Larvae: ARC uncat. (42.8 mm), 42°27'N 58°27'W, 0-300 m, 4 
Apr. 1979, USSR Belogorsk, sta. 35-4. ARC 6729 (37.3 mm), 40° 
22'N 62°38'W, 0-200 m, 20 Mar. 1979. ARC 6732 (28.0 mm, as.), 40° 



22 BREVIORA No. 487 

51'N 61°34'W, 0-50 m, 21 Mar. 1979. ARC 6845 (41.2 mm), 39°44'N 
56°59'W, 0-200 m, 1 Apr. 1979. ARC 6879 (80.0 mm), 42°27'N 58° 
27'W, 0-300 m, 4 Apr. 1979. MCZ 58531 (28.5 mm), JEC 7755, 1 1° 
12'N 53°49'W, 60 m, 27 Mar. 1977. MCZ 62397 (38.0 mm, c.s.), JEC 
7715, 0° 13'N 35°44'W, 90-100 m, 15 Mar. 1977. MCZ 62398 (30.0 
mm, c.s.), JEC 7750, 10°48'N 52° 17'W, c.80 m, 26 Mar. 1977. MCZ 
62399 (30.6 mm), JEC 7753, 10°48'N 52°17'W, c.140 m, 27 Mar. 
1977. MCZ 62400 (15.4 mm), MOC 10-118, 3, 34° 12'N 75°00'W, 
70-0 m, 14 Aug. 1978. MCZ 63055 (40.0 mm), JEC 7737, 38°55'N 72° 
25'W, 0-825 m, 3-4 Sept. 1976. MCZ 63056 (31.0 mm), JEC 7738, 
38° 5 1 'N 72° 27'W, 0-750m, 4 Sept. 1 976. Plus an additional 32 uncat- 
alogued MCZ specimens (11.7-49.1 mm) from the WHOI collec- 
tion. USNM 258610 (20.0 mm), 31°50'N 63°56'W, 40 m, 23 Feb. 
1972. 

(ZMUC) DANA 1166V (11, 14.6-24.1 mm), 100 meters of wire 
out (mw) and DANA 1166VI (2, 14.3-21.2 mm), 50 mw, both 10° 
16'N 40°41'W, 11 Nov. 1921. DANA 11681V (41.4 mm), 300 mw, 
and DANA 1 168VI (22, 13.8-23.9), 50 mw, both 9°30'N 42°41'W, 12 
Nov. 1921. DANA 1 190VIII (13.1 mm), 17°58'N 64°45'W, 100 mw, 
13 Dec. 1921. DANA 1 1941 (10.6 mm), 17°58'N 64°41'W, 320 mw, 
12 Dec. 1921. DANA 1202IV (12.5 mm), 100 mw, and DANA 1202 
(1 1.3 mm), 50 mw, both 9°40'N 79°56'W, 10 Jan. 1922. 

Transforming larvae: MCZ 57922 (75.5 mm), 1 1° 36'N 62°46'W, 
530m, 19 Apr. 1960. MCZ 62401 (85.0 mm) MOC 20-19, 0, 39° 13.51M 
71° 17.6'W, 0-1027m, 15 June 1982. 

Adults: MCZ 39968 (9, 130-190 mm), 07° 10'N 53°07'W, 360 m, 
6 Nov. 1957. MCZ 40561 (22, 90-175 mm), 07°34'N 54°50'W, 
360 m, 6 Nov. 1957. MCZ 40561 (3, 90-102.2 mm c.s.), 07°34'N 
54°50'W, 6 Nov. 1957. MCZ 41840 (85.0 mm), 18° 16'N 67° 17'W, 
450 m, 6 Oct. 1959. UMML 15608 (120 mm, c.s.) no data. 

P. fraser-brunneri 

Larvae: MCZ 62402 (31.5 mm), RHB 2053, 16°32'N 19°35'W, 
50-56 m, 13 Nov. 1970. 

Adults: USNM 245646 (3, 84-138), 04°26'N 08°29'W, 200 m., 
31 Oct., 1963, BBC 843. MCZ 63153 (3, 97-105, one specimen 
c.s.), same data as USNM 245646. 



1986 LARVAL PARASUDIS RELATIONSHIPS 23 

Comparative material 

STOMIIFORMES 

Gonostomatidae — Diplophos taenia: MCZ 52562, MCZ uncat. c.s.; 

Gonostoma elongatum: MCZ 62404; Gonostoma sp.: MCZ 

uncat. c.s. 
Sternoptychidae — Maurolicus muelleri: MCZ 62598, MCZ uncat. 

c.s. 
Photichthyidae — Photichthys argenteus: MCZ 56953; Polymetme 

corytheola: MCZ 56968, MCZ uncat. c.s. 

AULOPIFORMES 

Aulopidae — Aulopus nanae: MCZ 40516; A. japonicus: MCZ 

45169 c.s. 
Chlorophthalmidae — Chlorophthalmus agazzisi: MCZ 40539, MCZ 

40509 c.s., MCZ 41444 c.s.;C bicornis: BMNH 1939.5.24:457; 

C. brasiliensis: MCZ 51365, MCZ 40564 c.s.; C. chalybeius: 

MCZ 62155, MCZ 40564 c.s.; C. nigripinnis: BMNH 

1887.12.7:207; Chlorophthalmus sp.: MCZ 62403 c.s., MCZ 

62591, MCZ 62592, MCZ 62593, MCZ 62597. 
Ipnopidae — Ipnops murrayi: MCZ 41133; Bathypterois phenax: 

MCZ 57624; B. quadrifilis: MCZ 35598; B. viridensis: MCZ 

40567 c.s.; Bathysauropsis gracilis: BMNH 1887.12.7:209-210. 
Notosudidae — Scopelosaurus argenteus: MCZ 62405 c.s., MCZ 

62406 c.s.; S. harryi: MCZ 40512. 
Scopelarchidae — Scopelarchus analis: MCZ 62599 c.s. 
Bathysauridae — Bathysaurus agazzisi: MCZ 55305 c.s., MCZ 62409 

c.s.; B. mollis: MCZ AWAQ 
Synodontidae — Synodus synodus: MCZ 47378, MCZ 47490 c.s.; S. 

jaculum: MCZ 46972 c.s.; Saurida brasiliensis: MCZ 62408 

c.s.; Sa. tumbil: MCZ 59273; Sa. undosquamis: MCZ 56111 

c.s. 
Harpadontidae — Harpadon sp.: MCZ uncat. c.s. 
Alepisauridae — Alepisaurus brevirostris: MCZ 60345, MCZ 43153 

c.s. 
Evermanellidae — Evermanella sp.: MCZ uncat. c.s. 
Paralepididae — Paralepis elongata: MCZ 43140; P. speciosa: MCZ 

60332 c.s.; Lestrolepis intermedia: MCZ 62407 c.s.; Sudis 

atrox: MCZ 60336 c.s. 



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