(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Breviora"



/'•.v 



MP'S 



'."■Gi'- 














,^>'\ 




' i ■/ 




/V  '1 














.■A.>) 



HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



B R E V I O R A 

Miiseiim of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 21 October 1987 Number 488 



A NEW FLYING LIZARD FROM THE SANGIHE 
ARCHIPELAGO, INDONESIA 

James D. Lazell, Jr.' 

Abstract. A new species of Draco, characterized by small size (to 75 mm 
SVL), reduced sexual dimorphism, somber coloration, five ribs in patagium, and 
eight to ten postrostrals, is described from Pulau Biaro, southernmost isle of 
Kepulauan Sangihe, ca 60 km north of the northeast tip of Minahasa, Sulawesi 
Utara. 

Spanning some 450 km between Sulawesi and Mindanao, form- 
ing the northwestern limit of the Molucca Sea, are more than 40 
islands on 15 submarine banks. Those closest to Sulawesi are 
named for their largest member: Kepulauan Sangihe— the Sangihe 
Archipelago. The southernmost of these, some 60 km northeast 
of Ponto do Celebres, and about 25 km from the next nearest 
land (Ruang), is the isle of Biaro. Like its sisters, Biaro is of 
volcanic origin. I suspect it arose just where we find it today, did 
not drift there from somewhere else, and has never had any ter- 
restrial connection to any other land area. 

The flying lizards, genus Draco, recently have been reviewed 
by Musters (1983) and Inger (1983). Their views are disparate. 
Only Musters admits Draco in the Sangihe Archipelago. He says 
D. volans boschmai is "perhaps on the Kepulauan Sangihe." He 
examined no specimens from these islands and only one volans 
from Sulawesi (that from Macassar in the extreme southwest). 



' Associate, Department of Herpetology, Museum of Comparative Zoology, Har- 
vard University, Cambridge, Massachusetts 02138, and The Conservation Agen- 
cy, 6 Swinburne Street, Jamestown, Rhode Island 02835. 

MCZ 

OBT 1 i 1187 



124' 



BREVIORA 



126' 



No. 488 
128° 



2°- 




MINDANAO 



:&. 



'Miangas 



KAWIOi^f": 



^:ofNENUSA 



Manado 



0' 



124° 



'P'- p "■. 



^:'. 




TALAUD 



ISJSANGIHE 



ii-0 
* 



'o^ 




Ruang 
Biaro 



Batu Putih 
SULAWESI 



t 

N 



100 Km 



126"= 



0' 



128' 



Figure 1. The northwestern Molucca Sea showing the islands and banks 
(200 meter depth) and the locations of places mentioned in the text. 



1 987 ANEW FL YING LIZARD 3 

After carefully studying both papers and examining many spec- 
imens at the Museum of Comparative Zoology, I conclude that 
our knowledge of Draco today is similar to our knowledge of 
Anolls in 1950. Nevertheless, I have returned from Biaro with 22 
specimens of a Draco so distinctive that I have no hesitation in 
describing it as new. 

Draco biaro sp. nov. 
(Fig. 2) 

Type. MCZ 170898. 

Type Locality. Pulau Biaro, Kepulauan Sangihe, Indonesia. 
Bernard F. Page coll., 4 April 1986. See Figure 1. 

Paratypes. MCZ 170899-170919, same locality as the type; B. 
Page, J. Lazell, and local children coll., 4 April 1986. 

Diagnosis. A small species of Draco, adults to 75 mm SVL; 
five ribs in patagium; nostrils pointing laterally; most (90%) with 
small scales covering tympanum; four or five incisors (Inger's 
method); six to eight supralabials (73% have seven); eight to ten 
scales bordering rostral (64% have nine); male throat fan small, 
55 to 82% of head length (Musters' method) in adult males; col- 
oration in life of both sexes somber, patagium sooty to slaty grey 
above with 15 to 20 sublongitudinal to subradiate narrow light 
grey streaks. 

Description of the Type. Adult male, 73 mm SVL, with a com- 
plete tail 145 mm (199% SVL). There are eight weakly keeled 
supralabials and nine scales border the rostral. The tympanum is 
clothed in small scales. The throat fan is 71% of head length, 
hooked distally, and blunt at the tip. See Figure 2. 

The dorsals are smooth to weakly keeled; there are 15 in the 
standard distance (tip of snout to center of eye) at midbody. The 
ventrals are sharply keeled; there are 16 in the standard distance 
at midbody. There are 27 subdigital lamellae beneath the fourth 
toe of the pes, counting from that toe's separation from the third 
toe. 

There are no thomlike supraciliaries although a few anterior 
supraciliaries are enlarged and keeled. The Y-shaped arrangement 
of keeled snout scales is interrupted and only vaguely discernible. 
The lappets are edged by somewhat enlarged scales, but the throat 



BREVIORA 



No. 488 





Figure 2. Draco biaro sp. nov.. The type: MCZ 170898, from the island of 
Biaro, Sangihe Archipelago, Sulawesi Utara. Indonesia. 



fan bears only small ones. The caudal scales are subequal; there 
is no caudal crest. A low nuchal crest consists of about 22 enlarged, 
tectiform middorsals. The adpressed hindlimb just reaches the 
forelimb insertion. 

Teeth were not counted in this specimen because the number 
is inconsequential in relevant species and opening the mouth may 
entail damage. 

In life the type was largely grey. There was a beige-tan wash on 
the cheeks and jowls where the pattern was of irregular mottling. 
There are two bold, sooty nuchal blotches; between and flanking 
these is strongly contrasting pale ash-grey, making a roughly 
H-shaped figure, viewed from above. The dorsum is irregularly 
banded with light and dark grey in a lichcnate pattern. The chest 
was washed with dull yellow. The throat fan was pale lemon- 



1987 A NEW FLYING LIZARD 5 

yellow— almost while— with light grey barring. The dorsal surface 
of the patagium was sooty to slaty grey with a series of narrow 
ash-grey lines. The translucent, membranous skin of the flanks 
and ventral surfaces of the patagia looked light greenish with the 
patagia folded; this results from a pale bluish ventrolateral wash 
shading to pale yellow distally. When the patagia are expanded 
the dark dorsal color dominates, especially as sooty anterolateral 
blotches. The lappets were dull grey above, paler and yellowish 
grey below. 

Variation. The sexes are quite similar. Six adult males measured 
70 to 75 mm; seven adult females measured 71 to 75 mm; both 
sexes averaged 73 mm SVL. Eight of these 13 adults show some 
degree of supralabial carination. There may be low, somewhat 
irregular ridges (e.g., MCZ 170912, a male) or prominent, strong 
keels (e.g., MCZ 170910, a female). 

I elected to quantify trunk scale size by the standard distance 
count method used in iguanid work (e.g., Lazell, 1972). In Draco 
biaro the middorsals are juxtaposed, subimbricate, and smooth, 
keeled, or weakly tectiform; there are 12-17 (average 15) in the 
standard distance. The ventrals are always sharply keeled and 
fairly well-aligned in transverse rows; there are 14-21 (average 
16) in the standard distance. 

There are 26-29 (average 28) subdigital lamellae on the fourth 
toe of the pes, distal to its separation from the third toe. 

I could detect no sexual dimorphism in any mensurable or 
meristic characters. 

Twenty specimens of Draco biaro have small scales, arranged 
in a whorl, over the tympanum. Two depart from this norm: 
MCZ 1 70899 has large scales over the tympanum. In MCZ 1 709 1 9 
the condition seems intermediate between a large scale and the 
thin, smooth skin of a typical tympanic membrane. 

All specimens were very similar to the type in coloration in 
life. Females show little of the beige-tan wash on cheeks and jowls. 
Females have very small, unmarked grey throat fans, but the 
lappets are like those described for the male. The patagia of both 
sexes are similar. Juveniles tend to have a more strongly con- 
trasting lichenate dorsal pattern in shades of grey than do large 
adults. 

Inger and Musters concur that Draco of both the lineatus and 



6 BREVIORA No. 488 

volans complexes have four or five incisiform teeth. Musters sub- 
tracts two from the total between the caniforms to approximate 
real incisors while Inger gives the total count; I used Inger's meth- 
od because I cannot readily see which teeth are actually socketed 
in the premaxillaries. I checked five paratypes of Draco biaro, 
MCZ 170913-17. Only MCZ 170914 lacks a median tooth or 
socket and really seems to have four teeth. MCZ 1 709 1 7 has four 
teeth, one median, and one empty socket. MCZ 170915 has only 
three teeth, one median, and at least one empty socket. Five is 
probably the normal count for the species. 

Comparisons. In the key provided by Musters (1983), Draco 
biaro goes to the D. lineatus complex. Both Musters and Inger 
(1983) agree that D. volans normally has six ribs in the patagium, 
a number not seen in any D. biaro. A close reading of both texts 
renders the case more equivocal, however. There seems to be no 
absolute distinction between these nominal, polytypic species. 
Diagnoses are compromised by the great variation exhibited with- 
in both the lineatus and volans assemblages. 

In Table 1 I list some characters used by either Musters or 
Inger, or both, in diagnoses. My caveat is that many of the given 
character states are not absolute. Species recognition in Draco 
may well depend on finer grained analyses, including extensive 
knowledge of coloration in life, and field knowledge of ecology 
and behavior. This sort of knowledge helped Inger (1983: 8-15) 
separate D. maximus and D. quinquefasciatus at Nanga Telakit, 
Sarawak. 

On 28-29 April, 1986, 1 collected two series of Draco near Batu 
Putih in northeastern Minahasa, Sulawesi Utara. This locality is 
about 36 km northeast of Manado, type-locality of Draco lineatus 
spilonotus (taxonomy agreed by all workers). Batu Putih is about 
60 km south of Biaro. Field knowledge and fresh specimens from 
Minahasa made me sure I was seeing a new species on Biaro. 

Because the Minahasa series differs from key characters given 
by Musters (1983:34), and because coloration in life is so rarely 
known (in proportion to its probable extreme value in species 
recognition), I provide a brief description of D. I. spilonotus here. 

My series, MCZ 170922-933, includes five adult males, four 
adult females, and three juveniles. There is striking sexual di- 
morphism. The largest female is 72 mm (MCZ 1 70930), the larg- 



1987 



ANEW FL YING LIZARD 



Table 1. Seven ways in which species of Draco from Sulawesi differ. 





volans 


lincatus 


hiaro 


Ribs 


6 


5 


5 


Snout Y 


yes 


yes 


no 


Thorn 


distinct 


weak 


absent 


Postrostrals 


4-6 


5-7 


8-10 


Hindlimb 


no 


yes 


yes 


Tympanum 


skin 


scales 


variable 


Size 


96 


91 


75 



Ribs are those within and supporting the patagium. The snout Y is composed of 
continuous, enlarged, keeled scales. The thorn is an enlarged, pointed, anterior 
supraciliary. Postrostrals are the small scales in contact with the rostral, counting 
the first supralabials. The hindlimb is adpressed to determine if it is as long as 
the distance to the forelimb insertion ("yes") or not ("no"). The tympanum may 
be covered with undifferentiated small scales or smooth skin (see text). Size is 
maximum snout-vent length (SVL); the number is for a female in both lineatus 
and volans. but in hiaro the sexes are equal. Size varies geographically in both 
wide-ranging species. 



est male 64 mm (MCZ 170925) snout to vent. The male throat 
fan is relatively long, 96 to 102% (average 99) of head length; it 
is nearly triangular, gradually tapering, and acutely pointed. 

The males are brilliantly colored. The entire head and neck 
region is boldly spotted and marbled with chartreuse, aquamarine, 
and copper-tarnish green on an olive-beige ground. On the trunk 
this ground color is marbled with grey. The patagia are bright 
salmon pink, orange, or orange-yellow. The belly is green. Both 
lappets and throat fan vary from brilliant lemon to sulfur yellow. 

The females are darker and duller. The head and neck mottling 
is in shades of olive green and olive brown. The patagia are deep 
rich yellow or orange-yellow spotted or barred with near black. 
The lappets and throat are light yellow. Both sexes have some 
power of color change, to lighter or darker. This change does not 
seem to affect the patagia, lappets, or throat fan. 

My specimens differ most notably from those described by 
Musters (1983) in patterning of the head and neck. They have 
retained their bold patterns in alcohol (three months at time of 
writing), though the bright colors have faded. The significance of 
the differences cannot be judged without far more extensive 
knowledge of populations in life. 



8 BREVIORA No. 488 

I have examined six specimens o^ Draco lineatus bimaculatus, 
MCZ 26178-82 and 43640, from Mindanao. In these the rostral 
is tiny compared to that of D. biaro. The eye is roofed by large, 
plate-like, keeled supraciliaries. The enlarged, aligned, keeled scales 
on the frontal region form an arrow-shaped pattern, not a Y. 
There are 10 to 12 supralabials (60% have 1 1). A more cursory 
look at all other Philippines material in MCZ further convinces 
me that the relationships of Draco biaro do not lie with known 
Draco from that area. 

On balance, the affinities of Z). biaro seem to lie with the lineatus 
complex not the volans group. I predict the discovery of many 
more island forms in the Sangihe, Kawio, Nenusa, and Talaud 
archipelagos between Sulawesi and Mindanao. 

Comments. Draco biaro is common on its small island, fre- 
quenting coconut palms and other smooth-barked trees. Most 
were encountered two to four meters from the ground and noosed 
with a long pole. Often they fled up the trunks and children 
climbed after them. They sometimes ascended more than 20 me- 
ters. Eventually, when pursued, they would launch and glide. Then 
one could observe two large, middle-aged men and several dozen 
children racing through the grass and brush after the flying lizard, 
which sometimes landed low enough to be caught by hand. 

Courtship was often observed. The male rapidly extends the 
throat fan and lappets several times and then fans the patagia. 
Most adult females palpably contain eggs. Two eggs, MCZ 1 70920- 
21, were laid in a collecting bag with several females during the 
hours between capture and pickling. One egg was broken, but 
MCZ 170920 measures 14.7 by 7.8 mm. It is white and leathery. 

Pulau Biaro is subtended to the south by at least one small 
coastal cay. Coconut palms and other trees grow on this cay, but 
I did not visit it. Draco biaro may occur there. 

Six other species of reptiles were collected on Biaro on 4 April: 
the vine snake Ahaetul/a prasina, the skinks Mabiiya multifasciata 
and Lamprolepis smaragdinus, and three geckos. Hemidactylus 
frenatus and Gehyra miitilata are abundant Indo-Pacific human 
commensals. Gynmodacty/us Jellesmae is rare in collections and 
seems to have been previously known only from Sulawesi. 



1987 A NEW FL YING LIZA RD 9 

ACKNOWLEDGMENTS 

I am indebted to Bernard Page, William Disher, Karen Phil- 
lipps, and Mark Hopkins, my companions in the field during most 
of our Indonesia expedition. Mark Hopkins took excellent color 
photographs of living Draco lineatus near Batu Putih. The people 
of Biaro, Minahasa, and the other areas we visited were enthu- 
siastically hospitable and helpful. Franklin Ross curated and 
accessioned the material into MCZ in a most expeditious manner. 
Mauyra Twitchell contributed Figure 2, drawn from photographs 
by Greg Mayer. The entire expedition was funded by The Con- 
servation Agency. 

LITERATURE CITED 

Inger, R. F. 1983. Morphological and ecological variation in the flying lizards 
(genus Draco). Fieldiana, Zoology, New Series, 18: vi + 35 pp. 

Lazell, J. D. 1 972. The anoles (Sauria: Iguanidae) of the Lesser Antilles. Bulletin 
of the Museum of Comparative Zoology, 143(1): 1-1 15. 

Musters, C. J. M. 1983. Taxonomy of the genus Draco L. (Agamidae, Lacertilia, 
Reptilia). Zoolische Verhandelingen 199: 1-120 + 4 plates. 



B R E V I O R A 

Museum of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 2 September 1988 Number 489 



NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. V. 

ANOLIS DANIELI, A NEW SPECIES OF THE LATIFRONS 

SPECIES GROUP AND A REASSESSMENT OF 

ANOLIS APOLLINARISBOV\J£.NG¥.K, 1919 

Ernest E. Williams' 

Abstract. A new giant anole, Anolis danieli, is described from northern and 
western Antioquia, Colombia. Formerly confused with A. apollinaris Boulenger, 
1919, of Cundinamarca and southeastern Antioquia, the new species differs in 
the presence of a dewlap of moderate size in the female (absent in A. apollinaris) 
and in minor scale characters. A. danieli, A. apollinaris and A. propinquusV^ iWiams, 
1 984, are demonstrated to be a distinct subgroup within the latifrons species group 
defined by distinctly keeled head scales, relatively short limbs and a green ground 
color. Previous confusions regarding the taxonomic placement of ^. apollinaris 
and A. propinquus are corrected. 

INTRODUCTION 

In 1970 I redescribed Anolis apollinaris Boulenger, 1919, pri- 
marily on the basis of a series of six specimens from San Pablo, 
Department of Cundinamarca, Colombia, in the Munich collec- 
tion (ZSM 427—432) and the type specimen in the British Museum 
(BMNH 1946.13.22). I referred to the species' three additional 
specimens from Cundinamarca— two from Antioquia and one 
from Caldas. I compared the species only with^. biporcatus Wieg- 
mann. 

Material since made available to me from Cundinamarca and 
Antioquia now makes it quite clear that my 1970 material was 



' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 



2 BREVIORA No. 489 

composite, and that only the Cundinamarca specimens were A. 
apoUinaris. Fortunately, the illustrations in Williams (1970) are 
of Munich specimens from Cundinamarca, which are true A. 
apoUinaris. The specimens from Antioquia represent a new, al- 
though very closely related species. (The specimen from Caldas 
has not been re-examined.) Further, it can now be established 
that both A. apoUinaris and the new species belong to the alpha 
section of the genus (Etheridge 1960) and must be referred to the 
AnoUs latifrons species group {sensu Williams 1976); they are not 
at all close to A. biporcatus, which is a member of the beta section. 

Confusion as to the placement and affinities of ^. apoUinaris 
and as to the affinity or lack of affinity of the biporcatus and 
latifrons species groups has had a long history. Boulenger (1919) 
in his description of A. apoUinaris expressed no opinion about 
its relationships. Burt and Burt (1931) referred a number of Co- 
lombian anoles to the species, but were quite wrong in their iden- 
tifications as Dunn (1944) demonstrated. Dunn's own judgment 
was most importantly based on size (SVL of the type specimen 
o^ A. apoUinaris 106 mm) and he compared the species with A. 
soUfer and A. copei (both synonyms of ^. biporcatus), which are 
approximately this size. Unfortunately, Dunn did not compare 
apoUinaris with the other group of species, well known in western 
Colombia, that is comparable in size, in spite of the fact that he 
had previously reviewed this group— his "mainland giant anoles" 
(Dunn 1937). 

Confusion of the biporcatus and latifrons species groups first 
occurred when Giinther (1 859), dcscnhing AnoUsfraseri, included 
a specimen o{ A. biporcatus in the type series. Boulenger (1885) 
corrected the error at the species level, but apparently, as I have 
commented earlier (Williams 1966), still believed that the two 
taxa were close relatives. Dunn (1944) committed a parallel error 
in the reverse direction by associating A. apoUinaris with the two 
synonyms of /I. biporcatus. 

In fact, none of the standard external characters used in anole 
taxonomy permit the placement of the biporcatus and latifrons 
species groups as widely separate taxa. Species characters are clear 
enough, but there is quite obviously marked convergence in eco- 
morphic features (sensu Williams 1972, 1983). The significant 
internal character of the caudal vertebrae— anteriorly pointing 



1988 ANOLIS DANIELI 3 

transverse processes on these vertebrae in the beta section of 
Anolis and the absence of these processes in the alpha section of 
the genus— was discovered by Etheridge (1960) only with the aid 
of X-rays. 

This character was, of course, known to me in 1970, but by 
misfortune in 1 970 no suitable X-ray equipment was conveniently 
available to me or to Etheridge (on whom I usually relied for 
assistance in this particular) nor were any dry skeletons available 
(there are still none). Therefore, I contented myself at that time 
with externals. Influenced by minor aspects of color pattern— 
green with some white spotting, which appeared to eliminate A. 
fraseri as a close relative— and by the short limbs, shared with A. 
biporcatus as well as A. fraseri, and quite unlike the long limbs 
of the frenatus subgroup of the latifrons assemblage, I accepted 
Dunn's (1944) allocation. 

The recognition of a second species related to A. apollinaris 
and of the alpha affinities of both species began when two large 
Anolis from Antioquia belonging to the collections of the Museo 
de Historia Natural at the Colegio San Jose in Medellin (CSJ 1 1 1 
and 168, now ICN 5997-98) were turned over to me by Stephen 
Ayala for examination. The female, CSJ 1 1 1 from Yarumal, had 
a very evident large dewlap and bore a paper label in Niceforo 
Maria's handwriting: ^'Anolis purpurescens ." A caudal vertebra 
teased from its broken tail showed that it belonged to the alpha 
section of the genus, yet the scale counts routinely taken on Anolis 
specimens were disturbingly similar to those of A. apollinaris. 
The latter, however, was not only believed to be a beta anole, 
but, in the Munich series I had studied in 1970, five of the six 
were females, and they had shown at most a vestigial gular fold 
and not a dewlap. 

The absence of transverse processes on the caudal vertebrae of 
the Yarumal female and in A. apollinaris was verified by X-ray. 
The two specimens from Antioquia attributed to A. apollinaris 
in 1970 were re-examined: MLS 81, recatalogued as MLS 926, a 
female, and AMNH 38725, a male. Marco Antonio Sema pro- 
vided three additional specimens from Urrao, Antioquia, all males, 
from the Colegio San Jose collection. 

For comparison with the specimens from Antioquia, new ma- 
terial of verified v4. apollinaris has been required. Ten specimens 



4 BREVIORA No. 489 

collected by Juan Manuel Renjifo at Sasaima, Cundinamarca (IN- 
DERENA 2853-62) and two collected by Jose Vicente Rueda at 
Charala, Santander (ICN 2865, 6017— a new record) have been 
available, as well as additional Cundinamarca specimens from 
the Museum of Comparative Zoology (MCZ) and the Instituto 
de Ciencias Naturales (ICN) in Bogota. These comparisons fully 
established the distinctness of the Antioquian population, which 
may now be formally described as a new species to be named in 
honor of Hermano Daniel Gonzalez, now Director of the Museo 
de Historia Natural at the Instituto de La Salle, Bogota, in rec- 
ognition of his 37 years of association with the Colegio San Jose 
in Medellin, Antioquia: 

Anolis danieli, new species 

Holotype. ICN 5997 (formerly CSJ 111), adult male. 

Type Locality. Urrao, Antioquia, Colombia. Collector and date 
of collection unknown. 

Paratypes. Antioquia, Puerto Antioquia, Baja Rio Cauca: MLS 
926, Hno. Ignacio Saza coll., January 1963. Sabanalarga: AMNH 
38725, Hno. Niceforo Maria coll., no date. Urrao: MCZ 164894 
(formerly CSJ 278), Marco A. Sema coll., 28 May 1972; CSJ 441, 
M. A. Sema and H. Echeverri coll., 23 March 1983; CSJ 720, M. 
A. Sema and H. Echeverri coll., 26 July 1985. Yammal: ICN 
5998 (formerly CSJ 168), collector and date of collection un- 
known. 

Referred Specimen. "Westem Colombia:" AMNH 4844, col- 
lector and date of collection unknown. 

Diagnosis. A species very close to A. apollinaris but differing 
in the presence of a moderately large dewlap in the female (rather 
than a mere longitudinal fold indicating the position of such a 
structure), and in the possession of a differentiated anterior nasal 
(rather than a circumnasal separated from the rostral by a post- 
rostral). Also, by having the keels in the frontal depression with 
keels oriented anteroposteriorly (rather than keels radiating from 
the center of the depression); by having a distinct parietal depres- 
sion usually bounded by ridges (rather than a shallow depression 
never distinctly set off from the occiput); and by having the scales 
anterior and anterolateral to the interparietal subequal in size to 
those posterior to it, except for the scale row that abuts on the 



1988 ANOUS DANIELI 5 

semicircles, which is abruptly larger (rather than a// scales anterior 
and anterolateral to the interparietal markedly larger than those 
posterior to it). 

Description. Head. Head scales moderate to small, rugose or 
obtusely to strongly keeled. Eight to 12 scales across the snout 
between the second canthals. A moderate frontal depression, the 
scales within it slightly smaller than the surrounding scales and 
with keels oriented anteroposteriorly (flat in MLS 926). Five to 
8 scales border the rostral posteriorly. An anterior nasal scale 
differentiated (in ICN 5998, on one side, divided into upper and 
lower portions), in contact with the sulcus between rostral and 
first supralabial. About 7-8 scales between the supranasals dor- 
sally. 

Supraorbital semicircles separated by 3 scales, the middle one 
smallest, or (in AMNH 38725 and MLS 926) separated by 4 scales 
equal in size. Supraocular disk ill-defined but the medial scales 
longer and bluntly keeled, in contact with the supraorbitals or 
separated by one row of small scales. About 7 scales across the 
supraocular area between the supraorbitals and superciliaries. One 
to 3 elongate superciliaries anteriorly, flanked medially by mod- 
erately enlarged polygonal scales and continued posteriorly by 
granules. About 5-6 rather narrow canthal scales, the second larg- 
est, decreasing regularly in size forward. Five to 7 loreal rows, 
subequal or irregular in size. 

Temporal scales granular. An indistinct double line of slightly 
enlarged intertemporal scales. Supratemporals increasing in size 
laterally toward the margins of the parietal depression. Interpa- 
rietal round, slightly to much smaller than ear (indistinct or absent 
in MCZ 164894 and AMNH 4844). Two to 5 scales on each side 
between interparietal and semicircles. Three to 5 rows of scales 
behind interparietal larger than nape scales. 

Suboculars separated from supralabials by 1 scale row or nar- 
rowly in contact. Seven to 9 supralabials to below the center of 
the eye. 

Mental divided or nearly so, each half about as wide as long. 
Five to 8 scales behind the mental and between the infralabials. 
Two of these may be differentiated sublabials; if differentiated, 
as many as six moderately enlarged scales in sequence with the 
sublabials may be in contact with the infralabials. Central throat 



6 BREVIORA No. 489 

scales small, swollen, smooth or obtusely keeled, becoming grad- 
ually larger adjacent to the infralabials. 

Dewlap. Large in male, extending onto first third of belly, nearly 
as large in female, extending past axilla. With crowded scale rows 
in both sexes, and scales on the skin between the rows; lateral 
scales irregular and weakly keeled in males, flatter and more reg- 
ular in females; edge scales larger than ventrals and bluntly keeled 
in males, smooth and subequal to ventrals in females. 

Trunk. Middorsals distinctly keeled, to 4 rows slightly en- 
larged. Flank scales bluntly conical or pyramidal, separated, with 
each scale conspicuously surrounded by minute granules. Ventrals 
larger, squarish, subimbricate, smooth or slightly keeled. 

Limbs. Upper arm scales swollen, unicarinate or smooth, sur- 
rounded by minute granules like the trunk scales. Lower arm 
scales more crowed, sometimes larger, imbricate and multicari- 
nate. Thigh scales crowded, swollen, imbricate, unicarinate an- 
teriorly, small, subconical, separated posteriorly. Tibial scales 
larger anteriorly, distinctly or weakly unicarinate, separated, pos- 
teriorly smooth, subimbricate. Supradigitals of hand and foot 
multicarinate. Twenty-three to 27 lamellae under phalanges ii 
and iii of fourth toe, pad rather narrow. 

Tail. Long, about 3 x snout-vent length, slender, slightly com- 
pressed, fragile, but breaks apparently not across vertebrae. 

Size. The largest specimen of the type series is the male holotype 
(SVL 117 mm, tail length 331 mm). AMNH 4844 is a larger 
specimen (SVL 125 mm) but has not been made a paratype be- 
cause it has an obscure dorsal pattern of broad transverse bands 
not seen in the type material and has only the inexact locality 
"western Colombia." The largest female, from Urrao, like the 
holotype, has an SVL of 104 mm. A. apollinaris may be a slightly 
smaller species. The largest male (INDERENA 2856) has an SVL 
of 1 12 mm, the largest female (MCZ 156308) 94 mm. 

Color in Life. For most of the few specimens o{ A. danieli there 
is no data on color in life. The best information (translated) has 
been provided by Marco Antonio Sema for CSJ 820, a male: 

Back completely green with a few elongate spots of even 
brighter green dorsolaterally. A broad yellowish band extends 
from behind the eye to the dorsal crest, and a second band 



1988 ANOLIS DANIELI 7 

of similar color extends from behind the ear to a more pos- 
terior position on the dorsal crest. A yellowish white band 
above the forelegs. Pale yellow around the eye. Gular region 
yellowish green. Dewlap yellow with whitish scales. All the 
belly greenish yellow with a little blue ventrolaterally. Tail 
green with blackish bars. Palms of fore and hind feet whitish. 
Fore and hind legs green with barely perceptible bars of slight- 
ly darker green. At least twice during its life in captivity the 
animal changed to brown. When killed, it immediately began 
to change to rust brown. 

This description may be compared with three descriptions of 
color in life for A. apollinaris that I have been able to obtain. W. 
W. Lamar reports for a female specimen from Sasaima, Cundi- 
namarca: 

Top of head yellow green. Eyelids bright saffron yellow. A 
broad tan stripe continuous from neck to well down on tail 
where it is replaced by black bars. Side of head behind eye 
blue green to intense green. A pale greenish white line across 
upper labials to ear. Dewlap rather small and yellow green. 
Venter bluish green becoming more so distal to hind limb. 
A few poorly developed ocelli on sides of body. 

Stephen Ayala, reporting on animals from the same general 
locality, gives the following details: 

Anolis apollinaris is a green lizard, with a prominent white 
line or zone under the eye between the snout and the sides 
of the neck. The green changes to dark brown in less than 
half a minute. Small white spots or thin diagonal lines may 
be seen on the sides of the female, and some females have a 
broad tan vertebral stripe covering the entire back and tail. 
Light brown, saddle-shaped spots or bands may appear across 
the back of the male (especially in the dark phase) and small 
or large blue or reddish spots occur on the shoulders or sides 
of the neck. The eyelids stand out because of their contrasting 
color: yellow in female and yellow orange in the male. The 
dewlap of the male is pale yellow green, with rows of green 
scales (brown scales in the dark phase). 



8 BREVIORA No. 489 

For the animals from Charala, Santander, a description by Jose 
Vicente Rueda is available (translated): 

Dorsally head and body senf. green (olive green), edge of 
supraobital semicircles and postparietals black. Middorsal 
body spot chestnut. Irregular symmetrical spots black with 
a bluish cast above the insertion of the forelegs. Symmetrical 
and irregular brown spots on the base of the hind legs. Tail 
with well-spaced transverse black bands. Sides: a white band 
extending from posterior supralabials to shoulder. Eyelids 
burnt yellow (rust yellow). Ventrally mental, gular, dewlap, 
chest and forelegs yellowish green. Belly, tail and hind legs 
chartreuse (cream yellow). 

It is clear from these and other descriptions and slides that both 
species change color readily and show different elements of the 
pattern at different times. Both are predominantly green anoles, 
and it may not be easy to distinguish them on color alone. 

Color in Preservation. Most of the few preserved A. danieli are 
dull dark gray-blue, lighter below, with obscure traces of cross 
bars middorsally and of light lines on the nape. The Yarumal 
female is a faded brown. Only AMNH 38725, the male from 
Sabanalarga, shows any distinctive pattern (well-depicted in Fig. 
5). This specimen has mottled blue on the flanks, with the nuchal 
crest black, with faint and narrow yellowish cross streaks. The 
head is more brownish, mottled, the light patch on the labials 
whitish and the streak continuing it above the ear suffused with 
blue. The wider black streak parallel to this contains whitish spots 
as does the similar black streak in front of the shoulder. Between 
the two black streaks is an area that is grayish anteriorly, grading 
into a general darker coloration posteriorly. The posterior body, 
limbs and tail are essentially pattemless, the tail more olive than 
blue. In general terms, but not in detail, this animal matches rather 
well the description of the color in life of CSJ 720 above. 

A rather similar but distinguishably different head and nape 
coloration is seen in the most patterned of the preserved A. apol- 
linaris that I have examined (ICN 2865, Fig. 6). 

A. apollinaris, although the body patterns may often be some- 
what obscure, shows even in preservative the patterns mentioned 



1988 ANOLIS DANIELI 9 

by Ayala: the saddle markings of males, the broad dorsal stripe, 
the small white spots ("ocelli" of Lamar) or thin diagonal lines 
of females. No such patterns have been seen in A. danieli. Even 
AMNH 38725— the most patterned of the small type series- 
shows no comparable patterns. 

A. apollinaris, however, is now relatively well known, both in 
life and as museum specimens. A. danieli, as here described from 
only eight specimens, is still very inadequately understood. The 
relative absence of body pattern in A. danieli must for the present 
remain a poorly supported conclusion. AMNH 4844, which I 
have excluded from the type series and which has very imprecise 
locality, does show obscure broad banding. 

In the Parque de Las Orquideas, the borders of which are only 
1 5 km from Urrao, a population that in most respects is closely 
similar to A. danieli but is boldly patterned is known from a series 
of 5 specimens. It is, however, restricted to shaded forest. The 
body pattern, uniform in all specimens, of broad dark cross bands 
enclosing small light spots is quite unlike that of the most pat- 
terned known A. danieli, and the animals seem to have a slighter 
slenderer body build. I have provisionally excluded this series 
from the hypodigm of A. danieli as a distinct, though obviously 
sibling, species. 

Ecology. Almost nothing is known of the ecology of A. danieli. 
CSJ 720 is reported from a garden within the city limits of Urrao, 
1 ,850 m elevation. AMNH 38725 may be from a somewhat lower 
elevation (Sabanalarga, 1,250 m), while ICN 5998 from Yarumal 
is presumably higher (Yarumal, 2,265 m) Only MLS 996 from 
Puerto Antioquia may not be montane; Caceres near Puerto An- 
tioquia is given as 85 m elevation, but sites above 1,000 m are 
relatively close by. Lack of precision in the older locality records 
makes any comment on altitudinal range at best tentative. 

If A. danieli is like other members of the latifrons group, it 
should occur at low to moderate heights on large trees but not in 
canopy. A. danieWs sibling, A. apollinaris, is known to behave in 
this fashion (observations by Juan Manuel Renjifo and student). 
A danieWs occurrence in gardens indicates that it is not restricted 
to shaded forest, and A. apollinaris similarly occurs in rather open 
situations (J. M. Renjifo, personal observation). Stephen Ayala 
also reports that he has seen A. apollinaris in guava and several 



10 BREVIORA No. 489 

Other trees in rural household "gardens" in areas of low forest in 
Cundinamarca, usually on the vertical trunks. 

Distribution. A. danieli occurs in the northern regions of both 
the Western and Central Cordilleras in Antioquia. So far as is 
known, it is endemic to the Rio Cauca drainage, extending from 
Puerto Antioquia and Yarumal in the north, to Sabanalarga and 
Urrao in the south; perhaps over a considerable range of eleva- 
tions, but rather clearly montane. It is apparently replaced in the 
Western Cordillera in the Parque Nacional Natural "Las Orqui- 
deas" by the unnamed and more boldly patterned sibling men- 
tioned above. To the east and southeast, it is represented by the 
species with which I previously confused it, A. apollinaris. 

One juvenile but unmistakable A. apollinaris (CSJ 435) is known 
from El Retire, 23 km southeast of Medellin in Antioquia. It is 
a female without a dewlap, with the anterior nasal separated by 
one scale from the rostral, and with the keels of the scales in the 
frontal depression radiating from the center. It has a distinctive 
pattern of diamond-shaped light rhombs on the middle of the 
back that matches perfectly the dorsal pattern of a juvenile A. 
apollinaris (MCZ 46422) from La Mesa, Cundinamarca. 

The El Retiro specimen implies a close approach of these two 
closely related species, so similar structurally and not separated 
by any obvious physiographic or ecological barriers. What hap- 
pens in the potential range of contact or overlap remains an open 
question. 

Comparisons. Most of the characters of the species of the la- 
tifrons species group as I now understand it are summarized in 
Tables 1-3. I have added to the species in the group as listed 
by Savage and Talbot (1978) not only A. apollinaris (removed 
from the biporcatus species group of Williams 1970, 1976) but 
also A. propinquus Williams, 1984, described from a hatchling 
and in the description erroneously referred to the punctatus species 
group. 

A. apollinaris, A. danieli and the still unnamed danieli sibling 
from the Parque Las Orquideas, along with A. propinquus, appear 
to constitute a distinct subgroup within the latifrons species group 
defined by distinctly keeled head scales, relatively short limbs, 
and a green ground color. 



1988 ANOLIS DANIELI 11 

A. propinquus, on re-examination, seems clearly to belong here. 
Its size as a hatchling (41 mm SVL) implies a giant adult, and 
the lamellae number implies the same and fits well with counts 
found in the latifrons group. It lacks an interparietal— and this 
initially seemed significant— but absence of an interparietal occurs 
also, as an individual variation, in A. danieli (AMNH 4844; the 
interparietal is indistinct also in MCZ 164894) and in the Parque 
Las Orquideas sibling. Its dewlap or gular region was described 
in the field as "blue." From Lago Calima, Valle, it is geograph- 
ically distant from other members of this subgroup. The radiating 
keels of the scales of the frontal depression and the nasal separated 
by one scale from the rostral suggest a closer relationship to A. 
apoUinahs than to A. danieli. This unique specimen and the Par- 
que Las Orquideas sibling indicate that there may be still further 
surprises within the latifrons group. 

A. frenatus, A. purpurescens, A. latifrons, A. princeps, and A. 
squamulatus form a second subgroup. These species are relatively 
long-legged and share with A. apollinaris and A. danieli the char- 
acter of green background coloration, but always have a dorsal 
pattern of oblique bands or rows of spots, sometimes also an 
ocellus in front of the shoulder. Despite considerable morpho- 
logical variation in some features (most impressively in the swol- 
len superciliaries of typical A. latifrons), this is a tight knit subgroup, 
in which, in fact, the separate species status of some nominal 
species— .4. purpurescens and A. princeps— is still unconfirmed. 
(For this reason the latter species— cited by Savage and Talbot, 
1978— were not mentioned in Williams 1976.) 

A.fraseri is distinctive in head squamation— smooth head scales, 
the superciliaries squarish and flat, the suboculars always in con- 
tact with the supralabials. Its color— dark olive brown and green— 
is unlike that of any other species. It is short-legged like A. danieli 
and A. apollinaris, but it has more characters in common with 
the two Central American latifrons group endemics, A. insignis 
and A. microtus (not only short legs, but smooth head scales and 
suboculars in contact with supralabials and background color- 
ation not green), and it is best grouped with these. 

A. insignis and A. microtus may be, as Savage and Talbot (1978) 
suggest, relatives, but they are amply distinct from one another 



12 BREVIORA No. 489 

and, perhaps, end points of a former Central American radiation. 
A. microtus is the one latifrons group species, thus far described, 
that consistently lacks an interparietal scale. 

ACKNOWLEDGMENTS 

I am indebted to M. A. Sema and H. Echeverri for providing 
the newer specimens of ^. danieli, to Dr. Charles Myers and to 
Dr. George Zug for the privilege of examining the specimens under 
their care, and to Dr. Pedro Ruiz and to Juan Renjifo for loan of 
comparative material oi A. apollinaris. Dr. Stephen Ayala sent 
me the female from Yarumal that was the stimulus for the present 
investigation and has given much assistance and many essential 
comments. He has also generously donated the map that is here 
published as Figure 7. Laszlo Meszoely drew Figures 1-6. 

LITERATURE CITED 

BouLENGER, G. A. 1885. Catalogue of the lizards in the British Museum (Natural 
History), 2: ix + 497 pp. London British Museum (Natural History). 

. 1919. Descriptions of two new lizards and a new frog from the Andes 

of Colombia. Proceedings of the Zoological Society of London, 1919: 79-81. 

Burt, C. E., and M. D. Burt. 1931. South American lizards in the collection 
of the American Museum of Natural History. Bulletin American Museum of 
Natural History, 61: 227-395. 

Dunn, E. R. 1937. The giant mainland anoles. Proceedings New England Zoo- 
logical Club, 16: 5-9. 

. 1944. Herpetology of the Bogota area. Revista, Academia Colombiana 

de Ciencias Exactas, Fisicas y Naturales, 6: 68-81. 

Etheridge, R. 1 960. The relationships of the anoles (Reptilia: Sauria: Iguanidae), 
an interpretation based on skeletal morphology. Doctoral Dissertation, Uni- 
versity of Michigan. University Microfilms International, Ann Arbor. 236 pp. 

Savage, J. M., and J. J. Talbot. 1978. The giant anoline lizards of Costa Rica 
and western Panama. Copeia, 1978: 480—492. 

Wiluams, E. E. 1 966. South American anoles: Anolis biporcatus and Anolisfraseri 
compared. Breviora Museum of Comparative Zoology, No. 239: 1-14. 

. 1970. South American anoles: Anolis apollinaris Boulenger, 1919, a 

relative of .4. biporcatus Wiegmann (Sauria, Iguanidae). Breviora Museum of 
Comparative Zoology, No. 358: 1-11. 

. 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. Papeis Avulsos de 



Zoologia, Sao Paulo, 29: 259-268. 



1988 ANOLIS DANIELI 13 

. 1983. Ecomorphs, faunas, island size and diverse end points in island 

radiations of Anolis, pp. 326-370, 481-483. In R. Huey ct al. (eds.). Lizard 
Ecology: Studies of a Model Organism. Cambridge, Harvard University Press. 

. 1 984. New or problematic Anolis from Colombia II. Anolis propinquus, 

another new species from the cloud forest of Western Colombia. Brevoria 
Museum of Comparative Zoology, No. 477: 1-7. 



14 



BREVIORA 



No. 489 



Table 1 . Latrifrons group anoles with short legs and green bodies. 





apollinaris 


danieli 


propinquus 


Head scales 


keeled 


keeled 


keeled 


Number be- 


8-12 


8-12 


12 


tween second 








canthals 








Scales in frontal 


with keels radiat- 


with keels oriented 


with keels ra- 


depression 


ing from the 


anteroposteriorly 


diating from 




center 




center 


Circumnasal/ 


circumnasal or an- 


anterior nasal in 


anterior nasal 


rostral 


terior nasal sep- 


contact with sul- 


separated 




arated from ros- 


cus between first 


from rostral 




tral by one scale 


supralabial and 
rostral 


by one scale 


Scales between 


2-4 


3^ 


3 


supraorbital 








semicircles 








Superciliaries 


one very elongate 


one very elongate 


one extremely 




scale followed 


scale followed by 


elongate scale 




by one or two 


two shorter and 


{'/2 supracili- 




shorter and 


these by smaller 


ary) margin 




these by 


conical scales 


followed by 




subgranular se- 




granules 


Ear 


rics 
small 


small 


small 


Loreal rows 


5-7 


5-7 


7 


Interparietal 


small 


small 


not differentia- 
ted 


Scales in pari- 


large and rugose 


convex, weakly 


small, subequal. 


etal depres- 


anterior to inter- 


keeled or rugose. 


weakly keeled 


sion 


parietal, smaller 


largest next su- 






behind it 


praorbitals, 
slightly smaller 
behind interpari- 
etal 




Scales between 


2-A 


2-5 


no interparietal 



interparietal 
and semicir- 
cles 
Scales between 
suboculars 
and supralabi- 
als 



0-1 



0-1 



1988 



ANOLIS DANIELI 



15 



Table 1. Continued. 





apollinaris 


danieli 


propinquus 


Supralabials to 


7-8 


7-9 


1 


below center 








of eye 








Trunk scales 


swollen, keeled 


swollen, keeled or 


granular, con- 




with inter- 


pyramidal, sur- 


vex, subequal 




spersed granules; 


rounded by gran- 






middorsals 


ules; 1-^ mid- 






slightly enlarged 


dorsal rows 






or 2 rows dis- 


enlarged 






tinctly so 






Ventrals 


smooth or keeled, 


smooth, subimbri- 


smooth, juxta- 




juxtaposed to 


cate to imbricate 


posed or sub- 




imbricate 




imbricate 


Femoral scales 


unicarinate, multi- 


unicarinate, multi- 






carinate near 


carinate at knee 






knee 






4th toe lamellae 


23-29 


23-27 


25 


Dewlap 


in male only 


large in male and 


hatchling; no fe- 






female; smaller 


male known 






in female 




Dewlap scales 


densely scaled. 


densely scaled. 


scales crowded 




scales bluntly 


keeled 


but a series of 




keeled 




raised rows 
each two 
scales wide 


Postanal scales 


present in males. 


sometimes obscure; absent in females 


Scales posterior 


smooth or keeled 


keeled 


keeled 


to vent 








Tail crest 


never present in any species 


Tail SVL 


ca. 3x 


ca. 3x 


ca. 2x 


Maximum 


S 112 


S 117 


unique type, a 


SVL 


9 94 


$ 104 


hatchling 



16 



BREVIORA 



No. 489 



•a 
o 

c 
u 
u 
u 
&c 

•o 

c 

00 

JJ 

c 
o 



<u 

"o 

c 

a 

3 
O 

OO 

to 
C 
O 






S 



s: 






•o 




r- 


HJ 












u 




1 


(U 




<N 


.:<: 




"" 


•o 






JU 


<u 










!S 


5 




c 

•c 


3 




^ 






r 


X) 




jn 


3 




o 
o 


*-• 


in 




1 



E "^ 



T3 


U-) 


_4J 








"3 


1 


(U 


^^ 


^ 


^^ 



to 




1 


o 


S 


o 




6 


Cj, 


(A 


iT 


•-W. 


a 


p^ 


Ci. 






<u 




c 







1 


to 


OJ 


5 

1- 


s 


^ 


2 


u 


o 




O 


T3 


i; 


jr 


<U 






</5 

o 

(U 



o 
o 



u 

c 



u 

O w aj 
 <u 



1-1 « 
c 3 j: 

■C " - 



<u o 

c >. 

ca t: 

oi o 

"3 'C _ 

■^ to 

^ Q 



u 



o 
o 

6 

(A 



U ^ <U 

3 (U 5 

3 ii " 

• aj 

OX) O 

3 



C3 



U 

k> 

aj 
X) 

S 

3 
Z 



T3 
C 
O 



C 



.2 


o 


t^M 






tn 


c 


c« 




U 




u. 




a 


la 


T3 


o 




C/3 





u 




T3 










o 

t« 

ca 

c 


ca 

c 

u 
O 


ca 

ca 
o. 


la 

u 

o 


O 


VI 

ca 




■c 


m 


6 


c 


C/5 




e 


-4— » 


(A 


o 


O 




3 


c 
ca 




o 




*-• 


VO 


3 












f^ 



a « 

5^ 2 



ca 
o _ 



2 o 
5 w 



■c 

^-^ ca 

c 

ca 



i: o 



•o c *; 



ca aj 
c« 



8 p 



O. (U ■>-' 



ca £ c ^ 

c •-" •^ 

O £ w ^ 

^ VI "^ "* 

c ^ 
ca 



ca 



ca VI 

D. O 

« 2 

= T3 X) 



o 



3 



VI 

4> — ■:;; 
*^ ca ™ 
ca 2 o 

^ -^ VI 



ca (u 



ca 
o 



3 
O 



c 


o 


ca 


6 


u 


o 


la 

VI 


VI 


o 


c 


♦-* 


ca 


th 


o 


ca 

c 
<u 
C 
o 


ca 

c 

£ 

3 

o 

1_- 


c 

O 

■c 


-a 
<u 

2 

ca 


X) 

2 



3 « 
p O 

o 



I 



I 
en 



<N 



I 



1 




>, 


»- 




o 






_ca 

3 
c 




&> 




^ 


2 


CO 


> 


w 


_o 


X2 


3 




ca 


7^ 


3 


o 


a.> 


so t<=. 


tA 


40 


c 











I 



c 
ca 



(« 3 ^ 

y X) « 



Of) 3 
C X 

— " ."2 



K C .„ o i; r- — 



♦-t r- C 
C t 4J 



C 
ca 
■o 

c 

3 
X3 t3 w K C 1a 



o 

U5 



at ,P 






J3 



G X 



«5 



>. 3 

C 



aj 00 



. to 



•c 



Co *" 

O « u. 

S ca XI c 

I-' (« ^  



o <u 



U 00 






aj 
c ^ 



aj 



aJ ■;:; 



— -^ o ^ ca 

2. 2. £. ■^ ^ 

t« tn g 

C £ 2 

aj ii o _ 00 
o 



c — 

aj ca c/5 t/i 



D 


■«-• 


aj 


(U 


^ 


'B 


o 


•c 


4-» 


u 


k« 


2 


o 


o 






X 


ca 


o 


."^ 


v> 

_0J 


u 

o. 


e 


4J 


3 


3 

(/5 


aj 

VI 


a 


o 






3 


C/D 






C/D 



1988 



ANOLIS DANIELI 



17 



in 



to 

o 



^ 

3 






<3 

to 



to 
s; 



a 









>, 




"3 


c 


2 

■c 








4J 


lU 

C 

♦-• 


-C 


(/5 


3 


re 








*-• 
C3 


o 




O 


e- 




<u 

e? 


7 


•o 
o 

e 


o 
6 




C3 


*-• 

c 


1 








b. 


T3 
















u 


C 
















J= 


3 
















2 


2 


"3 
















— 


•c 












'4— • 


(73 


1 














s 


"« 


<u 










o 


O 


<-» 




c 






r- 


J2 


7 


re 










4 








tT 


3 


00 














10 


u 


■c 














o 


rt 














00 


u 


00 














s 


c 


c 
'•3 














x' 


lU 


c 












, 


4J 


00 


3 
O 
u 










7 

00 


13 
6 


> 

c 
o 
o 


J2 






00 






"w 






1 

C 


G 














ca 


o 


_« 












^— < 




CO 


4-* 














O 


<u 

•c 


T3 




c3 




cu 


<a 


o 


■c 


C 










c 


o 
S 


o 




'^ 




T3 




T3 




c 


(U 




O 




O 
c 




(rt 


n 




x> 









■^ 






3 "3 £? 










■7 T3 iC « 






, -rt U c 






ollen 
keelei 
tapos 
midd 
not e 


rs 


fS 


ci 


1 


g 



ON 



0= 



Ov 



3 = u, 
u « <u 



X) 

3 



5 oj 

U O -rt > 



00 — 



1- x) 



■c 

re 
a 



re " 
1- h 



6 giS re 



S S 



« re re y 



_ (U ~ D. 

•3 p. « — ^ c 

S Fii 2 2 

I- ^r ii re u ■« 

re o c o 

W J ^ V5 



I 



c 

u 
u 

X) 

re 
o 



re .is 

« .a 

■C c 

c = 
.- re o 



I 

o 



c 
? re 



o S <u 

^ 3 1 u. 00 
73 — 1 13 o  







•u ^ E c 






u ^ . c w 






■^ 3 -p « >> 






, cj (U — — 






■ollen 
tuber 
tapos 
dorsa 
slight 


<N 





ci 


oi 


^ 





1 




»-4 

c 


3 

c 
re 
k-i 
00 




.:^ 





c 


■0 




r 


a 


<D 


u 




c 


re 


3 


M 


7 




X 

3 






^ 


Ui 









o 
c o 






c o - ^ 

§ *" i" 

- ^ E c« 

c 



_re 
re 

a 

3 



o 

7 

00 

o ^ 

J2 c 
re <" 

o 



•c 




, 




^ 


XI 


re 


c 







E 




(U 




u 


3 

T3 


3 

Ui 






c 


00 


"re 
•T3 


13 


re 




a 


re 


E 


(U 










^ 











X) 

re 



<" i^ ^ 13 "TS re ii 

"^ re irt re X) 9- -^ 

o 3 

00 c/5 



(/I 
"re 

~ E 



18 



BREVIORA 



No. 489 



T3 
lU 

3 

c 
o 
U 



(N 












2 £ 

3 O 



■B^ 



CJ 



o 
o 



o 'C 



 X) 

W 3 

^ :! 

3 t; 



T3 r; 



. C 

(U 'C 

(JJ ™ w 

C y (U 

Cfl 3 -^ 

o c •- 



<u 



"3 <u 

o '" 
O 



o 'C 
a X) 



■* S CJ <u 
O 



■^ O t'l 

x; 2 nj C 

 w r; X) 

<u 2 c 

St 3 C 



o 
o 

6 



X 
3 
•' — > 

11 



«5 



•c 

X "^ 

•^ 2 



c 

> 



o 

Xi 



i! 

c " 

3 



25 



•r c 

§ -c 

3 



I " 

(N 



t/5 

X 



<N 



1^ 

(N 

I 



<N 



a 
>< 

3 
O" 

'S 

3 



13 



J. 2 

o 



ta 



,. 


c 


T3 




■H 


■^-» 


la 


o 


o 


c 


« 


W) 


>, 


_u 


2 


la 


ca 




» 





<L) I 
a <^ 

ea 

a si 

CO 



T3 



^ 2 



>-.•-" (« 



oo 
I " 



13 

G 

ja 

O " 

•5; Q 



ca 

& 
ca 



ca 



T3 



o 



o 



(fl 



o 



_i, 13 .S > -o 

^ ^u u I- ■' 



^§^12 
S? a o 2 o 

=2 ^ 



o 
o 



O « 

U <U 

^ 1 

!« **- O 
c o 

':: e 



•« .i: 



X) 



<u 



^ 3 

^ en 
> O 

- ?i T3 
o ^ — 

c ^ 



la 

e 

c 



« o 
o 



u 



a g 



ca 13 ii 
o oj ca 



ca 

c 
ca 

O 



J2 

*-• 
O 
O 

E 



u 
O 

•c 

♦-• 
</) 
o 
a 

t/5 

OJ 

la 
o 



E 

E 

00 
o 



ca ~ " 
u »o o 



X 



X 






r4 






c 
ca 


E 
E 


E 
E 










X 
<N 

§ E 

■B E 

4j in 

o ::^ 

E ^ 



c 
> 

3 



X 

<N 

C 

ca 

u 
u 
O 

E 












X > 

ca on 



1988 



ANOLIS DANIEL! 



19 



Table 3. Latifrons group anoles with short legs and not green. 





fraseri 


insignis 


microtus 


Head scales 


smooth 


smooth 


smooth 


Number be- 


6-10 


7-12 


7-9 


tween sec- 








ond canthals 








Scales in fron- 


smooth 


smooth 


smooth 


tal depres- 








sion 








Circumnasal/ 


circumnasal or an- 


circumnasal or 


circumnasal sepa- 


rostral 


terior nasal sepa- 


anterior nasal 


rated from ros- 




rated from rostral 


separated from 


tral by one 




by one or two 


rostral by one 


scale 




scales 


or two scales 




Scales between 


2-4 


2-6 


2 


supraorbital 








semicircles 








Superciliaries 


no very elongate 


3 short scales 


one scale longer 




scale, the ante- 


longer than 


than wide fol- 




riormost scale 


wide followed 


lowed by 




short but longer 


by a series of 


smaller smooth 




than wide and 


smaller flat 


oi subgranular 




followed by a 


scales, irregu- 


scales 




double series of 


lar in size 






smooth series of 








smooth squarish 








scales 






Ear 


moderate 


moderate 


moderate 


Loreal rows 


5-9 


5-8 


3-5 


Interparietal 


moderate 


moderate 


not diflerentiated 


Scales in pari- 


flat, smooth, large 


flat, smooth all 


flat, smooth. 


etal depres- 


all around inter- 


around inter- 


moderately 


sion 


parietal 


parietal 


large all around 
interparietal 


Scales between 


2-5 


2-5 


no interparietal 


interparietal 








and semicir- 








cles 








Scales between 











suboculars 








and suprala- 








bials 








Supralabials to 


6-9 


7-12 


7-9 


below center 








of eye 









20 



BREVIORA 



No. 489 



Table 3. Continued. 





fraseri 


insignis 


microtus 


Trunk scales 


smooth with inter- 


smooth, juxta- 


smooth or slight- 




spersed granules. 


posed one to 3 


ly rugose; flank 




none or two mid- 


middorsal 


scales rhomboi- 




dorsal rows en- 


rows enlarged 


dal; flat, mid- 




larged 




dorsal scales 
elongate, rather 
irregular in 
shape 


Ventrals 


smooth or keeled. 


smooth, juxta- 


smooth, juxta- 




juxtaposed to im- 


posed to sub- 


posed or imbri- 




bricate 


imbricate 


cate 


Femoral scales 


unicarinate, multi- 


smooth 


wrinkled, not 




carinate at knee 




keeled 


4th toe lamel- 


18-24 


23-27 


20-22 


lae 








Dewlap 


large in both sexes 


large in both 


large in both 






sexes 


sexes 


Dewlap scales 


small, smooth 


densely scaled, 


very weakly and 






scales small. 


densely scaled 



Scales poste- 


keeled 


rior to vent 




Tail crest 




Tail SVL 


ca. 2x 


Maximum 


$ 116 


SVL 


9 102 



very weakly 
keeled 

present in males, sometimes obscure, absent in females 
smooth smooth 



never present m any species 

ca. 2 X ca. 2 X 

■5153 .3111 

9 135 9 104 



1988 



ANOLIS DANIELI 



21 




Figure 1. Anolis danieli, new species, ICN 5997 (holotype). Dorsal view of 
head. 



22 



BREVIORA 



No. 489 




Figure 2. Anolis danieli. new species, ICN 5997 (holotype). Lateral view of 
head. 




Figure 3. Anolis danieli, new species, ICN 5997 (holotype). Ventral view of 
head. 



1988 



ANOLIS DANIELI 



23 




Figure 4. Anolis danieli, new species, ICN 5997 (holotype). Lateral view of 
whole animal. 







Figure 5. Anolis danieli, new species, AMNH 38725. The most distinct pattern 
seen in the type series. 



24 



BREVIORA 



No. 489 




Figure 6. Anolis apollinaris, ICN 2865. The most distinct pattern seen in the 
specimens of this species examined. 



1988 



ANOLIS DANIEL! 



25 




Figure 7. Map of distribution of the Anolis latifrons species group in Colombia. 



B R E V I O R A 

MuseiLioi of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 2 September 1988 Number 490 



NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. VI. 

TWO FUSCOAURATOID ANOLES FROM THE PACIFIC 

LOWLANDS, A. MACULIVENTRIS BOULENGER, 1898 

AND A. MEDEMI, A NEW SPECIES FROM 

GORGONA ISLAND 

Stephen C. Ayala' and Ernest E. Williams^ 

Abstract. Anolis maculivenths Boulenger, 1898, the widespread Pacific \ovj- 
\and fuscoaurat us group anole of northwestern South America, is redescribed on 
the basis of a series of specimens from the region of the type locahty in northern 
Ecuador. A second fuscoauratoid species, Anolis medemi, new species, is described 
from Gorgona Island, 56 km west of the Pacific Coast in Colombia. 

INTRODUCTION 

Anolis fuscoauratus-Wkt lizards are widespread and often com- 
mon at many sites in the Andean cordilleras and along the Pacific 
lowlands of northwestern South America. Williams (1976) rec- 
ognized and briefly defined a fuscoauratoid complex as part of 
his key to the species groups of South American anoles. Never- 
theless, accurate identification of individual specimens has often 
proved difficult: scale counts are very similar, distinctive colors 
fade soon after death, and only short descriptions of the type 
specimens of each species have been available for reference. 

Williams (1976) listed five species in the fuscoauratoid group: 
A. antonii Boulenger, 1908, A. fuscoauratus Dorbigny, in Dumeril 
and Bibron, 1837, A. maculiventris Boulenger, 1898, A. ortoni 



' 929 Pepperwood Lane, Petaluma, California 94952. 

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

LIBRARY 

OCT 3 1 1988 

f 

UAO\/ADnk 



BREVIORA 



No. 490 




Figure 1. Anolis maculiventris collection sites, and location of museum spec- 
imens: a-f, in the region of the type locality (a), specimens used for this description; 
g-x, other Colombian sites providing specimens assigned to this species, a— 
Hacienda Paramba, Imbabura (BMNH; NHMB, NHMW, ZMB); b-Lita, Im- 
babura (KU, NHMW); c-Cachabi, Esmeraldas (USNM), and Rio Cachabi near 



1988 TWO FUSCOAURATOID ANGLES 3 

Cope, \S6S, and A. trachydcrma Cope, 1875. Two additional taxa, 
A. tolitnensis Werner, 1916 and A. mariarum Barbour, 1932, were 
treated as synonyms o^ A. antonii but are now known to be valid 
species. A. ortoni has subsequently been considered a member of 
the pentaphon rather than the fuscoauratus species group because 
of its large, flat head scales and the low dorsal crest on its tail. 

Fuscoauratoids are part of the beta section of the genus, with 
transverse processes on the tail vertebrae. Adults are small to 
medium size, reaching 50-60 mm snout-vent length. Their dorsal 
scales are small, uniform and usually keeled; belly scales are usu- 
ally smooth, or sometimes bear a low keel; head scales are small 
and most often wrinkled or keeled, with those of the frontal 
depression sometimes much smaller than the others; the tail is 
round and slender, with no dorsal crest; toe lamellae are broad 
and well defined, totaling fewer than 20 on phalanges ii and iii 
of the longest hind toe. 

An eventual definitive review of this group will require multi- 
variate analysis, personal familiarity with each species' distinctive 
color patterns and behavior under natural conditions, and elec- 
trophoretic studies as well. Meanwhile, detailed knowledge of the 
characteristics and variation of the populations at each type lo- 
cality (the topotypic populations) is essential for reliable identi- 
fications and comparison with lizards from other areas. We begin 



Rio Basalito (USNM); d-Estacion Forestal "La Chiquita," 14.4 km S. of Lan 
Lorenzo, Esmeraldas (MCZ); e— San Lorenzo, Esmeraldas (USNM); f— Tangareal, 
Narino (ICN, MCZ. BMNH); g-La Guyacana, Narino (FMNH); h-Guapi, Cau- 
ca (UVC); i— Quebrada Guangui, Rio Patia, upper Rio Saija drainage, Cauca 
(AMNH); j — Rio Raposo, Valle (UMMZ); k— Anchicaya, Valle, and various sites 
in the vicinity (ICN, IND-R); 1- Llano Bajo, Valle, and Sabaletas (ICN); m- 
Hacienda Los Mangos, Rio Dagua, Valle (BMNH); n— Buenaventura, Valle 
(BMNH, UMMZ); o— Granja Forestal Experimental, bajo Rio Calima, Valle 
(BMNH, ICN, CSJ, KU); p— Quebrada Docordo, between Cucurrupi and Noan- 
ama, Choco (AMNH); q-Peiia Lisa, Condoto, Choco (BMNH, MCZ, ICN); r- 
Andagoya, Choco (BMNH, USNM, ICN); s-Rio Dubasa, Choco (MCZ); t- 
Pueblo Rico, "Caldas," now Risaralda (BMNH); u— area between Rio San Juan 
and Rio Atrato (LACM); v-Quibdo, Choco (CSJ, ICN); w- Alto del Buey, Choco 
(LACM); X — Serrania del Baudo, Choco (LACM). A list of the museums men- 
tioned is in the acknowledgments. 



4 BREVIORA No. 490 

here a series providing more useful descriptions, comparisons of 
the named tax within the fuscoaurat us group, and descriptions of 
some new species found during our work. Redescriptions of named 
species are based mainly on specimens recently collected at or 
near the type localities. Montane fuscoauratoids are discussed in 
a following report. 

The map (Fig. 1) includes an additional 16 Colombian localities 
for^. maculiventris, including six localities (93 specimens) visited 
by Ayala's group, and 1 based on specimens in various museums. 
The term "large adult" refers to our estimate of modal large adult 
size, based on snout-vent length (SVL) measurements. It may 
exclude the occasional very large individual and does not take 
into account sexual size differences mentioned in the text. Most 
measurements used were made at the time of preservation (e.g., 
all specimens collected by one of us (SCA) or Ayala's collaborators 
in Colombia), but some were taken after different times in pre- 
servative. Ayala has seldom found more than a 2-mm shrinkage 
following preservation for anoles that measured 50-55 mm in 
life. 

Anolis maculiventris Boulenger, 1898 

Syntypes. BMNH 1946.8.13.33, an adult male (SVL 45 mm), 
BMNH 1946.8.13.34, an adult female (SVL 50 mm). Hacienda 
Paramba, Imbabura, Ecuador (0°49'N, 78°21'W), W. F. H. Ro- 
senberg and assistants coll. 

This is a small brown lizard, the only fuscoauratoid yet de- 
scribed from the western lowlands of Ecuador and central and 
southern Colombia (Fig. 1). Its type locality was cited as Hacienda 
Paramba, a farm in the Mira River valley between Ipiales and 
San Lorenzo in northern Ecuador. Rio Mira briefly forms part of 
the Colombia-Ecuador border not far below the farm. The two 
maculiventris syntypes were donated to the British Museum by 
W. F. H. Rosenberg, who together with various assistants col- 
lected extensively up and down the valley between 1896-1899. 
Rosenberg was a collector-dealer, who frequently had some of his 
specimens first determined and described by Boulenger, and then 
sold the remainder of his series to other museums. Rosenberg's 
maculiventris must have come from the lower elevation, wet for- 
ested regions downstream (0°50-53'N, 78°20-30'W), possibly 



1988 TWO FUSCOAURATOID ANGLES 5 

in the Province of Imbabura (south bank of the Rio Mira at 
Paramba), or Carchi (north bank of the river at Paramba), or 
perhaps even in Esmeraldas (20 km below the farmhouse). Up- 
stream, the valley becomes an arid, almost treeless desert (tropical 
and premontane thorn woodland, in the Holdridge classification), 
unsuitable for forest lizards like A. maculiventris. 

The following description is based on our study of the two 
British Museum syntypes, plus 1 3 additional maculiventris from 
nearby sites in Imbabura, Esmeraldas and Naririo (Figs. la-O- 
COLOMBIA. Narino. Tangareal, along the Rio Mira: MCZ 
159587-89, 160216. ECUADOR. Esmeraldas. Immediate vicin- 
ity of Cachabi: USNM 234723; Rio Cachabi near Rio Basalito: 
USNM 234724; San Lorenzo: USNM 234725; Estacion Forestal 
"La Chiquita," 14.4 km S. San Lorenzo: MCZ 160248. Imbabura. 
Lita, 520 m: KU 133437, NHMW 12809; Paramba: BMNH 
1946.8.13.33-34 (syntypes), NHMB 5060, NHMW 12810-11, 
ZMB 16462 (the latter four specimens are probably Rosenberg 
material seen by Boulenger). Scale counts for the male and female 
syntypes, respectively, are given in parentheses. 

Description. Head. Head scales moderate to small, smooth, 
tuberculate or weakly keeled. Ten to 14 (m: 1 1, f: 14) across snout 
between second canthals. Frontal depression distinct, the scales 
within it posteriorly minute and tubercular, anteriorly slightly 
larger, smooth and flat. Six to nine (m: 8, f: 9) postrostrals, in- 
cluding the two anterior nasals. Anterior nasal in contact with 
the sulcus between rostral and first supralabial, rarely divided 
horizontally. 

Supraorbital semicircles well defined, separated by three or four 
(m: 3, f: 4) small scales, separated by circumorbitals from the 
scales of the supraocular disk. Supraocular disk rather well de- 
fined, 5-10 enlarged scales, usually 1-3 much larger than the 
others, smooth or weakly keeled, grading laterally, anteriorly and 
posteriorly into granules. One elongate superciliary covers about 
one-third of the superciliary border, followed by one or two much 
shorter scales and then by granules. Canthal ridge distinct, can- 
thals about nine, first 4-5 larger, the anteriormost in contact with 
the first supralabial or separated by one scale. Six to nine (m: 8, 
f: 8) loreal rows, the one or two lowest largest. Temporals and 
supratemporals granular. No distinct intertemporal zone or line 



6 BREVIORA No. 490 

of enlarged scales. Supratemporals grading into slightly larger 
scales lateral and anterior to the interparietal. Interparietal with 
well defined parietal eye, smaller, much smaller than or subequal 
to ear. Surrounding scales smooth, flat, pavement-like, much 
smaller than interparietal. Four to nine (m: 7, f: 9) scales between 
interparietal and semicircles. Scales behind interparietal tiny 
rounded granules, not or hardly differentiated from nape granules. 

Subocular in contact with supralabials. Six to nine (m: 8, f: 7- 
8) supralabials to below the center of eye. 

Mental almost completely divided, in contact with seven to ten 
(m: 9, f 8) scales between the infralabials. No differentiated sub- 
labials. 

Trunk. Dorsal scales subgranular, smooth or weakly keeled, 
subequal or two median rows very slightly enlarged. Ventrals 
larger than dorsals, smooth, flat or slightly swollen, separated, 
juxtaposed or subimbricate. Chest scales in female smooth (poorly 
visible in male because of dewlap). 

Dewlap. Extending well beyond insertion of forelimbs in males, 
absent in females. Lateral scales in rows separated by naked skin. 
Edge scales smooth, no larger than lateral scales. 

Limbs. Limb scales unicarinate anteriorly, granular behind. Su- 
pradigitals multicarinate. Twelve to 16 (m: 15, f: 16) scales under 
phalanges ii and iii of fourth toe. 

Tail. Round or slightly compressed, never with a crest, about 
2 X snout-vent length. Enlarged postanals almost always absent, 
but visible in occasional males. 

Measurements. Five males in this series measure 43-46 mm 
SVL (m: 43, Boulenger gave 45); six females measure AA-A9 mm 
(f: 49, Boulenger gave 50). Based on our series of nearly 100 
Colombian maculiventris >40 mm SVL, most large adults mea- 
sure 45-48 mm, with exceptional specimens reaching 50 mm, 
and with no apparent sexual difference. Tail almost twice snout- 
vent length. 

Color. Information on the color in life is available for only one 
of these specimens, KU 133437, a male from Lita, Imbabura— 
S. R. Edwards: "By night, sleeping on a leaf Dorsum light brown. 
Dewlap orange peripherally, red medially. Venter tan. Tail barred, 
tan and grayish brown." Other specimens assigned to this species 



1988 TWO FUSCOAURATOID ANGLES 7 

from sites farther south include MCZ 160249, a male from Tin- 
alandia, Pichincha, 16 km from Santo Domingo de los Colorados 
on the road to Quito— K. Miyata: 

Collected at night. Color light brown with hint of olive. 
Dark brown markings. Can change quickly to dark mahogany 
brown with almost black look. Dark brown line between eyes 
over top of head. Venter pale dirty brown, yellowish around 
cloaca. Dewlap bicolored, dull orange around edge, dirty brick 
red along throat. Scales on dewlap yellow anteriorly, dirty 
white posteriorly. Iris brown. 

KU 133709, a female from 4 km N. of Quevedo, Los Rios— 
T. H. Fritts: 

At base of elephant ear plant at edge of stream by day. 
Dorsum olive brown with few black flecks; venter gray-beige 
invaded laterally by olive brown flecks of lateral body. 

And a composite description drawn from several dozen spec- 
imens from the region to the north around Buenaventura, Valle: 

A brown or gray-brown lizard, most often pattemless or 
sometimes with diagonal series of small yellow spots on the 
sides, or occasionally in males with vague dark shadows or 
bands between the pale rows of spots. When frightened it can 
turn much darker, especially on the back and head. Some 
females have a striking pale tan or yellow vertebral stripe 
with dark borders. A dark interocular bar may sometimes be 
present. Both sexes often have a small but distinctive black 
spot on the back of the head. In occasional specimens several 
other less well-defined spots extend as a series along the back. 
The belly is pale tan or gray-white, speckled along the sides 
with brown spots. The gular region is white, yellow-white or 
sometimes pale yellow-green; in males the underside of the 
tail is yellowish. The male dewlap is reddish pink behind the 
brick red anteriorly, with longitudinal rows of white scales, 
and the tail has wide light and dark brown bands. 

Distribution. Anolis maculiventris ranges along the wet Pacific 
lowlands between central Ecuador and central Colombia (Fig. 1). 



BREVIORA 



No. 490 



■r\ 




o 

A 

"3 



c 



c 
o 

S? 

o 

O 

S 

o 









3 



1988 TWO FUSCOAURATOID ANGLES 9 

Status of the members of the fuscoauratoid complex farther north 
remains unresolved, perhaps involving other undescribed taxa. 
Whether the Central American fuscoauratoid A. limifrons enters 
Colombia from the Panamanian Darien is uncertain. Dr. Charles 
Myers is examining populations from that region. 

A second western fuscoauratoid anole from Gorgona and Gor- 
gonilla Islands, 56 km off the Colombian Pacific Coast, is de- 
scribed here as a distinct species, closely related to A. maculiven- 
tris. The earliest specimen we know of was collected in 1938. The 
species was discussed and illustrated, but not named, in a prelim- 
inary treatment of Gorgona Island lizards (Ayala et al. 1979: 234- 
5, figs. 16, 17). 

Anolis medemi, new species 

Holotype. ICN 4371, adult male, Isla Gorgona (2°59'N, 
76°12'W), La Esperanza, Cauca, Colombia. Stephen C. Ayala 
coll., 22 May 1979. 

Paratypes (22). COLOMBIA. Cauca. Isla Gorgona: SDNHM 
31122: C. S. Perkins coll., 22 February 1938. MCZ 78944-48: 
Federico Medem coll., 1961. MCZ 168519: Humberto Carvajal 
coll., 4 April 1977. IND-R-0468: Inge E. Morales, C. Chaparro 
and Pedro Rodriguez coll., 21 May 1978. IND-R-2226: Hum- 
berto Carvajal coll., 21 May 1979. IND-R-2899: Juan Manuel 
Renjifo coll. MCZ 168520-22: Humberto and Fanny Carvajal 
coll., 26 May 1 979. CSJ 690: Henry von Prahl coll., 26 May 1979. 
ICN 4364-65, 4367-71, S. C. Ayala, H. Carvajal and F. Carvajal 
coll., 19-24 May 1979. Isla Gorgonilla: ICN 4366: Olga Castaiio 
coll., 22 May 1979. 

Diagnosis. A h^XB. Anolis of the fuscoaurat us species group, most 
closely related to A. maculiventris but slightly larger, with an 
overall orange-brown color (rather than olive or gray-brown) in- 
cluding a well-defined pattern of darker brown bars and spots on 
the back and sides. 

Description. (Information on holotype in parentheses.) Head. 
Head scales moderate to small, weakly keeled or smooth. Snout 
moderately short, 9-15 (15) scales across snout between second 
canthals. Frontal depression moderately deep and well defined, 
scales within minute, granular, much smaller than surrounding 



10 BREVIORA No. 490 

scales, 7-10(10) across. Five to eight (7) border rostral posteriorly. 
Anterior nasal in contact with rostral and rostral-first supralabial 
sulcus. Eight scales between supranasals dorsally. 

Supraorbital semicircles well defined, separated by two or three 
(3) small scales, separated from supraocular disk by one or two 
rows of circumorbitals. Supraocular disk moderately well defined, 
5-1 1 (8) enlarged scales, usually 1-3 larger than others, smooth 
or weakly keeled, grading laterally and posteriorly into granules, 
anterolaterally into moderately large scales, posterolateral area 
minute granules. One elongate superciliary extends over anterior 
half superciliary border, followed by 0-2 (1) shorter scales and 
then by granules. Canthus distinct posteriorly, less defined an- 
teriorly, 8-10, usually 9 (10) scales to below nostril, first and 
second largest, anteriormost contacting first supralabial scale. Six 
to nine (9) loreal rows, lowermost largest. Temporals and supra- 
temporals granular, intertemporal area not distinctly differen- 
tiated. Supratemporals grading into slightly larger scales around 
interparietal. Parietal eye clearly defined. Interparietal elliptical, 
smaller than or subequal to oval ear. Surrounding scales smooth, 
flat or swollen, much smaller than interparietal. Three to eight 
(7) scales between interparietal and semicircles, 3-5 between in- 
terparietal and smaller rounded granules on nape. 

Suboculars in contact with supralabials. Six to nine (8) supra- 
labials to center of eye. Mental almost completely divided, in 
contact with six to nine (8) scales between infralabials; these scales 
larger laterally, but no differentiated sublabial rows. 

Trunk. Dorsal scales small, granular, weakly keeled, subequal 
or two to four median rows very slightly enlarged. Ventrals 4- 
5 X larger than dorsals, smooth, rounded, separate, juxtaposed or 
slightly subimbricate. Chest scales in female unkeeled (poorly 
visible in male because of dewlap). 

Dewlap. Extending onto anterior one-third of belly in males, 
absent in females. Lateral scales in well-spaced longitudinal rows, 
larger than ventrals; edge scales smooth, subequal to or slightly 
smaller than lateral scales. 

Limbs. Limb scales unicarinate dorsally and anteriorly, gran- 
ular behind. Supradigitals multicarinate. Thirteen to sixteen (15) 
lamellae under phalanges ii and iii of fourth toe. 

Tail. Round or slightly compressed, slender, with no crest, 



1988 



TWO FUSCOAURATOID ANGLES 



11 







O 

■c 

X) 
T3 



u 

U 

•c 

u 

u 

XI 
■*-• 

O 

e 

3 

a 
u 
u 
u 

o 

u 

■♦-> 

(A 

o 

<A 
00 

8 



s 



I 



N 
CO 



OS 



3> 



12 BREVIORA No. 490 

about 2 X snout-vent length. No enlarged postanal scales (except 
in MCZ 78947). 

Measurements. Holotype. Head length 1 5 mm, head width 7.0, 
snout-vent length 46, tail 83, foreleg 21, hindleg 37, reaching 
between eye and ear. Large adults 49-51 (largest of 20, 52 mm) 
SVL, with a 90-95 mm tail. 

Color. Color in life brown with a definite orange cast. Most 
individuals show some evidence of darker brown bands or bars 
across body, legs and tail. Those that, like the holotype, are more 
prominently marked have 9-1 1 dark brown spots or crossbars 
along the back, 5-6 becoming vertical or diagonal dark brown 
bands on flanks, separated by pale yellow-brown zones, lines or 
spots; in others, darker brown pattern limited to vertebral region. 
Pale zones toward end of tail sometimes very light tan or even 
almost white, accentuating contrast with darker brown bands. 
Several specimens have dark lines radiating forward, upward and 
back from eye region, and a dark brown supraocular crossbar. 
Occasional females show sex-linked vertebral stripe morph seen 
in females of many other anoles; here stripe golden yellow with 
dark brown edges. Clearly defined occipital spot behind parietal 
eye; round, dark brown and almost always present even when 
remaining pattern scarcely visible. Belly pale tan or gray-white, 
sometimes with tiny brown flecks toward sides. Dewlap new-brick 
reddish orange, brighter anteriorly, with 5-6 longitudinal lines of 
yellow scales. 

Background color of five paratypes collected by Federico Me- 
dem considerably faded, leaving darker pattern curiously accen- 
tuated, almost white with bold brown bars (Fig. 3). Most other 
preserved specimens similarly patterned, but pattern blends 
slightly, or almost completely in some females, into characteristic 
orange-brown background color. 

Etymology. This lizard is named in memory of Dr. Federico 
Medem, who first brought this species to our attention, and who 
was director for many years of the Villavicencio Field Station of 
the Universidad Nacional de Colombia. 

Comparisons. Anolis medemi is easily distinguished from the 
other three anoles known to occur on Gorgona Island: A. princeps 
Boulenger (some authors have cited it as A. latifrons) is much 
larger and green with brown diagonal stripes on the sides; A. 
biporcatus (Wiegmann) is larger, more robust and usually uniform 



1988 



TWO FUSCOAURATOID ANGLES 



13 





B 

Figure 4. Hemipenes: A. Anolis medemi. B. Anolis maculiventris. 



green— the regional race A. biporcatus parvauritus Williams may 
prove to be a distinct species; and A. chloris gorgonae Barbour 
(variously cited as A. chloris or A. gorgonae) is a sky-blue insular 
race of the common green mainland species A. chloris Boulenger. 

Parker (1926) listed two additional anoles from Gorgona Island 
whose presence there remains unconfirmed. His two specimens 
unfortunately seem to have been misplaced while in one of the 
authors' (SCA) care: BMNH 1926.1.20.106, Hsted originally as 
A. '^fasciatus,''' is a juvenile^, chocorum Williams and Duellman, 
a rain forest species found sporadically between Costa Rica and 
western Colombia; BMNH 1926.1.20.107, Hsted as A. ''lemnis- 
catus,'' appears indistinguishable from specimens of A. tropido- 
gaster, a bush and grassland anole found in Panama and northern 
Colombia. Peters and Donoso-Barros' (1970: 48) undocumented 
mention of A. ""hi not at us'' on Gorgona might refer to A. medemi. 

Anolis medemi is closely related to A. maculiventris. Scale count 
ranges overlap almost completely. A. medemi has 2-3 scales be- 
tween the supraorbital semicircles, whereas maculiventris usually 
has 3^, with no (Ecuadorian) or only occasional (Colombian 
specimens) counts of 2. Other scale counts cannot be distinguished 
even modally. The color of the male dewlap is nearly identical. 
Both species almost always have a round dark spot on the back 



14 



BREVIORA 



No. 490 



Table 1. Color and pattern difTerences between Anolis medemi and Anolis 
maculiventris. 



Character 


Anolis medemi 


A nolis maculiventris 


Basic color 


rufus, red- or orange 


brown, olive-brown. 




brown 


brown-gray 


Pattern 


usually evident, often 


sometimes prominent. 




prominent; dark bars on 


usually none at all; dor- 




sides and back of head, 


sal surfaces seldom 




body, tail and legs 


barred, if so not promi- 
nent 


Vertebral region 


series of 9-11 more or less 


often dark; usually little 




prominent spots or bars 


or no trace of dark 




between head and tail 


spots or bars between 
head and tail; some- 
times dark spots at top 
of lateral bars on flanks 


Flank region 


usually 5-6 darker brown 


bars if present may be 




bars, with no tendency 


nearly black; usually al- 




to black 


most completely absent 


Back of head 


Dark bar over head be- 


little or no pattern; bar 




tween eyes present, usu- 


between eyes often not 




ally prominent; occipital 


prominent; occipital 




spot present 


spot usually present 



of the head (smaller in maculiventris); both may have 5-6 vertical 
or diagonal bars on the flanks, sometimes separated by series of 
round yellow spots (much less often seen in maculiventris); and 
both have occasional uniform brown-gray individuals, especially 
females, with no visible pattern (much more common in macu- 
liventris). 

The main differences between the two species involve color and 
pattern (Table 1), adult size and microhabitat. Living and pre- 
served A. medemi are basically orange-brown, while A. maculi- 
ventris is brown, gray-brown or olive-brown, rarely with any or- 
ange cast. The darker brown markings are usually much less visible 
in A. maculiventris. Large adult A. medemi measure 49-51 mm 
SVL and weigh (live weight) about 2.0-2.3 g; large adult A. ma- 
culiventris measure 45-48 mm and weigh 1 5-1 7 g. The hemipenes 
of the two species are illustrated in Figure 4. 

Although vegetation and forest structure on Gorgona Island 
appeared similar to that on the adjacent mainland, most of the 



1988 TWO FUSCOAURATOID ANGLES 15 

14 specimens collected by our group were found on thin to in- 
termediate diameter tree trunks or larger branches, about 1.5-3.0 
m above the ground; one was on the ground. In contrast, A. 
maculiventris is usually seen on twigs, vines or slender branches 
0.5-1.5 m above the ground, or on green or dried leaves in open 
areas near the ground. 

ACKNOWLEDGMENTS 

Helen Chin, Fernando Castro, Humberto and Fanny Carvajal, 
Carlos Galvis H. and Dennis M. Harris provided many of the 
Colombian A. maculiventris specimens. Humberto and Fanny 
Carvajal collected most of the ^1. medemi. Work in Columbia was 
sponsored partially by the Universidad del Valle and the Tulane 
University International Center for Medical Research, grants from 
the U.S. Public Health Service (NIAID A1-A2151 1) and the Co- 
lombian National Science Foundation COLCIENCIAS (10006- 
1-07-76), and a COLCIENCIAS grant to Universidad de Los 
Andes and Henry von Prahl for a multidisciplinary study trip to 
Gorgona Island in May 1979. Museums listed include: AMNH— 
American Museum of Natural History, New York; BMNH— 
British Museum (Natural History), London; CSJ— Colegio San 
Jose, Museo de Historia Natural, Medellin; FMNH — Field Mu- 
seum of Natural History, Chicago; ICN— Instituto de Ciencias 
Naturales, Museo de Historia Natural, Universidad Nacional de 
Colombia, Bogota; IND-R-INDERENA, Bogota; LACM-Los 
Angeles County Museum of Natural History, Los Angeles; MCZ— 
Museum of Comparative Zoology, Harvard University, Cam- 
bridge; NHMB— Naturhistorisches Museum, Basel; NHMW— 
Naturhistorisches Museum, Vienna; SDNHM— San Diego Nat- 
ural History Museum, San Diego; UMMZ— University of Mich- 
igan, Museum of Zoology, Ann Arbor; USNM— United States 
National Museum, Smithsonian Institution, Washington, DC; 
UVC— Universidad del Valle, Cali, Departamento de Biologia; 
ZMB— Zoologisches Museum, Universitaet Humbolt, Berlin. 



LITERATURE CITED 

Ay ALA, S. C, H. Carvajal, F. Caro DE Carvajal, AND F. Castro. 1979. Los 
Saurios de la Isla de Gorgona, pp. 219-241. In H. von Prahl et al. (eds.), 



16 BREVIORA No. 490 

Gorgona. Bogota, Universidad de Los Andes, Special Publ., Futura Grupo 

Editorial. 
BouLENGER, G. 1898. An account of the reptiles and batrachians collected by 

Mr. W. F. H. Rosenberg in western Ecuador. Roy. Zool. See. London 1898, 

(1): 107-126, pis. x-xviii. 
Parker, H. W. 1 926. The reptiles and batrachians of Gorgona Island, Colombia. 

Ann. Mag. Nat. Hist., ser. 9, 16: 549-554. 
Peters, J. A., and R. Donoso-Barros. 1970. Catalogue of the Neotropical 

Squamata. Pt. 11. Lizards and Amphisbaenians. U.S. Nat. Mus. Bull., 297: 

1-293. 
Williams, E. E. 1976. South American anoles: The species groups. Pap. Avuls. 

Zool., Sao Paulo, 29: 259-268. 



B R E V I O R A 



useum of Comparative Zoology 

us ISSN OOo£^98 , 



.-_L 



Cambridge, Mass. 25 March 1991 Number 491 

m 1 1991 

u An\/APn 

LARVAL DEVELOPMENT, RELATIONSHIPS, AND 

DISTRIBUTION OF MANDUCUS MADERENSIS, 

WITH COMMENTS ON THE TRANSFORMATION OF 

M. GREYAE (PISCES, STOMIIFORMES) 

David G. Smith,' Karsten E. Hartel,' 
AND James E. Craddock'^ 

Abstract. Larval development of Manducus maderemis is described for the 
first time and additional information is presented on the development of M. greyae. 
Relationships of Manducus and its close relative, Diplophos, are discussed based 
on larval pigmentation, transformation size, and the degree of development of 
annular mucosal intestinal folds. Distribution of M. maderensis is updated with 
extensive new material. 

INTRODUCTION 

Manducus (Goode and Bean, 1896) occupies a position at or 
near the base of the teleostean order Stomiiformes; thus it is of 
considerable interest. Its morphology and relationships have been 
discussed by Fink and Weitzman (1982) and Ahlstrom et al. 
(1984). Of the two species currently recognized, the larva of only 
Manducus greyae Johnson, 1970 has been described (Ozawa and 
Oda, 1986). In this paper we describe pretransformation larvae 
of the second species, Manducus maderensis (Johnson, 1 890), and 
provide new information on its distribution. In addition, we de- 
scribe the transformation of M. greyae. 



' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. Present address for DGS: Division of Fishes, National Museum of 
Natural History, Washington, DC 20560. 
- Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543. 



2 BREVIORA No. 493 

trial, although a few definitions may require modification. The 
division of the original group VI into groups VI and IX follows 
Rivero and Sema, 1988. A description in English (expanded from 
the original in Spanish) of the Andean groups occurring from 
Colombia to Peru is provided here. 

Many of the proportions usually incorporated into descriptions 
have been omitted here (diameter of tympanum in relation to eye 
diameter, diameter of eye in relation to the distance between eye 
and nostril, etc.) as they can easily be determined from the mea- 
surements. When many specimens were available, averages and 
proportions are provided in a Variation section. 

Most measurements were taken with a compass; snout-vent 
length, head breadth, and length of tibiae, with calipers. Head 
length was measured between the posterior edge of the tympanum 
and the tip of the snout. 

The web between the toes is considered insignificant if it does 
not extend beyond the midpoint of the first subarticular tubercle 
in at least four toes, minimal if it extends to the anterior border 
of the first subarticular tubercle in at least four toes (V4-webbed), 
intermediate if it extends beyond the first subarticular tubercle 
but does not reach the last articulation (disk) in at least three 
toes, and extensive if it reaches the last articulation in at least 
four toes. The central and usually indented part of the web is the 
portion considered for determining its extension. Considering the 
individual variation, a more detailed description is unnecessary 
and may make comparisons more difficult. However, the pedal 
membrane of all holotypes is illustrated in the corresponding 
figures. 

The author wants to thank E. E. Williams and J. Rosado for 
all their courtesies and attentions during his stay at the MCZ. 

DEFINITION OF GROUPS 

Group I. Two pectoral spots present, dorsolateral and ventro- 
lateral stripes absent; oblique-lateral stripes present and usually 
complete (from eye to groin), rarely absent; pedal membrane ab- 
sent or insignificant, rarely extensive; third finger of males not 
dilated; cloacal funnel absent. 

Group II. Dorsolateral stripes present; oblique-lateral stripes 
absent or incomplete (not reaching the eye); ventrolateral stripes 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 3 

present or absent; pedal membrane absent or insignificant; paired 
pectoral spots absent; third finger of male not dilated; cloacal 
funnel absent. 

Group IV. Third finger of males dilated; dorsolateral stripes 
usually absent; oblique-lateral stripe present or absent; ventro- 
lateral stripe generally absent; pedal membrane absent or insig- 
nificant (except in Colombian C agilis); paired pectoral spots 
absent; cloacal funnel absent. 

Group V. Cloacal funnel present; dorsolateral stripe absent or 
indistinct; oblique-lateral stripe absent; ventrolateral stripe ab- 
sent; pedal membrane extensive; paired pectoral spots absent; 
third finger of male not dilated. 

Group VI. Pedal membrane usually extensive, at least V3 the 
length of the toes; first finger generally shorter than second; dorsal 
color usually blackish, sometimes marbled or spotted; dorsolat- 
eral stripes absent or not extending posteriorly beyond sacral 
hump; oblique-lateral stripes absent or incomplete; ventrolateral 
stripes generally absent; paired pectoral spots absent; third finger 
of male not dilated; cloacal funnel absent. 

Group IX. Dorsolateral stripes absent; oblique-lateral stripes 
present and usually complete (from eye to groin); ventrolateral 
stripes generally absent; pedal membrane absent or insignificant; 
paired pectoral spots absent; third finger of male not dilated (this 
last character distinguishes this group from group IV); cloacal 
funnel absent. 

DESCRIPTION OF SPECIES 

Colostethus mittermeieri, sp. nov. 
Figs, la-d 

Holotype. MCZ-A 100217, an adult female from Venceremos, 
394-395 km, on Marginal de la Selva Road, 1,620 m, Departa- 
mento de San Martin, Peru. Collectors: R. A. Mittermeier and 
H. Macedo Ruiz, 26 Sept. 1978. 

Paratypes. MCZ-A 100218-57. Forty specimens with the same 
data as the type. 

Etymology. Mittermeieri, in honor of Russell A. Mittermeier, 
one of the collectors of the species, and recent recipient of the 
N.Y. Zoological Society's Conservation Medal. 



4 BREVIORA No. 491 

peppering of small melanophores on the dorsal surface of the 
head posterior to the eyes, and a deep-lying longitudinal streak 
of pigment on the lateral surface of the head both anterior and 
posterior to the eye. Additional pigment is found dorsal to the 
anteriormost part of the esophagus, on the ventral edge of the 
pectoral-fin base, as a line midventrally on the isthmus, and as a 
series of minute chromatophores along the ventral edge of the 
mandible. There is a row of chromatophores along the anal-fin 
base and a scattering of chromatophores on the base of the tail 
opposite the hypurals. No photophores are visible, and in alcohol 
the ground color of the body is completely white (probably semi- 
transparent in life). 

The next larger specimen (14.5 mm, MCZ 82190) is damaged. 
It is untransformed and has no photophores. Pigmentation ap- 
pears similar to that of the 10-mm specimen. 

The third pretransformation specimen is 1 5.0 mm (MCZ 82 1 89). 
Like the 10-mm specimen, it shows no trace of photophores or 
scales and is white. The dorsal series of chromatophores is present 
from the occiput to the caudal peduncle; the posterior spots in 
this series are larger than the anterior ones. A row of chromato- 
phores is present internally along the anal-fin base. The remaining 
pigment is similar to that of the 10-mm specimen. 

Transformation occurs somewhere between 15 and 18 mm. 
Three specimens, 18.0-18.5 mm, are well into the process. An 
18.5 mm specimen (MCZ 82192) is still largely white and has no 
scales. The ventral photophores (the IC and OA series) have 
formed, but there is no sign of the other lateral photophores. On 
the head, the BR, SO, ORB, and OP photophores are present 
(Table 1). Larval chromatophores are still present, including the 
dorsal series. The adult pigmentation appears to be developing 
dorsolaterally around the larval chromatophores. The annular 
mucosal folds of the intestine are still apparent posteriorly. Two 
specimens, 1 8.0 and 1 8.5 mm (MCZ 82 1 94 [Fig. 1 B], and 82 1 93), 
resemble the preceding one, but the adult pigment is more ex- 
tensive. Larval chromatophores are still present but are partially 
obscured by the developing adult pigment. Ventral photophores 
are present, but there is no sign of lateral photophores. A 17.5- 
mm specimen (MCZ 82174), although slightly smaller than the 
preceding three, is further along in development. It is uniform 



1991 MANDUCUS. DEVELOPMENT, RELATIONSHIPS 5 

light brown in color and the larval chromatophores are no longer 
visible. Scale pockets are not visible. 

Our smallest specimen with lateral photophores is 18 mm (MCZ 
82158), and it has only the beginnings of the midlateral series 
(LLP). The 17-mm specimen figured in Grey (1964; fig. 23) ap- 
pears slightly more developed. The LLP series lengthens as the 
fish grows, but the small accessory photophores do not appear 
until about 30 mm (MCZ 82 1 78). At 24 mm (MCZ 82 1 77) scales 
are clearly visible. By about 48 mm, M. maderensis is fully adult 
in body form, pigment, and photophore development. 

Manducus greyae 

Ozawa and Oda (1986:80) described larvae of M. greyae based 
on 15 specimens of 7.5 to 21.4 mm. Their largest specimen was 
just beginning to transform; hence, they were unable to describe 
that process fully. The MCZ larval-fish collection contains spec- 
imens of M. greyae from 22 mm to about 46 mm, a range that 
encompasses the entire process of transformation. We are thus 
able to provide an account of transformation in this species. 

The earliest stage in transformation is represented by a speci- 
men of 22 mm (MCZ 82462). It is somewhat damaged, but the 
following photophores appear to be present: SO, ORB, OP, BR, 
IP, PV, VAV, and AC (Table 2). The OA series is not visible and 
there are no lateral photophores. The annular mucosal folds of 
the intestine are clearly visible. The dorsal series of chromato- 
phores is absent, and the only ventral chromatophores visible are 
located dorsal to the anal-fin base and as a longitudinal midventral 
streak on the isthmus. Pigment is present just posterior to the tip 
of the flexed notochord. Some lateral pigment is developing on 
the myosepta and near the midlateral line, but the fish as a whole 
is still white. 

In six additional specimens ranging from 22 to 24 mm (MCZ 
82463 [Fig. 2], 82465, and 82466) the photophore complement 
is somewhat more complete than in the previous specimen (Table 
2). In particular, the IP series increases from two to ten, and the 
OA series increases from zero to 52. Three of these specimens 
(23-24 mm) have begun to develop the midlateral photophores 
(LLP) and three (22-23.5 mm) have not. One of the 24-mm 
specimens (MCZ 82466) has begun to develop adult pigmenta- 



BREVIOR.4 



No. 491 



to 

s: 
-^ 



a 

c 
.a 



t 
ca 



a 



c 



<u 



c 



c 

3 

o 
o 

-o 

u 

■*-» 
u 
_u 

13 
en 



_aj 



N 






N 



rj 

00 



81 

•^ 00 



rsj '^ 

"^ 00 



o 

OS 



00 



ON 



N 

•^ oo 



o 
a 



OO — (N r~- 



— (Nmosr'ioo — Os>nO 
— — -^ (N "* 



C8 



00 f^l O i/^ 



rt 
u 



00 — — ^ 



0<Nr<^Ost^— 'COvO — 
(N — <N Tf 



— • 04 m OS r^j r- r^ t^ 









1/-) O O VO 






22 I I 



I I I I I 



I I I 



I I I 



I I I I I 



O r- 



O ^ 



CO 

u 



I I 



I I I I 



G -p 










e § 










^ 6 










£ ^-' 










00 -S 
















in 




i r 






o 




0] 03 




S 


a 




T3 C 


"ca 


15 


o 

o 


O 


2 u 


o 


c 


t/3 


yj a. 


Q < 


cu 





ca 
O 



Da 



> 



> 

< 
> 



< 

O 



1991 



MANDUCUS. DEVELOPMENT, RELATIONSHIPS 



5u 

I 

bo 

3 
a 



t 

c 



in 

C 
03 



C 






c 



c 

3 
O 

T3 

■«-» 
O 



s 



N 
U 




S 


00 


N 

U 




S 


00 


N 
U 




S 


00 


N 




S 


00 



ISl ^ 

■^ 00 



N ^ 

^ 00 



N '^ 

r 1 "^ 

5: (N 

■^ 00 



c 

s 

a 





q 




<N 








— T3 T3 -^ 



O 



+ I 



i/^ m ^ 

rn r'i (N 
(N — 



ro CO fN 






m m fN 






(N 
Tf (-0 (N 



'-I m rn ~- O-^ 



— TT ^ fN 
fN — CO ■^ 



— —  <N —  (^ in 



m m <N o^ 



o 



O — 
Tf in 



— c (^ (^ r<^ t^ 






^ 00 , 

m Tf + 
d 



(N r<-i <N 



— (^ m — I r^ 



r<^ m O 
— ro m 



C9 



(N (N (N 

rs — 



00 •^ w 

C 00 o. 

T3 — « 

(33 w aj 

C ?5 S 

c« JJ u 

c^D ci: a 



-H T3 m — ' 



r4 00 O ■* O 



<u 

u> 
O 

x; 
a 
o 

o 

Qlh 



73 
<u 

O 

a 

aj 






I I 



+ I 



I I 



I I 



o 
a 
>< 1 



T3 
(U 

O 

a 

X 

<u 



T3 

<u 

O 

a 

X 



73 

[A 

o 
a 

X 



o 
a 

X 

aj 



(U 

o 
a 

X 



a: 
o 






> 

< 
> 



0- 



c 
aj 



00 
'5. 



aj 3 
t; T3 
£ < 



c 

VJ 

a> 



o 

+ 



c 

C3 



IS 

X 
o 



T3 

u 

Vh 

3 

c« 
X 
O 

aj 

*-• 
o 

X 
T3 

aj 
00 
03 

E 
w 

T3 
C 



aj 

a 



o 

X 

E 

o 
>^ 



BREVIORA No. 491 



Figure 2. Transforming larva of Manducus greyae, 24 mm SL (MCZ 82463). 
Drawn by L. Meszoly. 



tion, the esophagus and stomach are black, and it is the only one 
of these specimens in which the intestine has withdrawn inside 
the contours of the body. The second 24-mm specimen (MCZ 
82463) is still white. The smallest specimen that shows the ac- 
cessory lateral photophores is 30 mm (MCZ 82464). By 46 mm 
the species is largely adult in body form, pigmentation, and pho- 
tophore development. 

Ozawa and Oda (1986:82) claimed that in M. greyae the mid- 
lateral photophores develop before the ventral photophores. This 
is not the case in the specimens examined by us. The ventral 
photophores, especially the IC series, are well developed and con- 
spicuous before the LLP series begins to appear. In this, M. greyae 
resembles all the other species in Manducus and Diplophos that 
we have examined. 

Manducus-Diplophos Relationships 

Opinions and evidence about the status o^ Diplophos Giinther, 
1873 and Manducus Goode and Bean, 1896 have differed among 
authors and over the years. Goode and Bean (1896:514) estab- 
Ushed Manducus to contain Gonostoma maderense Johnson, 1 890. 
While they recognized that this species was not a true Gonostoma, 
they did not comment on any resemblance to Diplophos taenia 
Giinther. Grey (1960:76) reduced Manducus to a subgenus of 
Diplophos because the distinction between the two nominal genera 
(i.e., differences in certain proportional measurements and num- 
bers of fin rays, photophores, and vertebrae) seemed trivial com- 
pared to differences among other gonostomatid genera. Johnson 
(1970:442) went further and concluded that even subgeneric rank 
was unwarranted. His new species, Diplophos greyae, seemed to 



1991 M/liVDt/Ct/S'. DEVELOPMENT, RELATIONSHIPS 9 

him to be intermediate between the type species of the two nom- 
inal genera. Mukhacheva (1978) reviewed Diplophos on a world- 
wide basis, recognizing four species {taenia, rebainsi Krefft and 
Parin, 1972, greyae, and maderensis) and agreed with Johnson 
( 1970) that subgenera are unnecessary. Fink and Weitzman (1982) 
described the osteology of taenia and maderensis and treated them 
both as Diplophos. Ahlstrom et al. (1984), however, resurrected 
Manducus based on the condition of the pectoral-fin radials. Man- 
ducus has the cartilages of the third and fourth proximal radials 
separate, and the ninth distal radial is in line with the others; 
Diplophos has the cartilages (though not the bones) of the third 
and fourth proximal radials fused, and the ninth distal radial is 
out of line with the others (see figs. 18 and 19 in Fink and Weitz- 
man, 1982). These authors recognized two species in each genus: 
Diplophos taenia, D. rebainsi, Manducus maderensis, and M. 
greyae. Ozawa and Oda (1986) recognized seven species, all in 
Diplophos. Ozawa et al. (1990) revised the D. taenia complex, 
which includes four species: D. taenia, D. proximus Parr, D. or- 
ientalis Matsubara, and D. australis Ozawa et al. 

The species of Diplophos and Manducus represent the most 
primitive of the stomiiform fishes (Fink and Weitzman, 1982; 
Ahlstrom et al., 1984; Fink, 1984). Diplophos shares one derived 
feature, the partially fused third and fourth proximal radials, with 
the Photichthyidae (an ill-defined group) and Stomiidae, but 
Manducus retains the primitive state of this character. Ahlstrom 
et al. (1984:198) were unable to find any derived characters that 
would unite Diplophos and Manducus as a stomiiform subgroup. 
Early-life-history characters could not be used because the larvae 
of M. maderensis and M. greyae were unknown at the time. 

Larvae of D. taenia are distinguished by their elongate body 
and the conspicuous series of dorsal and ventral chromatophores 
(Ahlstrom et al, 1984: fig. 98; Ozawa and Oda, 1986: fig. 2). The 
intestine has pronounced annular mucosal folds. The dorsal fin 
is located slightly anterior to the anal fin. Larvae of Z). orientalis 
closely resemble those of Z). taenia but are somewhat less elongate 
and transform at a smaller size (ca. 30 mm vs. 50 mm; Ozawa 
and Oda, 1986:77, fig. 3). Larvae of M greyae resemble those of 
D. taenia and D. orientalis but are shorter-bodied and have less 
conspicuous dorsal and ventral chromatophores (Ozawa and Oda, 



10 BREVIORA No. 491 

1986:81, fig. 5); in larger larvae, the dorsal series disappears. The 
intestine has pronounced annular mucosal folds as in D. taenia, 
D. orientalis, and M. maderensis. Manducus greyae transforms at 
about 21-24 mm. Larvae of M maderensis resemble those of M. 
greyae in their short body and inconspicuous dorsal and ventral 
chromatophores (Figs. lA-B). The dorsal chromatophores are 
better developed than those oi^ greyae and remain throughout the 
larval stage, but the ventral chromatophores are present only over 
the anal-fin base. Manducus maderensis transforms at a smaller 
size (15-18 mm) than any of the other species. 

The larvae of the species of Diplophos and Manducus, then, 
differ primarily in their relative body depth, the extent of dorsal 
and ventral pigmentation, and the size at metamorphosis. Even 
these characters show a certain amount of gradation among the 
species. Diplophos orientalis is intermediate between D. taenia 
and M. greyae in both body depth and size at metamorphosis. 
Manducus maderensis has a relatively deep body, like M. greyae, 
but its dorsal pigmentation is stronger and more persistent through 
growth. The most striking similarity between the larvae of all 
these species is the pronounced annular mucosal folds of the 
intestine. 

In assessing the significance of these larval characters, we face 
the same problem as in assessing the significance of adult char- 
acters: to find shared specializations that link Manducus and Di- 
plophos to each other or to other stomiiform genera. Ahlstrom et 
al. (1984) were able to find only one adult synapomorphy that 
links Diplophos to photichthyids and stomiids (the condition of 
the pectoral-fin radials) and no synapomorphies that link Man- 
ducus to any other stomiiform genus. The larval characters de- 
scribed in the present paper do not alter this situation. The re- 
semblances between larvae of the species of Diplophos and 
Manducus are considerable, but none of these characters is clearly 
specialized below the ordinal level. Similarities in body shape and 
pigmentation could simply represent the primitive stomiiform 
condition. Even the most conspicuous character shared by larvae 
of Diplophos and Manducus, the annular mucosal folds of the 
intestine, is a matter of degree of expression rather than presence 
or absence. Other stomiiformes have mucosal folds although they 
are not as pronounced. Indeed, a similar intestine is found in a 



1991 



MANDUCUS. DEVELOPMENT, RELATIONSHIPS 



11 




Figure 3. Distribution of Manducus maderensis based on all known material; 
O = specimens <18.5 mm,  = >I9 mm, and • = >95 mm. Symbols may 
represent more than one specimen or collection. 



variety of lower teleosts, including clupeids, engraulidids, and 
certain myctophids. Whether these mucosal folds are identical 
structurally and developmentally in all the taxa that possess them 
is unknown. A phylogenetic analysis of larval characters in sto- 
miiform fishes is clearly beyond the scope of the present paper, 
and without such analysis the significance of these characters 
cannot be determined. Clearly much work remains to be done 
before a useful phylogeny of stomiiformes can be constructed. In 
the meantime, we offer the present study as one more piece of a 
puzzle that may one day be assembled into a coherent picture. 

DISTRIBUTION 

At the time of Grey's (1960 and 1964) revisions of the "Gono- 
stomatidae" M. maderensis was known from fewer than 50 spec- 
imens. Of these, 31 were adults, all from near land— at Madeira 
in the eastern Atlantic, and off Suriname and Mississippi in the 
western Atlantic. The 16 juveniles (<90 mm) were from the cen- 
tral North Atlantic (1 1 specimens), near the Bahamas (4), and the 



12 BREVIORA No. 491 

South Atlantic off Brazil (a single specimen from 0°22'S). Based 
on these records and five additional specimens (two of them from 
between 1° and 2°S off Brazil), Mukhacheva (1978) published a 
map of the distribution of M. maderensis and considered the 
species to be distant neritic, being "endemic to the western and 
eastern parts of the central Atlantic . . . but . . . absent in the open 
waters." She was correct in that it is probably land-associated, 
especially when adult, but, according to more recently collected 
data, it also occurs in the open ocean both when young and as an 
adult (Fig. 3). Manducus maderensis is endemic to the Atlantic 
Ocean, occurring primarily in the tropics and the equatorward 
halves of the subtropical gyres (tropical-semisubtropical pattern 
of Backus et ai, 1977); it is now known from 37°39'N to 23°02'S. 

There are only 13 specimens known from the South Atlantic, 
none of them from the poleward half of the subtropical gyre. Its 
rarity there is probably a reflection of the low fishing effort. In 
the North Atlantic, however, there are many specimens from the 
poleward half of the subtropics; most of these are adults from 
near Madeira (Maul, 1 948; Grey, 1 964; ISH, lOS). In the northern 
Sargasso Sea, there are but five specimens— three Gulf Stream 
waifs (MCZ 82193, 82194, and 88254) and two specimens re- 
ported by Bond (1974) from Ocean Acre off Bermuda (USNM 
248766 and MCZ 9 1 350, ex URI). 

Since few specimens of M maderensis have been collected with 
opening/closing nets, we can say little with precision about its 
vertical distribution, especially at the deeper limit of its depth 
range. We can say, however, that it occurs in the upper meso- 
pelagic zone (about 450-600 m) at the edge of continental (and 
island) slopes and in the open ocean. The species makes a diel 
vertical migration into the upper 100 m at night at sizes between 
20 and 100 mm. The shallowest records of large adults are the 
177 mm individual at 200 m (ISH 748/66) and the 209 mm 
gravid female (MCZ 91350) from off Bermuda at 150 m. Of the 
193 known specimens, 55, between 18 and 64 mm, were collected 
with neuston nets at the very sea surface. 

LIST OF MATERIAL 

The following M. maderensis (141 specimens, 10 to 220 mm) 
have been collected since the papers of Grey (1964) and Mu- 



1991 M/iyVDl/Ct/S', DEVELOPMENT, RELATIONSHIPS 13 

khacheva (1978), or were not reported by them. Each entry con- 
tains the museum catalog number, the number and size(s) of 
specimens, and collection data (station number, position, maxi- 
mum depth reached by net, and the time of the beginning of the 
collection). Specimens not examined by us are marked with an 
asterisk. The collections at ANSP, GCRL, GMBL, MZUSP, 
ZMUC, TCWC, USF, SAM, and VIMS have no M. maderensis. 
Also included is the material of M. greyae used for the transfor- 
mation description. 



Manducus maderensis 

lOS Discovery 

7089#03* (2:31-42) 17°41'N, 25°23'W, surface, 0145 hrs. 
7089#12* (3:19-24) 17°34'N, 25°26'W, surface, 0245 hrs. 
7089#13* (2:18-21) 17°48'N, 25°29'W, 515-600 m, 0950 hrs. 
7089#21* (9:21-26) 17°52'N, 25°27'W, surface, 2145 hrs. 
7089#26* (5:22-41) 17°52'N, 25°25'W, surface, 2100 hrs. 
7089#27* (2:@ 22) 17°52'N, 25°25'W, 25-60 m, 0138 hrs. 
7089#32* (6:20-25) 17°45'N, 25°22'W, surface, 0100 hrs. 
7089#37* (1:28) 17°50'N, 25°29'W, surface, 0145 hrs.; uncat.* 
(1:220) off Madeira. 

ISH 

64/66* (1:163) WH 177/66, 33°45'N, 16°00'W, 600 m, 2110 
hrs.; 296/66* (3:145-167) WH 181/66, 19°11'N, 21°58'W, 460 
m, 2100 hrs.; 399/66* (8:52-82) WH 183/66, 6°30'N, 24°33'W, 
50 m, 2100 hrs.; 620/66* (1:163) WH 187/66, 5°34'S, 26°58'W, 
320 m, 2000 hrs.; 748/66* (1:173) WH 1 9 1/66, 2 TOO'S, 30°00'W, 
200 m, 2000 hrs.; 3 1 3/68* (1:177) WH 8-III/68, 26°1 0'N, 1 9°26' W, 
580 m, 2233 hrs.; 1125/68* (1:56) WH 20-III/68, 13°56'S, 
27°38'W, 580 m, 2255 hrs.; 1665/71*(1:157)WH 443/71, 2r35'S, 
2°00'W, 2,100 m, 2025 hrs.; 2742/71* (1:98) WH 498-1/71, 
17°22'N, 22°58'W, 105 m, 1955 hrs.; 2819/71* (1:125) WH 498- 
III/71, 17°27'N, 22°55'W, 610 m, 2203 hrs. 

MCZ 

52541 (5:48-52) SUN1207, 9°16'N, 27°55'W, surface, 0015 
hrs.; 52566 (1:44) RHB1290, 2ri7'N, 85°22'W, 124 m, 0020 



14 BREVIORA No. 491 

hrs.; 54303 (1:110) Oregon 2007, 7°34'N, 54°49'W, 445 m; 56952 
(1:145) RHB3052, 11°22'N, 65°01'W, 350 m, 1700 hrs.; 61476 
(1:96) RHB2290, 2°57'S, 8°05'E, 75 m, 2005 hrs.; 82170 (1:70) 
RHB2269, 18°33'S, 4°00'W, 100 m, 2005 hrs.; 82171 (1:49) 
RHB1207, 9°16'N, 27°55'W, 51 m, 0010 hrs.; 82172 (1:61) RHB 
1266, 12°44'N, 74°10'W, 575 m, 1255 hrs.; 82173 (1:130) Oregon 
4419, 1 1°43'N, 69°13'W, 455 m; 82174 (1:17.5) RHB966, 1°13'S 
34°35'W, 102 m, 0335 hrs.; 82175 (1:48) RHB1253, 16°38'N 
64°27'W, 133 m, 0038 hrs.; 82176 (1:29) RHB1286, 19°46'N 
83°07'W, 86 m, 0010 hrs.; 82177 (1:24) RHB2035, 22°25'N 
19°00'W, 500 m, 0845 hrs.; 82178 (1:30) RHB2069, 15°23'N 
24°28'W, 320 m, 0420 hrs.; 82179 (1:22) RHB2077, 15°30'N 
26°12'W, 95 m, 2135 hrs.; 82180(2:33 &41)RHB2084, 17°12'N 
27°59'W, 80 m, 0215 hrs.; 82181 (1:20) RHB2095, 25°52'N 
36°48'W, 140 m, 2110 hrs.; 82182 (1:19) RHB2930, 1 1°00'N 
41°3rW, 475 m, 0055 hrs.; 82183 (1:61) RHB2946, 9°03'N 
5r05'W, 510 m, 0220 hrs.; 82184 (1:73) RHB2979, 13°34'N 
50°50'W, 490 m, 0210 hrs.; 82185 (1:20) SUN2078, 15°43'N 
26°28' W, surface, 1 20 hrs.; 82 1 86 (2:22 & 52) SUN2083, 1 7°08'N 
27°55'W, surface, 000 1 hrs.; 82 187 (2:25 & 27) SUN2 10 1 , 26°37'N 
4ri8'W, surface, 0005 hrs.; 82188 (1:49) SUN1313, 23°55'N 
83°12'W, surface, 0034 hrs.; 82189 (1:15) RHB2924, 10°59'N 
40°22'W, 490 m, 2330 hrs.; 82190 (1:14.5) RHB2923, 1 1°00'N 
40°10'W, 500 m, 2045 hrs.; 82191 (1:10) RHB2966, 12°2rN 
59°34'W, 495 m, 0035 hrs.; 82192 (1:18.5) MOClO-137,4 
30°08'N, 79°30'W, 140-160 m, 0254 hrs.; 82193 (1:18.5) 
SUN9452, 37°36'N, 69°03'W, surface, 0115 hrs.; 82194 (1:18) 
same data as 82193; 82 197 (2:79 & 84) RHB982, 6°5 1 'S, 33°34'W, 
85 m, 2105 hrs.; 82198 (1:40) SUN2958, 9°13'N, 59°06'W, sur- 
face, 0115 hrs.; 82199 (1:57) RHB1222, 13°55'N, 57°00'W, 300 
m, 2300 hrs.; 82200 (1:64) same data as 82198; 88250 (1:18) 
SUN1431, 23°02'S, 32°15'W, surface, 0120 hrs.; 88251 (1:29) 
SUN3102, 22°57'N, 64°12'W, surface, 2020 hrs.; 88252 (1:19) 
SUN2966, 12°2rN, 59°34'W, surface, 0035 hrs.; 88253 (1:62) 
JEC7741, 8°33'N, 44°37'W, 100 m, 0155 hrs.; 88254 (1:48) 
KEH7716, 37°00'N, 65°38'W, surface, 0325 hrs.; 88255 (1:29) 
SUN 1253, 16°38'N, 64°27'W, surface, 0030 hrs.; 88256 (1:27) 
JEC7745, 9°15'N, 46°50'W, 100 m, 0300 hrs.; 88257 (1:24) 
JEC7712, 0°OrN, 37°40'W, 80 m, 2235 hrs.; 88258 (1:18) 



1991 M^A^Z)C/Ci7S, DEVELOPMENT, RELATIONSHIPS 15 

JEC7705, 3°08'N, 42°52'W, 25 m, 1 30 hrs.; 91350(1 :220) Ocean 
Acre 12-55N, 32°1 1'N, 64°10'W, 150 m, 2240 hrs. 

UMML 

14824 (1:32) Gerda 205, 23°20'N, 82°55'W, 1,000 m, 1843 
hrs.; 22740 (1:156) Pillsbury 455, 13°0rN, 71°55'W, 1,445 m; 
23074 (1:29) P-383, 10°19'N, 75°59'W, 70 m, 0101 hrs.; 27541 
(2:53 & 58) P-384, 10°24'N, 75°58'W, 40 m, 0302 hrs.; 27747 
(6:17-45) P-302, 2°26'N, 4°5rE, surface, 0230 hrs.; 29036 (1:22) 
P-821, 19°07'N, 65°28'W, 3,000 m, 1145 hrs. 

CAS 

61060 (1:124) Oregon II 46092, 18°27'N, 67°15'W, 1,499 m, 
0852 hrs. 

USNM 

186282 (5:85-128) Oregon 2007, 7°34'N, 54°49'W, 445 m; 
186364 (14:90-140) Oregon 2008, 7°38'N, 54°43'W, 490 m; 
248711 (1:27) Ocean Acre 1-18C, 32°10'N, 63°48'W, 100m,0145 
hrs. 

Manducus greyae 

MCZ 

75518 (1:43) GRH1046, 12°38'S, 148°55'E, 3,240 mwo, 1740 
hrs.; 82462 (1:22) GRHlOl 1, 6°25'S, 152°09'E, 2,380 mwo, 0000 
hrs.; 82463 (4:22-24) GRH1014, 4°55'S, 152°30'E, 2,380 mwo, 
0015 hrs.; 82464 (2:29 & 30) GRH1017, 6°54'S, 152°06'E, 2,380 
mwo, 2245 hrs.; 82465 (2:24 & 47) GRH1016, 6°43'S, 152°14'E, 
2,380 mwo, 1840 hrs.; 82466 (1:24) GRH1069, 7°44'S, 15r05'E, 
ca. 1,950 m, 0001 hrs. 

Comparative material 

Larvae and transforming specimens of the following taxa in the 
MCZ larval fish collection were examined (number of specimens 
examined is given in parentheses): Bonapartia pedaliota (215), 
Cyclothone spp. (1,092), Diplophos taenia (36), Gonostoma at- 
lanticum (1,157), G. denudatum (282), G. elongatum (547), Ich- 
thyococcus (97), Margrethia obtusirostra (54), Maurolicine cf. "al- 



16 BREVIORA No. 491 

pha" (4), Maurolicus muelleri (269), Photichthys argenteus (1), 
Pollichthys mauli (19), Valenciennellus tripunctulatus (553), Vin- 
ciguerria attenuata (l ,059), V. nimbana(\,S01), V. poweriae (S25), 
Yarella blackfordi (1). 

ACKNOWLEDGMENTS 

We thank A. Post and G. Krefft (ISH), C. R. Robins and K. 
Lindeman (UMML), J. Badcock and N. Merrett (unfortunately, 
no longer of lOS), S. Jewett (USNM), W. Krueger (URI), W. 
Eschmeyer (CAS), and M. E. Rogers (FMNH) for providing us 
with records and/or specimens. W. Krueger deposited a valuable 
Manducus specimen at MCZ. In addition, J. Nielsen (ZMUC), 
N. Menezes (MZUSP), B. Stender and W. Anderson (GMBL), W. 
Saul (ANSP), S. Poss (GCRL), J. Musick (VIMS), J. McEachran 
and F. Hendricks (TCWC), T. Hopkins (USE), and M. Bougaardt 
(SAM) searched their collections for records of M. maderensis. 
G. R. Harbison kindly provided a large collection of mesopelagic 
fishes from the western South Pacific which included the speci- 
mens of M. greyae. Laszlo Meszoly prepared the figures. W. Fink, 
G. Moser, T. Ozawa, and S. Weitzman kindly reviewed drafts 
and made numerous suggestions. Support for the curation of the 
MCZ larval specimens was supplied by the National Science 
Foundation (BSR 86 1 7845). Contribution number 7404 from the 
Woods Hole Oceanographic Institution. Publication costs were 
covered in part by a grant from the Wetmore Colles Fund. 

LITERATURE CITED 

Ahlstrom, E. H., W. J. Richards, AND S. H. Weitzman. 1984. Families Gon- 
ostomatidae, Stemoptychidae, and associated Stomiiform groups: Develop- 
ment and relationships, pp. 184-198. //; H. G. Moser, W. J. Richards, D. 
M. Cohen, M. P. Fahay, A. W. Kendall, Jr., and S. L. Richardson (eds.). 
Ontogeny and Systematics of Fishes. Special Publication of the American 
Society of Ichthyologists and Herpetologists, No. 1, ix + 760 pp. 

Backus, R. H., J. E. Cradix)CK, R. L. Haedrich, and B. H. Robison. 1977. 
Atlantic Mesopelagic zoogeography, pp. 266-287. In R. H. Gibbs, Jr. (ed.), 
Fishes of the Western North Atlantic. Sears Foundation for Marine Research 
Memoir 1, Part 7, xv + 299 pp. 

Bond, G. W. 1974. Vertical distribution and life histories of the gonostomatid 
fishes (Pisces: Gonostomatidae) off Bermuda. Report of the U.S. Navy Un- 



1991 M4Af£)(7C(/5, DEVELOPMENT, RELATIONSHIPS 17 

derwater Systems Center. Contract N00140-73-C-6304, Smithsonian Insti- 
tution, 276 pp. 

Fink, W. L. 1984. Stomiiforms: Relationships, pp. 181-184. //; H. G. Moser, 
W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., and S. L. 
Richardson (eds.). Ontogeny and Systematics of Fishes. Special Publication 
of the American Society of Ichthyologists and Herpetologists, No. 1, ix + 
760 pp. 

Fink, W. L., and S. H. Weitzman. 1 982. Relationships of the Stomiiform fishes 
(Teleostei), with a description of Diplophos. Bulletin of the Museum of Com- 
parative Zoology, 150(2): 31-93. 

GooDE, G. B., AND T. H. Bean. 1896. Oceanic Ichthyology. U.S. National 
Museum, Special Bulletin Number 2:1-553 + 1-26, 123 pis., 417 figs. 

Grey, M. 1960. A preliminary review of the family Gonostomatidae, with a 
key to the genera and the description of a new species from the tropical Pacific. 
Bulletin of the Museum of Comparative Zoology, 122(2): 57-125. 

1 964. Family Gonostomatidae, pp. 78-240. In H. B. Bigelow (ed.). Fishes 

of the Western North Atlantic. Sears Foundation for Marine Research Memoir 
1, Part 4, xix + 599 pp. 

HuBBS, C. F., AND K. F. Lagler. 1964. Fishes of the Great Lakes Region. Ann 
Arbor: University of Michigan Press, 213 pp. 

Johnson, R. K. 1970. A new species of Diplophos (Salmoniformes: Gonosto- 
matidae) from the Western Pacific. Copeia, 1970(3): 437-443. 

Leviton, a. E., R. H. Gibes, Jr., E. Heal, and C. E. Dawson. 1985. Standards 
in herpetology and ichthyology: Part 1. Standard symbolic codes for insti- 
tutional resource collections in herpetology and ichthyology. Copeia, 1985 
(3): 802-832. 

Maul, G. E. 1948. Monografia dos peixes do Museu Municipal do Funchal. 
Ordem Isospondyli. Boletim do Museo Municipal do Funchal, 3(5): 5-41. 

MuKHACHEVA, V. A. 1978. A review of the species of the genus Diplophos 
Giinther (Gonostomatidae, Osteichthyes) and their vertical and geographical 
distribution. Trudy Institute of Oceanology, 111: 10-27 (English translation). 

OzAWA, T., AND K. Oda. 1986. Early ontogeny and distribution of three species 
of the gonostomatid genus Diplophos in the Western North Pacific, pp. 74- 
84. In T. Ozawa (ed.). Studies on the Oceanic Ichthyoplankton in the Western 
North Pacific. Fukuoka-Shi: Kyushu University Press, 430 pp. 

Ozawa, T., K. Oda, and T. Ida. 1990. Systematics and distribution of the 
Diplophos taenia species complex (Gonostomatidae), with a description of a 
new species. Japanese Journal of Ichthyology, 37(2): 98-1 15. 

Weitzman, S. H. 1986. Order Stomiiformes, Introduction, pp. 227-229. In M. 
M. Smith and P. C. Heemstra (eds.). Smiths' Sea Fishes. Berlin: Springer- 
Verlag, xx + 1047 pp. 



MCZ 



B R E,XJ O R A 

useiim ofj_^(gf^jyi|v||t|[jative Zoology 

I I K I 1 V / r- r-) :-? I -T' \/ 

US ISSN 0006-9698 



Cambridge, Mass. 25 March 1991 Number 492 



A PERUVIAN PHENACOSAUR 
(SQUAMATA: IGUANIA) 

Ernest E. Williams' and Russell A. Mittermeier^ 

Abstract. A small lizard from Venceremos, Department of San Martin. Peru, 
is identified as a hatchling Phenacosaurus and possibly the third known specimen 
of Phenacosaurus orcesi Lazell, 1969. It is the first known specimen of the genus 
from Peru. 

INTRODUCTION 

The anoline lizard genus Phenacosaurus was initially known 
only from Colombia. Its type species, P. heterodermus, was de- 
scribed by A. Dumeril, 1851, in Dumeril and Dumeril (1851), 
from numerous specimens from "Nouvelle Grenade," the name 
of Colombia at that time (including Panama). Dunn (1944) added 
two more Colombian species, P. nicefori ("vicinity of Pamplona, 
Norte de Santander") and P. richteri ("Tabio, Cundinamarca"), 
and Hellmich (1949) still another, P. paramoensis ("Paramo de 
Sumapaz" at the border between Cundinamarca and Meta). The 
latter two have since been synonymized with P. heterodermus 
(Lazell, 1969). A new giant Colombian species has very recently 
been described {P. inderenae Rueda and Hemandez-Camacho, 
1988, from Gutierrez, Department of Cundinamarca). 

Specimens or species known or suspected to be from adjacent 



' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 
- President Conservation International, 1015 18th Street NW, Washington, D. C. 

20036. 



2 BREVIORA No. 492 

countries have, however, been reported. A specimen from the 
Sierra de Perija (Museo de Historia Natural La Salle, Caracas 
4477), regarded by both Aleman (1953) and Lazell (1969) as P. 
nicefori, is from a peak (Cerro Tetari) in Zulia, Venezuela. (It is 
probably an undescribed species.) The Field Museum's P. nicefori 
(FMNH 5684) from "Paramo de Tana," cited by Lazell (1969), 
may, as Rueda and Hemandez-Camacho (1988) have comment- 
ed, be from Venezuela and not Colombia, since the locality given 
is precisely at the border between the two countries. The latter 
problem is rendered moot by more recent collections, since P. 
nicefori is now known from unpublished material from Betania, 
State of Tachira, further inside Venezuela (specimens in the col- 
lections of the Museo de Ciencias Naturales, Caracas, and the 
Museum of Natural History, Kansas), and a small series of an 
undescribed phenacosaur has been collected by the expeditions 
to the Cerro de La Neblina, State of Amazonas, in the extreme 
south of Venezuela. (These are under study by Charles Myers.) 
Still another phenacosaur, a single specimen in the collection of 
the Museo de Ciencias Naturales La Salle, Caracas, has been 
collected by S. Gorzula and A. Farrera on the Massif de Chimanta, 
a tepuy in the State of Bolivar, in southeastern Venezuela (to be 
described by Williams, Prasiderio, and Gorzula). 

From Ecuador, Lazell, in his 1969 revision, has described the 
very distinctive species P. orcesi on the basis of two specimens, 
the type from "Mt. Sumaco," Napo Province, and a paratype 
from "between L'Alegria [sic] and La Bonita," both localities in 
the Sucumbios Province (formerly the northwest part of the Napo 
Province). Only recently, specimens of another giant species of 
phenacosaur have been collected at La Alegria and adjacent lo- 
calities (specimens in the Museo Ecuatoriano de Ciencias Natu- 
rales, the Escuela Politecnica Nacional, the National Museum of 
Natural History, and in the Museum of Comparative Zoology, 
to be reported on by FEW and collaborators). 

Now a juvenile phenacosaur (MCZ 16521 1) has been collected 
in Peru at Venceremos, in the northern part of the Department 
of San Martin, very near the Department of Amazonas border. 
This juvenile, both because it is small and immature and because 
it is not ideally preserved, is conservatively regarded as the third 
known specimen of Phenacosaurus orcesi despite the great dis- 



1991 A PERUVIAN PHENACOSAUR 




Figure 1. Phenacosaurusorcesijuv., MCZ 16521 1. Lateral view of head. (Right 
side reversed.) 



tance between it and the nearest other specimen of that species, 
the type from Mt. Sumaco. The new specimen is important enough 
to deserve detailed description, provided below in a format elab- 
orated from the one that has been used by the senior author in 
descriptions oi Anolis over many years (see also Figs. 1^). 

DESCRIPTION 

Head 

No trace of a casque, not even the ridges that bound the parietal 
region in adults of the smaller species. All scales smooth. 

Dorsal Head Scales. Antorbital region: Rostral much wider than 
long. Four postrostrals, these defined as all those scales posteriorly 
in contact with the rostral and therefore including the left cir- 
cumnasal that has a narrow contact on that side. The right cir- 
cumnasal is excluded from the rostral by a postrostral. Circum- 
nasals round or ovoid, the nostril nearly central. No differentiated 
anterior or inferior nasals. Each circumnasal broadly in contact 
with the first supralabial of its side. Three scales between the 
circumnasals dorsally. Scales posterior to the circumnasals much 
smaller than the anteriormost canthals, the scales of the frontal 
area, or the median series of three scales anterior to the frontal 
area. 



BREVIORA 



No. 492 




Figure 2. P. orcesi'}u\., MCZ 16521 1. Dorsal view of head. 




Figure 3. P. orcesi ']\iw., MCZ 16521 1. Ventral view of head. 



Frontal depression very shallow. Scales of the frontal area mod- 
erately large, polygonal, markedly varying in size. No rosette of 
larger scales surrounding smaller central scales. Four to six scales 
between the anterior canthals depending upon where the count is 
made. 



1991 A PERUVIAN PHENACOSAUR 




Figure 4. P. orcesi}u\., MCZ 165211. Lateral view of body scales. Arrow 
points anteriorly. 



Canthals five on each side, gently arched, not keeled, the first 
(= the posteriormost) largest on both sides, those on the right side 
grading smaller anteriorly, on the left side the third and fifth larger. 

Orbital region: Scales of the supraorbital semicircles large, two 
scales on the left side in broad contact with three on the right. 
The two largest supraocular scales in contact with the semicircles 
on each side. The next largest supraocular on the right side sep- 
arated from the semicircle of its side by granules; the comparable 
scale on the left in contact with a lateral supraocular scale. The 
other scales of the supraocular areas variable in size, smaller 
laterally. Two to four rows separate the largest supraocular scales 
from the superciliaries. On each side the two anteriormost su- 
perciliaries larger and elongate, the remainder subgranular. 

Parietal region: A parietal eye indicated by a light spot. The 
interparietal apparently fused with other scales: as indicated by 
an anterior median partial sulcus, by the marked asymmetry of 



6 BREVIOR.4 No. 492 

this, the largest scale in the parietal region, and by the slight 
depression containing the parietal eye, which does not at all cor- 
respond to the scale boundaries. The scales lateral to the inter- 
parietal strikingly larger than those posterior to it, although these 
again are sharply distinct from the nape scales. About five rows 
in the approximate midline between the interparietal and the nape 
scales. A penultimate row of the posterior parietal scales markedly 
enlarged. 

Lateral Head Scales. Loreal rows three on the right side, two 
on the left. Total loreals on the right side 1 1, on the left 10. Two 
preoculars (defined as the scales below the anterior comer of the 
eye) on each side, the uppermost in contact with the sulcus be- 
tween the first and second canthals. Four suboculars on the right 
side, three on the left. Postoculars ill-defined, grading into the 
lower temporals. 

Temporals in two areas, upper and lower, separated by the 
double row of slightly enlarged scales on the low ridge that in- 
dicates the lower border of the skeletal supratemporal fenestra. 

Lower temporals smallest centrally; the upper temporals more 
nearly subequal but slightly larger anteriorly. 

Ear many times the size of any adjacent scale, but much smaller 
than the (probably compound) interparietal. 

Supralabials more or less elongate rectangles, seven to nine 
below the center of the eye. 

Ventral Head Scales. Mental very deep, as deep as wide, almost 
wholly divided by a median sulcus, slightly indented by two very 
small medial gulars between the very large first sublabials. Three 
sublabials on each side in contact with the infralabials. Six to 
eight infralabials to below the center of the eye. 

The anterior gulars (those posterior to the medial gulars that 
are in contact with the mental) small, elongate, slightly swollen, 
larger than the central gulars posterior to them, but not as wide. 
The latter becoming more granular and more imbricate near the 
median insertion of the dewlap but larger and still juxtaposed 
next to the sublabial series of each side. Lateral gulars intervening 
between the sublabials and the infralabials at the level of the third 
sublabials, after which it becomes impossible to distinguish be- 
tween lateral gulars, sublabials, and the lateralmost central gulars. 
All gulars subgranular posteriorly alongside the dewlap. 



1991 A PERUVIAN PHENACOSAUR 



Trunk 



No trace of a middorsal crest. Dorsal and flank scales subequal, 
smooth or subimbricate, or (flank scales) sometimes with tiny 
granules visible between them. Ventrals larger, smooth, slightly 
convex, very weakly imbricate, in transverse rows. 

Limbs 

Scales smooth, anteriorly larger and imbricate on lower arm 
and lower leg, separated by naked skin on upper arm and thigh, 
posteriorly granular on upper arm and thigh but not so on lower 
arm and leg. Supradigitals smooth or very weakly carinate, wid- 
ened transversely, lamella-like. Lamellae under phalanges ii and 
iii of fourth toe ca. 21. 

Tail 

Curving at tip as though prehensile, weakly compressed, all 
scales weakly keeled, without a dorsal crest, but the middorsal 
row imbricate and weakly dentate. Enlarged postanals (male) small 
but distinctly larger than surrounding scales. 

Dewlap 

Strongly indicated (juvenile male), distinguished by the longi- 
tudinal orientation of its scales and extending onto belly beyond 
the insertion of the arms. Edge scales smaller than ventrals, lateral 
scales larger than edge scales but, perhaps, smaller than ventrals. 

Size 
Snout-vent length 32 mm; tail length 43 mm. 

Color in life 
(from kodachromes by Russell Mittermeier) 

Ground color cream mottled with brown. Dark brown streaks 
radiating from eye onto supralabials and toward ear. Dorsum 
with three broad dark brown bands variegated with lighter brown. 
Interstices of bands more or less vermiculate with darker brown. 
Limbs banded brown and cream. Small dewlap pinkish or or- 
angish with sparse black spotting. 



8 BREVIORA No. 492 

Locality 

Collected 14 December 1983 by Russell Mittermeier near Ven- 
ceremos ("few houses along the road"): "km 390-39 1 on the road 
between Rioja (6°05'S, 77°09'W) and Pedro Ruiz Gallo (= In- 
genio, ca. 5°56'S, 77°59'W), approximately 91-92 km from Rioja; 
downhill into the forest about one km from the road, on the forest 
floor, within ca. 100 m of a small rainforest stream; elevation 
4,750 ft in steep montane terrain, forest floor well covered with 
moss and humus." The locality is cloud forest with moss-covered 
trees and a springy, mossy floor. Much of the World Wildlife 
Fund film Monkey of the Clouds was shot in the area. The juvenile 
phenacosaur was the only herpetological specimen taken at Ven- 
ceremos in 1983. In 1978, frogs were collected: a number of frogs 
not yet identified, a species of Eleutherodactylus, two undescribed 
species of Colostethus, and toads of the Bufo granulosus and B. 
typhonius complexes. 

Discussion 

Because this specimen is a juvenile it lacks the casque that is 
one of the defining characters of adult phenacosaurs, but a casque 
\sabsQn\\n2i\\]\x\Qm\QPhenacosaurus{MCZ 14165:/*. heteroder- 
mus, 30 mm SVL; EPN 2218: P. sp. 38 mm SVL). It also lacks 
the diflerentiated large round flat flank scales separated by gran- 
ules, characteristic of P. hetewdermus, the type species of the 
genus, and present also in P. nicefori and P. inderenae (but these 
are quite absent in the third described species, P. orcesi). In this 
juvenile there is no trace of the median crest present in most 
phenacosaurs (but absence as an individual variation has been 
demonstrated [in adults] for P. hetewdermus [Lazell, 1969] and 
occurs also in the adult paratype of P. orcesi). The Peruvian 
juvenile has two loreal rows on each side and only nine loreal 
scales on one side, 10 on the other, the unmodified circumnasal 
scale broadly in contact with the first supralabial, and a short tail, 
very little longer than snout-vent length, with a curvature sug- 
gestive of prehensility. All are characters congruent with deter- 
mination as a member of the genus Phenacosaurus. It is clearly 
closest to the two known adult specimens of P. orcesi, which it 
matches in the absence of enlarged flat round flank scales. Figures 
5-7 display the head of the paratype of that species, which has 



1991 



A PERUVIAN PHENACOSAUR 




> 

T3 



c5 









u 



a 






in 

3 



10 



BREVIORA 



No. 492 




T3 
cd 

u 

o 

Vi 

l-> 
O 



CO 

ro 

IT) 

so 



a 
>. 

*-* 

Oh 
5J 



3 



1991 



A PERUVIAN PHENACOSAUR 



11 




Figure 7. P. orcesi Paratype, USNM 166533. Ventral view of head. 



12 BREVIORA No. 492 

Table 1 . Comparative scale variations in the Peruvian phenacosaur and orcesi 
and heterodermus. 





Peruvian 








phenaco- 




57 hetero- 




saur 


orcesi 


dermus 


Scales between second canthals 


4 


4 


3-6 


Postrostrals 


4 


2-3 


3-6 


Scales between supraorbital semicircles 








0-1 


Scales in supraocular disk 


4 


5-6 


1-6 


Elongate supraciliaries 


2 


1 


0-2 


Loreal rows 


2-3 


2 


1-3 


Total loreals 


10-11 


6-7 


3-12 


Scales between interparietal and semicircles 








0-2 


Scales between interparietal and nape scales 


5 


4 


3-8 


Postmentals (including sublabials) 


4 


5-6 


2-6 


Scale rows in vertebral crest 





0-1 


0-2 


Lamellae under phalanges ii and iii 








fourth toe 


21 


17-19 


18-24 



not been previously illustrated. (The head and body characters 
of the type off. orcesi are figured in Lazell, 1969.) 

It should be parenthetically mentioned that the issue of the 
validity or non-validity of the genus Phenacosaurus does not arise 
in the present context. That this Peruvian specimen belongs in 
the lineage that includes the species heterodermus is not, for us, 
in question. The validity of the genus Phenacosaurus depends 
upon osteological characters not observable in the present spec- 
imen and upon hypotheses of the phylogenetic significance of 
those characters. The placement of the Peruvian animal within 
the postulated lineage orcesi, nicefori, heterodermus depends upon 
external phenetic resemblances, unknown or very unusual in Ano- 
lis, that, in our judgment, demonstrate that these species are a 
clade. 

There are seven differences between the juvenile and the two 
adult P. orcesi, none of these such that they could not be ascribed 
to the sort of individual variation that is rampant in the one well- 
collected species, P. heterodermus. These are in bold face in Table 
1, which records counts for the Venceremos juvenile, the two 
Ecuadorian orcesi, and for 57 heterodermus. It is clear that vari- 
ation in these counts exceeds species boundaries. However, cer- 



1991 



A PERUVIAN PHENACOSAUR 



13 



COLOMBIA 




I 1 r 

100 200 



Figure 8. Map showing the Ecuadorian type locality of P. orcesi (Mt. Sumaco) 
and the locality for the Peruvian juvenile (Venceremos). 



tain counts generally tend to be associated with body size in 
anoles; thus the count of 2 1 for the fourth toe lamellae for the 
Peruvian juvenile may, possibly, indicate that the adults of this 
population might be nearer heterodermus size (maximum SVL 
ca. 80 mm) than orcesi size (known maximum SVL 67 mm). 

There are, therefore, only two reasons for hesitation for rec- 
ognizing the Peruvian juvenile as P. orcesi: (I) the fact that it is 
a juvenile, and (2) the very considerable range extension (more 
than 500 km; Fig. 8) from the southernmost (type) locality Mt. 



14 



BREVIOR.4 



No. 492 




U1 




O 




S 




(U 




k> 




(U 




(J 




c 




OJ 




> 




^ 




o 




A 




v> 




O 




>-• 




a 




a 




E 




1 








c« 




*-» 




u 




Q 




♦-» 




J3 




00 








a^ 




<D 


3 


o 


<1> 


sz 


cu 


^ 






tn 


03 


« 


«i 


c 


W 


o 


5 


N 


F 


Ql. 


< 


o 




^^ 


c 


c 


<u 


n 


o 




m 






rt 


Tl 


(U 


« 


k. 


T5 


c 


c 


■^ 


w 




c 


b 
W 


C 




ta 


O 


"P 




c 


u 


C/5 


o 

c 

> 


e 

x: 


<u 


C 


XI 


o 


■*-» 


c 


? 


c 


o 




Xi 


en 


(« 


(U 






o 


■*-» 






a 


W 


m 


n 


^ 


u. 




QJ 


tS 


J3 


<[) 


■^- 


hJ 


O 




o 



as c. 
" 2 

00 — 

Pu <- 



1991 



A PERUVIAN PHENACOSAUR 



15 





Figure 10. Photograph of the Peruvian phenacosaur in life. Photo by R. A. 
Mittermeier. 



Sumaco, Napo Province, Ecuador (0°34'S, 77°09'W), to Vencere- 
mos, Department of San Martin, Peru (ca. 5°45'S, 77°45'W) (see 
Fig. 9). While P. orcesi is quite distinct from P. heterodermus and 
its relatives, P. nicefori, P. inderenae, and the undescribed giant 
species from Ecuador, the latter are a complex in which the species 
are not very sharply delimited morphologically; it is a possibility 
that P. orcesi is a complex also, and that the Peruvian juvenile is 
a distinct species. Provisionally we assign the Peruvian specimen 
(Fig. 1 0) to the species P. orcesi, but new material and much more 
careful collecting in the montane areas of Peru and Ecuador are 
clearly much to be desired. 

ACKNOWLEDGMENTS 

The figures of the phenacosaurs were done by Laszlo Meszoly, 
the maps by Stephen D. Nash and Laszlo Meszoly. 



16 BREVIORA No. 492 

LITERATURE CITED 

Aleman G., C. 1953. Contribucion al estudio de los reptiles y batracios de la 

Sierra de Perija. Memorias de la Sociedad de Ciencias Naturales La Salle 

[Caracas], 13(35): 205-225. 
DuMERiL, C, AND A. DuMERiL. 1851. Catalogue methodique de la collection 

des reptiles du Museum d'Histoire Naturelle de Paris. Paris: Gide et Baudry, 

224 pp. 
Dunn, E. R. 1944. The lizard genus /*/ze«aco5(3M/-M5. Caldasia, 3(1 1): 57-62. 
Hellmich, W. 1949. Auf der Jagd nach der Paramo-Echse. Deutsche Aquarien- 

und Terrarien-Zeitschrift, 2(5): 89-91. 
Lazell, J. D., Jr. 1969. The genus Phenacosaurus (Sauria: Iguanidae). Breviora, 

Museum of Comparative Zoology, 325: 1-24. 
RuEDA A., J. v., AND J. I. Hernandez-Camacho. 1988. Phenacosaurus indere- 

nae (Sauria: Iguanidae), nueva especie gigante, proveniente de la Cordillera 

Oriental de Colombia. Trianea (Acta Cientifica y Tecnologica INDERENA), 

2: 339-350. 



^ ^ MCZ 



LIBRARY 

B R E.y.iLiO R A 

usemm of Comparative Zoology 

UlNii ^ / ^-^ 
US ISSN 0006-9698 

Cambridge, Mass. 25 March 1991 Number 493 



NEW COLOSTETHUS (AMPHIBIA, DENDROBATIDAE) 
FROM SOUTH AMERICA 

Juan A. Rivero' 

Abstract. Six new species of Colostethus are described from South America: 
C. mittermeieri, C. idiomelus, and C poecilonotus from Peru; C. maculosus and 
C. paradoxus from Ecuador; and C . faciopunctulatus from Colombia. C. maculosus 
and C. faciopunctulatus belong to group VI (Rivero and Sema, 1988), C. mitter- 
meieri and C. idiomelus to group I, C. poecilonotus belongs to group IX, and C 
paradoxus to group IV. 

The relationship of the various species is discussed. C poecilonotus is the first 
member of group IX described from Peru, while C. paradoxus is the second 
member of group IV known from Ecuador. Group IV only extends south to the 
latitude of Quevedo in northwestern Ecuador. C paradoxus extends the range of 
the group to southern Ecuador. However, the possibility that the dilated third 
finger, which characterizes male members of group IV, may have arisen indepen- 
dently on more than one occasion is discussed. 

INTRODUCTION 

Examination of the Colostethus collection at the Museum of 
Comparative Zoology revealed a number of undescribed species 
from Peru, Ecuador, and Colombia. From these undescribed spe- 
cies a great deal is learned regarding their mutual relationships 
and the delimitation of the groups the genus has been divided 
into (Rivero, 1988). 

All the described species are well characterized and one of them 
is quite unique, so unique indeed that it cannot easily be assigned 
to any of the known groups of Colostethus. Yet, most of the groups, 
as originally suggested (Rivero, 1988), seem to have withstood 



Biology Department, University of Puerto Rico, Mayagiiez, Puerto Rico 00708. 



2 BREVIORA No. 493 

trial, although a few definitions may require modification. The 
division of the original group VI into groups VI and IX follows 
Rivero and Sema, 1988. A description in English (expanded from 
the original in Spanish) of the Andean groups occurring from 
Colombia to Peru is provided here. 

Many of the proportions usually incorporated into descriptions 
have been omitted here (diameter of tympanum in relation to eye 
diameter, diameter of eye in relation to the distance between eye 
and nostril, etc.) as they can easily be determined from the mea- 
surements. When many specimens were available, averages and 
proportions are provided in a Variation section. 

Most measurements were taken with a compass; snout-vent 
length, head breadth, and length of tibiae, with calipers. Head 
length was measured between the posterior edge of the tympanum 
and the tip of the snout. 

The web between the toes is considered insignificant if it does 
not extend beyond the midpoint of the first subarticular tubercle 
in at least four toes, minimal if it extends to the anterior border 
of the first subarticular tubercle in at least four toes ('A-webbed), 
intermediate if it extends beyond the first subarticular tubercle 
but does not reach the last articulation (disk) in at least three 
toes, and extensive if it reaches the last articulation in at least 
four toes. The central and usually indented part of the web is the 
portion considered for determining its extension. Considering the 
individual variation, a more detailed description is unnecessary 
and may make comparisons more difficult. However, the pedal 
membrane of all holotypes is illustrated in the corresponding 
figures. 

The author wants to thank E. E. Williams and J. Rosado for 
all their courtesies and attentions during his stay at the MCZ. 

DEFINITION OF GROUPS 

Group I. Two pectoral spots present, dorsolateral and ventro- 
lateral stripes absent; oblique-lateral stripes present and usually 
complete (from eye to groin), rarely absent; pedal membrane ab- 
sent or insignificant, rarely extensive; third finger of males not 
dilated; cloacal funnel absent. 

Group II. Dorsolateral stripes present; oblique-lateral stripes 
absent or incomplete (not reaching the eye); ventrolateral stripes 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 3 

present or absent; pedal membrane absent or insignificant; paired 
pectoral spots absent; third finger of male not dilated; cloacal 
funnel absent. 

Group IV. Third finger of males dilated; dorsolateral stripes 
usually absent; oblique-lateral stripe present or absent; ventro- 
lateral stripe generally absent; pedal membrane absent or insig- 
nificant (except in Colombian C. agilis); paired pectoral spots 
absent; cloacal funnel absent. 

Group V. Cloacal funnel present; dorsolateral stripe absent or 
indistinct; oblique-lateral stripe absent; ventrolateral stripe ab- 
sent; pedal membrane extensive; paired pectoral spots absent; 
third finger of male not dilated. 

Group VI. Pedal membrane usually extensive, at least Vs the 
length of the toes; first finger generally shorter than second; dorsal 
color usually blackish, sometimes marbled or spotted; dorsolat- 
eral stripes absent or not extending posteriorly beyond sacral 
hump; oblique-lateral stripes absent or incomplete; ventrolateral 
stripes generally absent; paired pectoral spots absent; third finger 
of male not dilated; cloacal funnel absent. 

Group IX. Dorsolateral stripes absent; oblique-lateral stripes 
present and usually complete (from eye to groin); ventrolateral 
stripes generally absent; pedal membrane absent or insignificant; 
paired pectoral spots absent; third finger of male not dilated (this 
last character distinguishes this group from group IV); cloacal 
funnel absent. 

DESCRIPTION OF SPECIES 

Colostethus mittermeieri, sp. no v. 
Figs, la-d 

Holotype. MCZ-A 100217, an adult female from Venceremos, 
394-395 km, on Marginal de la Selva Road, 1,620 m, Departa- 
mento de San Martin, Peru. Collectors: R. A. Mittermeier and 
H. Macedo Ruiz, 26 Sept. 1978. 

Paratypes. MCZ-A 100218-57. Forty specimens with the same 
data as the type. 

Etymology. Mittermeieri, in honor of Russell A. Mittermeier, 
one of the collectors of the species, and recent recipient of the 
N.Y. Zoological Society's Conservation Medal. 



BREVIORA 



No. 493 









5 mm 




Figure 1. MCZ-A 100217, holotype of Colostelhus mittermeieri. (a) dorsal 
view; (b) ventral view of hand; (c) throat and chest; (d) ventral view of foot. 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 5 

Diagnosis. A fairly large member of group I (Rivero, 1 988) with 
'/4-webbed toes, first finger shorter than second, fingers without 
lateral fringes, toes with distinct lateral fringes, venter almost 
always marbled, especially on the anterior half, males without 
vocal slits, no ventral sexual dichromatism, flanks with contrast- 
ing and sometimes elongated spots, digital disks broader than 
distal segments, and no dorsolateral, oblique-lateral, or contin- 
uous and distinct ventrolateral stripe. 

Description ofHolotype. Tip of snout broadly triangular beyond 
nostrils, almost vertical when seen from the side; nostrils antero- 
lateral, slightly protuberant; tongue spatulate, nicked behind, about 
% free; choanae rounded; canthus rostralis well defined, angular, 
curved; loreal region flat, vertical; tympanum moderate, covered 
posterodorsally by skin; external metacarpal tubercle rounded, 
prominent; internal tubercle elongate, prominent; palm of hand 
smooth, with a slight ridge along outer margin; proximal subar- 
ticular tubercles of first two fingers large, prominent; proximal of 
outer two fingers smaller, less distinct; distal of outer two fingers 
reduced, inconspicuous; first finger shorter than second, second 
shorter than last; fingers without lateral fringes; disks large, broad- 
er than distal digital segments; first disk slightly smaller than 
second and fourth; third disk about ^4 size of tympanum; a dis- 
tinct, oblique tarsal fold extending to inner subarticular tubercle; 
three metatarsal tubercles, inner more elongated, central less 
prominent, than outer; plantar surfaces smooth, with a distinct 
ridge along outer margin; toes with a minimal web; toe disks 
broader than distal digital segments; first disk smaller than others; 
all toes with distinct lateral fringes; heel of adpressed hind limb 
extending anteriorly to middle of eye. 

Head smooth; dorsal surfaces behind head covered with flat, 
minute warts sometimes fusing to form short ridges; a few tu- 
bercles behind sacral hump, in the proximity of cloacal opening; 
two or three tubercles between tympanum and arm; flanks with 
small, flat warts and glandular ridges; loreal region smooth; a few 
tubercles on dorsal surface of arms and a few others on antero- 
ventral surfaces of upper arm; venter smooth except for some 
indistinct granules on the posterior lateral margins of abdomen; 
ventral surface of thighs smooth. 



6 BREVIORA No. 493 

Color. Dorsum solid dark grayish brown; loreal region, upper 
lip and temporal area lighter than dorsum; flanks the same color 
as dorsum but with a few distinct white spots near groin and a 
white streak that extends from axilla to % distance from axilla to 
groin; white streak shorter and not extending to axilla on left side; 
a white or discolored area at attachment of forelimb; black lon- 
gitudinal streaks along anterior and posterior aspects of forearms 
or thighs absent; posterior aspect of thighs with indistinct blackish 
marbling on a greenish yellow background; dorsal aspect of thigh 
with indistinct transverse blotches or bars. 

Throat and anterior part of belly infuscated, this color more 
concentrated on throat and chest, with two extensive black spots 
discernible; sides of belly with some infuscation and marbling; 
dorsolateral, continuous ventrolateral, and oblique-lateral stripes 
absent; a few indistinct whitish spots or bars on anterior aspect 
of thighs. 

Variation. Almost always the dorsal color of adults is solid dark 
grayish brown or blackish, but it may be lighter gray in an oc- 
casional specimen, and indistinct darker spots may be discerned 
in some specimens, particularly the young ones. Transverse bars 
on the limbs may also be distinct in juveniles but are rarely so 
in adults. 

The white spots of the flanks may vary in distinctness and may 
not be apparent in some juveniles. The ventrolateral streak may 
be continuous between groin and axilla, it may be broken into a 
series of longitudinal spots, or it may be limited to a few anterior 
spots. It never has either the smooth margins or the continuity 
of the ventrolateral streak in members of groups II and IV, nor 
does it extend anteriorly beyond the origin of the arm. The dis- 
colored area at the attachment of the forelimb is present in most 
specimens but is not apparent in those in which the limbs are of 
a light color. 

A marbled and spotted ventral pattern may not be apparent in 
a few specimens (N5) but it is usually present. Infuscation of the 
throat and chest is generally present, but again, an occasional 
specimen may be of a plain white color, except for the two spots 
on the chest. In some specimens with a distinct marbled pattern 
on the anterior venter, the knee may also be marbled and/or 
spotted. 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 7 

The white spots on the anterior aspect of the thigh may be quite 
distinct, and in some specimens there is a row of white spots along 
the posterior margin of the thigh and tibial segment. 

In one female specimen, the dorsum is quite tubercular; in 
others there are small tubercles on the anterolateral area of the 
dorsum. The indistinct warts and rugosities of the dorsum are 
generally present but may be absent in young individuals. In the 
latter the disks are not usually broader than the distal digital 
segments. 

There is some slight variation in the amount of webbing, es- 
pecially of the first finger, which may be from fully webbed to 
about half-webbed. 

The tubercles on the forearm may be quite abundant and dis- 
tributed all along the anterior and posterior surfaces, or limited 
to a few, which form an irregular row on the posterior face of the 
arm. The belly is usually smooth, but a few specimens have gran- 
ules on the distal third of the belly, on the sides, and/or on the 
ventral surface of the thighs. 

None of the 10 recognizable males in the group is larger than 
20 mm and none has vocal slits. The latter characteristics may 
be a sign of immaturity; however, these specimens' testicles seem 
to be well developed. There may be a distinct size dimorphism 
in this species. 

Measurements and Proportions. See Tables la and lb. 

Discussion. Colostethus mittermeieri is a member of group I, 
whose most distinctive feature is the presence of two dark pectoral 
spots. 

Group I has 1 2 species, six of which are yet to be described 
(Edwards, 1974; Rivero, 1988). The group ranges from southern 
Colombia to Peru south to Cerro de Pasco. In the Peruvian Andes 
it is the dominant group and the only one in their highest ele- 
vations, but it occurs, as do groups VI and II, on the eastern flank 
of the Cordillera. 

The only member of group I so far known from the lowland is 
C. littoralis, described from Lima, but this form is apparently 
identical with a species from Ancash and may have been taken 
to the coast, either intentionally or accidentally. The small coastal 
population seems to have disappeared now but the name C. lit- 
toralis prevails for the mountain form. 



BREVIORA 



No. 493 



Table la. Colostethus mittermeieri males, measurements and proportions. 



Catalog No. 


100228 


100224 


100233 


Average 


SV 


19.00 


17.45 


15.30 


17.25 


HB 


6.65 


6.20 


5.40 


6.08 


HL 


7.70 


6.30 


5.50 


6.50 


ETS 


3.60 


3.35 


2.80 


3.25 


EN 


1.95 


1.70 


1.80 


1.82 


lOS 


2.10 


2.30 


2.05 


2.15 


UE 


1.80 


1.60 


1.50 


1.63 


ED 


2.80 


2.45 


2.50 


2.58 


DT 


1.20 


0.95 


1.20 


1.12 


LP 


8.70 


7.90 


7.20 


7.93 


LT 


8.90 


8.30 


7.35 


8.18 


LFl 


8.70 


7.60 


6.50 


7.60 


HB/SV 


0.35 


0.36 


0.35 


0.35 


HL/SV 


0.41 


0.36 


0.36 


0.38 


UE/IOS 


0.86 


0.70 


0.73 


0.76 


DT/ED 


0.43 


0.39 


0.48 


0.43 


ED/ETS 


0.78 


0.73 


0.89 


0.80 


ED/EN 


1.44 


1.44 


1.39 


1.42 


LF/SV 


0.46 


0.45 


0.47 


0.46 


LT/SV 


0.47 


0.48 


0.48 


0.47 


LFT/SV 


0.46 


0.44 


0.42 


0.44 


LF/LT 


0.98 


0.95 


0.98 


0.97 



Key: SV = snout-vent length; HB = head breadth; HL = head length; ETS = 
distance between eye and tip of snout; EN = distance between eye and nostril; 
lOS = breadth of interorbital space; UE = breadth of upper eyelid; ED = eye 
diameter; DT = tympanic diameter; LF = length of femur; LT = length of tibia; 
LET = length of foot. 



C. mittermeieri is distinguished from all other members of the 
group, with the exception of an undescribed species from Dos 
Rios in Pichinga, Ecuador, by lacking an oblique-lateral stripe. It 
is also the most extensively webbed species, as in all the others 
the toes are either free or have an insignificant web. 

Most of the Peruvian members of group I are distinctly spotted 
above. C. mittermeieri is not usually spotted, but when spotted, 
the spots are not distinct and contrasting. On the other hand, the 
white lateral spots are usually distinct and very contrasting, and 
one of the lower ones may form a usually discontinuous, undu- 



1991 



NEW COLOSTETHUS FROM SOUTH AMERICA 






O 

o 



O 
O 



in 

(N 

o 
o 



O 

o 



ON 

(N 
JN 

O 

o 



O 

o 



(N 

o 
o 



o 

o 



OS 

O 
O 



IT) 

(N 

O 

o 



o 

(N 

o 
o 



o 
o 



QO 

o . 

u 



<^ CX5 so 

o -^ r- 

u-i od 00 Tf 
rsi 



0<^or<^TfTi-ir^d«mmooTj- 

di di d> d> d> -^ 



[^ 00 VO ON "T^ 

OO Tt Tt Tt TJ- 



ON 



«N<N<Nm — — '(N — 



o o o o 



0'noo'/^0'/^00«^o«^NOoomov'^i^oom<^) — 

O^ONiO(N'^Of*^rOO — ONrnr<-ioor^ON'^-^>OU-iOs 

--'r-'r-~"rorvir^rNir<S — o—'OOOOOO — oooo 



o o o o o o o 

fN « ro Tj- o ■^ O 



>r^ m o "/^ O 


li-i vo 


r«1 oo O 


O — ro (N -^ 


r<-i m 


00 f^ On 



m vo — I t^ — ' 
m 'i- >r^ -^ Ov 



t^t^rniNfNJNr'i — OvOOnOOOOO — OOOO 



O >/^ "/^ O O O O 

00 (N r~ O «^ 'i- ^ 



(N 


OS 


OS 


O 

o 


o 
oo 






r-- 


00 



ooooo — oooo 



Ti-(Nr^<Nrn — m^a-fN 



O 

o 



i/-iir)minoOOmTrooir)i/-iTj- — tt — vo^ 
mvOfNrom — oO'Or^rooOTtoom'^m'^oo 



■<1-<N(N(Nm — O — OOOOOO — OOOO 



OOOOOOin>O>r>i/-ii/-ii/-ifNr-r-v00N<Ns000 — so 
OO<^>/^'/^(N'*>ns00NTj-r^f0<^r^Tri^Tl-TfTj-Tt0N 

odosO'^'rsi<^r^irO---^(Nrn--^00000--0000 



■r^OOOOOOOOOO 
moosmoomo— '<Ni/-iO 



lOvOsOOOv-^rslOvtN 
■Ot^rnoOf^osl^Tj-ir^ 



00 OS OS -"I- 



OO'Oi/^O'^OOOO'/^Of^'^Os — 
'Tir^fN'^m — mt^insor^Osf^ror^Tf 

r~^odos 

fN 



oooooooo 

O'^w^sO'^O'^O 



o 
o 



00 OS 



moooo(^(N<N<Nm 



in o 



OS OS 00 
fN 



Tf 00 00 Tl- 
fN 


























> 


> 


O 


D 


C/5 


Z 




















Ih 


C/D 


or) 


LU 


UJ 


LU 


J 




7. 


O 


U 


Q 


H 


Uh 


H 


oa 


-J 


u 


H 


Q Q 


X 


U 


w 


J 


u 


U 


J 


J 


J 


K 


K 


J 


U 


u 


W 



OS 



Ost^soro — rs|<NrO— lOsOOOOOOOO— 'OOOO 



ooooomininoooo'tf^oor^r^sooosof*^'^ 

Omr'^fNiOfNmsor^-TrosOf^'^l^'^oO'^'^Tr'^O 



(Nmrslr<1— •r'1(N(NOOOOO— 'OOO — 



rO 00 <N O <^ ■^ 
00 -"^ tT m •rf 00 

■'*(Nr<^fNm—' — m— 'OOOOO— 'OOOO 



iniriTtOOf^Ttt^t^SOOOSOt^ 
f^l^ror'loO'^oosOTfTtTtOs 



O'^rsimfNTf-Hcrir'lcNOOOOO— 'OOOO 



O'/^oomi'^'n'/^'noooin'^f^it^— '<^soosTj-Tr 

mfN — r^r<)l^<Nr<^fNOssO'^rOrO00rOOs-<a-Tr'^'<a-Os 



OOOOOO— 'OOOO 



00>oO>n'nOOlor^^OOoooomTJ■sor^so 
Tft^fNsOOOOOOsTfsOf^rnoOTj-oOTtTj-Ta-Tt'Os 

rfrsir<ir4r<^ — JNrOfNOOOOO— 'OOOO 



OOOOOO'/^'^^'/^'^'/^O'^'^ — OOs— 't^Ossoso 
r-inr<-)OTfr<irn — (Nso— 'rOrnroI^Tfl^m'rrTfTros 



rsimfNcO — — <N — OOOOO— 'OOOO 



^ > 

> > C/5 

C/5 C/5 > 

ti; H C 

hJ hJ J 



H 

tu 



173 

H 

<u 
u 
en 

en 

u 

aj 
u 

ca 



o 



10 BREVIOKA No. 493 

lating line from the axilla to the proximity of the groin. The streak 
may not be analogous to the ventrolateral streak of groups II 
and IV. 

Colostethus idiomelus, sp. nov. 
Figs. 2a-c 

Holotype. MCZ-A 100260, an adult female from Venceremos, 
394-395 km, on Marginal de la Selva Road, 1,620 m, Departa- 
mento de San Martin, Peru. Collectors: R. A. Mittermeier and 
H. Macedo Ruiz, 26 Sept. 1978. 

Etymology. Idiomelus, from the Greek idio, distinct, peculiar, 
and melos, limb, in reference to the strikingly colored hind limbs 
of the species. 

Diagnosis. A medium-sized member of group I with small pec- 
toral spots, no dorsolateral or ventrolateral stripes, oblique-lateral 
stripe present and extending to the eye, first finger shorter than 
second, a short basal web between toes I and II, and II and III, 
a narrow lateral fringe on the inner side of toes II and III, fingers 
and toes long and slender, the disks small, much smaller than the 
tympanum, and the thighs with distinct transverse blotches on a 
white background. 

Description of Holotype. Snout short, the tip rounded, more or 
less vertical when seen from the side; nostrils anterolateral, not 
protruding, very near end of snout; tongue spatulate, broad, nicked 
behind, about % free; choanae small, ovate; canthus rostralis 
rounded but angular, not appreciably curved; loreal region ver- 
tical, flat; tympanum flushed with surface, covered posterodor- 
sally by skin; external metacarpal tubercle rounded, obliquely 
ridged; internal tubercle smaller, elongate; palm of hand smooth, 
with a narrow ridge along outer margin; basal subarticular tu- 
bercles of fingers I, II, and III large, distinct; distal tubercle of 
finger III and basal and distal tubercles of finger IV smaller and 
less distinct; fingers long, slender; first finger slightly shorter than 
second, second shorter than last; fingers without lateral fringes; 
disks small, approximately equal in size and slightly broader than 
distal digital segments; disk of third finger about '/3 size of tym- 
panum; an oblique, internal tarsal fold extending to inner meta- 
tarsal tubercle; metatarsal tubercles prominent; outer tubercle more 
or less rounded; inner, elongate; plantar surfaces smooth and with 



1991 



NEW COLOSTETHVS FROM SOUTH AMERICA 



11 





IS mm 




1cm 



Figure 2. MCZ-A 100260, holotype of Colostethus idiomelus, (a) dorsal view; 
(b) ventral view of hand; (c) ventral view of foot. 



distinct outer ridge extending as narrow keel along outer edge of 
fifth toe; toes slender, with insignificant web; disks of toes small; 
first disk not broader than digit; others slightly broader; heel of 
adpressed hind limb extending anteriorly to middle of eye. 

Dorsum smooth on head and anterior part of body but with 
indistinct flat warts increasing in size near cloacal opening; loreal 
region and flanks smooth; a tubercle between tympanum and arm; 



12 BREVIORA No. 493 

three or four small tubercles along posterior surface of lower arm; 
ventral surfaces, including posterior surface of thighs, smooth. 

Color. Dorsum grayish brown with moderately contrasting 
blotches and spots which are much smaller on snout and head; a 
black streak on each side, from behind eye, crossing groin area 
and continuing along anterior border of thigh to knee; an oblique, 
white (red?) stripe crossing base of thighs and on one side con- 
tinuing as a longitudinal stripe along posterodorsal margin, on 
other, breaking into a series of spots; anterior part of thighs white, 
this color getting dusky at fusion with white streak or spots on 
posterodorsal margin; white or whitish area crossed by three very 
distinct and contrasting dark brown spots or bars; posterior part 
of thighs behind white streak (or series of spots) dark brown, 
lighter and profusely spotted with white at proximal end; rest of 
hind limbs light brown with darker crossbands; a short dark brown 
streak on anterior part of upper arm and a longer one on posterior 
portion of same segment; a small, dark brown spot around nostril; 
a dark canthal streak continuing to tip of snout but not meeting 
contralateral streak; exposed part of tympanum white; upper lip 
and temporal area dusky, with spots below anterior and posterior 
comers of eye and smaller spots between these and below canthal 
streak; a whitish oblique-lateral stripe from behind eye to prox- 
imity of groin; groin area very dark brown, with distinct and 
contrasting white spots, one of which extending into white, oblique 
streak at base of thighs; the brown color, on the other hand, 
extending into black streak along anterior aspect of thighs; ventral 
surfaces immaculate except for a dusky line along margin of lower 
jaw and two tiny pectoral spots. 

Measurements (mm). Snout-vent 25.3; head length 8.2; head 
breadth 8.1; eye diameter 3.0; eye-nostril 2.0; eye-tip of snout 
4.0; upper eyelid 2.0; interorbital space 3.0; femur 12.0; tibia 
12.2; foot 12.5. 

Discussion. Colostethus idiomelus shares with most of the other 
Peruvian members of group I the oblique lateral stripe and the 
spotted coloration of the dorsum. Colostethus sylvaticus, C. elach- 
yhistus, and C. littoralis, the other described members of group 
I, have a greater amount of webbing and a distinct lateral fringe 
on all toes. None has the striking coloration of the thighs nor the 
white spots on the flank that seem to characterize C. idiomelus. 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 13 

Two undescribed species were considered by Edwards in his 
thesis (1974). In one, from Ancash, there is no pedal webbing 
and the oblique-lateral line does not extend to the eye; in the 
other, from Huanuco, fingers and toes are fringed, the flanks are 
dark brown, with a number of yellowish flecks and the dorsal 
surfaces of the limbs are olive tan with brown bars or spots. 

A third species collected by Edwards near Zamora in Ecuador, 
but not yet described, has a yellow spot at the insertion of the 
arm and brown legs with small black spots and transverse bars. 

Colostethus idiomelus is more typical of group I than C mit- 
termeieri. It differs from that species, among other things, in hav- 
ing black streaks along the anterior and posterior aspect of the 
upper arms, in having a reduced amount of webbing, no black- 
ening of the throat, very small pectoral spots, and an oblique- 
lateral streak. 

Colostethus poecilonotus, sp. nov. 
Figs. 3a-c 

Holotype. MCZ-A 89108, an adult female from between Cha- 
chapoyas and Bagua Grande Alva, 500 m, Departamento Ama- 
zonas, Peru. Collector: R. A. Mittermeier, 3 May 1974. 

Paratypes. MCZ-A 89106-7, 89109, all adults and with the 
same data as the type. 

Etymology. Poecilonotus, from the Greek poikilos, variegated, 
spotted, mottled, and notos, back, in reference to the spotted 
dorsum of this species. 

Diagnosis. A small Colostethus probably belonging to group IX, 
with granular, spotted dorsum, no dorsolateral or ventrolateral 
stripes, oblique-lateral stripe present and not extending beyond 
level of axilla, first finger equal in length to second, fingers without 
lateral fringes, toes free and without lateral fringes, first and fifth 
toe disks scarcely broader than the respective distal digital seg- 
ments, venter immaculate, granular on the posterior portion. 

Description of Holotype. Tip of snout slightly convex beyond 
nostrils and slightly inclined inwards towards lip when seen from 
the side; nostrils anterolateral, slightly protruding; tongue spat- 
ulate, entire, % free; choanae small, rounded; canthus rostralis 
sharp and angular, curving towards nostrils; loreal region flat, 
vertical; tympanum conspicuous, posterodorsally covered by skin; 



14 



BREVIORA 



No. 493 






1cm 



Figure 3. MCZ-A 89 1 08, holotype of Colostethus poecilonotus. (a) dorsal view; 
(b) ventral view of hand; (c) ventral view of foot. 



external metacarpal tubercle rounded, very protuberant; inner 
tubercle elongated and smaller; palm of hand rugose but without 
supernumerary tubercles and with a ridge along outer margin; 
subarticular tubercles prominent, the second of third finger and 
the two of fourth, the smallest (with the latter more prominent 
than the former); first finger equal in length to second, second 
slightly longer than last; fingers slender and without lateral fringes; 
disks of fingers small, all of approximately equal size and slightly 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 15 

broader than corresponding digital segments; disk of third finger 
not more than 'A size of tympanum; a short, obhque tarsal tubercle 
extending as a thin, inconspicuous ridge to inner metatarsal tu- 
bercle; inner metatarsal tubercle elongate, longer than the smaller, 
rounded, and conical outer tubercle; plantar surfaces smooth and 
with a narrow external fold extending along outer margin of fifth 
toe to disk; subarticular tubercles of toes small, but conical and 
protuberant; toes long and slender, free; fourth toe with narrow, 
indistinct lateral keels on outer segments; first and last disks the 
smallest and not broader than the corresponding distal digital 
segments; heel of adpressed hind limb extending anteriorly to 
between eye and nostril. 

Dorsum granular, the granules tending to be more prominent 
towards posterior end; limbs granular and tubercular; upper eye- 
lids granular; loreal region smooth; flanks granular and tubercular, 
especially towards groin; two small tubercles between tympanum 
and arm; throat smooth; abdomen granular on posterior third; 
posterior aspect of thighs smooth; two or three tubercles along 
anterior margin of lower arm. 

Color. Brownish tan with distinct, darker spots and a distinct 
whitish oblique-lateral stripe from level of axilla to groin; a black 
canthal stripe continuing, in back of eye, to about level of arm 
insertion, after which, continuing posteriorly as a thin stripe above 
oblique-lateral stripe to groin; upper flanks brownish, with two 
or three whitish spots near groin; face, lower part of tympanum, 
and temporal area whitish, this color continuing above arm in- 
sertion to lower flanks; an indistinct, brown longitudinal line along 
anterior face of upper arm and also along anterior face of thighs; 
hind limbs with narrow cross-bars and spots; posterior aspect of 
thighs approximately of same color as dorsal surfaces; venter 
immaculate. 

Variation. Paratype MCZ-A 89109 is very similar to the type 
in coloration, but the black streak above the oblique-lateral line 
is not easily discernible, the longitudinal line of the upper arm is 
very short, there are no white spots on the posterior aspect of the 
flanks, and all disks are broader than the respective digital seg- 
ments. 

In MCZ-A 89107 the dorsal spotting is less contrasting, the 
loreal region and face are more infuscated or spotted, and the 



16 




BREVIORA 




No. 493 


Table 2. 


Colostethus 


poecilonotus 


females, measurements and proportions. 


Catalog 












No. 


89106 


89107 


89108 


89109 


Average 


SV 


21.70 


24.65 


20.50 


19.70 


21.64 


HB 


7.20 


7.65 


7.00 


6.20 


7.01 


HL 


8.80 


9.45 


8.35 


7.60 


8.55 


ETS 


3.80 


3.85 


3.65 


3.35 


3.66 


EN 


2.10 


2.20 


2.10 


1.85 


2.06 


lOS 


2.75 


3.00 


2.80 


3.90 


2.86 


UE 


1.85 


1.90 


1.55 


1.40 


1.68 


ED 


3.00 


3.20 


2.95 


2.80 


2.99 


DT 


1.50 


1.70 


1.45 


1.40 


1.51 


LF 


9.80 


10.30 


9.80 


8.90 


9.70 


LT 


10.85 


11.05 


10.80 


9.55 


10.56 


LEI 


10.10 


10.00 


10.10 


9.00 


9.80 


HB/SV 


0.33 


0.31 


0.34 


0.31 


0.32 


HL/SV 


0.41 


0.38 


0.41 


0.39 


0.40 


UE/IOS 


0.67 


0.63 


0.55 


0.48 


0.59 


DT/ED 


0.50 


0.53 


0.49 


0.50 


0.51 


ED/ETS 


0.79 


0.83 


0.81 


0.84 


0.82 


ED/EN 


1.43 


1.45 


1.40 


1.51 


1.45 


LF/SV 


0.45 


0.42 


0.48 


0.45 


0.45 


LT/SV 


0.50 


0.45 


0.53 


0.48 


0.49 


LF17SV 


0.47 


0.41 


0.49 


0.46 


0.46 


LF/LT 


0.90 


0.93 


0.91 


0.93 


0.92 



For key to characters see Table la. 



dark lateral band is broad and has the oblique-lateral line within 
its confines. The dorsum of this specimen is more tubercular than 
in the type and there is a central tubercle between inner and outer 
metatarsal tubercles. 

In MCZ-A 89 106 there is more spotting on the flanks and some 
marbling behind the axilla. In both this specimen and MCZ-A 
89107 the thighs are marbled dark brown on a lighter brown 
color. 

In most specimens, the oblique-lateral line tends to become 
whiter, broader, and more distinct as it approaches the groin, and 
the elbow, the knee, and the heel tend to show a discolored area 
or spot, but it could not be ascertained if this was the result of 
erosion or if it is a natural spot. 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 17 

The lateral keels on the distal segments of the fourth toe are 
not evident in some specimens and cannot be described as fringes. 

Measurements and Proportions. See Table 2. 

Discussion. Colostethus poecilonotus is the first member of group 
IX described from Peru. Yet, its presence here is not unexpected 
as the group is known from southeastern Ecuador, where it is 
represented by at least three species. One of these, C. festae is 
little known but it is supposed to have a short web and no oblique- 
lateral stripe. The others, reported by Edwards (1974) but not yet 
described, may have a web, fringes on the toes, or a marbled 
venter. 

Sometimes, individuals of species belonging to group I may 
not have the pectoral spots that are diagnostic of the group. Thus, 
C. poecilonotus may actually be a member of group I, which is 
the most typical group in Andean Peru. There is no way of know- 
ing, however, until more specimens become available. 

Only one undescribed member of group I reported from Peru 
(Edwards, 1974) lacks toe webbing, but in this species there are 
fringes on the fingers and toes, and the first finger is longer than 
the second. 



Colostethus maculosus, sp. nov. 
Figs. 4a-c 

Holotype. MCZ-A 91558, an adult male from Puyo, between 
Turingia and theatre, 950 m, Provincia Pastaza, Ecuador. Col- 
lectors: K. Miyata and H. Weed, 22 July 1976. 

Paratype. MCZ-A 104946, an adult male from El Reventador 
(ca. 1,200 m), Provincia Napo, Ecuador. Collector: Giovanni 
Onore, Leg. June, 1983. 

Etymology. Maculosus, from the Latin maculosus, spotted, 
mottled, in reference to the color of the dorsum in this species. 

Diagnosis. A medium-size Colostethus referable to group VI, 
with mostly smooth and spotted dorsum, first finger shorter than 
second and second much shorter than fourth, vesicular inflam- 
mation at base of third finger, extensively webbed toes, no dor- 
solateral or ventrolateral stripes, but variously distinct, oblique- 
lateral line present, black lateral band not extending posteriorly 
behind arm, and immaculate venter. 



18 



BREVIORA 



No. 493 





5mm 




Figure 4. MCZ-A 91558, holotype of Colostethus maculosus. (a) dorsal view; 
(b) ventral view of hand; (c) ventral view of foot. 



Description of Holotype. Tip of snout almost truncate, slightly 
inclined inwards towards the lip when seen from the side; nostrils 
anterodorsal, scarcely protruding; tongue narrow, ovoid, nicked 
behind, and nearly Vi free; choanae small, rounded; canthus ros- 
tralis sharply angular; loreal region vertical, flat; tympanum mod- 
erate, covered posterodorsally by skin; external metacarpal tu- 
bercle large, rounded, protuberant; internal tubercle smaller, 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 19 

elongate, and less prominent; palm of hand smooth, with a distinct 
pad or cushion at base of third finger; an indistinct outer ridge 
along outer margin of hand; subarticular tubercles rather small, 
the ones in outer finger the smallest; first finger much shorter than 
second, second considerably shorter than fourth; fingers flat, with- 
out lateral fringes; disks large, the first the smallest, all broader 
than distal digital segments; disk of third finger about % size of 
tympanum; an oblique tarsal fold continuing along outer margin 
of first toe to its disk; inner metatarsal tubercle small, elongate; 
outer tubercle smooth, with a ridge along outer margin; plantar 
surfaces smooth and with an outer ridge continuing to disk of last 
toe; subarticular tubercles of toes small, inconspicuous; first toe 
disk the smallest, followed in size by fifth and second; all disks 
broader than distal digital segments; toes with an intermediate 
web; all toes with broad distinct lateral fringes; heel of adpressed 
hind limb extending anteriorly to middle of eye; a vocal slit on 
each side, not too close to angle of jaw, and two compact, rugose 
pouches behind each jaw. 

Dorsum smooth except for a few small tubercles at posterior 
end (under high magnification, dorsum covered with flat, incon- 
spicuous warts); flanks and loreal region smooth; ventral surfaces 
minutely granular; posterior aspect of thighs smooth. 

Color. Dorsum light brown, spotted and mottled with darker 
brown; an indistinct canthal streak; loreal region, face, and tem- 
poral areas light brown, lighter than dorsum; upper flanks ap- 
proximately the same color as dorsum but with a black band from 
posterior comer of eye to base of the upper arm; an indistinct, 
whitish oblique-lateral line from groin to about halfway along 
flank; hind limb with dark, narrow transverse bars; posterior as- 
pect of thighs same color as dorsum; ventral surfaces immaculate 
but with some infuscation on throat and chest, especially at base 
of forelimbs. 

Measurements (mm). Snout-vent 21.0; head length 10.0; head 
breadth 6.8; eye diameter 3.0; eye-nostril 2.0; upper eyelid 2.0; 
interorbital space 2.3; femur 9.6; tibia 10.3; foot 10.0. 

Variation. Specimen MCZ-A 104946 is considered with some 
misgivings as a paratype, but the presence of the hand pad in 
both hands of both specimens and the sharing of a very short 
second finger, much shorter than the fourth, led the author to 



20 BREVIORA No. 493 

believe that they belong to the same species although they may 
eventually be found to be subspecies of each other. 

The paratype is a little larger than the type (22.0 mm), slightly 
less webbed and more distinctly spotted above, but its most dis- 
tinctive feature is an oblique-lateral stripe that extends from eye 
to groin. However, the short oblique-lateral stripe of the holotype 
may have extended to the eye in the living animal and the anterior 
portion may have faded in preservation. Until more specimens 
are collected it is not possible to come to a conclusion. 

Discussion. Only three members of group VI have been reported 
from Ecuador and one of the three is still undescribed (Edwards, 
1974). This last species is from Rio Azuela, in the same river 
system and relatively close to Reventador, but in this species there 
is no oblique-lateral stripe, the first finger is said to be longer or 
equal to the second, the venter is spotted, and there are lateral 
fringes on the fingers. 

The other members of group VI in Ecuador, C fuliginosus and 
C. nexipus, are quite different from C. maculosus, and so is a 
species from western Ecuador, which will be described elsewhere. 

Colostethus paradoxus, sp. no v. 
Figs. 5a-d 

Holotype. MCZ-A 1 03924, an adult male from Lamtac, Cuenca, 
2,535 m, Provincia Azuay, Ecuador. Collector: Giovanni Onore, 
April 1982. 

Etymology. Paradoxus, from the Greek paradoxos, strange, 
contrary to all expectations, in reference to the combination of 
characters in this species. 

Diagnosis. A relatively small Colostethus referable to group IV, 
with dorsolateral and ventrolateral stripes, no oblique-lateral stripe, 
a dark-colored and well-defined lateral band; a short web between 
toes II and III, and III and IV, finger disks not broader than the 
distal digital segments, first finger longer than second, three outer 
fingers and all toes with a lateral fringe, and male with a dilated 
third finger. 

Description of Holotype. Tip of snout more or less rounded 
beyond nostrils; rounded when seen from the side; nostrils an- 
terolateral, not protruding; tongue spatulate, entire, not quite '/a 
free; choanae small, rounded; canthus rostralis rounded but an- 



NEW COLOSTETHUS FROM SOUTH AMERICA 




21 





5nnm 




Icm 



Figure 5. MCZ-A 103924, holotype of Colostethus paradoxus, (a) dorsal view; 
(b) ventral view of hand; (c) lateral view; (d) ventral view of foot. 



gular, straight; loreal region vertical, flat; tympanum moderate, 
its upper half covered by skin; external metacarpal tubercle con- 
ical, protuberant; inner tubercle slightly more elongate, less pro- 
tuberant; palm of hand smooth, with a ridge along outer margin; 
subarticular tubercles large, proximal of first finger and distal of 
fourth, smaller than others; first finger longer than second, second 
slightly shorter than last; third finger dilated; three outer fingers 



22 BREVIOR.A No. 493 

with distinct lateral fringes; disks not broader than distal digital 
segments and all more or less of same size; disk of third finger 
not more than Vs size of tympanum; a transverse, short, tarsal 
fold; metatarsal tubercles conical, prominent, with inner tubercle 
slightly more elongate than outer tubercle; plantar surfaces smooth, 
with a slight ridge along outer edge that continues to disk of fifth 
toe; a minimal web between toes II and III, and III and IV; except 
for first disk, all others slightly broader than distal digital seg- 
ments; toes with lateral fringes; heel of adpressed hind limb ex- 
tending anteriorly to between eye and nostril; a pair of vocal slits 
not too close to angle of jaw. 

Dorsum smooth except for a few tubercles between insertion 
of hind limbs, and a fringe of tubercles margining fold above 
cloaca; loreal region and flanks smooth; ventral surfaces, including 
posterior aspect of thighs, smooth; posterior aspect of upper arm 
smooth. 

Color. Above, solid light brown; two lighter colored dorsolateral 
stripes from posterior comer of eye crossing to groin and extend- 
ing for short distance on thighs; black lines along the anterior or 
posterior aspects of forearms or thighs absent; loreal and temporal 
regions, including lower edge of tympanum, cream, this light color 
extending posteriorly as a ventrolateral stripe; a short brown streak 
below eye; thigh with a brownish bar between whitish area of 
proximal portion and with another one closer to knee; two elon- 
gate, whitish spots on posterodorsal aspect of thighs; forelimbs 
and rest of hind limbs uniform light brown; a canthal streak; 
flanks with a distinct and well-defined dark brown band from 
behind eye to groin; white ventrolateral stripe margined below 
by an irregularly margined brown streak; ventral surfaces infus- 
cated and marbled on throat and limbs, but much less so on belly 
(except for the brown lateral streaks described above); whitish or 
unpigmented areas present in feet, metatarsal segments, and tibial 
segments. 

Measurements (mm). Snout-vent 19.5; head length 6.2; head 
breadth 6.0; eye diameter 2.7; eye-nostril 2.0; eye-tip of snout 
3.0; upper eyelid 3.4; interorbital space 3.1; femur 7.8; tibia 9.9; 
foot 8.2. 

Discussion. The swollen third finger of the male places Colo- 
stethus paradoxus in group IV. However, this species is very 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 23 

similar to some members of group II, particularly C. kingsburyi 
and a species from Cochabamba, Bolivia, reported by Edwards 
(1974). It also has the dorsolateral and ventrolateral stripes and 
the distinct and continuous dark lateral band that characterize 
members of group II, but in this respect, it is not different from 
C. pratti, which also has a dilated third finger and is a clear 
member of group IV. C. talamancae on the other hand, has dor- 
solateral and ventrolateral stripes and a dark lateral band but the 
males do not have a dilated third finger, which is the reason why 
Rivero (1988) suggested that it could be a member of group II in 
spite of its distribution, west of the Andes. This only serves to 
confirm the close relationship between groups II and IV, but 
whether a dilated third finger has arisen independently in different 
members of group II cannot be confirmed at present. 

Rivero and Sema (1988) indicated that group IV was typical 
of western Colombia (and Central America) and that it did not 
extend southwards beyond Quevedo in northern Ecuador. Yet, 
C. paradoxus is from Southern Ecuador and it doesn't show any 
relationship to the only Ecuadorian member of group IV (still 
undescribed; Edwards, 1974). In spite of its dilated third finger 
it seems to be closer to the East Andean members of group II. If 
it is to be related to any member of group IV, it is to C. pratti 
and perhaps C. latinasus, but these are only found in Central 
America and in Colombia, west of the Oriental Cordillera. 

Colostethus faciopunctulatus, sp. nov. 
Figs. 6a-c 

Holotype. MCZ-A 94751, an adult male from Puerto Narifio 
(3°46'N, 71°23'W, 15 km W. of Leticia), Departamento Ama- 
zonas, Colombia. Collector: R. Bleiweiss, 19 July 1977. 

Paratypes. MCZ-A 94746-50, 94552-6, 93782, 94757-61, 
96016-7, all from the same locality and collected by R. Bleiweiss, 
but 93782 collected on July 9, 94757-60 on July 19-21, 96016- 
7 on July 27, and 94761 on July 31. 

Etymology. Faciopunctulatus, from the l^dilin fades, face, and 
punctulatus, dotted, in reference to the white dots on the loreal 
region, under the eye, and in the temporal region of this species. 

Diagnosis. A medium-size Colostethus referable to group VI, 
with extensive webbing between the toes, generally with contrast- 



24 



BREVIORA 



No. 493 





5mm 




Figure 6. MCZ-A 9475 1 , holotype of Colostethus faciopunctulatus, (a) dorsal 
view; (b) ventral view of hand; (c) ventral view of foot. 



ing white dots on the loreal area, under the eyes, and in the 
temporal region, distinct ventral sexual dichromatism, the males 
having a blackened throat with white dots, short fingers, the sec- 
ond considerably reduced, no dorsolateral, ventrolateral, or 
oblique-lateral stripes, and a gray dorsum, usually with contrast- 
ing, black, V-shaped or transverse markings in back of the eyes 
and behind the sacral hump. 



1991 NEW COLOSTETHUS FROM SOUTH AMERICA 25 

Description of Holotype. Tip of snout truncate, almost vertical 
when seen from side; nostrils lateral, slightly protuberant; tongue 
spatulate, indented behind, and about % free; choanae small, 
ovate; canthus rostralis somewhat rounded, curved; loreal region 
slightly slanting and concave; tympanum moderate, not partic- 
ularly distinct, covered posterolaterally by skin; external meta- 
carpal tubercle rounded; inner tubercle narrow, elongate; palm of 
hand smooth, with no apparent ridge or fold on outer edge; sub- 
articular tubercles moderate, not too prominent; fingers short; 
first finger slightly longer than second, second shorter than last; 
fingers without lateral fringes; disks moderate, of approximately 
equal size, broader than distal digital segments; disk of third finger 
a little more than ~h size of tympanum; a short, oblique tarsal 
fold extending as a fringe along outer margin of first toe; metatarsal 
tubercles prominent, with outer tubercle rounded and inner tu- 
bercle elongate; plantar surfaces smooth, with a ridge along outer 
edge; pedal web intermediate; disks of toes broader than distal 
segments; first and last toe disks smaller than others; a distinct 
lateral fringe on toes; heel of adpressed hind limb extending an- 
teriorly to middle of eye; a pair of vocal slits close to angle of 
jaw. 

Dorsal surfaces smooth; loreal region and flanks smooth; ab- 
domen and throat granular. 

Color. Above, gray with a black, contrasting bar between the 
eyes, another V-shaped bar in back of the eyes, and a few smaller 
spots in back of the sacrum; a canthal streak; a short black bar 
between eye and base of arm; loreal region, area under eye, and 
temporal region blackish, with contrasting white dots; flanks a 
little darker than dorsum and with one or two white spots near 
inguinal region; a white or very little pigmented area in axilla; 
thighs and tibiae with indistinct dark blotches; posterior aspect 
of thighs indistinctly marbled black and tan; throat blackish (this 
color more intense on sides) and with white dots; abdomen white, 
with milky white dots on posterior end; arm tubercles white; 
ventral aspect of arms and posterior proximal aspect of thighs 
dark gray with white dots; no dorsolateral, ventrolateral, or 
oblique-lateral stripes; longitudinal black lines on the anterior 
and posterior aspects of arms or thighs absent. 

Variation. There is a distinct ventral sexual dichromatism, the 



26 BREVIORA No. 493 

males having a blackened throat with white dots while the females 
are uniformly colored. The white dots on the side of the head and 
the ventral portion of the thighs and forelimbs are either absent 
or inconspicuous in the females. 

The first finger is slightly longer than the second in two male 
specimens (including the type), shorter than the second in both 
females and one adult male, and equal to the second in one male. 
The first finger is shorter than the second in two juveniles and 
longer in one. 

The dots on the side of the head are present, in various degrees 
of distinctness, in all individuals except in one female, but the 
dots on the flanks are absent in four juveniles, and those on the 
lower aspect of the thighs are absent in most. 

The dorsal color may be light gray or brownish gray, with 
contrasting dorsal markings, or very dark gray with imperceptible 
markings. The discolored area of the axillae is present in all spec- 
imens but may not be noticeable in those in which it is continuous 
with the ventral coloration. 

Juveniles tend to be of a tan or light yellowish brown color with 
contrasting dorsal spots and better-defined bars on the legs. The 
white ventral dots are present in most individuals, and certainly 
on the throat of all males, but the abdominal dots may only be 
evident under a lens. 

Granules are present on the abdomen but not evenly distributed 
in most cases and absent in some cases. 

Measurements and Proportions. See Table 3. 

Discussion. Colostethus faciopunctulatus is a clear member of 
group VI (as restricted), although in all members of that group 
the first finger is almost always shorter than the second, while in 
a few specimens of C faciopunctulatus the first finger is slightly 
longer and in others it is equal to the second. 

Distinctive features of C. faciopunctulatus are the white dots 
on the face, the very reduced second finger, and the usually con- 
trasting dark bars or splashes on the dorsum. 

The only member of group VI reported from eastern Colombia 
is C fuliginosus, and the only members from eastern Ecuador 
are, besides C. fuliginosus, C. nexipus and an undescribed species 
from Rio Azuela (between Quito and Lago Agrio), 1 ,740 m, Napo, 
Ecuador (Edwards, 1974). 



1991 



NEW COLOSTETHUS FROM SOUTH AMERICA 



27 



15 
E 



a 

> 
< 



OS 



O 

ON 



Ov 



4J 
QO 

m 

k> 

< 



ID 

■<^ 

ON 



OD 

O . 



P3 

u 



mr-mro</^>/^'/^oot^t^i^'/^oor-<N>/^ONr^ON>o — r~- 

OOOsOsON'^"'1<^'S"''^''^00(^r*^<~OON'^00'^T}-'<^'^0 

rnc>dodr-i(N<NfNrO--'--'OOvOOOOO--'000— ' 



iriir)000«^00>'^000 
fNododf^rNrNifNc^ — — 'OON 



mroos'/^av'^'/^'^'^o 

OOOOO— 'OOO-^ 



OOOOOi'^'/^OOOOnO^-- 
— rOfN'^ON'/^sDNO^^sOrOr^ 



00 -rr t^ (N NO 


<N CO On 


00 Tt 00 i/^ •<a- 


Tj- (^ O 



m ON On 



Tt (N (N fN r«-i — 



OONOOOOO— 'OOO— ' 



Oi'lOO'OOOO'/^OO 



"/^ On 00 NO — 1/^ <N 
NO f*^ rn On ■rf 00 (^ 



O ON i/^ (^ 
m -rj- tt o 



m ON ON 



Tf<NfNrNlrO" — — 'OOOOOO — OOO — 



rv)oo— ir-miri(Nr<^m^'ri>r^mroON'^ON'n-Ti--^'^0 

r^r-'odrOfN'«N'<NrO-^OOONOOOOO--'000— ' 



ooooooooo — o 

<NNOOONr«-irN)(NOO(NOI^ 

— ^ P~- r~-- r*^ <N (N (N rn — O O 

fN — < -H 



O NO rn 
t^ r<-i ro 



r-l r^ i/^ t^ O — ' ■^ 
r<-i ON vO ■rf "/^ Tj- ON 



OOOO— 'OO— 'OOOO 



OOOOOOOOOO'/^O'^NOmOONOO'^NOr^ 
OOi/^fNOOOOONOO-^"^— ""^'^'^ONt^I^i/^i/^'^'^ — 



I^00rO(N(NfNm(N— 'OOOOOOO— 'OOO — 



OOOO'/^'/^''^'/^ 
t~~NO<NfNfNTf — m 

r^ 00 ON 



OOO^ONOONOOONirioONOt^rNlON 
TtO — ONrnmoorOoomrfTj-TfON 

<N(N(Nrn — — — 'OnOOOOO — OOOO 



OOOOOOOOOOOONOr^ONt^t^ONt^t^r-lON 
Tft^OOt^00<N<N>/^O — ONf'^r'^t^'^O— '■^"^'^ON 

— 'r-'o6rO(NfN<Nfn— 'OOOOOOOO — — OOOO 



> > 



t/2 



z > > ^ 



> ? 



H Z 



Q 









yjv^SWtUUJj^c/^ 



CQ J W 

X X D 



Q 



Q Q Uh H 

W LU -J J 






X5 

H 

C/5 



ha 

u 



(U 

o 



28 BREVIORA No. 493 

The Rio Azuela species also has white specks on the upper Up, 
but the dorsum is chestnut brown with faint white spots, adult 
females grow to 28-31 mm, and the color of the belly is light 
with darker spots. 

LITERATURE CITED 

Edwards, S. 1974. A phenetic analysis of the genus Colostethus (Anura, Den- 
drobatidae). Unpublished Ph.D. thesis, University of Kansas, Lawrence, Kan- 
sas, 417 pp. 

RivERO, J. A. 1988. Sobre las relaciones de las especies del genero Colostethus 
(Amphibia, Dendrobatidae). Memorias Sociedad Ciencias Naturales La Salle, 
48(129): 3-32. 

RivERO, J. A., AND M. A. Serna. 1988. La identificacion de los Colostethus 
(Amphibia, Dendrobatidae) de Colombia. Caribbean Journal of Science, 24(3- 
4): 137-154. 



B R E V,..I,,1)'R a 



^Nivk^ro 



ISlTy 

MmseiLim of Comparative Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 31 January 1992 Number 494 



ON SOME OVERLOOKED SPECIES OF THE GENUS 
LIOLAEMUS WIEGMANN (REPTILIA TROPIDURIDAE) 

FROM PERU 

R. F. Laurent' 

Abstract. Three new species of the genus Liolaemus, L. robustus, L. polystic- 
tus, and L. williamsi, from the upper western slopes of the Cordillera Central and 
of the eastern slopes of the Cordillera Occidental of Peru are described. They were 
previously confused with L. multiformis Cope, 1856, which is here synonymized 
with L. signifer (Dumeril and Bibron, 1841) and seems to be restricted to the 
northern part of the Altiplano. The status of other northern species oi Liolaemus 
is discussed. 

INTRODUCTION 

The bewildering diversity of the genus Liolaemus Wiegmann 
has been well documented by L. Miiller, W. Hellmich, R. Donoso- 
Barros, and J. M. Cei for the southern part of its range in Chile 
and Argentina, more or less south of the 30th parallel. In contrast, 
this diversity has been largely neglected in the northern part of 
its range, in northwestern Argentina, Bolivia, and Peru. Many of 
the forms that occur in this region have been inadequately de- 
scribed, and a number of names have been placed in synonymy 
with little or no documentation. Thus, the discovery of three 
undescribed species from the upper western slopes of the Cor- 



' Investigador Principal del CONICET, PRHERP-Fundacion Lillo, Miguel Lillo 
251, 4000 Tucuman, ARGENTINA. 



BREVIOHA 



No. 494 



Table 1 . Characters of the subgenera Liolaemus and Eulaemus. 



Liolaemus 
(395 3<5, 356 99) 



Eulaemus 
(313^3, 313 99) 



Preanal pores 



Nostrils 

Distance between upper 
border of subocular 
and lip compared 
with distance between 
nasal plates 

Upper labials 



Range 



0-7 

x= 2.19 

<5 in 91% of speci- 
mens, the series gen- 
erally shorter than the 
1st toe (12.4% of ex- 
ceptions) 

lateral 

inferior (3.37% of ex- 
ceptions) 



generally flat and long, 
the 4th below eye, 
with posterior border 
oblique 

Chile (> 50 taxa) 

southern and western 
Argentina ( 1 8 taxa) 

Bolivia (3 taxa) 

Peru (3 taxa) 



3-12 

X = 6.40 

>4 in 92% of speci- 
mens, the series gen- 
erally longer than the 
1st toe (2.29% of ex- 
ceptions) 

latero-dorsal 

superior (7.03% of ex- 
ceptions) 



generally high and 
short, 5th, 6th, or 7th 
below eye, with pos- 
terior border vertical 

northern Chile (6 taxa) 

southern and western 
Argentina (25 taxa) 

Bolivia (6 taxa) 

Peru (9 taxa) 



dillera Central and the upper eastern slopes of the Cordillera 
Occidental of Peru necessitates an evaluation of the status of other 
taxa from the region, before the new forms can be adequately 
diagnosed. 

Elsewhere (Laurent, 1983), I have pointed out that the great 
majority of Liolaemus species, including all that occur in the 
northern part of its range, can be referred to one or the other of 
two large groups: 1) a primarily Chilean group (subgenus Liolae- 
mus), and 2) a primarily Argentinian group (subgenus Eulaemus). 
Distinguishing characteristics of these two groups are provided 
in Table 1 . 

Two groups o^ Eulaemus may be recognized: 1) a fitzingeri 
group in which there is a patch of enlarged scales on the posterior 
surface of the thigh, and 2) a signifer group in which the patch of 
enlarged scales is lacking. The species allocated to the subgenus 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 3 

Liolaemus and to the two subgroups of Eulaemus are listed in 
the Appendix. 

Members of the Chilean group (subgenus Liolaemus) are few 
in the northern part of the range oi^ Liolaemus. Liolaemus tacnae 
(Shreve), originally described in the genus Stenocercus, is appar- 
ently a local species from the department of Tacna in southern 
Peru. Liolaemus alticolor Barbour and L. walkeri Shreve are names 
that have been applied to a large set of Andean populations from 
Peru southward to Catamarca Province, Argentina. The form 
walkeri was considered by Hellmich (1961) and Donoso-Barros 
(1966) to be a subspecies of alticolor. It is uncertain whether this 
form represents a valid species or subspecies. In fact, several taxa 
may be represented by specimens now referred to walkeri. 

The majority of northern Liolaemus, including the three new 
species described below, are members of the Argentinian group 
(subgenus Eulaemus). A number of Koslowsky's names have been 
revived (Laurent, 1982a) for members of this group, and new 
species have been recently described (Laurent, 1982a, 1984, 1985, 
1986), but there still are problems with certain other names in 
the group. 

Liolaemus ornatus Koslowsky, 1898, is an abundant species of 
the fitzingeri group that occurs from low to high altitudes from 
Catamarca Province, western Argentina, northward to the Lake 
Titicaca region in southern Peru and northern Bolivia. Pellegrin 
(1909) described Liolaemus pulcher and L. mocquardi from Ti- 
ahuanaco, Depto. de La Paz, Bolivia. Examination of the syntypes 
reveals that those of L. pulcher are males and those of L. moc- 
quardi are females of the same form. Peters and Donoso-Barros 
(1970) correctly placed L. pulcher in the synonymy of L. ornatus, 
thus L. mocquardi may now be added to that synonymy. 

Liolaemus simonsi Boulenger, 1902, based on specimens from 
Potosi, Challapata, and Uyuni, Bolivia, was considered to be a 
subspecies of multiformis by Burt and Burt (1931), an allocation 
followed by Peters and Donoso-Barros (1970). The syntypes of 
Liolaemus simonsi {BM 1902.5.29.74-79 [RR 1946.8.12.20-23], 
1902.5.29.85-87 [RR 1946.8.12.24-26]), kindly lent by Dr. C. 
MacCarthy, possess a patch of enlarged scales on the posterior 
aspect of the thigh, a fact not mentioned in the type description, 
but which excludes simonsii from the synonymy of multiformis. 



4 BREVIORA No. 494 

However, a comparison of the syntypes with Bohvian specimens 
of the widespread Liolaemus ornatus confirm the synonymy of 
simonsi with ornatus. 

The remaining northern species of Liolaemus, including the 
three new forms described below, are members of the signifer 
group of Eulaemus. Liolaemus dorbignyi Koslowsky, 1898, from 
Catamarca Province in western Argentina and L. jamesi Boulen- 
ger, 1891, from west of the Andes in Tarapaca Province, northern 
Chile, are large-scaled members of the signifer group, similar to 
one another in scalation and proportions, and m^ay represent vi- 
cariant forms on opposite sides of the Andes, an hypothesis to 
be investigated in a later paper. 

Liolaemus signifer (Dumeril and Bibron, 1841) is one of the 
two oldest names available for members of the Argentinian group, 
the other is L. fitzingerii. The type locality of L. signifer -was given 
as "Chile," but recently Cei, Lescure, and Ortiz (1980) have 
mapped the route taken by d'Orbigny, its collector, in Chile, Peru, 
and Bolivia, and restricted the type locality of signifer to the 
highlands of Peru and Bolivia. For the most part, the route taken 
by d'Orbigny passed through the range of the species that most 
subsequent authors refer to as Liolaemus multiformis Cope with 
a very short stretch passing through the range of Liolaemus an- 
nectens Boulenger, 1901, in Arequipa Province, Peru. There are 
significant statistical differences between these two forms, but the 
presence of a zone of intergradation indicates that a single species 
with two geographic races is involved. When the holotype of L. 
signifer (MNH Paris 6890) is compared with the two races, it falls 
always with the population of the intergrade zone or with mul- 
tiformis, never with annectens. It is on this basis that Liolaemus 
multiformis Cope, 1856, is here considered to be a synonym of 
L. signifer {Y)\xmhv\\ and Bibron, 1841). 

L. multiformis was based on a series of specimens (Acad. Nat. 
Sci. Phila. 13064-6, 13098, 13104, 13168-70) from Lake Titi- 
caca, Peru. A number of forms described subsequently have been 
placed in its synonymy, some correctly, but some apparently not. 
I have examined all of the relevant type material and consider 
the following synonymies to be correct: lenzi Boettger, 1891 (fide 
Burt and Burt, 1931), type locality "Bolivianische Ufer des Ti- 
ticaca-Sees"; tropidonotus Boulenger, 1901 (fide Burt and Burt, 



1992 OVERLOOKED LIOL.4EMUS SPECIES FROM PERU 5 

1931), type locality "Tirapata, E. Peru, 13,000 feet"; bolivianus 
Pellegrin, 1909 (fide Hellmich, 1962), type locality "Tiahuanaco, 
Depto. de La Paz, Bolivia"; variabilis crequii Pellegrin, 1909 (fide 
Hellmich, 1962), type locality "Tiahuanaco, Depto. de La Paz, 
Bolivia"; variabilis courtyi Pellegrin, 1909 (fide Hellmich, 1962), 
type locality "Tiahuanaco, Depto. de La Paz, Bolivia '; variabilis 
neveui Pellegrin, 1909 (fide Hellmich, 1962), type locality "Tia- 
huanaco, Depto. de La Paz, Bolivia." 

Since multiformis has been shown to be a synonym of signifer, 
all of the above forms are properly referred to the synonymy of 
the latter. In addition, L. pantherinus Pellegrin, 1909 (syntypes 
MNH Paris 05-344-05-345), for which no locality was given, also 
cannot be distinguished from L. signifer. 

Two forms that have been synonymized with Liolaemus mul- 
tiformis appear to be valid: L. annectens Boulenger, 1901 (syn- 
onymized by Hellmich, 1962), type locality "Caylloma and Sum- 
bay, 1 1,300 to 13,600 feet," and L. annectens orientalis Miiller, 
1923 (synonymized with multiformis simonsi Boulenger, 1902, 
by Burt and Burt, 1931), type locality "Oberer Pilcomayo, zwi- 
schen Tarija and S. Francisco, Bolivien." As pointed out above, 
Liolaemus annectens is probably a geographic race of Liolaemus 
signifer. Evidence for the validity of orientalis will be presented 
at another time. 

As indicated earlier, the signifer group of Eulaemus is defined 
by the absence of a patch of enlarged postfemoral scales, while 
the fitzingerii group is defined by their presence. A patch of en- 
larged postfemoral scales is unique within tropidurine iguanids 
and is almost certainly derived. However, if a patch of enlarged 
postfemoral scales was derived only once within Liolaemus, then 
the fitzingerii group is paraphyletic, because the patch is also found 
in species excluded from the group and placed in the subgenus 
Ortholaemus (Laurent, 1984), i.e., wiegmanni, cranwelli, multi- 
maculatus, scapularis, salinicola, occipitalis, lutzae. Since the 5/^- 
nifer group, at present, is defined only by the absence of these 
enlarged scales, this subgroup also may be paraphyletic. Although 
the question of their monophyly cannot be resolved at this time, 
the signifer and fitzingerii groups provide a useful means for the 
diagnoses and identification of the new forms described below. 



BREVIOR.4 



No. 494 



/^ iSi-^^?^^^^ 



;#^ 




^jZ/ 




-^y 




Plate I. Liolaemus robustus. sp. nov. Paratype male: MCZ 45811. Depto. 
Junin, Peru. 1. Dorsal view of head. 2. Ventral view of head. 3. Dorsal view. 4. 
Ventral view. 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 




Plate II. 1. Liolaemus polystictus, sp. nov. Holotype male: MCZ 45845. Side 
view of head. 2. Liolaemus polystictus, sp. nov. Paratype female: MCZ 45849. 
Dorsal view. 3. Liolaemus robustus. sp. nov. Paratype female: MCZ 4581 1. Side 
view of head. 



BREVIORA 



No. 494 



dSo, 



^ X ^8 



.rfo X X 
x^ 
X O X V • 



X O o 

O 



• s 



Figure 1 . Scatter-diagram of minimum width of frontal region (ordinates) and 
length of 5 dorsal scales (abscissa). Measurements in tenths of a millimeter, x = 
Liolaemus signifer, = L. polystictus, sp. nov. • = L. rohustus, sp. nov. S = Type 
of L. signifer. ® = Lectotype of L. multiformis. 



Liolaemus robustus, new species 
(PI. I, Figs. 1^; PI. II, Fig. 3) 

Holotype. One male (FMNH 'iAlAl/W) from Junin, Depto. 
Junin, collected by K. P. Schmidt. 

Paratypes. PERU: Depto. Junin: Same data as holotype: FMNH 
34242/1-23, 34247, 1 1 males, 4 females, 9 juveniles. Huayre, N 
of Junin: FMNH 34253, 2 males, 1 female, 4 juveniles. Huawhay 
(=Huayre ?): UMMZ 89484, 1 male. Ondores on Lake Junin: 
MCZ 157226, 1 male. "Dept. of Junin" only: MCZ 45809-12, 
16155-56, 1 male, 5 females, W. F. Walker, 14 April 1939. Depto. 
Lima: Yauricocha: MCZ 45830, 1 male. 

Diagnosis. A species of the Liolaemus signifer group, differing 
from L. signifer by the lower number of scales around the body 
(47-61, instead of 66-82), the frontal azygous, generally divided 
in two parts, anterior and posterior, instead of divided into many 
scales (at least 3, generally 5-8, and even more), by a narrower 
frontal zone, bigger head, more robust general proportions, and 
by its characteristic color pattern with black spots or dots. 



1992 OVERLOOKED L/OL.4£:M(y5 SPECIES FROM PERU 9 

Other differences are: 1) 44-59 scales (rather than 65-87) be- 
tween occiput and level of the front border of the thighs; 2) 1 2- 
1 9 dorsal scales (instead of 1 8-30) in head length; 3) 63-78 ventral 
scales (instead of 74-92) between postmentals and vent; 4) 49- 
70 lateral scales between the legs (instead of 65-89); 5) minimum 
width of frontal region 1 3-25% (rather than 25-49%) of the length 
of 5 dorsal scales (see Fig. 1). 

Description of the Holotype. Head length (from posterior edge 
of ear opening) (HL = 20.13 mm) 26.8% of snout-vent length 
(SVL = 77 mm). Two vertical antehumeral folds and a longitu- 
dinal oblique and sinuous fold on the side of the neck, bifurcated 
behind the ear on the right side. Scales on the upper side of head 
markedly convex, a count of 15 on the midline. Rostral scale 
about 2.5 times as wide (W = 4.0 mm) as high (H = 1.63 mm). 
Nasal triangular, separated from rostral, surrounded by 7 scales. 
Nostril round, in the posterior part of the nasal, a little nearer to 
the point of the snout (2.94 mm) than to the posterior extremity 
of the canthal (3.40 mm). Intemasals 2 anterior + 2 azygous + 
2 posterior. Ear opening somewhat oblique, nearly rectangular, 
surrounded by granular scales that are smaller behind than in 
front of the ear. Temporals convex, 7 between the postsubocular 
and the ear. Interparietal small, pentagonal, surrounded by 5 
scales, the anteriormost median. Frontal region occupied by 2 
azygous scales. Five supraoculars, 7 supraciliaries, the 5th below 
the 4th and 6th. Five scales between the rostral and frontal region. 
Semicircles simple, 4 scales between the frontal region and the 
supraciliaries. 

In prefrontal zone, between the posterior intemasals and the 
frontal region, 1 1 scales, 3 intercanthal scales. Across the snout, 
1 1 scales between the labials at the postnasal level, 1 1 also at the 
canthal level. Four scales between nasal and subocular. Subocular 
divided in two. Paralabials 8, 4 in contact with subocular. Su- 
pralabials 8. Infralabials 5-6, followed by 9-7 granules as far as 
the comer of the mouth. Mental fan-shaped, in contact with 4 
scales. 

Lateral scales of neck granular, 4 1 between ear opening and 
forelimb. About 30 scales between ear openings. Fifty-five scales 
around the body. Fifty-four between occiput and level of anterior 
border of thigh. Dorsal scales juxtaposed or imbricate on the sides, 



10 BREVIORA No. 494 

very faintly keeled or smooth, 14 in head length. Rank scales 
smaller, erect, granular at armpit and groin, about 61 between 
legs. Ventrals smooth and imbricate, 68 between mental and pre- 
anal pores. Caudal scales similar to body scales, 22 in 1 5th verticil. 
No patch of enlarged scales behind the thigh. Fourteen to fifteen 
infradigital lamellae beneath 4th finger, 1 9 beneath 4th toe. Tail 
(83 mm) 107.8% of snout-vent length. 

Color (in Alcohol). Above, olivaceous gray, with some scales 
blackish, these often clustered in small groups, which give a more 
or less punctate appearance. Belly whitish with grayish pigmen- 
tation on most scales; throat with gray dots, without definite 
pattern. 

Variation (see Table 2). Upper labials generally 8 (38 sides), 
sometimes 9(16 sides) or 7 (9 sides), rarely 10 (5 sides) or 11 (2 
sides), the first 5 to 6 without small scales below and inside. Lower 
labials generally 6 (46 sides), sometimes 5(14 sides), rarely 7 (6 
sides), 8 (3 sides), or 4 (1 side). Supraoculars generally 4 (34 sides) 
or 5 (25 sides), rarely 6 (7 sides) or 3 (8 sides). Supraciliaries 
usually 7, the 5th below the 4th and 6th, but 6 on 6 sides and 8 
on 9 sides. Temporals between the postsubocular and the ear 
most often 8(31 sides), not infrequently 7 (24 sides), sometimes 
9(11 sides), exceptionally 6 (4 sides). Plates between the rostral 
and the frontal normally 5 (43 sides), sometimes 6 (20 sides), 
rarely 7 (3 sides) or 4 (2 sides). Scales between the frontal and the 
supraciliaries 4 (46 sides), sometimes 3 (19 sides), rarely 2 (3 
sides) or 5 (2 sides). Scales around the interparietal usually 5(11 
cases) or 6 (14 cases), sometimes 7 (7 cases), rarely 8 (3 cases), 
symmetrical (14 cases) or irregular (21 cases). Scales in contact 
with the nasal generally 7 (35 sides), sometimes 6(19 sides) or 8 
(1 1 sides), rarely 9 (2 sides) or 5 (1 side), rarely adjacent to the 
rostral (2 sides). Paralabials usually 8 (28 sides), often 7 (22 sides), 
sometimes 9(13 sides), rarely 6 (4 sides) or 10 (1 side). Paralabials 
in contact with subocular generally 4 (4 1 sides), sometimes 5(18 
sides), rarely 3 (7 sides), exceptionally 2 or 6 ( 1 side each). Plates 
between the upper labials over the snout posterior to the nasals 
7 to 12 (mean = 9.97), at canthal level 9 to 14 (mean = 1 1.60). 
Usually 4 scales between nasal and subocular. Preanal pores in 
males 3 (2 cases), 4 (8 cases), 5 (6 cases), or 6 (3 cases). A single 
female has one vestigial pore. Almost always, the frontal is di- 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 11 

Table 2. Meristic characters of Liolaemus robustus. 

(7 (5<5, 7 99) 



(5(5 99 



Scales around midbody 48-61 (x = 53.05) 50-60 (Jc = 54.43) 
Dorsal scales between occiput 
and levels of anterior border 

of thighs 44-59 (x = 52.28) 51-56 (Jc = 53.14) 
Ventral scales between post- 
mentals and vent 63-74 (x = 68.59) 72-78 (.v = 74.14) 
Lateral scales between anterior 

and posterior lim^s 49-68 (.v = 58.40) 56-68 (Jc = 62.14) 

Scales in the 1 5th verticil of tail 1 8-22 {x = 20.28) 1 9-23 (x = 20.86) 

Gular scales between ears 29-35 (Jc = 32.04) 28-32 {x = 30) 

Hellmich's index 14-17 (Jc = 15.40) 13-15(.v=14) 

Lamellae under 4th finger 14-17 (Jc = 15.63) 14-16 {x = 15) 

Lamellae under 4th toe 18-21 (Jc = 19.78) 19-21 {x = 19.57) 



vided into two plates, one anterior and one posterior. In only 
three cases are there 3 plates with 2 anterior, one posterior. In 
one case there are 3 plates in a longitudinal series, in another the 
anterior plate is asymmetrically located on the left. In three spec- 
imens there is a single undivided frontal. 

The dorsal coloration does not appear very variable in pre- 
served material. The black dots or spots may be more or less 
distinct. They have a tendency to concentrate in two laterodorsal 
zones in some specimens. The ventral pigmentation may be al- 
most absent, uniformly distributed or scattered into ill-defined 
spots. Intact tails vary from 106 to 123% of snout-vent length in 
males (mean = 1 1 1.47), from 109 to 126% (mean = 1 17.73) in 
females. 

Size. Snout-vent length of the largest male (from Yauricocha) 
85 mm, of the largest female 82 mm. 

Geographic Variation. One specimen (MCZ 45830) from Yaur- 
icocha, Lima Department, is somewhat different from the other 
specimens from Junin Department. The belly and throat are black 
with white dots; the frontal is divided in three; there are 5 scales 
between the frontal and the supraciliaries, 22 lamellae beneath 
the 4th toe. The supraciliaries are only 5, the 3rd below the 2nd 
and 4th. The last may be an anomaly, since the formula 7 (5) is 



12 



BREVIORA 



No. 494 



Table 3 . Comparative variation of meristic and morphometric characters 
in llolaemvs robustus. sp. nov., l. polystictvs. sp. nov., l. willi amsi, sp. 

NOV., AND L. SIGNIFER (DUMERIL AND BiBRON). 











signifer 




robustus 


polystictus 


williamsi 


{= multiformis) 




(N= 15) 


(N= 17) 


(N= 15) 


(N = 32) 


Frontal divided 


2(1 + 1), 


1 to 5 (2 + 


1 to 5 (2 + 


3(1 + 2) to 


into plates 


rarely 3 


1 +2) 


1 + 2or 


9 (3 + 2 + 




(2 + 1) 




2 + 2 + 
1) 


2 + 2) 


Scales around mid- 


48-61 


57-70 


54-67 


66-82 


body 










Dorsal scales be- 


44-59 


55-70 


48-65 


65-87 


tween occiput 










and level of 










front borders of 










thighs 










Ventral scales be- 


63-78 


62-71 


67-78 


74-92 


tween postmen- 










tals and vent 










Lateral scales be- 


49-68 


54-76 


66-77 


65-89 


tween legs 










Hellmich's index 


12-17 


14-21 


17-22 


18-30 


Minimum width of 


13-23 


24-38 


18-29 


25^9 


frontal region as 


(A- = 16.9) 


(x = 31) 


(Jc = 23.8) 


(Jc = 35.72) 


% of length of 5 










dorsal scales 










Width of head in 


19.6-24.6 


19.70-23.7 




18.3-22.3 


% of snoul-vent 


(A- = 21.68) 


(Jc = 21.25) 




(Jc= 19.98) 


length 










Distance between 




85.2-110.1 




70.9-107.6 


posterior borders 




(A- = 95.49) 




(Jc = 84.9) 


of eyes in % of 










head height 










Length of 4th toe 




20.5-38.6 




31.6-50 


nail in % of 




(Jc = 30.02) 




(Jc = 40.81) 


width of 5 ven- 










tral scales ((53 










only) 










Distance between 




80-124 




62-99 


the pubic sym- 




(Jc = 98.84) 




(Jc = 78.65) 


physis and the 










vent as % of ear- 










eye distance (99 










only) 











1992 



OVERLOOKED LIOLAEMUS SPECIES FROM PERU 



13 



Table 3. Continued. 











signifcr 




robustus 


polvsl ictus 


williamsi 


(=mullijbrmis) 




(N= 15) 


(N= 17) 


(N= 15) 


(N = 32) 


Minimum distance 


90-142 


56-87 


65-139 




between nasals 


{x = 


(X = 70.8) 


(.V = 95.7) 




in % of mini- 


113.31) 








mum distance 










between supra- 










ocular scales 










Rostral height in % 


43-65 . 


50-74 


48-62 




of eye-lip dis- 


(.V = 50.57) 


(.X- = 60.62) 


(.V = 55.85) 




tance 










Length of 5 dorsal 






67-104 


29-71 


scales in % of 






(x = 79.4) 


(X = 49.58) 


ear-eye distance 










Eye-lip distance in 


47-70 




37-58 




% of subocular 


(Jc = 54.83) 




(X = 49.05) 




length 










Length of 1st finger 




51-83 


40-65 




(without claw) in 




(X = 65.95) 


(x= 54.14) 




% of length of 5 










dorsal scales 











the norm for the entire genus Liolaemus, but the other features 
might characterize a vaUd subspecies if confirmed for a majority 
of the specimens from the region. 

Relationships. All of these specimens had been identified as L. 
multiformis (Cope). However, they are clearly different from the 
syntypes of the species and series collected around Lake Titicaca. 
The most obvious differences are indicated in Table 3 and Fig- 
ure 1. 

There is also a large and clear-cut morphometric difference: the 
width of the frontal region at its narrowest point is less than 25% 
of the length of 5 dorsal scales (lowest value 13%) in robustus; in 
signifer the same measurement is more than 25% (highest value 
48.9%). The name of the species has been inspired by its robust 
appearance. While it is expected to be most significant in this 
respect, the width of head/snout-vent length ratio is not diagnostic 
at all: 18.31 to 24.57% (mean = 21.73%) in robustus versus 18.78 
to 22% (mean = 20.54%) in signifer (see Fig. 2). 



14 



BREVIORA 



No. 494 









O 








-o 








o 








o 


*^-^ 






"o 


* 








'■■■■■■■\..,.. 








- ® "-■•■■-■.. 








•> 








X \ 








\ " ""-■■■: d "'■■- 






o 


' ® • %, ■■. 






-o 
















\ ® ■•■:^ \ 
















"-•.. '•• "\ 








• "■••••. \« 0® " "'■■■■ 
■■■■■% ® \ 

■■■■' >.■■■: •■•.. \ 














'■•>\, ■■•-.. \ "O 

'^ -■■■-.. .->c--.. <. ® \. ^ 






o 


« ;^ " -v ® - ■■.. \ <i. 






CO 


'*■■: '■■■■■:. ®«--.. «© \A ^ 








•;••.. •-.. -o* -o « o. ••.:•.. .^ 








JV "^^ ° * • ® ''■■■ * ® 








'■•-_ 






■x ■■-...>■•... ..--'v « « i 


\ '■.. 






■■■-■'■<..„ \ o / ® 


's. 






•-•-ir-- .0®. 






o 


«« ■••. . 






-o 


■■•■■... ' ^ 


® ', 




r* 


'* 








\ ® 


-6 
® 








® 






'•■.. 


® 






'■•,_ 


® 




o 

"(0 




"■••■, ® 


Q 






"* 


■■,. 






'■■■".'■•  








K 








s.. 


X 










o 








o 






ft 


Jf 


O 4n b 


m 


r 


" 



c 


^ 


in 


^ 






c 


.§3 


6 


^ 


^ 


-J 


3 


(*. 


« 


O 


CO 




4> 


(U 


:s 


a 




H 






w 


II 


!« 




!« 


OT! 


'o 




1/5 


0+" 


X> 


II 


^ 


£ 


® 


So 

c 


T3 

C 








«o 


«-< 




c 




> 




1 






■S? 


3 

O 


^ 


c 

W5 


-J 


T3 


II 


C 


X 


"^ 


o' 


(U 


II 




c 


o 






'•5 


-a 


w 


c 


3 


a 






■a 


*o 


(U 


^ 


j= 


io 






t« 




o 


-C 




P 


j: 




^j 


^ 


T3 




1 


s 




5j 


t— 


a 


O 






]o 


S 


5 


« 




w. 


II 


00 


5 


• 


■5 




1 


u* 


u 


(U 


OJ 


^^ 


*-» 


«-> 




E 


o 




V) 


^7^ 








'e 


<^i 


w 


<u 


t— 


k- 


o 


3 


c/5 


oo 


.c 


U- 


c 




OJ 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 15 

L. robustus, as well as the two new species described below, is 
not compared here with other Eulaemus species. Other papers, 
which will remedy this lack, are in preparation. They include 
descriptions of other new species, one from Peru, one from north- 
em Chile, and 5 from northwestern Argentina, and the last of the 
series is intended to provide a key to all Eulaemus without en- 
larged femoral scales. 

The sexual dimorphism of L. robustus is not as conspicuous as 
that of L. signifer. The size difference is less marked, and the color 
pattern is about the same in both sexes, at least in alcohol. How- 
ever, the colors in life are probably brighter in males than in 
females. 

Liolaemus polystictus, new species 
(PI. Ill, Figs. 1-4; PI. II, Figs. 1-2) 

Holotype. One male (MCZ 45845) from Santa Inez (13°12'S, 
75°05'W), about 100 km S of Huancavelica, Depto. Huancavelica, 
Peru, W. F. Walker Sr., collected February 1939. 

Paratypes. PERU: Depto. Huancavelica: Same data as holotype: 
2 males MCZ 45844, 45846, 3 females MCZ 45847-49, 2 juve- 
niles MCZ 161157-58, 1 male, 1 female UMMZ 89482. Same 
locality and collector: MCZ 43782, collected 14 December 1936. 
Huancavelica: 5 males, 4 females, 2 juveniles FNHM 81453-63, 
no collector, no date. Six km SW Castrovirreyna, 3,650 m, KU 
163563, W. E. Duellman, collected 24 February 1975. 

Diagnosis. A species of the Liolaemus signifer group, differing 
from all other members of this group by the male color pattern, 
in which each dorsal scale is bicolor, pigmented at the base, clear 
behind, giving a striking appearance of fine punctation, and by 
having a greater sexual dimorphism in size. 

It can be distinguished from L. signifer by the following dif- 
ferences: 1) 62-75 ventral scales instead of 74-92 between post- 
mentals and vent; 2) 57-70 scales instead of 66-82 around mid- 
body; 3) 55-70 dorsal scales instead of 65-87 between occiput 
and level of front borders of thighs; 4) some morphometric dif- 
ferences only noticeable on scatter-diagrams because of allometry 
(Figs. 3-5). 

L. polystictus differs from L. robustus in the following charac- 
ters: 1) minimum width of frontal region 24-38% of length of 5 



16 



BREVIORA 



No. 494 










Plate III. Liolaemus polystictus. sp. nov. Holotype male: MCZ 45845. Huan- 
cavelica, Peru. 1. Dorsal view of head. 2. Ventral view of head. 3. Dorsal view. 
4. Ventral view. 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 17 



100 



90- 



80- 



70 



• .V 



X 



X XX 



X 
X ^ X 

• s 

• X 

X 



60 80 100 120 lAO 

Figure 3 . Scatter-diagram of distance between posterior eye borders (ordinates) 
and head height (abscissa). Measurements in tenths of a millimeter. • = Liolaemus 
polystictus, sp. nov. x = L. signifer. S = Type of L. signifer. 



dorsal scales instead of 1 3-23% (see Fig. 1); 2) minimum distance 
between nasals 56-87% of minimum distance between supraocu- 
lar scales instead of 90-142%; 3) 57-70 scales around midbody 
instead of 47-61; 4) 55-70 dorsal scales instead of 44-59 between 
occiput and level of front borders of thighs. 



30 



25- 



20- 



15 



X 



X 



X 



X X 



X 

X 
X 



45 50 55 60 65 70 75 80 85 

Figure 4. Scatter-diagram of length of claw of 4th toe (ordinates) and width 
of 5 ventral scales (abscissa). Measurements in tenths of a millimeter, x = Lio- 
laemus signifer. • = L. polystictus, sp. nov. Males only. 



18 BREVI0R.4 No. 494 



80- 



30 



X 



• IT" 



^0 50 60 70 80 90 100 

Figure 5. Scatter-diagram of distance between armpit and groin (ordinates) 
and ear-eye distance (abscissa). Measurements in tenths of a millimeter, x = 
Liolaemus signifer. • = L. polystictus, sp. nov. Females only. 



Description of the Holotype. Head length (from posterior rim 
of ear opening) (HL = 22 mm) 28.9% of snout-vent length (SVL 
= 76 mm). Vertical lateral folds of the neck overshadowed by a 
horizontal fold, which is sinuous and bifurcated just behind the 
ear. Scales on upper surface of head strongly convex, 1 6 on the 
midline. Temporals convex, some slightly keeled, keels higher 
behind than in front, 8-9 between the postsubocular and the ear. 
Frontal region occupied by two azygous scales, their front and 
rear borders oblique, five supraoculars. Rostral plate 2.76 times 
wider (WR — 4 mm) than high (HR = 1 .45 mm). Nasal triangular, 
separated from rostral, surrounded by 8 scales, nostril in the 
posterior part of nasal, nearer to point of snout (NS = 2.95 mm) 
than to hind border of canthal scale (NC = 3.7 mm). Intemasals: 
4 anterior + 4 posterior, all irregular. Ear opening oval and oblique, 
surrounded by granular scales, smaller behind than in front of 
the ear. Eight to seven supraciliaries, the 5th lateral to the others 
on the left, the 4th and 5th on the right. Six scales between the 
rostral and the frontal region. Five scales between the frontal 
region and the supraciliaries. 

The prefrontal zone between the posterior intemasals and the 
frontal region contains 13 scales, rather irregularly arranged. Five 
intercanthal scales. Across the snout between the supralabials 1 2 
scales behind the postnasal level, 14 at the canthal level. Four 
scales between the nasal and the subocular. Paralabials 9-7, 5-3 
in contact with subocular. 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 19 

Table 4. Meristic characters in Liolaemvs polystictvs. sp. nov. (9 66, 8 99, 

juveniles not included). 

66 99 

Scales around midbody 57-70 (.v = 6 1 .34) 63-68 (x = 64.87) 

Dorsal scales between occiput 
and level of anterior border 

of thigh 55-63 (Jc = 59.22) 57-70 {x = 62.62) 
Ventral scales between post- 
mentals and vent 62-67 (jc = 64.67) 64-75 (Jc = 68.12) 
Lateral scales between legs 54-70 (Jc = 62.78) 53-76 (Jc = 63.75) 
Scales in 1 5th verticil of tail 1 7-20 (Jc = 1 9.40) 1 9-22 (x = 20.87) 
Gular scales between ears 27-32 (x = 29.56) 28-32 (Jc = 29.88) 
Hellmich's index 16-21 (.v = 17.78) 14-21 (x = 18.38) 
Lamellae under 4th finger 1 7-1 8 (x = 1 7.67) 1 5-1 9 (x = 16.25) 
Lamellae under 4th toe 20-24 (Jc = 2 1 .56) 1 9-22 (Jc = 20.5) 



Nine supralabials on the left side, 8 on the right side. Seven to 
six infralabials, followed by 6-7 granules as far as the comer of 
the mouth. Mental fan-shaped, in contact with 4 scales. 

Lateral scales of the neck granular, 41 between ear opening and 
front leg, 32 scales between ear openings, 63 scales around the 
body, 6 1 between occiput and level of front border of thighs. 
Dorsal scales imbricate and keeled. Lateral scales smaller, erect 
and granular, smaller still at armpit and groin, 58 between legs. 
Ventral scales smooth and imbricate, 63 between mental and 
preanal pores. Five preanal pores. Caudal scales similar to body 
scales, 20 in 1 5th verticil of tail. Seventeen subdigital lamellae 
beneath 4th finger, 22 beneath 4th toe. 

Color (in Alcohol). Above, blackish with a very dense and fine 
punctation resulting from the fact that each dorsal scale is pig- 
mented at the base, unpigmented at the tip. Below, the belly is 
light, with black pigmentation on the borders of the scales, which 
gives a reticulate appearance especially marked on the throat. On 
the lateral scales, the pigmented and unpigmented areas are about 
equal, so that the general effect is checkered. Upper surface of 
head blackish. 

The color in life is unknown, but it is surmised that the un- 
pigmented parts are actually vividly colored (white, yellow, or- 
ange, red, green, or blue). 



20 BREVIORA No. 494 

Variation (see Table 4). Upper labials generally 9 (15 sides), 
sometimes 8 (6 sides) or 10 (5 sides), rarely 6-7 (1 side each) or 
11 (4 sides), the first 5 to 8 without small scales below and inside. 
Lower labials generally 5(14 sides) or 6 (12 sides), sometimes 7 
(5 sides), rarely 4 (1 side). Supraoculars generally 5 (15 sides), 
sometimes 4 (8 sides), rarely 3 (3 sides), 6 (5 sides) or 7 (1 side). 
In one individual, only one plate on the left and two on the right 
are enlarged enough to be termed supraoculars. Supraciliaries as 
usual 7, the 5th below the 4th and the 6th, at least in females, 
except in one case where the number is 6, but anomalies are 
surprisingly common in males, where 6 is actually the most fre- 
quent number (9 sides), while 7 is less common (5 sides), 5 (2 
sides) and 8 (1 side) are rare; the lower supraciliary, which is 
generally the 5th, is the 4th on 4 sides (3 in males). Three males 
each show another anomaly on one side: 1) the 5th supraciliary 
is in front of the 6th, not below; 2) the 3rd supraciliary is also 
below the 2nd and the 4th; 3) the 4th and the 5th supraciliaries 
are both overhung by the 3rd and the 6th. Temporals between 
the postsubocular and the ear usually 8(10 sides) or 9 (9 sides), 
often 7 (8 sides, 6 sides in females), rarely 10 (3 sides, all in 
females) or 6 (3 sides), exceptionally 5 (1 side in a male). Plates 
between the rostral and the frontal most often 6 (16 sides), less 
often 5(12 sides), sometimes 7 (6 sides). Scales between the frontal 
and the supraciliaries 5, sometimes 4 (9 sides) mostly in females 
(8 sides), exceptionally 3 (1 side). Scales around the interparietal 
usually symmetrical (irregular in only 4 specimens), usually 6 (9 
specimens), sometimes 7 (5 cases) or 8 (3 cases). Scales around 
the nasal generally 7 (20 sides), sometimes 6 (7 sides, 2 in males) 
or 8 (6 sides, 5 in females), rarely 9 (1 side), rarely including the 
rostral (2 sides). Paralabials usually 9(18 sides), sometimes 8(13 
sides), rarely 7 (3 sides). Paralabials in contact with subocular 
generally 4(15 sides), sometimes 5(10 sides) or 3 (9 sides). Plates 
between the upper labials around the snout behind nasals 7-13 
(mean = 10.6), 10-15 at the canthal level (mean = 12.25). Scales 
between nasal and subocular usually 4, rarely 5 (7 sides) or 6 (2 
sides); in this last case, the subocular is actually divided, which 
is a presumed return to a primitive condition. Preanal pores 5 (6 
specimens), 4 (2 specimens), or 3 ( 1 specimen). In frontal, always 
at least one azygous element, most often 2 (7 cases), only single 



1992 



OVERLOOKED LIOLAEMUS SPECIES FROM PERU 



21 



30 



20 



10 



15 



"20" 



T" 

25 



30 



Figure 6. Scatter-diagram of minimum distance between nasals (ordinates) 
and minimum distance between supraoculars (abscissa). Measurements in tenths 
of a millimeter, x = Liolaemus robustus, sp. nov. • = L. polystictus, sp. nov. 



(frontal undivided) in one case, 3 in a longitudinal line (1 case), 
3 with 2 in front (2 cases), 3 with 2 behind (2 cases), 3 with 2 
laterals (1 case), 4 with 2 in front and 2 longitudinally behind (1 
case), 5 with 2 in front, 2 behind and 1 central (2 cases). 

The color pattern of males hardly varies at all; the fine and 
regular dorsal punctation and the ventral reticulation are always 



20- 



15 - 



10 



X 



• \ 



16 



26 



31 



Figure 7. Scatter-diagram of rostral height (ordinates) and distance between 
subocular upper border and mouth (abscissa). Measurements in tenths of a mil- 
limeter. X = Liolaemus robustus. % = L. polystictus, sp. nov. 



22 BREVIORA No. 494 

present but are obscured in the specimens belonging to the Field 
Museum of Natural History, presumably because of too long an 
exposure to formalin. The females and the young are gray to 
reddish brown (in alcohol) with the usual two latero-dorsal series 
of blackish blotches. 

Habitat. The only information we have is that of W. E. Duell- 
man, who found the juvenile paratype under a rock in a grassy 
river valley. 

Size. Snout-vent length of the largest male 86 mm (tail 102 
mm), of the largest female 69 mm (tail 77 mm). 

Systematic Position. Both L. robustus and L. polystictus are 
vicariants of L. signifer. They are considered different species 
because they are well-differentiated and separated by mountain- 
ous barriers that must have interrupted any gene flow over a long 
period of time. 

The most obvious differences between L. polystictus and L. 
signifer are the color pattern, and the great disparity in size be- 
tween the sexes in the former. 

There are other characters. The frontal in L. polystictus is more 
often divided than in L. robustus but less than in L. signifer. A 
similar intermediacy is apparent in scale numbers (see Table 3) 
except for the ventral longitudinal counts that are even lower, 
although very slightly, in polystictus than in robustus. 

Morphometrically, L. polystictus is more similar to signifer than 
to robustus in some respects but more similar to robustus in others 
(width of head). 

The pertinent data are indicated in Table 3, but the ratios 
somewhat understate the differences because of allometric dis- 
tortions. The scatter-diagrams give a better idea of the character 
differences (see Figs. 3-7). 

Liolaemus williamsi, new species 
(PI. IV, Figs. 1-4) 

Holotype. One male (LACM 9323), Pampas Galeras, between 
Nazca and Puquio, Depto. Ayacucho, Peru, x-1965, coll. S. W. 

Taft. 

Paratypes. Four males, 3 females, 2 juveniles (LACM 9319- 
22, 9324-28), same data. Four males, 3 females, 3 juveniles 
(LACM 9329-38), Lucanas, Pampas Galeras, 96 km from Nazca, 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 



23 




Plate IV. Liolaemus williamsi, sp. nov. Holotype male: LACM 9323. Pampas 
Galeras, Depto. Ayacucho, Peru. 1. Dorsal view of head. 2. Ventral view of head. 
3. Dorsal view. 4. Ventral view. 



24 BREVIORA No. 494 



37- 










© 

© 


® 










®® 












.v^' ■•■■ 




.®...-- 










<f > ® ......,,-l 


©.-•••■V" 












..■■•© .••■••■■■ ® ^ 


..••"■' ^ ': 




27- 


.•■•y 




■x 


X 

X ...-■•■ 
.©••■■■■ ®.x-"' X 


X ;■' 
X 




17  

2 


5 


30 




35 40 45 50 


55 


60 



Figure 8. Scatter-diagram of length of the first finger, without claw (ordinates) 
and length of 5 dorsal scales (abscissa). Measurements in tenths of a millimeter. 
X = L. williamsi. ® = L. polystictus. 



Depto. Ayacucho, Peru, iv-vii-1963, coll. S. W. Taft. One female 
(LACM 35867), Pampas Galeras, 300 miles south of Lima, Dep- 
to. Ayacucho, Peru, iii-1966, coll. S. W. Taft. One male (MCZ 
100435), Lucanas, Pampas Galeras, Depto. Ayacucho, Peru. One 
male, 2 females, 4 juveniles (MCZ 145335-41), Reserva Nacional 
de Pampas Galeras, 90 km from Nazca, Depto. Ayacucho, 21, 
iv-1974, coll. R. A. Mittermeier. One female (MCZ 157223), 
Pampas Galeras, Lucanas (exch. Mus. Javier Prado). 

Diagnosis. A middle-sized and somewhat melanistic species of 
the L. signifer group, differing from L. signifer by its larger and 
less numerous scales and the presence of preanal pores in some 
females. 

It differs from L. signifer in the following features: 1) 54-67 
scales around midbody instead of 66-82; 2) 48-65 dorsal scales 
instead of 65-87 between occiput and level of front borders of 
thighs; 3) 67-78 ventral scales instead of 74-92 between post- 
mentals and vent; 4) minimum width of frontal region 18-29% 
of length of 5 dorsal scales instead of 25-49%; 5) length of 5 dorsal 
scales 67-104% of ear-eye distance instead of 29-71%. 

It can be distinguished from L. robustus by the following dif- 
ferences, apart from color pattern: 1) 66-77 lateral scales between 



1992 



OVERLOOKED LIOLAEMUS SPECIES FROM PERU 



25 



w- 



^ 



C/7 ;■ 
O ." 



• • 



\. 



v^V 



.\N^ 



c^^" 



— r- 

15 



— I— 
20 



— T— 
25 



10 



Figure 9. Scatter-diagram of minimum distance between nasals (ordinates) 
and minimum distance between supraoculars (abscissa). Measurements in tenths 
of a millimeter, x = Liolaemus polystictus, sp. nov. • = L. williamsi. The difference 
is essentially allometric. 



legs instead of 49-70; 2) 17-22 dorsal scales instead of 12-19 in 
head length. 

It differs from L. polystictus by the larger number of ventral 
scales between postmentals and vent (67-78 instead of 62-71) 
and several morphometric characters mostly noticeable only on 
scatter-diagrams (Figs. 8, 9). 

Description of the Holotype. Head length (HL = 19.2 mm) 
25.6% of snout-vent length (SVL = 75 mm). Vertical lateral folds 
of the neck overshadowed by a horizontal and sinuous fold bi- 
furcated in front. Scales on upper face of head convex, 1 5 on the 
midline. Rostral plate 2.87 times wider (WR = 3.7 mm) than 
high (HR = 1 .29 mm). Nasal more or less trapezoidal, surrounded 
by 6 scales. Nostril round, in the posterior part of nasal, nearer 
to point of snout (NS = 2.55 mm) than to hind border of canthal 
scale (NC = 2.89 mm). Two anterior and 4 posterior intemasals. 



26 BREVIORA No. 494 

Table 5. Meristic characters in Liolaemus williamsi, sp. nov. (11 $S, 11 52, 

JUVENILES NOT INCLUDED). 

(5(5 99 

Scales around midbody 54-64 {x = 59) 57-67 (Jc = 60.09) 

Dorsal scales 48-62 {x = 54.64) 49-65 {x = 57.82) 

Ventral scales 67-77 (.v = 70.82) 71-78 {x = 74.55) 

Lateral scales 66-75 {x = 70.36) 66-77 {x = 72.36) 

Scales of tail's 15th verticil 17-21 (.v = 19.36) 18-22 (x = 19.36) 



Ear openings oval and very slightly oblique, surrounded by gran- 
ular scales smaller behind than in front. Temporals convex, 
smooth, and slightly imbricate, 6-7 between the postsubocular 
and the ear. Frontal region occupied by 2 azygous scales. Six 
supraoculars, 7 or 6 supraciliaries, the 5th or 4th below the 4th 
and 6th or the 3rd and 5th. Five to six scales between the rostral 
and the frontal. Five scales between the frontal and the superci- 
liaries. 

In prefrontal zone, between the posterior intemasals and the 
anterior frontal, 1 2 scales, more or less symmetrically arranged. 
Four intercanthal scales. Across the snout, between left and right 
labials, 1 1 scales just behind the postnasal level, 1 1 also at the 
canthal level. Four or 5 scales between nasal and subocular. Para- 
labials 8-9, 5 in contact with the subocular. 

Supralabials 8-9. Infralabials 5, followed by 6-7 granules as far 
as the comer of the mouth. Mental fan-shaped, in contact with 
4 scales. 

Lateral scales of neck granular, 46 between ear and fore limb. 
Thirty-three scales between ear openings. Fifty-nine scales around 
body. Fifty between occiput and level of insertion of thighs. 

Dorsal scales imbricate and keeled. Lateral scales smaller, im- 
bricate and feebly keeled, still smaller at axilla and groin. Seventy- 
two scales between legs. Ventral scales smooth and imbricate, 68 
between mental and preanal pores. Four preanal pores. Caudal 
scales similar to body scales, 1 7 in the 1 5th verticil. Sixteen sub- 
digital lamellae under 4th finger, 20 beneath 4th toe. 

Color (in Alcohol). Blackish with numerous tiny gray dots (one 
on each scale). Below, darkish gray, with light areas on cloacal 
region and thighs. 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 27 



60 



40- 



20 



<.'.=!-■'•■•■■■ 


X '■. 


-^w.'tec. 






X 


" ..--■ % 


• t 




\ 


X 
X 


4 '^•» 


• •• ll • 




•, 


/ 


• 


•••• .• 


• 




■'.' • 


i. 


. •• •• 






K 


• 






• 


• - 









40 50 60 70 80 90 100 

Figure 10. Scatter-diagram of length of 5 dorsal scales (ordinates) and ear-eye 
distance (abscissa). Measurements in tenths of a millimeter, x = Liolaemus wil- 
liamsi. % = L. signifer. S = Type of L. signifer. 



Color (in Life). Unknown. 

Variation (see Table 5). Upper labials generally 8(18 sides) or 
9 (20 sides), sometimes 10 (9 sides), rarely 7 (2 sides), 11(1 side) 
or 12 (1 side), the first 4 to 8 without small scales below and 
inside. Lower labials generally 6 (26 sides) or 5 (2 1 sides), rarely 

7 (4 sides) or 4 (1 side). Supraoculars generally 5 (30 sides), some- 
times 6 (15 sides) or 4 (9 sides), rarely 3 (1 side) or 7 (1 side). 
Supraciliaries as usual 7, the 5th below the 4th and 6th, but 
anomalies are frequent: 6 (5) (5 sides), 6 (4) (1 side), 8 (6) (3 sides), 

8 (5) (1 side), 9 (6) (1 side), or 8 (4 and 6) (1 side), 6 (0) (1 side). 
Temporals between postsubocular and ear generally 7 (32 sides), 

sometimes 8 (9 sides) or 6 (7 sides), rarely 9 (4 sides). Scales 
between rostral and frontal most often 5 (27 sides), less often 6 
(20 sides), rarely 7 (3 sides), 4 (1 side) or 8 (1 side). Scales between 



28 



BREVIORA 



No. 494 



35 



25 



robustus 






• • : 






■•;;VUiamsi 



45 



Figure 11. Scatter-diagram of minimum distance between upper border of 
subocular and lip (ordinates) and supraocular length (abscissa). Measurements in 
tenths of a millimeter. • = L. robustus. x = L. williamsi. 



20 



10- 



3fO 



" «' (S)^4*» 






n 



• t 



/ 



40 



Figure 1 2. Scatter-diagram of minimum width of the frontal region (ordinates) 
and length of 5 dorsal scales (abscissa). Measurements in tenths of a millimeter. 
X = Liolaemus williamsi, sp. nov. • = L. robustus, sp. nov. 



1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 29 

frontal and supraciliaries normally 4 (29 sides), often 5(19 sides), 
rarely 3 (5 sides), exceptionally 2 (1 side). Scales around the in- 
terparietal often asymmetrical (13 cases), most often 7 (13 cases), 
sometimes 8 (7 cases), rarely 9 (3 cases) or 6 (3 cases), excep- 
tionally 10 (1 case). Scales around nasal normally 6 (33 sides), 
sometimes 7(13 sides), rarely 5 (2 sides) or 8 (3 sides). Paralabials 
most often 8 (23 sides), often 7 (13 sides) or 9 (14 sides), rarely 
6 (1 side) or 10 (1 side). Paralabials in contact with subocular 
generally 4 (22 sides), often 3(13 sides) or 5 (1 1 sides), rarely 2 
(3 sides) or 6 (1 side). Scales between supralabials 8-14 at the 
postnasal level (mean = 10.75), 11-15 (mean = 12.53) at the 
canthal level. Preanal pores, 5 in 5 males, 4 in 4 males, 6 in 2 
males, present also but poorly developed in some females. Frontal 
not divided in 1 case, divided into 2 median scales in 12 cases, 
into 4 symmetric scales in 1 case, into 5 scales symmetrically 
arranged (2 anterior, 1 azygous, 2 posterior) in 2 cases, into 5 
scales with the azygous posterior in 1 case, into 3 scales with the 
azygous one posterior in 2 cases, into 3 scales with the azygous 
one anterior in 2 cases. 

Tail, when not broken or regenerated, longer than snout-vent 
length, 127 to 138.5% (mean = 131.45%). 

Size. Largest male 76 mm SVL, largest female 77 mm SVL. 

Color (in Alcohol). In males, as shown by specimens other than 
the holotype, the color is generally blackish, sometimes with light 
markings that can fuse and form more or less oblique and dis- 
continuous transverse lines. In one specimen, the color is lighter 
and allows about 14 pairs of blackish spots to be seen. The belly 
can be light with a few black dots, but is more often dark with a 
few light areas or spots. In females and juveniles the two longi- 
tudinal series of dark blotches on a light background are distinct. 
The belly is light with dark markings. 

L. williamsi differs from L. signifer, L. robustus, and L. poly- 
stictus by its smaller size and dark coloration (possibly artificial 
in some specimens), the presence of preanal pores in some fe- 
males, the frequencies of its meristic characters, and some of its 
proportions (see Table 3 and Figs. 8-12). As described above, the 
species represents the differentiation of populations of relatively 
smaller size in the upper parts of Pacific Andean valleys isolated 
from larger-size populations of the Altiplano. 



30 BREVIORA No. 494 

LITERATURE CITED 

Burt, C. E., AND M. D. Burt. 1931. South American lizards in the collection 
of the American Museum of Natural History. Bulletin of the American Mu- 
seum of Natural History, 61: 227-395. 

Cei, J. M., J. Lescure, and J. C. Ortiz. 1980. Redecouverte de Tholotype de 
Proctotretus signifer Dumeril et Bibron, 1837 (Reptilia, Iguanidae). Bulletin 
du Museum national d'histoire naturelle, 4'' Serie, 2A(3): 919-925. 

Donoso-Barros, R. 1966. Reptiles de Chile. Santiago: Ediciones de la Univ- 
ersidad de Chile. 458 + cxlvi pp. 

Hellmich, W. 1961. Bemerkungen zur geographischen Variabilitat von Liolae- 
mus alticolor Barbour (Iguanidae). Opuscula Zoologica, 58: 1-6. 

. 1962. Bemerkungen zur individuellen Variabilitat von Liolaemus mul- 
tiformis (Cope) (Iguan.). Opuscula Zoologica, 67: 1-10. 

Laurent, R. F. 1 982a. Description de trois especes nouvelles du genre Liolaemus 
(Sauria, Iguanidae). Spixiana, 5(2): 139-147. 

. 1982b. Las especies y "variedades" de Liolaemus descritas por J. Kos- 

lowsky (Sauria Iguanidae). Neotropica, 28(80): 87-96. 

. 1983. Contribucion al conocimiento de la estructura taxonomica del 

genero Liolaemus Wiegmann. Boletin, Asociacion Herpetologica Argentina, 
1(3): 16-18. 

. 1984. Tres especies nuevas del genero Liolaemus (Reptilia, Iguanidae). 

Acta Zoologica Lilloana, 37(2): 273-294. 

. 1985. Description de Liolaemus huacahuasicus spec. nov. (Iguanidae, 

Reptilia) des Cumbres Calchaquies, Province de Tucuman, Argentine. Spix- 
iana, 8(3): 241-249. 

. 1986. Descripciones de nuevos Iguanidae del genero Liolaemus. Acta 



Zoologica Lilloana, 38(2): 87-105. 

Pellegrin, J. 1 909. Description de cinq lezards nouveaux des hauts-plateaux du 
Perou et de la Bolivie, appartenant au genre Liolaemus. Bulletin du Museum 
d'histoire naturelle, 6: 324-329. 

Peters, J. A., and R. Donoso-Barros. 1970. Catalogue of the Neotropical 
Squamata: Part II. Lizards and amphisbaenians. United States National Mu- 
seum Bulletin, 297: 1-293. 



APPENDIX 



Subgenus Liolaemus Subgenus Eulaemus 



signifer group 



L. alticolor Barbour 1 909 L. andimis 

L. atacamensis Miiller & Hellmich a. andinus Koslowsky 1895 

1933 a. poecilochromus Laurent 1986 

L. austromendocinus Cei 1974 L. aymararum Veloso, Sallaberry, 



1992 



OVERLOOKED LIOL.4EMUS SPECIES FROM PERU 



31 



APPENDIX. Continued. 



Subgenus Liolaemus 



Subgenus Eulaemus 



signifer group 



L. bellii 
b. bellii Gray 1845 
b. araucaniensis Miiller & Hellmich 

1932 
b. moradoensis We\\m\c\i 1950 

b. neuquensis Miiller & Hellmich 
1938 

L. bibronii Bell 1 843 

L. bisignatus Philippi 1860 

L. bitaeniatus Laurent 1984 

L. buergeri 'Werner 1907 

L. capillitas Hulse 1979 

L. ceii Donoso-Barros 1971 

L. chiliensis Lesson 1828 

L. coeruleus Cei & Ortiz 1983 

L. constanzae Donoso-Barros 1 96 1 

L. copiapensis Miiller & Hellmich 

1933 
L. curicensis Miiller & Hellmich 1938 
L. curis Nunez & Labra 1985 
L. cyanogaster 

c. cyanogaster Dumeril & Bibron 
1837 

c. brattstroemi T>or\o%o-Barros 1961 
L. donosoi Ortiz 1975 
L. duellmaniCei 1978 
L. elongatus 
e. elongatus Yjo^Xov^sky 1896 
e. petrophilus Donoso-Barros & Cei 
1971 
L. exploratorum Cei & Williams 1984 
L. fitzgeraldi Bo\x\enger 1899 
L. /w5a/5 Boulenger 1885 
L. gracilis BeW 1843 
L. gravenhorstii Gray 1845 
L. hellmichi Doixoso-Barros 1974 
L. hernani Sallaberry, Nunez & 

Yaiiez 1982 
L. kriegi Muller & Hellmich 1939 
L. kuhlmanni Miiller & Hellmich 
1932 



Navarro, Iturra, Valencia, Penna 
& Diaz 1982 
L. disjunctiis Laurent 1990 
L. dorbignyi Koslowsky 1898 
L. eleodori Cei, Etheridge & Videla 

1985 
L. fabiani Yaiiez & Nunez 1983 
L.famatinae Cei 1980 
L.fittkaui Laurent 1986 
L. forsteri Laurent 1982 
L. griseus Laurent 1984 
L. huacahuasicus Laurerw 1985 
L. islugensis Ortiz & Marquet 1987 
L. jamesi Boulenger 1891 
L. montaniis Koslowsky 1898 
L. multicolor Koslov/sky 1898 
L. nigriceps Philippi 1860 
L. orientalis Miiller 1923 
L. ortizi Laurent 1982 
L. polystictus Laurent 1 99 1 
L. puritamensis Nunez & Fox 1989 
L. robust us Laurerw 1991 
L. ruibali Donoso-Barros 1961 
L. schmidti Marx 1 960 
L. signifer 
s. signifer Dumeril & Bibron 1837 
5. annectens Boulenger 1901 
L. williamsi Laurent 1 99 1 



ftzingerii group 



L. boulengeri Koslowsky 1898 
L. chacoensis Shreve 1 948 
L. darwinii BeW 1843 
L. donosobarrosi Cei 1974 
L. fitzingerii 

f fitzingerii Dumeril & Bibron 1837 

/ canqueli Cei 1973 
L. irregularis Laurent 1986 
L. melanops 

m. melanops BurmeisXer, 1888 



32 



BREVIORA 



No. 494 



APPENDIX. Continued. 



Subgenus Liolaemus 



Subgenus Eulaemus 



L. le>n?iiscatiis GravenhoTsl 1838 
L. leopardinus 
I. leopardinus Miiller & Hellmich 

1932 
/. ramonensis Miiller & Hellmich 

1932 
/. valdesianus HeWmich 1950 
L. lorenzmuelleri HtWmich 1950 
L. monticola 
m. monticola Miiller & Hellmich 

1932 
m. chillanensis Miiller & Hellmich 

1932 
m. villaricensis Miiller & Hellmich 
1932 
L. nigromaculatus Wiegmann 1834 
L. nigroviridis 
n. nigroviridis Miiller & Hellmich 

1932 
n. c amp anae HtWmich 1950 
n. minor Miiller & Hellmich 1932 
n. nigroroseus Donoso-Barros 1966 
L. nitidus Vsl'iegmann 1834 
L. paulinae Donoso-Barros 1961 
L. pictus 
p. pictus Dumeril & Bibron 1837 
p. argentinus Miiller & Hellmich 

1939 
p. chiloeensis Miiller & Hellmich 

1939 
p. mayo/- Boulenger 1885 
p. talcanensis Urbana & Zungia 
1977 
L. platei Werner 1898 
L. robertmertensi Hellmich 1964 
L. sanjuanensis Cei 1982 
L. schroederi Miiller & Hellmich 1938 
L. tacnae Shreve 1941 
L. tenuis 
t. tenuis Dumeril & Bibron 1837 
/. punctatissimus Miiller & Hell- 
mich 1933 



m. xanthoviridis Cei & Scolaro 

1980 
L. ornatus Koslowsky 1898 
L. rot h i Koslowsky 1898 
L. uspallatensis Macola & Castro 

1982 



1992 OVERLOOKED LIOLAEMUS SPECIES FHOM PERU 33 

APPENDIX. Continued. 

Subgenus Liolaemus Subgenus Eulaemus 

L. vahegatus Laurent 1984 
L. velosoi Ortiz 1987 
L. walkeri Shreve 1938 
L. zapallarensis 

z. zapallarensis Miiller & Hellmich 
1933 

z. ater Muller & Hellmich 1933 

z. sieversi Donoso-Barros 1954 



B R E V I R"A 

_, FFR ■•! in.n.n 

Museum of Comparative Zoology 

us ISSN 0006-9698 HARVARH 

Cambridge, Mass. 31 January 1992 j^umber 495*^ 



NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. 

VII. ANOLIS LAMARI, A NEW ANOLE FROM THE 

CORDILLERA ORIENTAL OF COLOMBIA, WITH A 

DISCUSSION OF TIGRINUS AND PUNCTATUS 

SPECIES GROUP BOUNDARIES 

Ernest E. Williams • 

Abstract. A new species, Anolis lamari, is described from the eastern slopes 
of the Cordillera Oriental of Colombia. It is compared with the species previously 
assigned to the tigrinus group. The characters separating the tigrinus and punctatus 
species groups are reassessed. A plea is made for the temporary use of informal 
groupings— recognized as such— whenever sufficient grounds for the recognition 
of formal taxa do not exist. 

INTRODUCTION 

A small anole from the eastern slopes of the Cordillera Oriental 
of Colombia is recognized as another new species apparently allied 
to the tigrinus species group. It is named Anolis lamari after its 
discoverer, W. W. Lamar. 

DESCRIPTION 

Anolis lamari, new species 
Figures 1-4 

Holotype. ICN 6762 (Instituto de Ciencias Naturales, Universi- 
dad Nacional de Colombia, Bogota) (formerly WWL 1576). 

Type Locality. Portachuelo, about 2 miles (by air) north of 
Manzanares, a police inspection station in the Municipio de Aca- 



Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138. 



BREVIOR.4 



No. 495 




Figure 1. Anolis lamari, holotype. Dorsal view of head. 



1992 



ANOLIS LAMARI 




o 



2 
u 

♦J 

o 

JS 






^ 






e 



BREVIORA 



No. 495 




Figure 3. Anolis lamari, holotype. Ventral view of head. 



cias, Meta, Colombia. W. W. Lamar coll. July 10, 1980. Elevation 
ca. 1,600 m. 

Diagnosis. Close to Anolis solitarius, tighnus, menta, and ruizi, 
apparently differing from all in details of coloration. It differs 
from tigrinus in the absence of minute tubercles on the head scales, 
from menta in that the dewlap is uniform yellow-orange rather 
than bicolor white and lemon-yellow, from ruizi in the possession 
of a small knob at the posterior edge of the parietal, and from 
solitarius in body pattern and in the presence of a line of enlarged 
scales continuing back onto the parietal knob. The pattern of the 
only known specimen, a male, is closest to that of male ruizi, but 
lacks the distinctly reticulate flanks of that species. 



1992 



ANOLIS LAMARI 




E 
t 

o 

Xi 
in 






2 






o 



^ 






s 
op 



6 BREVIORA No. 495 

Description. Head: Dorsal head scales smooth, small anteriorly, 
much larger from the anterior margin of orbits posteriorly. Eight 
scales across snout between second canthals. The scales within 
the moderate frontal depression as large as or larger than those 
anterior to them. Five postrostrals. Large anterior nasal in contact 
with the suture between rostral and first supralabial. About eight 
scales between the circumnasals dorsally. 

Scales of supraorbital semicircles very large, broadly in contact 
with each other and with the supraocular disks. About 1 6 enlarged 
scales in each supraocular disk, the largest medial, the disk bor- 
dered anteriorly, laterally, and posteriorly by granules. One elon- 
gate superciliary on each side, occupying about one half the lateral 
supraocular margin. The more posterior superciliaries minutely 
granular. 

Canthus blunt, six canthal scales, the first and second largest 
becoming gradually smaller anteriorly. Three to four loreal rows, 
subequal; 18-22 total loreals. 

Lower temporals finely granular, subequal. Two intertemporal 
rows on the bony bar that is the lower margin of the upper tem- 
poral vacuity. Supratemporals subgranular near the intertemporal 
rows, becoming abruptly larger near the interparietal. Scales just 
lateral to the interparietal large, ranging to one third the size of 
the interparietal, the latter very large, much larger than the ear, 
in broad contact with the semicircles. Four scale rows posterior 
to the interparietal relatively large but not as large as those lateral 
to it, and behind these a further series of four smaller scales leading 
to a bony boss on the parietal bone, i.e., a total series of eight 
scales larger than the nape scales behind the interparietal. Nape 
scales grading rapidly into the granular dorsals. 

Suboculars in contact with the supralabials, grading anteriorly 
into the loreals, posteriorly becoming abruptly smaller, grading 
into the lower temporals. About seven supralabials to below the 
center of the eye. 

Mental semidivided, in contact with six scales, in a gentle con- 
cave arc, between the infralabials: Four medial postmental gran- 
ules and, lateral to them, a first sublabial on each side at least 
five times the size of the postmental granules. Behind the first 
sublabials on each side, three to four additional sublabials in 



1992 ANOLIS L.4MARI 7 

contact with the infralabials. Medial gulars granular, convex, 
smooth, becoming larger near the sublabial rows. 

Trunk: Dorsals smooth, juxtaposed, the two middorsal rows 
slightly larger, smooth or very weakly keeled. Flank scales almost 
as large as dorsals, smooth, juxtaposed. Ventrals much larger, 
subquadrate, flat, smooth, subimbricate to imbricate, in trans- 
verse rows. 

Dewlap: Very large, extending posteriorly nearly to middle of 
belly. Edge scales smooth, imbricate, somewhat smaller than ven- 
trals. Lateral scales narrow, in widely spaced rows separated by 
naked skin. 

Limbs and Digits: Forelimb scales smooth, larger anteriorly, 
granular posteriorly. Upper thigh scales very weakly keeled, pos- 
terior thigh scales granular, lower leg scales smooth. Supradigitals 
of hand and foot weakly keeled. Nineteen lamellae under pha- 
langes ii and iii of fourth toe. 

Tail: Weakly compressed, scales small and smooth at base, 
becoming larger and keeled distally. A wider row dorsally, some- 
times double, keeled. Lateral caudals smaller, weakly keeled. A 
midventral double row abruptly larger, sharply keeled. Postanals 
enlarged (male, one hemipenis extruded, bifid at tip). 

Measurements. (Before preservation, provided by W. W. La- 
mar.) SVL 42.8 mm, total length 131 mm, dewlap 19 mm long, 
8.5 mm deep. 

Color. (As preserved.) Purplish to yellow-brown. Occipital area, 
including interparietal, dark. Snout anteriorly smudged with darker 
pigment. An irregular hollow triangle behind eye, narrowly bor- 
dered by black. Black oblique broken lines on nape in series with 
similar lines on flanks. A middorsal black line widest and most 
intense on the nape. 

Color in Life. (Adapted from W. W. Lamar's notes.) Overall 
pattern complex because of scattered speckling. 

Snout to eyes weak green. Lips pale creamy-tan. A bold white 
to gold stripe extending from below eye to ear at the level of the 
angle of the mouth. Distal portion of head golden tan. Body tan 
becoming green on posterior third. Beginning just posterior to 
eyes, four or five bars consisting of pairs of wavy dark green lines 
slanting backward and downward to venter. Middorsum with six 



8 BREVIORA No. 495 

brown rectangles all poorly defined but increasing in intensity as 
they approach the tail. Tail boldly banded in maroon brown and 
green, the brown bands fading to tan posteriorly. Limbs finely 
barred like sides. 

Venter pinkish cream with indistinct specks, but midventer 
ft-om chest to vent very pale yellow-green. Sides of belly pinkish. 
Vent and inner thighs and first third of tail pale yellow-green. 
Limbs brownish beneath, palms and soles maroon brown. Tail 
below banded brown and tan with some greenish cast. 

The very large dewlap pale yellow-orange with pale greenish 
white raised scales. 

Eye iris sooty-bronze, pupil narrowly ringed with gold. Eyelids 
translucent, tan like body. 

Tongue pink. 

Habitat. (From Lamar's notes.) Portachuelo is a "cuchilla de 
la Cordillera Oriental" of the Andes, a ridge that, at its highest 
point, is about 1,800 m. The area collected by Lamar was in the 
vicinity of 1 ,640 m, with the area of collection of the Anolis closer 
to 1,600 m. When collected, the unique type specimen was crawl- 
ing over the mossy bank of a mountain stream on the property 
of Sefior Chucho Cortez. The air temperature was 17.5°C. The 
animal was very slow-moving at that temperature but became 
very active when warmed up. 

RELATIONSHIPS 

The narrow relationships o^ Anolis lamari seem clear; its wider 
relationships involve some confusion. 

Narrow Relationships 

Apparently Corroborated Colombian- Venezuelan Relatives. A. 
lamari appears, on phenetic grounds, to be a close relative of the 
one Venezuelan and the three Colombian species with which it 
has been compared in the diagnosis, so close in all characters that 
this relationship seems corroborated. All five may then be con- 
sidered geographically replacing forms within the tigrinus species 
group, as currently understood (Rueda and Williams, 1986): ti- 
grinus from various localities in the coastal range of Venezuela, 
solitarius from the northern slopes of the Santa Marta Range in 
Colombia, menta from the southwestern slopes of the same range, 



1992 ANOLIS LAMARI 9 

niizi from the eastern slopes of the Cordillera Oriental in the 
Departments of Boyaca and Casanare, and lamah, also from the 
eastern slopes of the Cordillera Oriental, but in the Department 
of Meta. All members of this northwestern South American com- 
plex are relatively small species (maximum known size 57 mm 
SVL in tighnus) in montane forest. All share or tend to share 
certain features of squamation: (1) a large interparietal, (2) bor- 
dered laterally by large scales, and (3) usually in contact with the 
supraorbital semicircles, which (4) are almost always in contact, 
(5) relatively few scales across the snout between the second can- 
thals (four-ten), (6) relatively few scales in the supraocular disk 
(11 or less, except in lamari [16]), (7) usually, a series (five or 
more) of relatively large scales between the interparietal and the 
nape scales, which are granular like the dorsals, (8) suboculars in 
contact with the supralabials, (9) large well-differentiated subla- 
bials, except in two of 13 tigrinus, (10) dorsal scales uniform in 
size, (11) smooth ventrals larger than the dorsals, (12) lamellae 
under phalanges ii and iii of the fourth toe no fewer than 1 6 nor 
more than 22. 

A further external feature unites lamari with tigrinus, solitarius, 
and menta: a small parietal knob coincident with the border 
between post-interparietal scales and the nape scales. A skeleton 
recently made of Anolis solitarius (ICN 6153) shows that the 
external parietal knob is, as expected, underlain by a bony spur 
on the parietal bone (Fig. 5). This feature is absent in A. ruizi. 

Within this complex, differences are not sharp. Color, including 
dewlap color, is important. The problems within the tigrinus group. 





Figure 5. A. solitarius (ICN 6153). Left: The skull in profile. Right: The parietal 
bone to show the "parietal knob" and associated ridging. 



10 



BREVIORA 



No. 495 



as presently understood, are problems of the delimitation of taxa, 
not of features that unite them. It is, furthermore, not clear that 
all the populations referable to the group are yet known. Three 
of the five species, menta, ruizi, and lamari, have only recently 
been discovered, and no contact zones are known. The closest 
approach is that of menta and solitarius, which are less than 30 
km apart in the west and north respectively of the Sierra de Santa 
Marta. A. lamari is the southernmost described Colombian spe- 
cies, but a single unnamed specimen is known from Vista Her- 
mosa in the Sierra de Macarena that is, with high probability, the 
veritable southernmost Colombian record for the species group. 




/O 



0' 



-zA-IO' 



Figure 6. Colombian distribution of the ligrinus superspecies. MACARENA 
marks the presumed locality of an undescribed member of the tigrinus superspe- 
cies. 



1992 



ANOLIS LAMARI 



11 



The locality has previously been regarded as doubtful, because it 
was thought to be too distant from the other Colombian species 
of the tighnus group. No details of collection nor description of 
colors in life exist. (The distribution of Colombian species of this 
complex is shown in Fig. 6; A. tighnus, as mentioned, is widely 
distributed in the coastal range of Venezuela.) 

Of this northwestern South American complex lamari and ruizi 
are probably closest relatives. They are both on the eastern side 
of the Cordillera Oriental and show clear similarities in color and 
pattern (Fig. 7). They differ, however, as mentioned in the di- 
agnosis, in the absence, in ruizi, of the parietal knob that is present 
in lamari. 

Questionable Relationships with Two Brazilian Species. Two 
species, nasofrontalis and pseudotigrinus, in the Atlantic Forest 






Figure 7. Male dorsal patterns in three species of the tigrinus superspecies. A. 
A. solitarius. B. A. ruizi. C. A. lamari. 



12 BREVIORA No. 495 

in the state of Espirito Santo in Brazil, thus as widely disjunct 
from the Colombian-Venezuelan complex as the width of the 
South American continent permits, have been tentatively referred 
to the tigrinus species group. On size {nasofrontalis reaches 59 
mm SVL, pseudotigrinus 54 mm SVL) and all scale characters 
they do so belong. If these two Brazilian species do belong to the 
same species group as the Colombian-Venezuelan complex, then, 
most probably, this is an old group. Relict populations may very 
possibly exist elsewhere, and the evidence on the present and past 
distribution of the group must at present be presumed to be very 
incomplete. 

Remoter Relationships 

Problems of Distinction of the Tigrinus Group from the Punc- 
tatus Species Group. When A. lamari is compared with the punc- 
tatus species group as presently defined, a problem arises. This is 
typified by the confusion that occurred even during the discovery 
of the type specimen. 

Because a somewhat atypical adult of Anolis huilae (assigned 
to the punctatus species group) was found very close to the locality 
where the A. lamari type specimen was found, the small animal 
here described was at first thought to be a juvenile of that species. 
There is, in fact, no similarity in color or in pattern. The new 
species entirely lacks the bold spotting and nape ocellus of male 
huilae. There is some general similarity in squamation; in par- 
ticular the interparietal of huilae may often be in contact with 
the supraorbital semicircles. Figure 8A shows the interparietal in 
huilae, but also that there is but one row of enlarged scales behind 
the interparietal in huilae, while in the midline in lamari there 
are eight such enlarged scales in front of the nape scales. 

The source of the initial confusion between A. lamari and A. 
huilae was, in fact, the conspicuous very large interparietal scale 
in contact with the supraorbital semicircles. This condition is 
relatively unusual in Anolis, perhaps never invariable, but in cer- 
tain groups or species characteristic to the point of being almost 
or quite diagnostic. It is one of the features apparently primitive 
for the Anolis roquet series of the southern Lesser Antilles, lost 
only, and then only sometimes, in the two giant species, A. griseus 
and /I. richardi. In continental South America I count 1 8 described 



1992 



ANOLIS LAMARI 



13 







Figure 8. Scales of the parietal area in some members of the '"punctatus species 
group." A. A. huilae. paratype: MCZ 159122. B. A. caquetae, holotype: MCZ 
131 176. C. A. boettgeri: MCZ 1731 1 1. D. /I. deltae. holotype: MCNC 2031. 



species of Anolis and three described species of Phenacosaurus 
that are known to have the interparietal in contact with the semi- 
circles in at least some specimens. The three species of Phena- 
cosaurus and five species belonging to the beta section of Anolis, 
ibague, lineatus, macrolepis, rivalis, and meridionalis, require no 
comparison with lamari. They and the groups to which they be- 
long show abundant differences from the present species. Of the 
remainder, one, laevis, is known from a poorly preserved single 
specimen; there are not many characters to check, but laevis is a 
proboscis anole and lamari is clearly not. 

The other species in which a large interparietal in contact with 
the semicircles is known are currently allocated to two species 



14 BREVIORA No. 495 

groups, the punctatus group and the tigrinus group (Williams, 
1976). 

In the paper just cited I provided a key in which the punctatus 
group and the tigrinus group were separated by couplets 5 and 6: 

5. Ventrals smooth and/or dorsal squamation quite uniform 6 

Ventrals keeled, middorsals noticeably larger than flank scales 11 

6. Small anoles (ca. 50 mm snout-vent length) with large flat head scales 

tigrinus group 

Anoles large or small but not with large head scales punctatus group 

The confusion, however momentary, of lamari, presumed on 
the basis of general similarity to be a member of the tigrinus 
group, with huilae, assigned to the punctatus group anoles, dem- 
onstrates that the distinction provided by the key is inadequate. 
Clearly the reality of the distinction between XhQ punctatus group 
and the tigrinus group needs to be better demonstrated. 

The type species of the two species groups, A. punctatus and 
A. tigrinus, do seem to differ impressively. Three characteristics 
only need be mentioned: A. punctatus is primarily green in color, 
A. tigrinus primarily lichenate; A. punctatus reaches a maximum 
size of 89 mm SVL, A. tigrinus only 57 mm SVL; and in A. 
punctatus the parietal area of the head is devoid of any median 
prominence, in A. tigrinus there is posteriorly a distinct parietal 
knob (Ayala et al, 1984). Differences of the first two sorts, color 
and size, however, are ecomorphic (Williams, 1972, 1983) and 
imply neither relationship nor lack of it. In the West Indies A. 
punctatus would fit the classic definition of a trunk-crown eco- 
morph, and A. tigrinus that of a classic twig dwarf (Williams, 
1983). The character of the parietal knob is equivocal both in the 
South American complex that has been assigned to the A. tigrinus 
group and in the West Indies twig dwarfs: three of the West Indian 
twig dwarf species, insolitus, sheplani, and placidus, have such a 
knob; occultus does not; four of the South American species that 
are inferentially twig dwarfs have an analogous knob— tigrinus, 
solitarius, menta, and lamari; ruizi does not, and such a knob is 
not obvious in the two Brazilian species that have been referred 
to the tigrinus group. 

Is it possible that the "'tigrinus species group" is at best an 
ecomorphic (sensu Williams, 1972, 1983) subgroup of the punc- 
tatus assemblage? Even in squamation there is overlap between 



1992 ANOLIS LAMARI 15 

the two species typical for the supposed species groups. Counts 
across the snout between the second canthals are eight to 14 in 
A. punctatus, five to eight in A. tigrinus. This is confirmation that 
head scales tend to be smaller in punctatus than in tigrinus, but 
there is also overlap. Despite the difference in maximum size 
there is overlap in fourth toe lamellae, 22-32 under phalanges ii 
and iii in A. punctatus, 18-22 in A. tigrinus. While in the 11 
specimens of .4. tigrinus there is no example in which the supra- 
orbital semicircles are not in contact medially, there are 14 ex- 
amples of separation in 1 10 specimens examined o^A. punctatus. 
In the nine specimens of A. tigrinus in which the interparietal can 
be seen, eight have the interparietal in broad contact with the 
semicircles; only the type of the inferred tigrinus synonym, im- 
petigosus, has it separated by two scales. However, in none of the 
110 specimens of punctatus is the interparietal even in point 
contact with the supraorbital semicircles. 

And, while these two type species of the two "species groups" 
are on balance rather sharply distinct, the additions made to our 
knowledge of old species placed in the two groups and the recent 
referral of new species to one or the other of the two assemblages 
have greatly reduced the distinctiveness of one assemblage as 
compared with the other. 

In the case of the tigrinus group, solitarius and menta are green 
rather than lichenate. A parietal knob is not visible externally in 
ruizi, nor in Brazilian nasofrontalis and pseudotigrinus. Some 
species currently referred to the punctatus group are nearly as 
small as members of the tigrinus set, and some of these are species 
that have or frequently have a large interparietal broadly in con- 
tact with the supraorbital semicircles: caquetae (maximum SVL 
58 mm), deltae (maximum SVL 58 mm), dissimilis (maximum 
SVL 56 mm). 

The one character that, on inspection of all species belonging 
to both species groups, appears to separate all members of the 
tigrinus grouping from all sptcits— except one, Anolis santamar- 
/<3^— referred to the punctatus grouping is the enlarged scales lat- 
eral to the very large interparietal (Fig. 9). Such enlarged scales 
are not seen in even those species (with the exception of santa- 
martae) now referred to the punctatus group that have a large 
interparietal and are similar in size to members of the tigrinus 









Figure 9. Scales of the parietal area in members of the ""tigrinus species group." 
A. A. nasofrontalis. B. A. pseudotigrinus. C. A. tigrinus. D. A. solitarius. E. A. 
menta. F. A. ruizi. 



1992 



A NO LIS LAMARI 



17 



group (Figs. 8, 10). (It is noteworthy that the members of the 
roquet series, which primitively show the interparietal in contact 
with the supraorbital semicircles, also lack these large scales lateral 
to the interparietal.) 

A. santamartae: An Anomalous Species. The one exception 
within what I have called the punctatus species group, Anolis 
santamartae Williams, 1982 (the significant parietal area is shown 
in Fig. 11), is the only species of alpha anoles known to occupy 
the southeast comer of the Sierra de Santa Marta, the north end 
of which is occupied by A. solitarius and the southwest by A. 
menta. It might be plausible to consider A. santamartae on geo- 
graphic grounds alone as a possible member of the tigrinus group. 







Figure 10. Scales of the parietal area in some members of the ^^ punctatus 
species group." A. A. sp. ? near transversalis: MHNJP 1 192. B. A.jacare: AMNH. 
C. A. calimae, holotype: MCZ 158392. D. A. dissimilis: FMNH 81369. 



18 BREVIORA No. 495 




Figure 1 1. Scales of parietal area in A. santamartae, holotype: CAS 1 13922. 

I did consider this possibility when I described it: it is the right 
size, approximately the right habitus. Its habits and habitat are 
undescribed. 

Why did I reject this hypothesis? On the basis, first, of the 
distinctive pattern, in particular the light line from the lower jaw 
onto the upper arm (not emphasized in the rather muted figure 
of the type in the original description, but stressed in the text and 
very evident in one of the paratypes, MCZ 15631 1, the pattern 
of which is diagrammed in Fig. 12). This is a singular and dis- 
tinctive pattern for any anole. I was much more comfortable in 
assigning santamartae to the punctatus species group, which I 
knew to be quite varied in pattern, rather than to the tigrinus 
species group, which I then believed to be rather uniform in 
pattern, its members differing among themselves primarily in 
dewlap color. 

When santamartae was described in 1982, menta Ayala, Harris, 
and Williams, 1984, and ruizi Rueda and Williams, 1986, were 
not yet recognized. I had not yet seen the large new series of 
solitarius, collected by Pedro Ruiz and John Lynch, and I was 
not aware that the ground color of that species in life was green. 
I was confident that the tigrinus series was ecologically the equiv- 
alent of the twig dwarf species of the West Indies. The West Indian 
twig dwarf anoles are basically cryptic in pattern, lichenate, as 
tigrinus was known to be. I did not expect a relative of tigrinus 
to have the pattern o^ santamartae, even H santamartae did have 



1992 ANOLIS LAMARI 19 



Figure 12. Male dorsal pattern in A. santamartae, after MCZ 1 563 1 1 , a para- 
type. 

much of the size and habitus of tigrinus. Unhappily, the extrap- 
olations as regards pattern from the West Indian twig dwarf eco- 
morphs have proved quite wrong. Additionally, the distinctly, 
although rather weakly keeled, dorsals and ventrals influenced 
me. Keeled ventrals are unusual even in the punctatus species 
group, but they are known in punctatus itself The boulengeri 
morph of that species, characteristic in western populations, has 
keeled ventrals. There was no indication of comparable condi- 
tions in the known species of the tigrinus species group. There is 
also no evident parietal knob in santamartae. If santamartae 
belongs to the tigrinus species group, it is the most distinctive of 
the included species. 

Remaining Problems in the Recognition of the Tigrinus Group. 
Even if the tigrinus group is phenetically recognizable, is it phy- 
logenetically a unit? There are several difficulties here. One is the 
possibility, mentioned above, that the tigrinus group is an eco- 
morphic grouping: The tigrinus group may be the South American 
twig dwarfs. On the limited present evidence they seem likely to 
be so in an ecological sense. However, the scale character, the 
enlarged scales lateral to the parietal, by which the group may 
possibly be recognized morphologically, is not an attribute of twig 
dwarfs as an ecomorphic category. It is not present in any of the 
classic West Indian twig dwarfs. This possibility may provision- 
ally be dismissed. 

Unfortunately another possibility cannot be so readily dis- 
missed. The polarity of the diagnostic scale character is in doubt. 
One of the two possibilities in the analysis of head scale characters 
in the Squamata is that small undifferentiated head scales are 
primitive, and that they have repeatedly united in larger units. 
The other is that large scales like parietals, frontals, postparietals, 
etc., are primitive. In the first hypothesis the larger scales lateral 



20 BREVIORA No. 495 





Figure 13. A. punctatus (MCZ 155994). Left: The skull in profile. Right: The 
parietal bone to show absence of the parietal knob. 

to the interparietal are genuinely synapomorphic, and the tigrinus 
group is a genuinely monophyletic unit that includes the two 
widely disjunct Brazilian species. In the second hypothesis, which 
I favor on general grounds rather than the specifics of this case, 
the larger scales lateral to the interparietal are remnants of prim- 
itively present parietal scales in this area. Then, the tigrinus group 
may not be monophyletic, but merely an assemblage of possibly 
only remotely related species that happen to remain plesiomor- 
phic in the size of the scales lateral to the interparietal. The Co- 
lombian set of species might be genuinely a superspecies, but the 
Brazilian members of the assemblage would be only species that 
by a combination of ecomorphology and symplesiomorphy have 
come to resemble their relatively distant relatives on the other 
side of the continent. 

The parietal knob seen in tigrinus, solitarius, menta, and now 
in lamari seems certainly a derived character. Figure 5 shows the 
bony structure underlying the external parietal knob in A. soli- 
tarius. Figure 1 3 shows the complete absence of such a structure 
in A. punctatus. An approach to the solitarius condition is seen 
in A. jacare (Fig. 14), and in no other of the South American 
alpha anoles examined {punctatus, agassizi, chloris, peraccae, 
gemmosus, ventrimaculatus, aequatorialis, princeps, squamulatus, 
latifrons, and frenatus). 

Etheridge in his thesis (1960) discussed the ontogenetic and 
phylogenetic history of parietal crests in Anolis. He was able to 
show that in the ontogeny o^ Anolis carolinensis (well illustrated 
in his fig. 9) the parietal crests first delimit a distinctly trapezoidal 
area, then a triangular area, and finally have a Y shape with the 



1992 



ANOLIS LAMARI 



21 





Figure 14. A. jacare (MCZ 112096). Left: The skull in profile. Right: The 
parietal bone to show similarity to A. solitarius in the incipient or convergent 
crest structure of the parietal in this species. 



arms bounding the triangular area continued backward as a me- 
dian ridge. He asserted that this ontogenetic sequence was pre- 
cisely parallel to the sequence seen in phylogeny: A. carolinensis, 
a relatively derived species on other characters, had the Y crests 
in adults, whereas the South and Central American alpha anoles 
and the southern Lesser Antilles anoles, which are primitive in a 
number of other osteological characters, retain a trapezoidal crest 
pattern. My own observations confirm Etheridge's statements with 
minor revisions. Derived anoles definitely have Y-shaped parietal 
crests with a relatively long and narrow median posterior ridge. 
However, in relatively basal anoles the crests bound a trapezoidal 
area consistently only in the roquet series of the southern Lesser 
Antilles. In this set of species the lateral ridges do turn inward 
posteriorly to provide a raised transverse boundary to the parietal 
roof (called the occipital crest by Etheridge). However, in the 
primitive mainland alphas (e.g., A. punctatus, Fig. 1 3) the Y stem 
frequently is broad and short, a condition transitional to the fully 
derived Y shape. The parietal roof anterior to this posterior ridge 
is still trapezoidal, but lacks the distinct posterior boundary of 
an occipital crest that is present in the ontogenetically and phy- 
logenetically primitive condition. A. jacare (Fig. 14) is still more 
derived. The stem of the Y is narrow— a single ridge, differing 
from the more advanced condition only in being short. A. soli- 
tarius (Fig. 5) retains the short narrow single ridge oi A. jacare, 
but adds a small bony knob at the end of it, the skeletal under- 
pinning of the external parietal knob. 

The similar bony parietal knob according to this analysis occurs 
in some of the dwarf West Indian species (Williams, in Ayala et 



22 



BREVIORA 



No. 495 





Figure 15. A. insolitus (MCZ 107018). Left: The skull in profile. Right: The 
parietal bone to show a parietal knob convergent with that of ^. solitarius. 



al, 1984; Fig. 15 in this paper). (In A. sheplani and A. placidus 
the skull is not known.) In A. insolitus, at least, the skeletal knob 
is at the end of a triangular parietal; there is not even a short- 
stemmed Y. Clearly this condition is convergent and not relevant 
to the issue of the monophyly of the tigrinus species group. 

The parietal knob, defined as the condition seen in solitarius, 
might then be a synapomorphy of the tigrinus species group, but 
only //this feature has been lost in ruizi, nasofrontalis, and pseu- 
dotigrinus, as well as santamartae, if the latter belongs in the 
group. The similarity of pattern in ruizi and lamari and their 
geographic proximity suggest that this hypothesis of loss may be 
true for ruizi. The widely disjunct range of the two Brazilian 
species does not rule out this possibility for those two species, 
but it is clearly not as well supported. The Brazilian species were 
always problematic members of the tigrinus species group. They 
remain so. 

CONCLUSION 

I see no means to resolve the tangle presented here. I am content 
to speak of a tigrinus species group, provided it is recognized as 
a convenient means to call attention to phenetic resemblances 
that may or may not be phylogenetically meaningful. 

As currently used, the punctatus group is clearly the residue of 
those South American alpha Anolis believed or known to have 
arrow-shaped interclavicles (Williams, 1989) that are not placed 
in the presumed tigrinus lineage. It might be a rescue of the 
punctatus group concept if the tigrinus lineage were placed within 



1992 ANOLIS LAMARI 23 

it. !t would then consist of all South American a\pha. Anolis known 
or believed to have arrow-shaped interclavicles and known to 
have non-autotomic caudal vertebrae as adults. Even this would 
be a dubious rescue, again because of a question of polarity. I see 
no objective grounds for deciding whether arrow- or T-interclav- 
icles, sensu Etheridge (1960), are primitive. This leaves me again 
with the punctatus group as a cluster of convenience, intended 
not to formally decide a phylogenetic question but to informally 
raise that question. 

The problem of which the present case is an example is a 
pervasive and difficult one, and very clearly not limited to the 
genus Anolis. A useful recent discussion, with a summary of the 
pertinent literature, is that of Bauer et al. (1988). / very much 
concur with their point that "species groups" are not formal taxa, 
but often, perhaps usually, operational clusters, phenetic groupings 
of convenience, intended at best to suggest possible affinities but 
not pretending at all to their demonstration. I disagree with Bauer 
et al. in being less optimistic than they that data sufficient for the 
analysis that they hope for will soon be available. 

I plead for extensive periods of use of informal groupings in 
cases in which taxonomic decisions must be based on evidence 
that is less than conclusive. The levels of confidence for every 
taxon erected or changed need not be quantified— there may be 
no plausible means of doing so— but the grounds for these levels 
of confidence should always be spelled out in detail, as I have 
attempted to do in this paper. 

ACKNOWLEDGMENT 

The illustrations were done by Laszlo Meszoly. 

LITERATURE CITED 

Ayala, S. C, D. M. Harris, and E. E. Williams. 1984. Anolis menta, sp. n. 
(Sauna, Iguanidae), a new tigrinus group anole from the west side of the Santa 
Marta Mountains, Colombia. Papeis Avulsos de Zoologia, Sao Paulo, 35(12): 
135-145. 

Bauer, A. M., A. P. Russell, Ahfo H. I. Rosenberg. 1988. Formal taxa, species 
groups, and perception of the genus Diplodactylus (Reptilia: Gekkonidae). 
Zeitschrift fur zoologischen Systematik und Evolutionsforschung, 27: 44-48. 

EtheriekjE, R. 1 960. The relationships of the anoles (Reptilia: Sauria: Iguanidae): 
An interpretation based on skeletal morphology. Ann Arbor, Michigan, Uni- 
versity Microfilms, xiv + 236 pp. 



24 BREVIORA No. 495 

RuEDA, J. v., AhfD E. E. Williams. 1986. Una nueva especie de saurio para la 
Cordillera Oriental de Colombia (Sauna; Iguania). Caldasia, 15(7 1-75): 5 1 1- 
524. 

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. Papeis Avulsos de 

Zoologia, Sao Paulo, 29(26): 259-268. 

. 1982. Three new species of the Anolis punctatus complex from Ama- 
zonian and inter-Andean Colombia, with comments on the eastern members 
of the punctatus species group. Breviora, Museum of Comparative Zoology, 
467: 1-38. 

. 1983. Ecomorphs, faunas, island size, and diverse end points in island 

radiations of Anolis. pp. 326-370, 481^83. In R. B. Huey, E. R. Pianka, 
and T. W. Schoener (eds.). Lizard Ecology. Studies of a Model Organism. 
Cambridge, Harvard University Press. 501 pp. 

. 1989. A critique of Guyer and Savage (1986): Cladistic relationships 

among anoles (Sauria: Iguanidae): Are the data available to reclassify the 
anoles?, pp. 433-478. In C. A. Woods (ed.), Biogeography of the West Indies. 
Past, Present, and Future. Gainesville, Florida, Sandhill Crane Press, Inc. xii 
+ 878 pp. 



B R E V I R A. 

Museum of Comparative Zoology 



us ISSN 0006-9698 , , ^^ a nT-> 



Cambridge, Mass. 2 February 1994 Number 496 ' 

A NEW SPHAERODACTYLUS (SAURIA: GEKKONIDAE) 
FROM BEQUIA, GRENADA BANK, LESSER ANTILLES 

James Lazell' 

Abstract. A new species of Sphaerodactylus of small size (25 mm SVL) is 
described from Bequia, Grenada Bank, Lesser Antilles. It is without keeled scales; 
with large, subimbricate lateral dorsals (10-12 in standard distance at midbody) 
and slightly smaller, convex middorsals (12-15 in standard distance); with large, 
imbricate, cycloid ventrals (8 in standard distance); and with a blotchy, obsolete 
pattern in somber colors. Its discovery fills a long-standing biogeographical gap. 

/ believe sufficient collecting will demonstrate sphaerodactyls to be present. 

Wayne King (1962) 

INTRODUCTION 

The small geckos of the genus Sphaerodactylus are nearly ubiq- 
uitous in the West Indies. They occur on tiny fragments of land 
less than a hectare in area (e.g., Watson Rock in the British Virgin 
Islands: Museum of Comparative Zoology [MCZ] 176729). There 
are dozens of species on some of the larger Greater Antilles (Haas, 
1991). The absence of a species o{ Sphaerodactylus from the Gre- 
nada Bank, southernmost of the Lesser Antilles, has long been a 
sore point for biogeographers (King, 1962; Williams, 1989; Haas, 
1991). 

In 1964, I first went down through the Grenadines— small is- 
lands on the northern two-thirds of the Grenada Bank— collecting 
specimens. I travelled on the sloop Flamingo, a St. Vincent gov- 
ernment fishing vessel. My trip was arranged by Dr. I. Earle Kirby, 
then St. Vincent government veterinarian, and always an avid 
naturalist. In November 1989 I returned with Thomas Sinclair, 
of The Conservation Agency, and Christopher Luginbuhl, of the 



'Associate, Department of Herpetology, Museum of Comparative Zoology, and 
The Conservation Agency, 6 Swinburne St., Jamestown, Rhode Island 02835. 



2 BREVIORA No. 496 

David B. Luginbuhl Foundation, with the intention of exploring 
possibilities for a biological station on Luginbuhl's property on 
Bequia, Grenadines. We obtained specimens of a distinctive 
Sphaerodactylus, reported as "Sphaewdactylus cf. vincenti" by 
Lazell and Sinclair (1990). Dr. Kirby, who had spent the inter- 
vening years working to develop science, education, and conser- 
vation in St. Vincent and the Grenadines, advised us on proce- 
dures and arranged meetings and logistics that insured our success. 
The contributions of Earle Kirby to progress and enlightenment 
in the Caribbean are legion; see Gibson, in Jinkins and Bobrow, 
1985. I thus take pleasure in naming: 

Sphaerodactylus kirbyi, sp. nov. 

Type. MCZ 175141, adult male, collected by Thomas Sinclair, 
30 November 1989. See Figure 1. 

Type Locality. Slope above Friendship Bay, Bequia, Grena- 
dines. See Figure 2. 

Diagnosis. A small species (to 25 mm SVL) of moderate rather 
than attenuate proportions: axilla to tip of snout ca. 44 percent 
(40-48, av. 43.8 ± 2.9) of snout-vent length (SVL). Standard 
distance (STD), tip of snout to center of eye, 14-16 (av. 14.7 ± 
0.8) percent of SVL. No keeled scales, but lateral dorsals at mid- 
body convex to subtectiform, subimbricate, and large: 10-12 (av. 
1 1.3 ± 0.8) in STD; middorsals slightly smaller, convex, 12-16 
(av. 14 ± 1.7) in STD; ventrals imbricate, cycloid, and large: 8 
in STD; 10-12 (av. 10.3 ± 0.8) subdigital lamellae, counting the 
terminal spheroid. Subcaudals at least twice the width of lateral 
caudals. Coloration somber, shades of gray-brown, with irregular 
small blotches and dim cephalic stripes. 

Description of the Type. MCZ 175141 is an adult male 23.6 
mm SVL. The standard distance (STD) is 3.4 mm, 14 percent of 
SVL. There are 1 1 large, convex, subtectiform, slightly imbricate, 
lateral dorsals contained in STD at midbody. The middorsals are 
rather similar, convex, juxtaposed scales, and slightly smaller: 12 
contained in STD. The ventrals are large, flat, and cycloid: 8 in 
STD. The gulars are smooth, mostly granular, but enlarged and 
elongate close to the mental; counting from immediately posterior 
to the mental, there are 17 gulars in STD. There are 10 subdigital 
lamellae, counting the terminal spheroid, under the fourth toe of 



1994 



GRENADA BANK SPHAERODACTYLL'S 





O 



03 



I 

U 



N 

U 



'S. 

O. 

m" 

c 



N 

U 



S ^ 



a 


o 

c 


>-* 


> 


J3 


4-* 


■t-» 


V3 


< 


cT 


. , 


u 


^ 


"H 


■^ 


C<3 


5 





to 


<u 


a 


C 




rt 


b 


u 


'o 




1 




s 


t~~ 


S3 


OS 


<3 


r^ 




N 
U 




S 


^ 


..^ 




■•^ 


P 




3 


i 


00 


.c 



BREVIORA 



No. 496 




Figure 2. Type locality of Sphaerodaclylus kirbyi. A, the Caribbean basin; 
arrow indicates the Grenada Bank. B, the major islands of the Grenada Bank. 
Bequia is shaded. The dotted line indicates the approximate land area of greater 
Grenada at glacial maximum, ca. 1 2,000 yr BP. Bar is 1 km. C, Bequia: contours 
are approximately 100 m; 1, Friendship Bay; 2, Princess Margaret (formerly 
Admiralty) Bay; 3, Spring Bay. Bar is 1 km. Dot indicates area in which type 
series was collected. 



1994 GRENADA BANK SPHAERODACTYLUS 5 

the pes. There are three large supralabials, the third subtending 
the eye, followed by three small, granular supralabials to the com- 
missure of the mouth. 

In contrast to the gracile, attenuate, long-necked habitus of 
Sphaewdactylus v. vincenti (and 5". molei of Trinidad), the pro- 
portions are more ordinary by generic standards. The distance 
from the axilla to the tip of the snout is 44 percent of SVL. 

The escutcheon consists of pale scales in contrast to those sur- 
rounding it, which are peppered with sooty gray-brown. It spans 
five scales along the ventral midline and extends out the thighs 
to a maximum width of 24 scales: The escutcheon is configured 
essentially as in most races of S. vincenti; see, for example, Schwartz 
(1964), figures 19 {S. v. festus), 21 {S. v. adamas), 22 (S. v. psam- 
mius), and 28 (S. v. vincenti). 

In life, the type was somber, dark, shades of gray-brown. The 
belly was dark gray apart from the pallid escutcheon. There was 
a yellow wash on the chin, throat, and sides of the neck. The 
underside of the tail was mottled with rosy-orange. Color change 
was to velvety near-black. Apart from the small slate blotches 
(freckles in my notes), no pattern was apparent. Some pattern, 
however, is visible in the preserved specimen. 

The pattern consists of irregular, small, dark blotches on the 
trunk. There is a rather faint linear head pattern consisting of a 
broad gray-tan band extending caudad through the eye and con- 
verging from each side to make a short middorsal area on the 
shoulders clear of blotches. This gray-tan band is bordered above 
and below by fine slate stripes. The dorsal of these begins on the 
upper eyelid and fades on the temple. The ventral begins on the 
snout and extends to the shoulder. Below this stripe is another, 
finer, irregular streak extending from the posterior edge of the eye 
to break up into speckles on the cheek. 

Dark, slate-gray stripes extend caudad from the sacrum and 
converge to form a V on the tail base. This figure is subtended 
by an ash-brown band extending from above each hindlimb in- 
sertion to converge and form a short area clear of blotches on top 
of the tail. This band is, in turn, subtended by another slate-gray 
stripe which breaks up into blotches on the tail. 

There is a light pattern of longitudinal ash-gray streaks on the 
yellowish gray throat. 



6 BREVIOR.A No. 496 

Paratopes. MCZ 175 142, another adult male 23 mm SVL, and 
MCZ 175143, a juvenile 17.5 mm SVL, were also collected by 
Sinclair above Friendship Bay on 30 November 1989. Two adult 
females, MCZ 175144, 23.7 mm SVL, MCZ 175146, 25 mm 
SVL, and a juvenile, MCZ 175145, 14.7 mm SVL, were collected 
by Sinclair above Friendship Bay on 6 December 1989. 

Apart from the escutcheon of the male, MCZ 175142, which 
is 4 by 24 scales, there is no evidence of sexual dimorphism in 
meristic characters. Counts for the paratypes, with average and 
standard deviation for the entire series (including the type) in 
parentheses are as follows: Lateral dorsals in STD at midbody 
10-12 (av. 11.3 ± 0.8); middorsals 12-16 (av. 14 ± 1.7); mid- 
ventrals in STD 8 in all specimens; gulars in STD, counted from 
immediately behind the mental posteriorly at midline of throat, 
15-18 (av. 15.7 ± 1.5); subdigital lamellae under fourth toe of 
pes, including the terminal spheroid, 10-12 (av. 10.3 ± 0.8). 

In all specimens there are three large supralabials to the eye. 
Posterior to these are three to five much smaller, granular su- 
pralabials. In all specimens there are broad subcaudal plates, at 
least twice the width of the lateral caudals. 

I saw and annotated the type and paratypes MCZ 175142-43 
alive, on 30 November. Both paratypes showed faint cephalic and 
sacro-caudal patterns when alive; these are visible in the type 
after preservation. Otherwise both were very similar to the type 
in colors except that the smaller, MCZ 175143, had cream-color 
subcaudal blotches and the other adult male, MCZ 175142, had 
orange-brown subcaudal blotches; it is intermediate in size be- 
tween the smaller and MCZ 175141, the type. 

I did not see MCZ 175144-46 alive, but they are similar in 
coloration and pattern to the others as preserved; coloration and 
pattern seem little affected by preservation. The pattern elements 
vary in clarity and development. The male MCZ 175142 is very 
similar to the type, but the throat streaking is virtually absent 
and the throat color was cream and faintest ash-gray. The cephalic 
pattern breaks up into blotches on the jowls. In the juvenile MCZ 
175143 longitudinal throat streaks are well developed but the 
color was even more somber, with no yellowish throat tint. This 
juvenile has a well-developed cephalic pattern, and throat with 
streaking as in the type, but has broad, dark sacro-caudal stripes 



1994 GRENADA BAUK SPHAERODACTYLUS 7 

reducing the light V on the tail base. The two females set extremes 
of variation. One, MCZ 175144, is the palest individual seen; 
preserved, the ground color is tan-brown to ash-gray. The blotches 
and the cephalic pattern are especially bold, but the sacro-caudal 
figure is irregular and weakly demarcated (Fig. 1). 

The female MCZ 175146 is the darkest specimen seen. The 
blotching is heavy and amalgamates to marbling. Both cephalic 
and sacro-caudal patterns are largely obliterated by blotch amal- 
gamation. Throat streaking is especially dark and prominent in 
this specimen. 

Comparisons. Sphaerodactylus kirbyi resembles its closest geo- 
graphic neighbor, S. v. vincenti of St. Vincent. St. Vincent lies a 
mere 8 km to the north of Bequia. However, the diiferences in 
the geckos are striking. The Vincentian form is larger, has a longer 
neck and more gracile proportions, and is absolutely distinct in 
smaller scale size and rich cephalic coloration. 

I have examined 2 1 specimens of Sphaerodactylus v. vincenti 
from St. Vincent: MCZ 10788, the type, no precise locality; MCZ 
20550, Grand Sable Estate; MCZ 38190-94 plus one untagged 
juvenile. Diamond Beach; MCZ 797 1 6-19, Brighton; MCZ 79720- 
28, Cane Garden; MCZ 96032, Colonarie. Of these, five were 
examined meticulously for all measurable and meristic characters; 
MCZ 79720-24. Eight more were checked for both of the dorsal 
scale counts and midventrals in STD. I agree with King (1962) 
that STD counts can be skewed by allometric growth, and did 
not count juveniles less than 19 mm SVL. However, I believe 
STD counts are more useful than long counts along and around 
the body. The smallest Sphaerodactylus kirbyi seen, MCZ 175145, 
only 14.7 mm SVL, agrees in proportions and counts with the 
larger individuals. Because of its proportionately longer body, S. 
kirbyi might yield axilla-to-groin counts overlapping those of the 
much smaller scaled S. v. vincenti, but STD counts show no over- 
lap. 

In S. V. vincenti STD is 14-15 (av. 14.6 ± 0.5) percent of SVL, 
essentially identical to S. kirbyi. However, eight of 20 Vincentian 
specimens exceed 25 mm SVL; King (1962) gives the maximum 
length as 27 mm SVL; this fits the type specimen, MCZ 10788. 
S. V. vincenti is slender and elongate in proportions; axilla to tip 
of snout is relatively long, 46-53 (av. 48 ± 2.8) percent of SVL. 



8 BREVIORA No. 496 

The sample sizes, however, are too small for statistical signifi- 
cance. 

Scale size provides clear distinctions. In S. v. vincenti there are 
13-15 (av. 13.8 ± 0.8) lateral dorsals at midbody in STD. There 
are 16-22 (av. 18.6 ± 2.4) middorsals in STD. There are 9 or 10 
(av. 9.4 ± 0.5) midventrals in STD. There are 19-22 (av. 20 ± 
1 .4) gulars in STD counted along the midline of the throat be- 
ginning immediately posteriorly to the mental. Because of small 
sample sizes, I simply summed lateral dorsal and middorsal counts 
for the two species and obtained statistical significance (Student's 
/-test) at 95 percent level of confidence. The sum of lateral dorsals 
and middorsals in STD is 23-28, av. 25.3 ± 1.9 in S. kirbyi and 
31-36, av. 32.4 ± 2.2 in S. v. vincenti. 

Important quantitative characters are shown in Table 1 . 

Sphaewdactylus kirbyi resembles King's (1962) figure 12B and 
C of S. V. vincenti in pattern, but is far more spotted; the spots 
are larger and I refer to them as blotches (see Fig. 1). In coloration 
the two species are strikingly distinct. S. v. vincenti has a yellow 
head and a bluish-green iris (Schwartz, 1964:404). In my field 
notes of 4 and 8 June 1964, I describe individuals as "orangish 
on heads" with throats "orange-yellow" or "yellow," and iris 
"pale blue-green." 5'. kirbyi has undistinguished head coloration, 
with only the faintest yellowish throat tinges in some individuals; 
the iris is not notably colorful. 

More cursory comparisons are required to Sphaerodactylus kir- 
byi 's closest relative to the south, S. molei of Trinidad, Tobago, 
and the adjacent mainland of South America. I have examined 
19 specimens of 5. molei. The scales are much smaller even than 
in S. V. vincenti (King, 1962) and there are four supralabials to 
the eye. Like S. kirbyi the gulars of S. molei are smooth. The 
median subcaudals are not so large or well differentiated (illus- 
trated by Harris, 1982:13). The sacro-caudal pattern of 5". molei 
consists of subparallel, longitudinal, light, dark-bordered bands, 
not converging to form a sacral V as in S. kirbyi and S. vincenti. 
The well-developed longitudinal stripes from head to tail in S. 
molei are also distinctive. 

Sphaerodactylus kirbyi has much larger scales than either S. 
vincenti or S. molei, which flank it north and south, respectively. 



1994 GRENADA BANK SPHAERODACTYLUS 9 

It is in no sense intermediate between these species, and cannot 
be argued to unify them taxonomically. Similarly, S. kirhyi does 
not fit into the elaborate pattern of geographic variation in S. 
v/«c£'/7// extending from St. Vincent to Dominica. Some diagnostic 
and meristic characters indicative of aspects of this geographic 
variation are shown in Table 1 . 

Despite its diversity, S. vincenti hangs together rather well. 5". 
V. diamesus of St. Lucia is comfortingly intermediate in scale size 
between S. v. psammius and S. v. vincenti, which flank it north 
and south, respectively. S. v. psammius of extreme southern Mar- 
tinique is similarly intermediate in scale size between diamesus 
and josephinae. The trend to larger scales from nominate vincenti 
through diamesus and psammius Xojosephinae reaches its extreme 
in the isolated S. v. adamas, very slightly differentiated from 
josephinae. 

There is something of a break across central Martinique, where 
sphaerodactyls are as yet undocumented. To the north occur the 
very large, ocellate, heavily carinate forms pheristus, ronaldi, and— 
on Dominica— /^20/7/7z7^r. However, ocellate S. v. festus effeclivQly 
bridges the gap between these forms in pattern and scale size. S. 
V. pheristus and ronaldi intergrade and monilifer seems in most 
respects an extension of the trend from pheristus to ronaldi. At 
glacial maximum sphaerodactyls like ronaldi, or pheristus x ron- 
aldi, would presumably have occurred closer to Dominica than 
is possible today (Fig. 3). Sphaerodactylus kirbyi does not fit into 
this picture of variation in S. vincenti at all. In accord with my 
long-held views on evolutionary species (Lazell, 1972 and cited 
therein), I grant kirbyi full species status. 

Relationships. S. /czr^y/ agrees with 5. v. ronaldi and S. v. monili- 
fer in meristics, but is utterly unlike these very large, boldly pat- 
terned, ocellate forms with heavy gular (and even chest) scale 
keeling. The distinctions from smaller, similarly proportioned, 
southern Martinique forms josephinae and adamas are weak. They 
have larger scales than kirbyi and the sum of lateral dorsals plus 
ventrals in STD separates them from kirbyi, but it is a close thing 
(see Table 1). 

My sample sizes are so small that I had to sum the lateral- 
dorsal-plus-ventral counts for josephinae and adamas to dem- 



10 



BREIIOR.4 



No. 496 



u w 

X U 

H 2 

« "J 



D 
O 
oi 
O 

S 

s 
I 






o 
a; 
til 

I 



Co w 

03 



H 



Z 

o 

< 

> 



D 
Z 

< 

C/) 
UJ 



111 

H 
< < 



o 
u 



H 
D 
O 

z 

1/5 

u. 
O 
cu 
H 

5q 

+ ^ 
+ ^ 



oi 
u 
Q 



CO 
PU 

(- 



< 

o 

uu 

o 

X 
H 
UJ 

H 

f_ C/5 
O 

z 



UJ 

< 

< 

< 

< 

>J 
D 
O 
w 
X 
H 

z 

z 
o 



O 5 

. Q 

— a! 

w < 

5 Q 



a: 
< 
o 

UJ 

< 

o 

(/5 
PL, 

o 

+? 

UJ 
UJ 

Pi 
O 

UJ 

Q 

OS 

O 



I- -J 



a: 

UJ 

H 

z 

UJ 

> 

OS 

o 

H 

C/5 

UJ 

X 

u 
o 

H 

z 

o 



OS 

o 
T 2 

— ^ H 

UJ »} 



Q 
OS 
< 

Q 
Z 

< 

H 
en 

UJ 

X 
H 

- Z 



UJ 

.J 
< 
u 

< 

c/5 
OS 

o 

Q 

UJ 

i- . 
< UJ 

-J Z 

II J 

Q i 

-J s 



f- 
< 

U 
< 

u 

(/5 



Q 

ui 

>■ 

UJ 



+ 



Q 

+ 
Q 



Q 

H-l 



Q 



c 
o 

X 



0^ 


r) 


r~- 


so 


* 




^^ 


' — ' 


(^1 


o 


— ' 


(^1 


, ^ 


^ 


+ 1 


+1 


+1 


+1 


— ' 


fN 


— ' 


n-i 


■* 


>n 


00 


+1 


+1 


+1 


iri 


r4 


rr 


m 


_ 


_ 


m 


ri 


r<-i 


r-i 


fN 


(N 


(N 


(^4 


00 


so 


IT) 


iri 


(N 


m 


TT 


f^l 


ro 


1 


1 


(N 


1 


fN 


m 




^ 




OS 


OS 


rN 


r-J 


m 


r-j 


r^l 


1 — 1 


^- 


fN 



00 ^^ r-- 

H — — ' 

+1 +1 +1 

■ri t^ m 

ro <N rN 



r~~ 00 -^ 

CO <N (^J 

I I I 

rn so — 

CO 04 D 



oo 


00 


„ 


* 


* 




* 


so 




o 


o 


OS 


00 


so 


rj- 


rj- 


O 


r- 


+1 


+1 


o 


o 


o 






+1 


— i 


ro 


oo 


+1 


+1 


+1 


+1 


+1 


rn 


+1 


_-■ 


m 


'^ 


00 


t^ 


o 


ir-i 


_ 


o 


^^ 


^^ 


OS 


t^ 


I^ 




^~^ 


_ 




^— ^ 


^".^ 








^^ 


^.^ 


^-^ 


^— ' 


(N 


m 


o 






_ 


so 


(^4 


_ 


1 


7 * 


7 


^ 


oo 

1 


T 


7 


7 


7 



OfOoooor~~r-osTt — 00 



■* r-- OS — ' ~- * so 

r~- rsi o O — <^i ■^ O 

" +1 +1 +1 +1 +1 -- +1 

+1 so "/^ ■^ fN m +1 r- 

Tfodsd''J'r^r^ 0«^ 

— __—,_—. (N—  

'/-) r^l r-- m tT iri — ^ so 

7 I 7 7 7 7 I 7 
risosor^ri — mosin 



« * 

* * # * + * +1 

+ + +1 +1 +1 +1 + +1 +1 



so ro n in iri ro r) r-l (^) ro 



-- to 

-5 s ^ 



>) S S3 



5i ^' 



5; 2 



1994 



GRENADA BANK SPHAERODACTYLUS 



11 



t3 

(U 

3 

C 

o 
U 



u 

< 



> 



Q 
Q 



c 
o 

X 



00 

d 


00 

d 


d 




« 


* 


^ 




ri 


G" 


+1 


+1 


+1 


+1 


o 
+1 


+1 
1^ 


+1 




+1 


+1 


O 




•^ 


r- 


IT) 


u-i 


00 




ON 


oo 


O 




in 


ON 


vO 


t~- 


ON 




rl 


o 


00 




^ 


U-) 


rr 


in 


r- 




Os 


m 



—I rsi — ' — ' <N * '"i '^. ~: "^1 

+1 +1 +1 +1 +1 "=' +1 +i +1 +1 

TT m fN — — ' +1 r<^ r- Tt (N 

7~'T7777i7T 

r<-ir<-i — — 'OOOn-^'*'^— _ 



in 

d 


d 


00 

d 


d 




'I- 


d 




+1 


+1 
m 


+1 


+1 


+1 


+1 


+1 


+1 


o^ 


S- 


oo 


r- 


00 


^ 


00 


00 


o 


r- 


ON 


00 


00 


o 

1 


ON 

1 


ON 

1 


ON 


nO 


r- 


r~- 


t^ 00 


oo 


oo 


r~- 



(^) 


rT 


^^ 


fN 


m 


r<-) 


^ 


""^ 


^ 


ri 


o 


O 


o 


o 


o 


O 


o 


o 


O 


O 


+1 


+ 1 


+1 


+! 


+1 


+1 


+1 


+1 


+1 


+1 


m 


'i; 


<N 


in 


r- 


00 


m 


^_ 


■^_ 


ro 


in 


VO 


— 


i^ 


o 


o 


■^ 


'^ 


in 


NO 


— ' 


— ' 


»N 




<N 


n 




1 


1 


1 


q 




q 


m 


r<^ 


rr 


r| 


r<-i 


r^ 


r] 



^. 'iv s 









S -5 



I 



to -2 



C a c 
"^ s: o 
5 '2 ~ 



12 BREVIOJ14 No. 496 

onstrate statistical significance (despite lack of overlap) for the 
scale size distinction from kirbyi (Student's r-test; 95 percent level 
of confidence). 

If a subjective assessment of overall resemblance be evidence 
of actual relationship, one might argue that S. kirbyi is more 
closely related to the S. vincenti forms of southern Martinique 
than to any other extant taxa. At least I cannot disprove that 
hypothesis with available evidence. 

EVOLUTION AND BIOGEOGRAPHY 

On both morphological and geographic grounds Sphaerodac- 
tylus kirbyi fits into the group including S. vincenti and S. molei 
(King, 1962). However, Haas (1991) has shown that molei is 
remote from the Antillean species biochemically. It is not sur- 
prising, therefore, that S. kirbyi is in no appreciable way inter- 
mediate between vincenti and molei: it does not unite these spe- 
cies. Within Sphaerodactylus vincenti there are nine forms named 
as subspecies. S. kirbyi resembles S. molei in lacking keeled gulars, 
but S. vincenti josephinae, from southwestern Martinique, and S. 
V. adamas from adjacent Rocher du Diamant, may also have 
smooth (or virtually smooth) gulars. Indeed, Schwartz (1964) ar- 
gues that sphaerodactyls from this area have given rise to all the 
rest of the vincenti complex. A depiction modified from Schwartz's 
(1964) deployment scheme is my Figure 3 A. 

King (1962) presented a far simpler view, arguing that sphaero- 
dactyls had moved up into the Lesser Antilles from an ancestral 
stock represented by S. molei today. As long as nominate S. 
vincenti from St. Vincent was the proximate form, closest in both 
geography and morphology to S. molei, this position was readily 
defensible. Insertion of morphologically divergent 5". kirbyi into 
an intermediate geographic position immediately complicates this 
picture. Indeed, many of us over the years have been uncom- 
fortable with a South American origin for southern Lesser An- 
tillean sphaerodactyls (E. E. Williams, A. Schwartz, G. Mayer, 
all personal communication). Our feeling seems vindicated by the 
biochemical approach of Haas (1991). 

Many might suggest a cladistic analysis would resolve at least 
the temporal sequence of lineage divergence in this group. This 
seems to me to be a superb example of just why and how cladistic 



1994 



GRENADA BANK SPHAERODACTYLUS 



13 




Figure 3. Scenarios for the dispersal and deployment of Sphaerodactylus of 
the vincenti-kirhyi-molei complex. Only the banks, which are islands at glacial 
maximum, and the continental shelf, mainland at glacial maximum, are shown. 
SA is South America. G is Grenada; V, St. Vincent; L, St. Lucia; M, Martinique; 
and D, Dominica. Some small banks, now submerged, but potentially important 
islands at glacial maximum, are shown. A is elaborated from the notions of 
Schwartz (1964) and B derives from the view of King (1962). 



analysis fails to satisfactorily resolve relationships within the spe- 
cies group. There is simply no rational way of assigning polarities 
to the relevant characters. One may assume that small, granular 
dorsals are primitive in Sphaerodactylus, and larger granules, 
eventually evolving to imbricate scales, are derived. Barbour (1921) 
explicitly stated this and King (1962) followed suit. Accepting 
this notion would mean I could assume with facility that the large 
scales of kirbyi (and those southern Martinique forms ofvincenti) 
are derived within the vincenti complex. Common sense, in com- 
bination with knowledge of habitat and of evolutionary trends in 
other lizards, leads me to reject this notion in this case. Large 
scales seem to be selected for in dry habitats (MacLean, 1985). 



14 BREVIOR.4 No. 496 

Thus, kirbyi occupies a more xeric terrain than nominate vincenti 
or molei. To the north, the dry country races of S. vincenti tend 
to have large scales, but the picture is not perfect. 

The largest scales in any 5". vincenti race are those of -S. v. 
adamas of Rocher du Diamant, off the southwest comer of Mar- 
tinique, and an arid little cay. Adjacent S. v. josephinae, of more 
mesic but still fairly dry southwestern Martinique, has scales near- 
ly as large. However, S. v. psammius, of xeric Point des Salines 
and adjacent southeastern Martinique, and S. v. ronaldi, of Mar- 
tinique's arid Presqu'ile de la Caravelle, have significantly smaller 
scales. The form from northern, upland country on Martinique, 
originally rain forest, is 5". v. pheristus, and it has by far the smallest 
scales. 

Thus, among allopatric variants on Martinique we see that a 
very dry country form has large scales and the form from the 
wettest terrain has the smallest scales, but two forms from the 
most xeric habitats are intermediate. Gene exchange probably has 
retarded selection towards large scales in S. v. psammius and S. 
V. ronaldi —IsLVgely peninsular forms— while isolation has per- 
mitted S. V. adamas on its dry, steep, little islet to evolve to the 
extreme of large scale size for the group. 

Further, S. v. pheristus of the erstwhile rain forest has probably 
been selected for small scales. In other respects such as elaborate 
color pattern, large size, and extreme carination, it may be the 
most modified ("derived") of the lot. Thus scale size cannot be 
assigned polarity; it is highly plastic and selected to fit specific 
needs in a geographically varying range. 

Off Martinique, 5". v. diamesus of the very dry northeast coast 
of St. Lucia has rather large scales, but they are only slightly 
smaller in S. v. monilifer from the sodden interior of Dominica. 
In all, the situation is suggestive of selection, but not unequivocal. 

Similarly, scale carination presents no clear picture. Gular keels 
seem the norm for S. vincenti, but S. v. josephinae may lack them, 
and they may be virtually absent in S. v. adamas. It is facile to 
argue (as does Barbour, 1921, and by implication King, 1962) 
that keels are advanced and smooth scales primitive. Forms in 
genetic continuity vary— as do individuals in some panmictic 
populations— with respect to the presence and degree of keeling. 



1994 GRENADA BANK .S7'//.^£-/?OAfrr)7.C'S 15 

SO selection may go in either direction, and apparently has 
done so. 

Color patterns, and even specific pattern elements, defy clear 
assignments of polarity. The light, dark-bordered cephalic bands 
amalgamating into a scapular V, and the light, dark-bordered 
sacro-caudal V, are characteristic of widespread, scattered, and 
seemingly unrelated Sphaewdactylus. They are present in S. kir- 
byi, nominate S. v. vincenti, and variably present in the other 
races of vincenti. Both pattern components are conspicuously 
present in such geographically and morphologically remote spe- 
cies as 5". macrolepis in the Virgin Islands of the Greater Antilles, 
and both are quite absent from 5'. molei, which has bands that 
do not amalgamate to form Vs. 

Ocellate patterns facilitate individual recognition in dark hab- 
itats, notably in rain forest Anolis lizards. Ocellae are present in 
those 5". vincenti from the original rain forest areas: S. v. pheristus 
of upland Martinique and S. v. monilifer of Dominica. In rudi- 
mentary form, ocellae are also incipient in some S. v. vincenti 
from mesic St. Vincent (King, 1962, fig. 12A). They are promi- 
nently present in S. v. ronaldi from arid, xeric Presqu'ile de la 
Caravelle and in 5". macrolepis (at least females) from the most 
parched islets in the Virgins, such as Watson Rock. Whether 
ocellae characterized the very first Sphaewdactylus or not is pres- 
ently unknowable (but not terribly unlikely). 

Sphaewdactylus kirbyi may be derived from S. v. vincenti, or 
more directly from an ancestral stock on Martinique. Of the two 
scenarios depicted in Figure 3, I prefer A, the one derived from 
Schwartz (1964). Although Haas (1991) did not have S. vincenti 
(or S. kirbyi) available for analysis, her evidence strengthens this 
view. The genus Sphaerodactylus seems an Antillean autochthon. 
Of course an aboriginal, ancestral sphaerodactyline must have 
first colonized the Antilles; probably the Greater Antilles. It must 
have departed from a continent; possibly tropical North America 
tens of millions of years ago, well before emergence of the Panama 
Land Bridge. Sphaerodactyls may have subsequently dispersed 
outward, over water, as waifs, to Central America, and the Lesser 
Antilles (Haas, 1991). Martinique, with its geologic mix of pre- 
Miocene to Recent igneous extrusive strata (Smith and Roobol, 



16 BREVIORA No. 496 

1990), looks like a fine probable colonization site for the ancestral 
vincenti-kirbyi stock of Sphaerodactylus. It makes a good staging 
area for Lesser Antillean banks southward. Several other Greater 
Antillean stocks have demonstrably colonized the Windward Is- 
lands. Anolis lizards of the bimaculatus group, certainly of Greater 
Puerto Rican ancestry, have extended as far down-chain as Dom- 
inica (Lazell, 1972). Iguana delicatissima, whose closest relative 
seems to be Iguana pingiiis of the Puerto Rico Bank (Lazell, 
1989a), has colonized all the way to Martinique (Lazell, 1973). 
Anolis of the roquet group, a Lesser Antillean and Windward 
Island autochthon, have colonized some banks just off the con- 
tinental shelf of South America (Lazell, 1972). 

Sphaerodactylus and other Antillean forms have demonstrably 
colonized outward across water to mainlands. For example, ex- 
Antillean natural colonizers outnumber ex-continental colonizers 
of the Antilles at least three to two in Florida and southeastern 
North America. Sphaerodactylus n. notatus is endemic to Florida. 
Crocodylus acutus has invaded the Florida peninsula, and Anolis 
carolinensis has expanded all the way to North Carolina and Texas 
(Lazell, 1989b). Other species, often thought to be human intro- 
ductions, may in fact involve natural colonizations: Sphaerodac- 
tylus elegans and the frogs Eleutherodactylus planirostris and Hyla 
septentrionalis are good candidates (Lazell, 1989b). The water 
snake Matrix compressicauda has gone the other way, from Florida 
to Cuba (Lazell, 1989b), and turtles of the genus Chrysemys have 
colonized the Greater Antilles (Williams, 1989). 

At least two stocks of Sphaerodactylus, the species argus and 
S. rosaurae of the copei group, have colonized the Central Amer- 
ican main or shelf islands (Harris and Kluge, 1984). Harris and 
Kluge (1984) say little about the origin of other Central American 
species. Their remark that "more than one line of evolution from 
the Caribbean" may be required for this modest assemblage of 
nine species might be taken as support for a notion of Antillean 
origin (as opposed to just the Caribbean coastal region of the 
main). 

Similarly, Anolis lizards have colonized the Central American 
main: A. mayensis of the sagrei group and A. allisoni of the por- 
catus (or carolinensis) group (Williams, 1976). 

My view oi Sphaerodactylus dispersal and deployment cannot 



1994 GRENADA BANK SPHAERODACTYLUS 17 

be proved or refuted on grounds of present evidence, but it sets 
a clear target for future work. The lines of evidence which need 
to be pursued are likely to be far more expensive than catching 
lizards and counting scales, and will certainly involve biochem- 
istry. Haas (1991) is a firm step in the right direction, but the 
pattern for 5". kirhyi, S. vincenti, and S. molei remains unexam- 
ined. 

Sphaerodactylus kirbyi may have been derived from S. molei 
or a stock ancestral to them both. It would then retain smooth 
scales and have been selected in dry terrain for large scales. Its 
somber pattern might then be either the ancestral condition (with 
molei subsequently evolving stripes), or have resulted from se- 
lection in isolation. 

Alternatively, S. kirbyi may have colonized the Grenada Bank, 
some 4,128 km- at glacial maximum, from southwestern Mar- 
tinique. If it arrived before S. vincenti reached St. Vincent, then 
it may retain ancestral character states. Conversely, it may have 
arrived subsequently (in a geographic sequence that seems efficient 
but is quite unnecessary) and resemble the southwest Martinique 
forms because of a more proximate common ancestry. The strong 
distinctions between S. kirbyi and S. v. vincenti, its closest neigh- 
bor, are closely comparable to the distinctions in the two Anolis 
lizard species pairs which occupy these islands on separate banks 
(Lazell, 1972). In the cases of both Anolis pairs and the Sphaero- 
dactylus pair, I ascribe the distinctions to adaptive selection in 
an arid versus a mesic habitat augmented by dichopatric character 
divergence resulting from failed invasions (Williams, 1969; La- 
zell, 1972:103). 

Why has it taken so long to find sphaerodactyls on the Grenada 
Bank? They are genuinely scarce. I spent only about six hours 
hunting, and found none. Sinclair hunted about six hours each 
day for a week and got only six, in two sets of three, from very 
small areas. Sphaerodactyls are often discontinuously distributed 
in the Windward Islands, and sometimes appear absent from 
seemingly suitable habitat (the gaps on Martinique are an excellent 
case in point). However, S. kirbyi on the Grenada Bank is extreme 
in these respects. The presence of other chthonian and leaf-litter 
lizards such as Bachia and Gymnophthalmus may partially fill the 
Sphaerodactylus niche and mitigate abundance with competition. 



18 BREVIOIL4 No. 496 

Such a view is inevitably controversial, the more so because Bach- 
ia is far from ubiquitous on the Grenada Bank, known only from 
a few islands. Gymnophthalmus may be a newcomer here. Our 
Bequia specimens were the first recorded from the Grenada Bank 
(Lazell and Sinclair, 1990) and G. underwoodi seems to be ex- 
panding its range in the Lesser Antilles. 

Our short sojourn on Bequia resulted in discovery of not just 
S. kirhyi and G. underwoodi, but the first Eleutherodactylus frog 
recorded for the Grenadines (Lazell and Sinclair, 1990): E.john- 
stonei. The biogeographically interesting details were lost in the 
editorially gutted form in which our geographic distribution notes 
appeared. 

Since the type-locality of Eleutherodactylus johnstonei is on 
Grenada, and it is present from St. Vincent to St. Martin, virtually 
throughout the Lesser Antilles (Schwartz and Henderson, 1988), 
its occurrence on the larger Grenadines seems predictable. How- 
ever, the range of this species presents numerous enigmas well 
described by Schwartz (1967). Briefly, there is testimony that this 
species was introduced to Grenada ca. 1885 from Barbados, to 
which it had been introduced ca. 1879. In four weeks search 
Schwartz himself (and Klinikowski— both superior collectors) 
failed to locate Eleutherodactylus in the Grenadines including 
Bequia. In five nights in the Grenadines (one on Bequia), 1 1-16 
June 1964, I heard no frogs. 

On 8 December 1989 Mr. and Mrs. Thomas Johnston, resident 
on Bequia, reported to Sinclair that they brought about 1 2 frogs 
from the Botanical Garden above Kingstown, St. Vincent, to 
Bequia in 1969. They testified that there were none on Bequia 
previously. These were reportedly released ca. 4 km west of 
Friendship Bay. Males were calling each night we were present at 
Friendship Bay at a density of ca. 5-6 per hectare. Despite this 
evidence, there is reason to doubt human introduction as the 
source of these populations: people catch and transport few, if 
any, females, and frogs, albeit native, may be cryptic for years. 
This species presents a singularly attractive opportunity to test 
molecular clocks such as mtDNA and elucidate biogeography. 

The Grenada Bank, largest in the Windward Islands, is struc- 
turally and ecologically very complex, requiring comparison even 
to the Puerto Rico Bank of the Greater Antilles. As our efforts of 



1994 GRENADA BANK SPHAERODACTYLUS 19 



a few days suggest, there is probably much left here to be dis- 
covered. 

ACKNOWLEDGMENTS 

This work is dedicated to the memory of Albert Schwartz. 

We are indebted to Mr. Brian Johnson, Forestry, and Mr. Alan 
Cruickshank, Minister of the Interior, Government of St. Vincent 
and the Grenadines for permits to collect, and to Christian Lu- 
ginbuhl for assistance in the field. Our trip was funded in part by 
the David B. Luginbuhl Foundation and The Conservation 
Agency. 

LITERATURE CITED 

Barbour, T. 1921. Sphaerodactylus. Memoirs of the Museum of Comparative 
Zoology, 47: 217-278. 

Haas. C. A. 1991. Evolution and biogeography of West Indian Sphaerodactylus 
(Sauria: Gekkonidae): A molecular approach. Journal of Zoology, London, 
225: 525-561. 

Harris, D. M. 1 982. The Sphaerodactylus (Sauria: Gekkonidae) of South Amer- 
ica. Occasional Papers of the Museum of Zoology, University of Michigan, 
704: 1-31. 

Harris, D. M., and A. G. Kluge. 1984. The Sphaerodactylus (Sauria: Gek- 
konidae) of Middle America. Occasional Papers of the Museum of Zoology, 
University of Michigan, 706: 1-59. 

JiNKJNS, D., AND J. BoBROw. 1985. St. Vincent and the Grenadines: A Plural 
Country. New York, W.W. Norton and Co. 126 pp. 

King, W. 1 962. Systematics of the Lesser Antillean lizards of the genus Sphaero- 
dactylus. Bulletin of the Florida State Museum, 7(1): 1-52. 

Lazell, J. 1972. The anoles (Sauria: Iguanidae) of the Lesser Antilles. Bulletin 
of the Museum of Comparative Zoology, 143(1): 1-115. 

. 1973. The lizard genus Iguana in the Lesser Antilles. Bulletin of the 

Museum of Comparative Zoology, 145(1): 1-28. 

. 1989a. (Review) Phylogenetic systematics of Iguanine lizards. Copeia, 

1989(3): 807-809. 

. 1989b. Wildlife of the Florida Keys. Washington, D.C., Island Press. 

xvi + 253 pp. 

Lazell, J., and T. Sinclair. 1990. Geographic distribution: Eleutherodactylus 
johnstonei. Anolis trinitatis. Gymnophthalmus underwoodi. Sphaerodactylus 
cf Vincent t. Herpetological Review 21(4): 95-97. 

MacLean, W. p. 1985. Water loss rates oi Sphaerodactylus parthenopion (Rep- 
tilia: Gekkonidae), the smallest amniote vertebrate. Comparative Biochem- 
istry and Physiology, 82A: 759-761. 

Schwartz, A. 1 964. A review of Sphaerodactylus vincenti on the southern Wind- 
ward Islands. Caribbean Journal of Science, 4(2&3): 391-409. 



20 BREVIOR.4 No. 496 



1967. Frogs of the genus Eleutherodactylus in the Lesser Antilles. Studies 



on the Fauna of Curacao and Other Caribbean Islands, 24(91): 1-62. 
Schwartz, A., and R. W. Henderson. 1988. West Indian amphibians and 

reptiles: A checklist. Milwaukee Public Museum Contributions in Biology 

and Geology, 74: 1-264. 
Smith, A. L., and M. J. Roobol. 1990. Mt. Pele, Martinique, a study of an 

active island-arc volcano. Geological Society of America Memoir, 175: vii + 

105 pp. 
Williams, E. E. 1969. The ecology of colonization as seen in the zoogeography 

of anoline lizards on small islands. Quarterly Review of Biology, 44(4): 345- 

389. 
. 1976. West Indian anoles: A taxonomic and evolutionary summary. 1. 

Introduction and a species list. Breviora, 440: 1-21. 
. 1989. Old problems and new opportunities in West Indian biogeography, 

pp. 47-102. In C. A. Woods (ed.), Biogeography of the West Indies: Past, 

Present, and Future. Gainesville, Florida, Sandhill Crane Press, vii + 878 pp. 



B R E V I O R A. 

MuseiLim of Comparative Zoology o 

us ISSN 0006-9698 
Cambridge, Mass. 2 February 1994 Number 497 



CHELID TURTLES OF THE AUSTRALASIAN 

ARCHIPELAGO: I. A NEW SPECIES OF CHELODINA 

FROM SOUTHEASTERN PAPUA NEW GUINEA 

Anders G. J. Rhodin' 

Abstract. A new species of Chelodina (Testudines: Pleurodira: Chelidae) is 
described from the Kemp Welch River drainage basin. Central Province, south- 
eastern Papua New Guinea, where it occurs in a restricted distribution. It is 
endemic to Papua New Guinea and isolated from other members of the genus. It 
is superficially most similar to Chelodina novaegiiineae of southwestern Papua 
New Guinea, but osteologically more closely related to C. longicollis of eastern 
Australia. The recently described species Chelodina reimanni from southeastern 
Irian Jaya, Indonesia, is most closely related to C. novaeguineae. 

INTRODUCTION 

The side-necked turtles of the family Chelidae (Testudines: 
Pleurodira) from the New Guinean region of the Australasian 
Archipelago remain one of the most poorly known turtle faunas 
of the world. The zoogeography and the diversity of the snake- 
necked chelid turtle genus Chelodina that inhabits this region of 
eastern Indonesia and Papua New Guinea have received some 
attention, but are far from fully resolved. Until the latter part of 
this century only two species of Chelodina were known from the 
regions north of Australia: Chelodina novaeguineae Boulenger, 
1888 and Chelodina siebenrocki Werner, 1901. Chelodina no- 
vaeguineae was known only from southwestern Papua New Guin- 
ea and adjacent southeastern Indonesian Irian Jaya, as well as 
from Roti Island west of Timor in Indonesia, with C siebenrocki 
supposedly occurring only in "Deutsch-Neu-Guinea," the former 
German colony comprising the northern half of present-day Pa- 
pua New Guinea. In 1976 Rhodin and Mittermeier described the 



' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts, and Chelonian Research Foundation, Lunenburg. Massachusetts. 



2 BREVIORA No. 497 

new Species Chelodina parked from the inland grass-swamps of 
Lake Murray and the Aramia River of southwestern Papua New 
Guinea. They also redescribed and clarified the distributional 
status of C siebenrocki, which is endemic to the estuarine swamps 
of the southern trans-Fly region of southwestern coastal Papua 
New Guinea and adjacent southeastern Irian Jaya, and absent 
from the northern half of Papua New Guinea, where the type 
specimen had allegedly been collected. Subsequently, Philippen 
and Grossman (1990) described Chelodina reimanni from the 
coastal regions near Merauke, southeastern Irian Jaya, but did not 
compare their new species to the very closely related C. novae- 
guineae and provided no osteological description. Both C. rei- 
manni and C. novaeguineae belong to the subgeneric group of 
Chelodina species that have relatively narrower heads, shorter 
necks, and broader plastrons designated as Chelodina spp. "A" 
(Goode, 1967; Burbidge et al, 1974; Legler, 1985). Conversely, 
C. parkeri and C. siebenrocki are both members of the subgeneric 
group with relatively broader heads, longer necks, and narrower 
plastrons designated as Chelodina spp. "B". 

Very little other work has appeared on the taxonomy or natural 
history of New Guinean and Australasian Chelodina. Gaffney 
(1977) performed a phylogenetic analysis of all Chelidae based 
primarily on cranial osteology and included some New Guinean 
taxa. Cann (1978) contributed a photographic documentary of 
some of the species, and Rhodin and McCord (1990) documented 
some reproductive parameters of Chelodina siebenrocki. Wells 
and Wellington (1985) created a host of destabilizing nomencla- 
torial novelties in their contentious catalogue of Australian rep- 
tiles, which has been severely criticized by the International Com- 
mission on Zoological Nomenclature (ICZN, 1991). In addition, 
very little research has been done on the New Guinean short- 
necked chelid genera Emydura and Elseya. In particular, Mc- 
Dowell's (1983) recent work is not generally accepted because of 
its sweeping synonymies (King and Burke, 1989), and Legler and 
Cann (1980) restricted their work to continental Australian taxa. 

Over the last 20 years, I have been privileged in having access 
to the large Papua New Guinean chelid turtle collections obtained 
by Fred Parker and deposited at the Museum of Comparative 
Zoology by Ernest E. Williams. Early analysis of this material has 



1994 NEW CHELODINA FROM PAPUA NEW GUINEA 3 

led to the description of Chelodina parkeri and redescription of 
C. siebemocki (Rhodin and Mittermeier, 1976). Continued study 
of these collections has subsequently been combined with exten- 
sive comparative analysis of specimens obtained from other mu- 
seums and professional colleagues as well as from personal col- 
lecting trips to Papua New Guinea and Indonesia. To date, I have 
been able to examine personally approximately 700 chelid turtles 
from the eastern Indonesian and New Guinean regions of the 
Australasian Archipelago, and 450 specimens from continental 
Australia, for a total database of about 1,150 Australasian Chel- 
idae. Through this study, I have reached several taxonomic con- 
clusions regarding the chelid taxa of the Australasian Archipelago. 
In this, the first in a series of papers documenting these conclu- 
sions, I describe a remarkable new species of Chelodina from 
southeastern Papua New Guinea. In addition, I provide the first 
osteological description of C reimanni, and compare both these 
species with C novaeguineae and C longicollis. 

In 1985, through the courtesy of Dr. Peter C. H. Pritchard, I 
received two specimens of what had been assumed to be Chelo- 
dina novaeguineae from the vicinity of Hula, Kemp Welch River 
drainage, 90 km southeast of Port Moresby, southeastern coastal 
Papua New Guinea. These two specimens would have represented 
an enormous range extension for C novaeguineae, which occurs 
primarily in the Western Province of southwestern Papua New 
Guinea, southeastern Irian Jaya, and northeastern Australia. The 
Kemp Welch population is disjunct and separated from the reg- 
ular range of C novaeguineae by about 500 km of relatively well- 
collected coastal territory including the whole developed Port 
Moresby region. From a preliminary examination of Pritchard's 
two specimens I was convinced that they represented a new and 
distinct species. In 1987 I traveled to Papua New Guinea's Kemp 
Welch River area and succeeded in obtaining a third specimen 
of the same taxon at Bore, Kemp Welch River. In addition, I was 
able to examine a fourth specimen preserved in the Papua New 
Guinea Museum without locality data other than "near Port 
Moresby". Later, through the courtesy of Dr. William P. McCord 
I had the fortunate opportunity to examine an additional large 
series of 39 live specimens obtained from just east of Bore in the 
Kemp Welch River area, for a total study sample of 43 animals. 



4 BREVIOKA No. 497 

These 43 Kemp Welch River specimens were then compared 
to a series of 5 1 C. novaegidneae from southwestern Papua New 
Guinea, 10 C. novaeguineae from northern Australia, 54 C lon- 
gicollis from eastern Australia, 12 C. reimanni from southeastern 
Irian Jaya, Indonesia, and 7 C steindachneri from western Aus- 
tralia, for a total study series of 177 specimens. Analysis of ex- 
ternal morphology and cranial osteology demonstrated that the 
isolated Kemp Welch population of Chelodina was an unde- 
scribed species. It is much more closely related to C. longicollis 
of Australia than it is to either New Guinean or Australian pop- 
ulations of C. novaeguineae. I now describe this new species and 
name it after Dr. Pritchard, who obtained the first two specimens 
and brought them to my attention. 

TAXONOMY 

Chelodina pritchardi, sp. nov. 
(Figs. 1-3 and Table 1) 

Holotype. MCZ 173543, alcohol-preserved sub-adult male of 
129.5 carapace length, purchased from native villagers by Anders 
G. J. Rhodin on 14 August 1987 at Bore, Kemp Welch River, 
1 3 km southeast of Kwikila, Central Province, Papua New Guinea 
(9°53'S, 147°46'E); specimen is also former AGJR-T 1259 and 
bears old tags RZ Field- 13602 and AMNH 133079. 

Paratypes. MCZ 175813 (former PCHP 1343) and AMNH 
139735 (former PCHP 1342), obtained from natives by Peter C. 
H. Pritchard at Port Moresby in 1978, said to be from vicinity 
of Hula, Kemp Welch River drainage basin, 32 km south of 
Kwikila, Central Province, Papua New Guinea (10°06'S, 147°43'E). 

Referred Specimens. PNGM 23373, collected by natives "near 
Port Moresby" (possibly in Kemp Welch River drainage basin), 
Central Province, Papua New Guinea, died in captivity at Moi- 
takaZoo, Port Moresby; AGJR-T 1575-1609, 1643-6, ca. 10 km 
east of Bore, Kemp Welch River region. Central Province, Papua 
New Guinea (39 specimens from Dr. William P. McCord's private 
live collection, photographed and measured by Rhodin, 13 of 
these (AGJR-T 1601-9, 1643-6) preserved in Rhodin's personal 
collection, others (AGJR-T 1575-1600) recorded in Rhodin's 
turtle database; all preserved specimens eventually to be deposited 
in the MCZ or other museum collections). 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 



s 

E 



On 




3 « 
O 



BREVIORA 



No. 497 




Figure 2. Lateral head view of Holotype of Chelodina pritchardi (MCZ 1 73543). 
Note the striking Ught-colored iris. 



Distribution. The species is known definitively only from the 
Kemp Welch River drainage basin southeast of Port Moresby, 
Central Province, Papua New Guinea, an area of relatively high 
rainfall and mesic lowland alluvial forests (Fig. 4). It appears to 
be absent from the Port Moresby region itself, an area of low 
rainfall and xeric savannah vegetation. It is not yet known whether 
the range also extends further southeast along the coast toward 
Cape Rodney and Abau, where appropriate wetland habitat also 
occurs. The species may additionally occur in the mesic lowland 
coastal plain northwest of Port Moresby in the Laloki River area, 
where reports tentatively suggest its presence. 

Diagnosis. A medium-sized New Guinean snake-necked chelid 
turtle oi Chelodina subgeneric group "A" (Burbidge et al., 1974) 
with relatively narrow head and wide plastron (Fig. 5), superfi- 
cially resembling C novaeguineae but more closely related to 
Australian C longicoUis. Plastral and head widths intermediate 
between C novaeguineae and C. longicollis (Fig. 6). Skull oste- 
ology distinctive: narrow maxillary and mandibular triturating 
surfaces with decreased anterior skull robusticity; decreased tem- 
poral fossa muscular volume with increased parietal roof width. 
Skull most similar to C. longicollis, markedly divergent from C. 
novaeguineae and C. reimanni, both of which have significantly 
more robust skulls. Eye unicolor light tan, pale plastron often with 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 






Figure 3. Dorsal, ventral, and lateral views of skull of Chelodina pritchardi 
(Paratype AMNH 1 39735, adult female measuring 1 93 mm carapace length) from 
nr. Hula, Kemp Welch River, Central Province, Papua New Guinea. 



darker color pattern following midline sutures broadly. Eggs rel- 
atively smaller than any other Chelodina. 

Etymology. The specific epithet is a patronym honoring Dr. 
Peter C. H. Pritchard, well-known turtle authority, close profes- 
sional colleague, and personal friend, who serendipitously ob- 
tained the first two specimens while on a marine turtle consultancy 
in Papua New Guinea. 



BREMOR.A 



No. 497 



Table 1 . Basic external dimensions of Chelodina pritchardi. All mea- 
surements IN MM. CL = carapace LENGTH (STRAIGHT-LINE IN MIDLINE); CW = 
CARAPACE WIDTH (GREATEST); CD = CARAPACE DEPTH (GREATEST IN MIDLINE); PL-M 
= PLASTRON LENGTH (ALONG MIDLINE, NOT INCLUDING ANAL SPURS); PL-T = 
PLASTRON LENGTH (MAXIMUM, INCLUDING ANAL SPURS); PW = PLASTRON WIDTH 
(at axillary NOTCH); HW = HEAD WIDTH (TYMPANIC). 



Specimen 


















Number 


Sex 


CL 


CW 


CD 


PL-M 


PL-T 


PW 


HW 


AGJR-T 1643 


f 


125.0 


96.5 


40.0 


98.0 


104.5 


59.5 


20.2 


MCZ 173543 


m 


129.5 


100.0 


41.0 


103.5 


109.5 


60.5 


20.5 


AGJR-T 1583 


J 


134.7 


106.5 


44.4 


106.0 


112.4 


63.3 


21.0 


AGJR-T 1601 


m 


136.0 


106.0 


44.5 


108.0 


114.5 


64.5 


21.2 


AGJR-T 1608 


m 


139.0 


112.5 


45.5 


112.0 


117.5 


67.0 


21.8 


AGJR-T 1589 


m 


140.0 


111.0 


45.0 


111.5 


118.4 


66.5 


21.5 


AGJR-T 1593 


m 


140.5 


112.0 


47.0 


111.5 


118.0 


68.0 


22.0 


AGJR-T 1592 


m 


141.0 


108.0 


44.5 


112.0 


118.0 


65.0 


21.6 


AGJR-T 1600 


m 


142.5 


107.5 


45.5 








21.8 


AGJR-T 1602 


m 


143.5 


111.0 


47.5 


111.0 


119.0 


68.0 


22.0 


AGJR-T 1588 


m 


145.0 


113.5 


46.3 


115.0 


122.0 


68.2 


22.4 


AGJR-T 1590 


m 


145.5 


113.7 


48.2 


115.5 


122.0 


70.0 


22.5 


AGJR-T 1599 


m 


146.5 


115.0 


46.5 








22.5 


AGJR-T 1585 


m 


150.0 


117.0 


49.0 


115.0 


124.0 


70.5 


22.3 


AGJR-T 1607 


m 


150.0 


115.0 


48.0 


115.0 


123.0 


72.0 


22.6 


AGJR-T 1577 


m 


150.7 


117.0 


47.0 


116.0 


125.0 


71.0 


22.4 


AGJR-T 1591 


m 


152.0 


121.0 


50.5 


119.0 


127.5 


72.0 


22.5 


AGJR-T 1644 


m 


153.5 


125.0 


50.0 


121.0 


129.0 


73.5 


22.9 


AGJR-T 1580 


m 


156.0 


125.0 


51.4 


122.0 


131.0 


74.5 


23.0 


AGJR-T 1609 


m 


156.5 


124.5 


48.0 


122.0 


131.0 


74.0 


23.5 


AGJR-T 1575 


m 


156.8 


123.3 


52.8 


121.0 


129.8 


75.4 


23.9 


AGJR-T 1594 


m 


157.0 


125.0 


53.0 


121.5 


129.0 


75.0 


24.3 


AGJR-T 1595 


f 


157.0 


121.5 


52.5 


123.0 


131.0 


74.5 


23.8 


AGJR-T 1646 


f 


157.0 


128.5 


55.0 


125.5 


132.5 


76.5 


23.0 


AGJR-T 1587 


f 


157.5 


121.0 


49.0 


119.5 


128.0 


74.0 


23.4 


AGJR-T 1645 


m 


160.0 


126.5 


52.0 


124.5 


133.0 


79.0 


23.8 


AGJR-T 1582 


m 


160.2 


122.5 


51.2 


121.7 


131.2 


75.5 


24.3 


AGJR-T 1586 


m 


160.5 


123.5 


51.5 


122.5 


133.0 


75.5 


24.2 


AGJR-T 1598 


m 


161.0 


131.0 


52.5 








23.8 


AGJR-T 1606 


m 


161.0 


125.5 


53.0 


123.0 


133.0 


77.5 


23.8 


PNGM 23373 


f 


162.0 


118.0 


52.0 


126.0 




76.0 


25.0 


AGJR-T 1605 


m 


163.5 


131.5 


54.5 


128.5 


139.0 


79.0 


24.8 


AGJR-T 1581 


m 


163.8 


127.8 


54.2 


127.0 


137.7 


78.0 


24.6 


AGJR-T 1603 


m 


165.0 


128.0 


53.5 


126.5 


136.5 


77.0 


24.7 


AGJR-T 1576 


m 


165.4 


131.5 


51.8 


126.3 


137.3 


78.0 


24.5 


AGJR-T 1604 


m 


168.0 


128.5 


56.0 


127.0 


136.5 


78.0 


25.3 


AGJR-T 1596 


f 


171.0 


133.0 


57.5 








25.0 



1994 NEW CHELODINA FROM PAPUA NEW GUINEA 



Table 1. Continued. 



Specimen 
















Number Sex 


CL 


CW 


CD 


PL-M 


PL-T 


PW 


HW 


AGJR-T 1597 f 


175.5 


138.0 


59.0 








25.5 


AGJR-T 1584 m 


186.4 


143.8 


63.7 


140.0 


151.5 


87.8 


27.5 


AMNH 139735 f 


193.0 


156.0 


62.0 


154.0 




93.5 


27.0 


AGJR-T 1578 f 


199.3 


161.4 


65.5 


157.5 


166.0 


97.4 


28.4 


AGJR-T 1579 f 


206.0 


167.0 


69.5 


158.0 


169.0 


99.0 


29.0 


MCZ 175813 f 


228.0 


180.0 


90.0 


183.0 




107.0 


31.0 



Related Ta.xa. Chelodina pritchardi is most similar to the fol- 
lowing four chelid taxa from New Guinea and Australia (all mem- 
bers of Chelodina subgeneric group "A"). 

Chelodina longicolHs (Shaw, 1794). Original designation Testudo longicoUis. Type 
locality "New Holland" [=Australia]. Holotype BMNH 1947.3.5.86, a dry 
specimen of 1 34 mm carapace length. 

Chelodina novaegiiineae Boulenger, 1888. Original designation Chelodina novae- 
gidneae. Type locality "Katow, S.E. New Guinea" [= Mawatta, Binaturi River, 
Western Province, Papua New Guinea]. Original syntypes BMNH 1 946. 1 .22.36 
and MCG CE 8407, collected by L. M. d'Albertis. BMNH 1 946. 1 .22.36, a sub- 
adult female of 137 mm carapace length figured by Boulenger, 1889 (plates 5, 
6) and photographed in this paper (Fig. 7) is hereby confirmed as lectotype 
(previously designated by Wells and Wellington, 1985, p. 8). 

Chelodina reimanni Philippen and Grossman, 1990. Type locality "Merauke- 
River, West-Irian, Neuguinea" [= Merauke River, Irian Jaya, Indonesia]. Ho- 
lotype MTKD 29178, adult female of 180 mm carapace length, collected by 
Frank Yowono, 1988. 

Chelodina steindachneri Siebenrock, 1914. Type locality "Marloo Station am Grey 
River in Westaustralien" [Western Australia, Australia]. Holotype NMW 19798 
(Naturhistorisches Museum Wien). 

DESCRIPTION 

External Morphology 

Carapace. Carapace of C pritchardi smooth and broadly oval, 
width averaging 78.5% of length, moderately flared posteriorly 
with marginals 6-9 somewhat expanded. No lateral marginal re- 
curving. No vertebral knobs or keel. Slight vertebral flattening or 
shallow furrow in some specimens. No supracaudal notch or mar- 
ginal serrations. Dorsal nuchal long and broad, slightly protruding 
beyond anterior carapace edge in smaller specimens. Ventral un- 



10 



BREVIORA 



No. 497 




Figure 4. Top: Distribution of Chelodina pritchardi, C. novaegiiineae. and C. 
longicollis. 1. New Guinean distribution of C novaeguineae in Irian Jaya and 
Papua New Guinea. 2. Australian distribution of C. novaeguineae in Northern 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 



11 



5.5 



4.5 



fe 3.5 



X 



2.5 



1.5 



OChcItxJiiiJi 
• ( 'hckxJma 
 ChekxJiti;i 



spp "A" 
spp "B" 
pntchardi 



OOo 



m_^ o 



^ 



•:« 



= • . * • • •— ». *• • 

: • • .• • * . 



Narrower I lead 
Wider Plastron 



Wider Head 
Narrower Plastron 



50 100 150 200 250 

Carapace Length (mm) 



300 



350 



400 



Figure 5. Scattergram plotting the relationships of head width ratio (Carapace 
Length/Head Width) and plastron width ratio (Plastron Width/Plastron Length), 
expressed as a bivariate product, versus carapace length for the subgeneric groups 
of Chelodina {Chelodina "A" = C. longicollis, C. novaeguineae, C. pritchardi, C. 
steindachneri, and C. reimanni, Chelodina "B" = C expansa, C. oblonga, C. 
parkeri, C. rugosa, and C siebenrocki). Note the position of C. pritchardi within 
the Chelodina "A" generic grouping. Note also the three specimens of subgeneric 
group "A" within the group "B" area; these represent extremely broad-headed C 
reimanni. 



Territory and Queensland. 3. Distribution of C. pritchardi in Papua New Guinea. 
4. Approximate northern limit of the distribution of C longicollis in eastern 
Australia. Major watershed limits indicated as heavy dotted lines. Bottom: Area 
3 of top map enlarged here and showing distribution of Chelodina pritchardi in 
the Port Moresby region. Central Province, Papua New Guinea. The shaded area 
represents elevation above 200 m, the heavy dotted line shows the watershed 
limit of the Owen Stanley Ranges. Starred locality (1) is Port Moresby. Black dots 
represent localities for C pritchardi in the Kemp Welch River basin, as follows: 
(2) Bore, Kemp Welch River; (3) Hula; (4) ca. 10 km east of Bore. 



12 



BREVIORA 



No. 497 



5.5 



4.5 



0. 



4 



a 3 



2.5 



2 



1.5 



o C longict>llis 

A C novacguiiicac 

 C pritclKirdi 

A C reimanni 



Narrower I lead 
W idcr Plastron 



o o o 
o 



Wider Head 
Narrower Piaslroii 



o^ 



o 
o o 

OOo 






o 



o 



A 
A O 



^"k A 



o 



o 



o 



^AA' 



Aa^ 



o 



25 50 75 100 125 150 175 

Carapace Length (mm) 



200 



225 



250 



Figure 6. Scattergram plotting the relationships of head width ratio (Carapace 
Length/Head Width) and plastron width ratio (Plastron Width/Plastron Length), 
expressed as a bivariate product, versus carapace length in four species of Chel- 
odina. Note that C. pritchardi is essentially intermediate between C. longicollis 
and C. novaeguineae, but more similar to C longicollis. 



derlap of nuchal also relatively long and broad. Vertebral 1 widest, 
then 2, 3, 5, and 4 in descending order of width. Carapace some- 
what broader in C pritchardi than in New Guinean C novae- 
guineae, intermediate in C. longicollis, significantly narrower in 
C reimanni (see Fig. 8 and Table 2). 

Carapace moderately deep in older specimens, relatively flatter 
in younger ones. No sexual dimorphism in carapace depth in 
specimens examined. Carapacial scutes lightly rugose with par- 
tially retained growth lines until mid-adult size. 

Color dark chestnut brown. Carapace very similar to both C. 
novaeguineae and C longicollis, but generally more broad than 
C novaeguineae and slightly less broad than C. longicollis. Car- 
apace superficially more similar to C novaeguineae than to C 
longicollis (Figs. 1 and 7). 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 



13 




<-• 
00 




c 




_aj 








(U 




o 




« 




a 




« 




k. 




ca 




o 




s 




s 




[^ 




ro 




^^ 




00 




c 




•c 


• 


3 


a 


i« 


<u 




c 


£ 


'3 


O 


_aj 




13 


^ 




Z 


Ci 






03 


1— • 


3 


3 


O. 


■o 


^ 


rt 


CU 


X) 


iT 


3 


CJ 


(« 


c 






^' 


> 


m 


o 




u, 


j^i 


Cl, 


(N 






e 




aj 


^ 


-*-* 


OS 




X 


uT 


2 


> 




'cS 




'U 


<ij 


3 


!3 


■*-» 


s 


C3 


s: 


C 


■| 


m 


<;j 


oT 


a 


■*— » 


>: 


*-• 


o 






Ki 




S 






~« 


II 


_o 








"S 


rt 


-c 


!U 


U 


a 


<*- 


'3 


o 


O 


K 


^ 


>. 


(D 


o 


z 


^-» 




4J 


w 


hJ 


C/3 




sT 


r-' 


o 




♦-• 


S-* 


a 


3 


^ 


00 


B 
o 



14 



BREVIORA 



No. 497 



.8 



.78 



.76 



J. 74 
£.72 



.68 



.66 



.64 











 




 
 

 
 


 

 

 

 
 


 

 A 
 
A  


 


 




 




A C- novaegurneac 
 C- pritchardi 
A C- rcimaiiiii 






A A A 






A 


 A 

A 






A 














A 

A 


A 


A 

A ^ ^ 
A A 

A^ ^^^ 


A 
A 

A 


 
^A 

A 


A 

A 
A 

^ A 
A 




A 








Broader 


t 




Carapace 


I 




A 


A "^ 








▲ 




















A 






A 






A 





80 



100 



120 



200 



220 



240 



140 160 180 

Carapace Length (mm) 

Figure 8. Scattergram plotting carapace width ratio (CW/CL) versus carapace 
length in three species of Chelodina. utilizing data only from New Guinean spec- 
imens of greater than 100 mm carapace length. Note the broader carapace in C 
pritchardi. 



Plastron. Plastron broad, axillary width averaging 61% of mid- 
line length, anterior lobe moderately broad, intermediate between 
the relatively narrow tapered lobe of C. novaeguineae and C. 
reimanni, and the broadly expanded anterior lobe of C longicollis 
(Fig. 6 and Table 2). Slight secondary expansion of anterior plas- 
tral lobe at posterior border of humeral scutes, not present in C 
novaeguineae. Anal notch moderately deep, no sexual dimor- 
phism noted. Intergular broad, long, and recessed without mar- 
ginal contact. Plastral scute suture length formula: Ig > An > 
Abd > Pec > Fem > Gul. No axillary or inguinal scutes. 

Plastral color yellow with variable amount of light to moderate 
brown pigment following central portions of sutures broadly, of- 
ten expanding to cover much of central plastron. Holotype plas- 
tron oxidized to darker brown. Plastral color different from C. 



1994 NEW CHELODINA FROM PAPUA NEW GUINEA 15 



Table 2. Means and standard deviations for shell measurement ratios 

OF FOUR ChELODINA SPECIES. ABBREVIATIONS AS IN TaBLE 1 . DATA BASED ONLY ON 
specimens OF CARAPACE LENGTH GREATER THAN 1 00 MM. C N01.4EGUINEAE INCLUDES 

ONLY New Guinean specimens, no Australian ones. 









C. novae- 






C. longicollis 


C pritchardi 


guineae 


C reimanni 


Feature 


(n = 37) 


(n = 43) 


(n = 51) 


(n = 5) 


CW/CL 


.775 ± .042 


.785 ± .018 


.723 ± .028 


.690 ± .022 


PW/PL-M 


.618 ± .026 


.610 ± .012 


.576 ± .018 


.543 ± .028 


HW/CL 


.151 ± .012 


.150 ± .005 


.167 ± .010 


.202 ± .012 


CD/CL 


.310 ± .026 


.328 ± .014 


.327 ± .021 


.315 ± .007 



novaeguineae, which usually has an immaculate yellow plastron 
with occasional very thin pigment lines following the sutures, and 
from C. longicollis, which usually has broad black color zones 
along the sutures and sometimes over most of the plastron. Plas- 
tron superficially more similar to C novaeguineae than to C 
longicollis (Figs. 1 and 7). 

Head and Soft Parts. Head with small irregular scales covering 
temporal skin, smooth over parietal and interorbital roof. Neck 
with low soft tubercles, less pronounced than the larger raised 
firmer tubercles of C novaeguineae. Soft parts grayish-brown dor- 
sally, yellowish-white ventrally. Hands and feet with 4 claws each. 

Head width narrow, typical of Chelodina subgeneric group "A" 
species (Fig. 5), intermediate between the wider heads of C. no- 
vaeguineae and C. reimanni and the narrower head of C. longi- 
collis (Fig. 6). Relative width of head narrows ontogenetically. 

Eye color of C pritchardi primarily light tan with medium dark 
tan thin area at outer periphery of iris and very light tan thin 
inner rim, becoming nearly white along the pupillary edge of the 
iris. No color flecks or cross-bar. Eye color of C. novaeguineae 
from Papua New Guinea (personal observation) and Australia 
(Cann, 1978: plate 19) dark brown with more sharply distinct 
yellowish-white pupillary rim around inner iris, and dark area of 
iris with multiple small irregular flecks of darker and lighter pig- 
ment. Overall impression of eye color of C. pritchardi unicolor 
whitish-tan, of C. novaeguineae bicolor dark brown with inner 
yellow circle. 



16 



BREVIORA 



No. 497 






Figure 9. Dorsal, ventral, and lateral views of skull of Chelodina reimanni 
(AGJR-T 746, 199 mm carapace length female from Merauke, Irian Jaya, In- 
donesia). 



Size and Sexual Dimorphism. The largest specimen of C prit- 
chardi recorded is a female of 228 mm carapace length. The largest 
male examined has a carapace length of 1 86 mm, indicating prob- 
able sexual dimorphism, with females larger than males. Calcu- 
lating the sexual dimorphism index according to the method of 
Gibbons and Lovich (1990) yields an SDI value of approximately 
1.22 for C phtchardi. The SDI value for New Guinean C no- 
vaeguineae is approximately 1.37, with the largest confirmed fe- 
male measuring 207 mm and the largest male 151 mm. 

The largest specimen of New Guinean C novaeguineae I ex- 



1994 



NEW CIIELODINA FROM PAPUA NEW GUINEA 



17 






Figure 1 0. Dorsal, ventral, and lateral views of skull ofChelodina novaegnineae 
(AGJR-T 504, 1 78 mm carapace length female from Boze, Binaturi River, Western 
Province, Papua New Guinea). 



amined was a female measuring 207 mm, but Philip Hall (per- 
sonal communication) has photographed and measured a 2 1 8 mm 
specimen from the Irian Jaya-Papua New Guinea border. The 
largest specimen of Australian C. novaeguineae I examined mea- 
sured 279 mm, but Cann ( 1978) records 300 mm as the maximum 
size. The largest specimen of C. reimanni I examined measured 
199 mm, but Philippen and Grossman (1990) records 206 mm 
as the maximum size. In general, C. pritchardi is larger than New 
Guinean C. novaeguineae or C. reimanni, and smaller than Aus- 
tralian C novaeguineae. 



18 



BREMOR.A 



No. 497 






Figure 1 1 . Dorsal, ventral, and lateral views of skull of Chelodina longicollis 
(AMNH 108952, from Patho, Victoria, Australia). 



Osteology 

Skull The description of skull osteology is based on the ex- 
amination of 6 skulls of C pritchardi. Comparison is performed 
with skulls of 7 C. longicollis, 1 5 C. novaeguineae ( 1 2 from New 
Guinea, 3 from Australia), and 2 C reimanni. Refer to Figures 
3 and 9-1 1 for skull illustrations of the four species and Tables 
3 and 4 and Figures 12-14 for additional skull measurements and 
ratios. 

The skull of C. pritchardi is a typical Chelodina subgeneric 
group "A" type skull, not overly elongate, flattened, or wide as 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 



19 



.24 






















C longicollis 








a C novacgumcac 














.22 




 C pnlchardi 
A C reiniaiini 






A 




A 




£ 










A 


















C 






k 








^ .2 








A 


A 




^i* 






A A 








"3 


A ^ 


A 


A 








^.18 




























£ 














■^ 


















•a 




Broader 


i I 




















^ .16 




Tnturatino 
Surface 














U) 








_c 




 
















 


















 




U .14 














3 












 


■^^ 


O 


























L. 














H 


o 












.12 

1 





o 


% 









80 100 120 140 160 180 

Carapace Length (mm) 



200 



220 



240 



Figure 1 2. Scattergram plotting maxillary triturating surface width ratio (TW/ 
SL) versus carapace length in four species of Chelodina. Note the much broader 
triturating surface in C novaeguineae and C retmanni with C. pritchardi being 
somewhat intermediate and C longicollis much narrower. 



in subgeneric group "B". It is strikingly similar to the skull of C 
longicollis, from which it is differentiated by only a few features. 
It differs markedly from its more geographically proximate con- 
geners C novaeguineae and C reimanni. 

The major differentiating features involve the width and ro- 
busticity of the triturating surfaces and the relative volume of the 
muscular temporal fossa. C novaeguineae and C. reimanni have 
wide and robust maxillary and mandibular triturating surfaces, 
with correspondingly wide and robust homy rhamphothecae. C 
longicollis has very narrow and weak surfaces and C. pritchardi 
is intermediate (Fig. 1 2), but more similar to C. longicollis. The 
mandibular coronoid process is high and prominent in C. no- 
vaeguineae and C. reimanni, low and less prominent in C. lon- 
gicollis, and intermediate in C pritchardi. C. novaeguineae and 



20 



BREVIOR.4 



No. 497 



X. 

S 



180 



160 



140 



120 



100 



•J so 

1/1 

s 



40 



20 



oc 


longicotlis 


A f  


noxacguincac 


 C 


priltluirdi 


*c 


reimaiuu 


M 


ore 


< I 


Robust 




Skull 





A A 



O 



A 



OO 



o 



o o 



80 100 120 140 160 ISO 200 

Carapace Length (mm) 



220 



240 



Figure 13. Scattergram plotting skull Robusticity Index [RI = (TW x SWT 
X SD)/SL] versus carapace length in four species of Chelodina. Note the inter- 
mediate position of C. pritchardi with reference to the other species. 



C reimanni have a long midline maxillary suture, reflecting the 
increased width of the maxillary triturating surface; both C. lon- 
gicollis and C. pritchardi have short sutures. 

The relative position of the choanae is posterior in C novae- 
guineae and C. reimanni, anterior in C. longicollis and C prit- 
chardi, once again as a result of the widened triturating surface. 
The vomer is more robust, wider, and reaches further posterior 
in C. novaeguineae and C reimanni than in C. longicollis or C. 
pritchardi. It reaches the level of the palatine foramen and sep- 
arates the palatines widely in C. novaeguineae and C reimanni, 
does not reach the level of the palatine foramen and only barely 
separates the palatines in C. longicollis and C. pritchardi. In C. 
novaeguineae and C. reimanni the pterygoids do not extend an- 
teriorly along the midline to meet the vomer anterior to the pal- 
atine foramen, in C. longicollis and C. pritchardi the pterygoids 
do extend anteriorly. 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 



21 





.22 




















^^ 






















£ 


.2 




















▲ 




«-> 








ex) 






o C loiigicollis 




















e 






A c" no\ acguiiieac 




















.18 




 r pnlckirdi 


























* (■ reiniaiini 














































MM 








3 


.16 




















.:^ 






















t/} 






















E 


.14 


























S 






(ircatcr Rclali\c Mass 


' ^ 




















Skull plus .Mandible 


















1 ■> 




















E 


. 1 L 




















-25 
























.1 










▲ 


























A 






W 






















S 


.08 








A 










 


__ 














A 








"3 










A 






 






u 


.06 








A ^ 












m 






















> 


.04 




A 


 


 o" 

O 












'-3 




A 


O 






































— 


.02 


o o 


















0^ 


r\ 


O 



















80 100 120 140 160 180 

Carapace Length (mm) 



200 



220 



240 



Figure 14. Scattergram plotting relative skull mass (skull and mandible weight 
in gms/skull length in mm) versus carapace length in four species of Chelodina. 
Note the intermediate position of C. phtchardi with reference to the other species, 
with C. reimanni and C. novaeguineae having heavier, more robust skulls. 



The shape of the anterior skull tomial edge also differs, with 
C novaeguineae and C reimanni having a somewhat rounded, 
blunted shape, and C. longicoUis and C. pritchardi slightly more 
angular. The premaxillae are usually fused into a single small 
premaxilla in New Guinean C. novaeguineae (11 of 1 2 skulls) and 
C. reimanni (2 of 2); unfused but very small in Australian C. 
novaeguineae (3 of 3), and unfused and large in C. longicoUis and 
C pritchardi. Premaxillary foramina are absent in New Guinean 
C. novaeguineae and C reimanni, reduced but present in Aus- 
tralian C novaeguineae, and well-developed in C longicoUis and 
C. pritchardi. 

The ventral view of the skull reveals that the posterior extension 
of the quadrate beyond the posterior extension of the opisthotic 
differs between the species. C novaeguineae, C. reimanni, and C 
pritchardi have prominent quadrate extensions, C. longicoUis does 



Table 3. Basic skull measurements for four species of Chelodina. SL = 
skull length (snout-occipital condyle); swt = skull width, tympanic 
maximum; SWM = skull width, maxillary maximum; SDM = skull depth at 

POSTERIOR edge OF MAXILLAE; SD = SKULL DEPTH IN MIDLINE BETWEEN SUPRAOC- 
CIPITAL SPINE AND BASISPHENOID; lOW = INTER-ORBITAL WIDTH, MINIMAL; OW = 
ORBITAL WIDTH, SHORT AXIS; PtW = PTERYGOID WIDTH, MINIMAL; TW = TRITURAT- 
ING WIDTH, MAXILLARY (MEASURED IN MIDLINE FROM TOMIAL EDGE TO ANTERIOR 
CHOANAL BORDER). REFER TO TABLE 4 AND FIGURES 1 2 AND 1 3 FOR ANALYSIS OF 

SKULL MEASUREMENT RATIOS. 



Species 




















Mus. Sp. No. 


SL 


SWT 


SWM 


SDM 


SD 


lOW 


OW 


PtW 


TW 


Chelodina longicollis 


















AGJR-T 159 


25.2 


16.1 


12.8 


5.9 


7.1 


2.4 


6.0 


8.8 


2.6 


MCZ 8369 


26.4 


16.8 


12.4 


5.0 


6.7 


2.5 


5.5 


8.7 


3.5 


MCZ 8377 


27.1 


17.9 


13.0 


6.0 


7.0 


2.4 


5.8 


9.4 


3.4 


AGJR-T 158 


33.6 


21.6 


17.0 


7.3 


8.9 


3.4 


7.0 


11.1 


3.9 


MCZ 86783 


35.5 


23.4 


18.6 


7.6 


9.2 


3.5 


7.5 


11.0 


4.2 


AGJR-T 179 


36.3 


23.2 


18.9 


8.2 


9.2 


4.3 


7.8 


12.0 


4.4 


AMNH 108952 


40.5 


25.1 


20.2 


8.2 


10.1 


3.5 


8.3 


13.9 


4.6 


Chelodina novaegidneae 


















MCZ 134394 


26.3 


17.8 


13.3 


6.6 


8.8 


3.0 


6.3 


8.4 


4.9 


AMNH 57589 


31.2 


21.0 


16.3 


8.0 


10.3 


3.6 


7.5 


9.7 


5.9 


MCZ 134712 


32.5 


22.4 


16.5 


7.7 


10.2 


3.8 


7.3 


10.5 


6.0 


UU 14716 


33.7 


23.0 


18.2 


7.2 


9.5 






11.2 


6.3 


MCZ 134391 


35.0 


23.3 


17.7 


8.2 


11.7 


4.6 


7.0 


10.5 


7.2 


AMNH 117939 


35.7 




18.5 




11.5 


4.3 


7.5 


11.2 


6.9 


MCZ 134395 


35.7 


24.0 


18.0 


9.0 


11.2 


4.7 


7.8 


11.8 


6.9 


MCZ 134390 


36.5 


24.4 


18.9 


8.5 


11.3 


4.6 


7.6 


11.5 


7.6 


MCZ 134392 


37.0 


26.0 


20.0 


9.5 


13.0 


4.8 


8.6 


12.2 


7.9 


AGJR-T 504 


38.0 


26.4 


20.7 


9.0 


12.4 


4.8 


8.0 


11.4 


7.6 


MCZ 134393 


38.0 


26.0 


20.0 




12.0 


5.0 


8.0 


12.0 


7.1 


MCZ 134396 


38.0 


25.0 


19.0 




11.6 


4.4 


8.0 


11.0 


7.3 


MCZ 142495 


41.0 


27.6 


22.2 


9.6 


14.4 


5.6 


8.9 


12.5 


8.2 


AMNH 86547 


46.0 


34.0 


27.8 


10.6 


14.2 


5.6 


9.8 


15.5 


8.4 


AMNH 86544 


50.7 


37.3 


30.3 


12.2 


15.4 


5.4 


10.3 


16.2 


9.5 


Chelodina pritchardi 


i 


















AGJR-T 1608 


32.2 


20.8 


18.0 


7.6 


9.8 


3.3 


7.5 


10.6 


4.8 


AGJR-T 1607 


34.1 


21.6 


18.7 


7.7 


9.8 


3.6 


7.9 


11.3 


5.0 


AGJR-T 1606 


35.5 


22.8 


19.8 


8.3 


10.5 


4.1 


8.4 


12.1 


5.1 


AGJR-T 1605 


37.0 


23.8 


20.5 


8.8 


10.7 


4.0 


8.4 


12.1 


5.3 


AMNH 139735 


39.5 


26.5 


21.5 


8.8 


11.0 


4.5 


8.5 


13.6 


5.7 


MCZ 175813 


44.5 


29.0 


24.8 


10.0 


12.6 


4.5 


10.5 


15.8 


6.0 


Chelodina reimanni 




















AGJR-T 1614 


42.9 


30.7 


23.1 


10.6 


16.0 


6.5 


9.0 


13.4 


9.3 


AGJR-T 746 


47.7 


39.5 


28.4 


13.5 


19.4 


7.3 


10.6 


12.2 


10.3 



1994 NEW CHELODINA FROM PAPUA NEW GUINEA 23 



Table 4. Means and standard deviations for skull measurement ratios 
OF four Chelodina species. Abbreviations as in Table 3. 









C novae- 






r. longicollis 


C. pritchardi 


guineae 


C. reimanni 


Feature 


(n = 7) 


(n = 6) 


(n= 15) 


(n = 2) 


SWT/SL 


.642 ± .014 


.648 ± .013 


.687 ± .024 


.774 ± .076 


SWM/SL 


.501 ± .020 


.553 ± .006 


.532 ± .032 


.568 ± .039 


SD/SL 


.260 ± .011 


.290 ± .009 


.320 ± .018 


.388 ± .026 


TW/SL 


.119 ± .009 


.144 ± .005 


.194 ± .009 


.218 ± .003 


PtW/SWT 


.521 ± .027 


.522 ± .014 


.461 ± .018 


.372 ± .090 


lOW/OW 


.456 ± .052 


.470 ± .037 


.570 ± .055 


.705 ± .024 



not. This represents a major difference between the otherwise 
somewhat similar skulls of C. pritchardi and C. longicollis. The 
pterygoid trochlear processes are prominent and markedly di- 
vergent in C. novaeguineae and C reimanni, with New Guinean 
specimens exhibiting prominent flaring, while Australian ones 
exhibit none; the processes are minimally divergent and much 
less prominent in C. longicollis and C pritchardi. The ventral 
view of the skull shows the flared pterygoid processes are very 
prominent in New Guinean C. novaeguineae and C reimanni, 
less prominent in the other species. Also, on the ventral view of 
the skull, the postorbital portions of the jugal and postorbital are 
well seen in C. novaeguineae and C reimanni, but not in C. 
longicollis or C pritchardi. 

The parietal roof extent and shape differ markedly between the 
four species. Chelodina reimanni has an extremely narrow parietal 
crest, with nearly complete temporal emargination, C. novaegui- 
neae also has an extremely narrow parietal crest, but with very 
slightly less emargination, C longicollis has a fairly wide trian- 
gular parietal roof, with much less temporal emargination, and 
C. pritchardi is intermediate in both roof extent and temporal 
emargination. In C. reimanni the frontal enters the temporal 
emargination border, in C novaeguineae and the other species it 
does not. The height of the supraoccipital crest above the foramen 
magnum is extremely high in C reimanni, high in C novaegui- 
neae, low in C longicollis, and intermediate in C pritchardi. The 
volume of the temporal fossa (occupied by the mandibular ad- 



24 BREVIORA No. 497 

ductor muscle mass) is extremely large in C. reimanni, large in 
C novaeguineae, smaller in C longicoUis, and intermediate in C 
pritchardi. 

The lateral view of the skull reveals that the relative positions 
of the postorbital wall strut and of the anterior edge of the brain 
case differ in the four species. In C reimanni and C. novaeguineae 
the postorbital wall is relatively caudad and overlaps the anterior 
brain case, giving increased stability and strength to the anterior 
third of the skull. In C. longicollis the postorbital wall is further 
cephalad, making it possible to look directly through the skull 
between the wall and the anterior brain case, and providing less 
strength and stability to the anterior skull. In C. pritchardi this 
relationship of the postorbital wall and the anterior edge of the 
brain case is intermediate. 

All four species share the following skull osteological charac- 
teristics typical of other Chelodina: frontals fused, prefrontals 
separated by frontals, nasals present, dentaries sutured, splenials 
present, exoccipital contact above foramen magnum, temporal 
arch absent, and chelid foramen absent (variably present in ru- 
dimentary form in C longicollis) (see McDowell, 1 983; this "chel- 
id foramen" is also called the posterior pterygoid foramen by 
Legler, personal communication). 

Most of the differences in skull osteology reflect the increased 
robusticity of the skulls in C novaeguineae dind C. reimanni, with 
C. longicollis being the least robust, and C. pritchardi being in- 
termediate. These differences can be calculated and demonstrated 
graphically as a Robusticity Index (RI; see Fig. 1 3). This Index 
reflects the cumulative effects of increased triturating surface width, 
overall tympanic skull width, and skull depth for adductor muscle 
volume, and is determined by the formula that follows. 

_,^ _TW X SWT X SD 
^^' SL 

In this formula, TW = maxillary triturating surface width, SWT 
= tympanic skull width, SD = midline skull depth, and SL = 
skull length. The Robusticity Index increases ontogenetically with 
size and age, and is significantly different in the four species. 
Increased robusticity is directly related to the increased width of 



1994 NEW C//ELOD/M4 FROM PAPUA NEW GUINEA 25 

the maxillary and mandibular triturating surfaces and accom- 
panying skeletal modifications of the skulls. These modifications 
reflect the secondary requirements for increased posterior skull 
bracing resistance and increased muscular mass to allow for in- 
creased mandibular muscle adductor force generation. These skel- 
etal modifications have also created heavier and more massive 
skulls in the more robust species, which can additionally be dem- 
onstrated through an analysis of relative skull and mandible mass 
(grams per mm skull length) versus carapace length (Fig. 14), 
where C. novaeguineae and C reimanni have heavier skulls than 
C. longicollis, and C. pritchardi is once again intermediate. 

Based on skull osteology, C reimanni and C novaeguineae are 
probably dependent on a mollusciform and gastropod diet re- 
quiring extensive crushing of hard food matter. The diets of C. 
longicollis and C. pritchardi are probably more generalized car- 
nivorous or piscivorous, with less dependence on hard-shelled 
bivalves and snails. 

Some differences were noted between skulls of New Guinean 
versus Australian C. novaeguineae. However, full analysis of ex- 
ternal morphological differences was not undertaken, and only a 
few Australian specimens were available for complete study. It is 
premature to evaluate whether these populations are distinct or 
not, and they are treated here as conspecific. 

Cervical Spine. Central cervical articulation pattern is 
(2(3(4(5)6)7(8) in 5 specimens (4 by direct exam, 1 by X-ray), 
the only known pattern for all Chelidae as described by Williams 
( 1 950). Atlanto-axial (C 1 and C2) cervical morphology is identical 
in all four species: C. pritchardi, C. novaeguineae, C. reimanni, 
and C. longicollis. 

Shell. No neural bones in 7 specimens, all pleurals meeting in 
the midline. Axillary buttress moderately robust, articulating with 
lateral first pleural and posterior third peripheral, inguinal buttress 
less robust, articulating with postero-lateral edge of fourth and 
antero-lateral edge of fifth pleurals, and anterior seventh periph- 
eral. Suprapygal relatively wide, contacting tenth peripheral. One 
specimen with atypical ten peripherals on each side, rather than 
normal eleven. Broad contact between first peripherals and first 
pleurals. 



26 



BREVIORA 



No. 497 





30- 




sicbcnrocki A 










"expansa" 




28- 






A 


'i" 








A 


E 


2f)- 




ZA parkeri 


expansa 


'--^ 










4= 






A rugosa 














■5 


24- 






 


^ 












22- 




A oblonga 




u 










00 

w 

u 
u 

> 






"longicollis" 




20- 


prilchardi 


(^ ^) novacguincae 




< 




o 


longicollis 






18' 




o 






16- 




steindachneri 
• 1 ' 1 " 1 ' 1 f— 


1 > 1 



26 



28 



30 



32 



34 



3(1 



40 



Average Egg Length (mm) 



Figure 15. Plot of average egg width versus average egg length in all species 
of Chelodina. Circles represent subgeneric group "A" species; triangles, subgeneric 
group "B" species. Solid symbols are Legler's (1985) composite groups. See Table 
5 for supporting data. 



Ecology and General 

Reproduction. Two specimens, obtained from Hula, each had 
eggs. The larger female (CL 228 mm) laid 4 eggs in captivity in 
Florida, one was broken, the other three measured 27.9 x 19.2, 
27.8 X 18.0, and 27.0 x 18.6 mm. The smaller female (CL 193 
mm) had one shelled oviducal egg measuring 26.4 x 19.7 mm 
when dissected post-mortem. All eggs were white, oval, with hard, 
brittle shells. Eggs are smaller than any other species of Chelodina 
(Table 5 and Fig. 1 5), but similar in shape to most other Chelodina 
(Fig. 1 6). Compared to the size of the adult female, the eggs laid 
by C pritchardi are proportionately extremely small (Fig. 1 7). 

Growth. The sub-adult male holotype has prominent concentric 
wide growth zones evident on carapacial and plastral scutes in- 
dicating rapid juvenile growth. No larger adults noted with similar 
growth evidence. 

Sympatry. Chelodina pritchardi occurs sympatrically with 
Emydura subglobosa in the Kemp Welch River drainage basin. 
Both of these species are commonly eaten by the local inhabitants 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 



27 



q.^ 



.2 .85 






oo 



u 

J 75 



■5 ,7 



.55 



26 



/\ parke 



/\ siebenrocki 



prilchardi 

o 



/\njgosa 



oblonga 



"cxparisa" 

AA 



cxpansa 



steindachncri 



longicollis 

Q^"longicollis" 



novaegumeac 



28 



30 32 34 36 

Average Egg Length (mm) 



40 



Figure 1 6. Plot of egg width/length ratio versus average egg length in all species 
of Chelodina. Circles represent subgeneric group "A" species; triangles, subgeneric 
group "B" species. Solid symbols are Legler's (1985) composite groups. See Table 
5 for supporting data. 



and often kept in the villages. The giant softshell turtle Pelochelys 
bibroni occurs in the Laloki River in the Port Moresby region 
and may be sympatric with C. pritchardi, which has tentatively 
been recorded from the same area. 

Vernacular Names. In the inland regions of the Kemp Welch 
River area, the local language is Sinaugoro (Guise, 1985). All 
freshwater turtles are known as gaokori, but C. pritchardi and E. 
subglobosa do not have different names, despite the fact that the 
villagers readily distinguished them as being different. The Sin- 
augoro name for marine turtles is gaogao. In the coastal regions 
the local language is Keapara (Guise, 1985) and only one ver- 
nacular name, aoao, refers to both marine and freshwater swamp 
turtles (see also Rhodin et ai, 1980). 

DISCUSSION 

Chelodina pritchardi is in most ways more closely related to its 
geographically distant Australian congener C. longicollis than it 
is to the more geographically proximate New Guinean C novae- 



28 



BREVIOJ14 



No. 497 



u 

3 
U 



X 

u 
a 
u 

N 



60 
SO 



120- 






i ' 1 1 1 1 1 


T 


110- 




 novacguincac 
# pritchardi 






A rugosa 






O 100- 




▼ sicbcrirockj 




T 


V. 




n parkcri 

O steindachiicri 




T 

A 

O 


^ 90- 




3 oblonga 




^ 80- 






* 




D 




J 70. 




o 




OO 

> 

■< 60- 

50- 


 o 

°o 
o 


3 

 

• • 




40- 




1 • 1 I 1 I 1 





140 



160 



180 



200 



220 



240 



260 



280 



300 



Carapace Length (mm) of Female 

Figure 17. Plot of egg-size index of individual clutches (avg. egg length x avg. 
egg width/ 10) versus carapace length of female that laid the clutch. Smaller index 
indicates smaller eggs. Data based on information from sources in Table 5. Note 
that for its body size, C. pritchardi has eggs relatively smaller than the other taxa. 



guineae or C. reimanni. A number of shared osteological features 
of the skull suggest a close phylogenetic relationship between C 
pritchardi and C. longicollis, and I regard them as more closely 
related to each other than either is to any other species. Super- 
ficially, however, based on only external morphology, C prit- 
chardi appears more similar to C novaeguineae than to C. lon- 
gicollis. 

Chelodina novaeguineae is more similar to C pritchardi than 
it is to C longicollis, but its most closely related congener is C. 
reimanni. This latter species was described in 1990 by Philippen 
and Grossman, but they failed to describe the osteology or to 
compare their species to other New Guinean or Australian Chel- 
odina. I have had an opportunity to examine several specimens 
of C reimanni, including two osteological preparations (AGJR-T 
746, a female of 199 mm carapace length from Merauke, Irian 
Jaya obtained via Michael Reimann and Walter Sachsse, and 
AGJR-T 1614, a female of 170.5 mm carapace length from Mer- 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 



29 



o < 



o 
< 

0. 



< 

s 

X 

< 



u 

i 

Q 
O 

a 

O 

u 

U 
u 

</} 
yj 

D 
O 

< 
> 

o 



O w 



H 
Z 



m 



i 






I. 
1. 




5 




J 


Q 
Z 

< 


> 




UJ 


"^ 


V) 


^ 
^ 


s 


-J 





8 


g 



O 



< 

< 

Q 

UJ 

Z 

3 

o 
u 

u 



£ UJ 



UJ 

-J 
O 

UJ 

O O 



X 
H 
Q 

% 
Q 
Z 
< 



Q 
a; 
< 
Q 
Z 
< 

\ii \ri M 

m +1 UJ 



X 

H 
G 
Z 



o: 



UJ 

D 
-) 

w < 

-1 > 



O 
oa 

UJ 

X 



< 









c 
-J 



00 
00 

W 



-J 

u 



o 

a 
en 



3 
*— » 

C 



3 

■«-» 
C 
(D 



On 

O v," 
ON «J 

•a s 

o -^ 



OS 
00 
OS 



u 

3 



-g "^ s' i -^' 

eg C OS C no 

c -3 "" 



oo 

OO 

^ 00 

= °° .s s; . 

t« OS — ii, t« 

M o -- 



x: » ^ 
0^ UJ 



•C 



00 



^ 5 c 5 



I/-) 

oo 
OS 



"ob 



00 

oo 
OS 



oo 

c 
o 

3 



o 

00 
OS 



c 

C 
., « 

OS (J 

OS ^ 

C «J 

1) "^ 



OS 
OS 



> 



_«J 


ir> 




oo 


5 


OS 


^^ 


^^ 


u- 


, 


4J 


u 


s 


_aj 




"ob 


13 


<u 


cu 


J 



i^i^ooos— .t~~ooOir-iO Tt— "■^Ooo'/^<N<^ .00—; 

OOSOO— <'— 'OI~~-'^'^'_,OOrsi— 'OOso— '"-^^Os 

^^ rsi m (N rsl'—' r*^ ro — <Ni| (N 

+1 I +1 I +1 I +1 I +1 \ ^ M I +11+11 +1 I +' +1 I 

00— .Os-H00fNSO<N<N— . vOrsll^— 'OO— lOs — ii^O— ' 



^oo^O-~-Oi-vO 

oso'^t^'^fNOm 



r~- — I <N 

I +1 I 

Tt Csl 



00 fo csi — m 



+1 



I 

so 



CO 

I 



+1 

o 



» rn 



(^) — ^fNinmoofNrnm 



rxi 



r^OSTtOsosOoOOO 

■^ •^ ro f*^ XI 

+1 I +1 I +1 I +1 I +' 

1/-^ r^ "Ti o 

^ r-; ^i d ^ t-' ^ 

OO m OS CO OS <N O 

r<-) ^^ r<-i  — • oi  — - r«-i 



R (N 



OO ^ 

00 «^ 

<^ d 

O fN 



'^ 


m 


o 


^ so 


'^ 


O 


"* 


OS r-4 


so 


' — ' 


00 


-^ •<1- 


o 


rn 


in 


so oo 
— ' <^) 



00 
(N 
(N 



o 


r-- 


r~ 


Tf >n 


in 


as 


SO 


^ 


O r- 


r- 


CN 


rj 


<^4 


m m 


r<-i 



O 


o 


OS 


^ 


Tl- 


r~- 


(^) 


f^l 


oi 



I 

s 



U U 






o 



U 









u u 








^ 






c 


.to 


a 


-s: 




to 


^ 


^ 


K 


a 





a 


-<3 


u 


g. 


s: 


So 


^ 











u u u u u 



30 BREVIORA No. 497 

auke obtained via Frank Yowono and William McCord). Chel- 
odina reimanni is most similar to C novaeguineae and the features 
of increased skull robusticity seen in C novaeguineae are further 
amplified in C. reimanni. Its skull is massively enlarged and heavy, 
with wide triturating surfaces, a narrow parietal crest, large tem- 
poral fossa, and increased buttressing in a deep, wide skull. It has 
small fused premaxillas and lacks chelid foramina. I regard C. 
novaeguineae and C. reimanni as more closely related to each 
other than either is to any other species, and from my preliminary 
examination, I also recognize C reimanni as being distinct from 
C. novaeguineae. 

Phylogenetically, I consider enlarged maxillary and mandibular 
triturating surfaces and increased skull robusticity as derived fea- 
tures within the Chelodina lineage. Though clearly diet-related, 
these features represent a significant specialization by only a few 
members of the genus, notably C. reimanni and C. novaeguineae. 
These shared derived characteristics suggest a close phylogenetic 
relationship between these two species. The narrow triturating 
surfaces and less robust skulls of C. pritchardi and C. longicollis 
represent more primitive features within the genus and suggest 
retained plesiomorphic features in those two species. These prim- 
itive features are also present in the only other member of Chel- 
odina subgeneric group "A", the western Australian species C. 
steindachneri. Examination of skulls of this species demonstrates 
that it is most similar to C longicollis, with narrow triturating 
surfaces, a relatively wide anterior parietal roof, shallow temporal 
fossa, and markedly decreased skull robusticity. Unlike C. lon- 
gicollis, however, it usually retains fairly well-formed chelid fo- 
ramina (posterior pterygoid foramen of Legler), a relatively prim- 
itive feature among all Chelodina. The chelid foramen is absent 
in all other members of Chelodina subgeneric groups "B" and 
"A" except for C. longicollis, where it is variably present but 
usually absent. 

Of the five currently recognized taxa in Chelodina subgeneric 
group "A", I regard C. reimanni as the most derived, and C. 
steindachneri as the most primitive. The other three species fall 
out in a series between these extremes, with C novaeguineae most 
derived, C. longicollis most primitive, and C. pritchardi inter- 
mediate between the two. Two alternate phylogenetic hypotheses 



1994 



NEW CHELODINA FROM PAPUA NEW GUINEA 



31 



rciniiiiim 



novaeguineue pritchardi longicollis steindachneri \ Che lading "B 




Figure 18. Hypothesized relationships of the currently recognized species of 
Chelodina subgeneric group "A". See Figure 19 for an alternative phylogenetic 
hypothesis. 

expressing the possible relationships within Chelodina subgeneric 
group "A" are depicted in Figures 18 and 19, with Figure 18 
representing what I consider the more likely hypothesis. This 
hypothesis would be strengthened through the discovery of shared 



reimanni novaeguijieae pritchardi longicollis steindachneri \Che lading "B" 




Figure 19. Alternative phylogenetic relationships of Chelodina subgeneric group 
"A". See Figure 1 1 for preferred hypothesis. 



32 BREVIORA No. 497 

derived features between C pritchardi and C. longicoUis, and is 
encumbered by the less parsimonious double loss of chelid fo- 
ramina in the novaeguineae-reimanni lineage and the pritchardi- 
longicoUis clade. Clearly, a rigorous cladistic analysis of multiple 
morphologic features of all the species of Chelodina will be nec- 
essary to help further elucidate the phylogenetic history of the 
genus. 

These five taxa oi Chelodina subgeneric group "A" form a well- 
defined monophyletic assemblage clearly differentiated from 
Chelodina subgeneric group "B". Features of shell morphology, 
head width, skull osteology, and cervical spine length and mor- 
phology clearly define the two groups. The currently recognized 
genus Chelodina (sensu lato) is also a clearly defined monophyletic 
assemblage with a long list of shared derived characteristics (see 
Gaffney, 1977). Recognition of this monophyly needs to be in- 
corporated into whatever taxonomic arrangement provides the 
most specific nomenclatorial definition of the subgroups involved. 
Whether the "subgeneric" groups "A" and "B" are best recog- 
nized as subgenera of Chelodina or as full separate genera awaits 
full evaluation by Legler (in preparation). My own analysis of the 
phylogenetic relationships of all the Chelidae of Australasia and 
South America (in preparation) suggests recognition at the generic 
level for these two separate groups of Chelodina, with a new 
suprageneric category replacing our old concept of Chelodina. 

ACKNOWLEDGMENTS 

I thank Peter C. H. Pritchard for his donation of the two spec- 
imens which stimulated this study and William P. McCord for 
his generosity in allowing me to examine and measure his large 
series of live captive animals. I also thank Richard Zweifel, op- 
portunely present in Papua New Guinea when I collected the 
holotype, for facilitating export of the specimen via the American 
Museum of Natural History to the MCZ. Susan and Michael 
Rhodin provided valuable field assistance and Jose Rosado at the 
MCZ curated specimens and helped with logistics. I prepared all 
illustrations. For the loan of other study specimens and general 
assistance with the project, I also thank John Legler, Robert Wi- 
nokur, Arnold Kluge, Allen Greer, Ron Heyer, Sam McDowell, 
Alan Leviton, A. F. Stimson, Philip Hall, Walter Sachsse, Chuck 



1994 NEW C/IELODINA FROM PAPUA NEW GUINEA 33 



Crumly, John Iverson, John Carr, Russ Mittermeier, and Ernest 
WilHams. 

APPENDIX 

Comparative material examined; collection acronyms as fol- 
lows: AGJR-T = personal collection of Rhodin (including pre- 
served specimens, voucher photographs, and reliably documented 
database entries, including data from many live specimens mea- 
sured in the unnumbered personal collection of William P. 
McCord); AMNH = American Museum of Natural History; AMS 
= Australian Museum, Sydney; BMNH = British Museum of 
Natural History; CAS = California Academy of Sciences; FMNH 
= Field Museum of Natural History; MCZ = Museum of Com- 
parative Zoology; MTKD = Museum fur Tierkunde, Dresden; 
PCHP = personal collection of Peter C. H. Pritchard; PNGM = 
Papua New Guinea Museum; SMcD = collection of Samuel Mc- 
Dowell; UMMZ = University of Michigan Museum of Zoology; 
UNLV = University of Nevada, Las Vegas; USNM = United 
States National Museum; UU = University of Utah. 

Chelodina longicollis: Australia: A^^m' South Wales: Armidale (nr.), UNLV s/n 
(15); Sydney, AMNH 14151; Sydney, 60 mi S. UMMZ 1 30 1 6 1 , 1 30549; Talbnager 
R., 8 mi W Varbry, btw. Dunedo and Cassilis, AMS 40828; Victoria: Bright, 
FMNH 75317; Patho, 5 mi S Murray R., 20 mi W Echuca, UU s/n, AMNH 
108952, Queensland: Burnett R., FMNH 16885; Capella, 20 mi W, CAS 77809; 
Duaringa, 15 mi S, CAS 77808; Eidsvold, upper Burnett River, AMS 5979; No 
data: AMNH s/n (2), UU s/n, AGJR-T 133, 158-9, 179, AMNH 2323, MCZ 
8369, 8371-7, 86783-4, USNM 8894, AMNH 9002, 45079, 45085, 75165, 97629, 
110483, MTKD 14605, FMNH 22681, 31047, 35538, BMNH 1947.3.5.86 (ho- 
lotype of C. longicollis), BMNH 1947.3.5.87 (holotype of C. sulcata). 

Chelodina novaeguineae: Australia: Queensland: Alice River, 15 mi WSW 
Townsville. UMMZ 132328; Armraynald, 26 mi SE Burketown, AMNH 86545- 
7. Burdekin River, lower, BMNH 1908.2.25.1; Edward River. Cape York, UU 
14718; Greta Creek, PCHP 2385; Staaten R., 100 mi. N Normanton, AMNH 
86543-4; Northern Territory: Batten Creek, 13 mi WSW Borroloola, UU 14716; 
Indonesia: Irian Java: Kuprik, nr. Merauke (8°25'S, 140°28'E). SMcD 49-1, 49- 
2; Papua New Guinea: Western Province: Abam, Oriomo R. (8°57'S, 143°13'E), 
AMNH 117939, MCZ 120353, 127404, 134390-1, 134709-10, 134712; AH Vil- 
lage, Aramia River (8°05'S, 142°55'E), USNM 213490; Boze, Binaturi River (9°05'S, 
143°0rE), AGJR-T 504; Daru Roads (9°03'S, 143°12'E), MCZ 142500; Emeti, 
Bamu River (7°48'S, 143°15'E), MCZ 138102; Hy River at Strickland River 
junction (7°35'S, 141°25'E), MCZ 53758-61; Giringarede, Binaturi River (9°03'S, 
142°57'E), MCZ 153930; Katatai(9°01'S, 143°18'E), MCZ 138101, 142495, 154340; 



34 BREVIORA No. 497 



Katow(= Mawatta, Binaturi River) (9°08'S, 142°55'E), BMNH 1946.1.22.36 (lec- 
totype C. novaeguineae); Komovai Village, Fly River (7°33'S, 141°15'E) AGJR-T 
1338; Kuru, Binaturi River (8°55'S, 143°04'E), MCZ 134711; Lake Daviumbo 
(7°35'S, 141°17'E),AMNH 59874; Lake Murray (7°00'S, 14!°30'E), MCZ 134392; 
Mabaduane, Pahoturi River (9°! 7'S, 1 42°44'E), AMNH 57589-9 1 , MCZ 137516; 
Masingle, Binaturi River (9°07'S, 142°55'E), AGJR-T 501, MCZ 153046-8, 153906, 
153923, 153926; Morehead (8°43'S, 141°38'E). PNGM 23505; No data, USNM 
231527; Oriomo, Oriomo River (8°52'S, 143°10'E), PNGM 23510; Peawa, Ori- 
omo River (8°55'S, 143''12'E), AMNH 104010; Tarara, Wassikussa River (8°50'S, 
141°52'E). AMNH 58410; Togo, Pahoturi River(9°14'S, 142°40'E), PNGM 23502- 
3, 2351 1, MCZ 134393-6; Ume. Binaturi River (9°03'S, 143°03E), PNGM 22407, 
MCZ 127405; Wipim (8°51'S, 142°55'E), USNM 204856. 

Chelodina phtchardi: Papua New Guinea: Central Province: Bore, Kemp Welch 
River (9°53'S, 147°46'E), MCZ 173543; nr. Hula, Kemp Welch River basin (10°06'S, 
147°43'E). MCZ 175813, AMNH 139735; nr. Port Moresby, PNGM 23373; ca. 
10 km east of Bore, Kemp Welch River, AGJR-T 1575-1609, 1643-6. 

Chelodina reimanni: Indonesia: Irian Java: Merauke (8°25'S, 140°28'E), AGJR-T 
746, 1299-1300, 1325, 1614-1619, 1642; A^o ^a?a.- MTKD 14603. 

Chelodina steindachneri: Australia: Western Australia: Marloo Station, MCZ 
33501; MundabuUangana, MCZ 74871, 134469; 1 mi S. Minilya River on NW 
coastal hwy., MCZ 74872; Woodstock, AMNH 101977-9. 



LITERATURE CITED 

BouLENGER, G. A. 1888. On the chelydoid chelonians of New Guinea. Annali 

del Museo Civico di Storia Naturale di Genova, (2a)6: 449-452. 
BouLENGER, G. A. 1889. Catalogue of the Chelonians, Rhynchocephalians, and 

Crocodiles in the British Museum (Natural History). London, Trustees of the 

Museum. 311 pp. 
BuRBiDGE, A. A., J. A. W. KiRSCH, AND A. R. Main. 1974. Relationships within 

the Chelidae (Testudines; Pleurodira) of Australia and New Guinea. Copeia, 

1974: 392-409. 
Cann, J. 1978. Tortoises of Australia. Sydney, Angus and Robertson. 79 pp. 
Clay, B. T. 1981. Observations on the breeding biology and behaviour of the 

long-necked tortoise, Chelodina oblonga. Journal of the Royal Society of 

Western Australia. 64: 27-32. 
EwERT, M. A. 1985. Embryology of turtles, pp. 75-267. In C. Gans, F. Billett, 

and P. F. A. Maderson (eds.). Biology of the Reptilia. London, Academic 

Press, Vol. 14. 
Fritz, U. and D. Jauch. 1989. Haltung, Balzverhalten und Nachzucht von 

Parkers Schlangenhalsschildkrote Chelodina parkeri Rhodin & Mittermeier, 

1976 (Testudines: Chelidae). Salamandra, 25(1): 1-13. 
Gaffney, E. S. 1977. The side-necked turtle family Chelidae: A theory of re- 
lationships using shared derived characters. American Museum Novitates, 

2620: 1-28. 
Georges, A. 1986. Observations on the nesting and natural incubation of the 



1994 NEW r/ZELOD/A'/l FROM PAPUA NEW GUINEA 35 



long-necked tortoise Chelodina expansa in south-east Queensland. Herpe- 
tofauna, 15(2): 27-31. 

Gibbons, J. W. and J. E. Lovich. 1990. Sexual dimorphism in turtles with 
emphasis on the slider turtle (Trachemys script a). Herpetological Mono- 
graphs, 4: 1-29. 

GooDE, J. 1967. Freshwater Tortoises of Australia and New Guinea (in the 
Family Chelidae). Melbourne, Lansdowne Press. 1 54 pp. 

Guise, A. 1985. Oral tradition and archaeological sites in the eastern Central 
Province. Papua New Guinea National Museum Records, 9: 1-84. 

ICZN. 1991. Decision of the Commission. Three works by Richard W. Wells 
and C. Ross Wellington: proposed suppression for nomenclatural purposes. 
Bulletin of Zoological Nomenclature, 48(4): 337-338. 

King, F. W. AND R. L. Burke. 1989. Crocodilian, Tuatara, and Turtle Species 
of the World. A Taxonomic and Geographic Reference. Washington, DC, 
Association of Systematics Collections. 216 pp. 

KucHLiNG, G. 1988. Gonadal cycles of the Western Australian long-necked 
turtles Chelodina oblonga and Chelodina steindachneri (Chelonia: Chelidae). 
Records of the Western Australian Museum, 14: 189-198. 

Legler, J. M. 1985. Australian chelid turtles: reproductive patterns in wide- 
ranging taxa, pp. 117-123. In G. Grigg, R. Shine, and H. Ehmann (eds.), 
Biology of Australasian Frogs and Reptiles. Royal Zoological Society of New 
South Wales. 527 pp. 

Legler, J. M. and J. Cann. 1 980. A new genus and species of chelid turtle from 
Queensland, Australia. Contributions in Science of the Natural History Mu- 
seum of Los Angeles County, 324:1-18. 

McDowell, S. B. 1983. The genus Emydwa (Testudines: Chelidae) in New 
Guinea with notes on the penial morphology of Pleurodira, pp. 169-189. In 
A. G. J. Rhodin and K. Miyata (eds.). Advances in Herpetology and Evo- 
lutionary Biology: Essays in Honor of Ernest E. Williams. Cambridge, Mas- 
sachusetts, Museum of Comparative Zoology. 725 pp. 

Palmer-Allen, M., F. Beynon, and A. Georges. 1991. Hatchling sex ratios 
are independent of temperature in field nests of the long-necked turtle, Chel- 
odina longicollis (Testudinata: Chelidae). Wildlife Research, 18: 225-231. 

Philippen, H.-D. and P. Grossman. 1990. Fine neue Schlangenhalsschildkrote 
von Neuguinea: Chelodina reimanni sp. n. (Reptilia, Testudines, Pleurodira: 
Chelidae). Zoologische Abhandlungen, Staatliches Museum fur Tierkunde, 
Dresden, 46(5): 95-102. 

Rhodin, A. G. J. AND W. P. McCord. 1990. Reproductive data on the chelid 
turtle Chelodina siebenrocki from New Guinea. Herpetological Review, 21(3): 
51-52. 

Rhodin, A. G. J. and R. A. Mittermeier. 1976. Chelodina parkeri. a new 
species of chelid turtle from New Guinea, with a discussion of Chelodina 
siebenrocki Werner, 1901. Bulletin of the Museum of Comparative Zoology, 
147(11): 465-488. 

Rhodin, A. G. J., S. Spring, AND P. C. H. Pritchard. 1980. Glossary of turtle 
vernacular names used in the New Guinea region. Journal of the Polynesian 
Society, 89(1): 105-117. 



36 BREVIORA No. 497 



Shaw, G. 1794. Zoology of New Holland. Vol. I. London, J. Davis. 33 pp. 
SiEBENROCK, F. 1914. Einc neue Chelodina Art aus Westaustralien. Anzeiger der 

Akademischen Wissenschaften Wien, 17: 386-387. 
Vestjens, W. J. M. 1969. Nesting, egg-laying and hatching of the snake-necked 

tortoise at Canberra, A.C.T. Australian Zoology, 15(2): 141-149. 
Wells, R. W. AND C. R. Wellington. 1985. A classification of the Amphibia 

and Reptilia of Australia. Australian Journal of Herpetology, Supplemental 

Series, 1: 1-61. 
Werner, F. 1 90 1 . Ueber Reptilien und Batrachier aus Ecuador und Neu-Guinea. 

Verhandlungen der Zoologisch-Botanischen Gesellschaft Wien, 51: 593-603. 
Williams, E. E. 1 950. Variation and selection in the cervical central articulations 

of living turtles. Bulletin of the American Museum of Natural History, 94: 

510-561. 



B R E V I O R A 

useiim of Comparative Zoology 



US ISSN 0006-9698 



Cambridge, Mass. 2 February 1994 / Number 498 

CHELID TURTLES OF THE AUSTRALASIAN 

ARCHIPELAGO: 11. A NEW SPECIES OF CHELODINA 

FROM ROTI ISLAND, INDONESIA 

Anders G. J. Rhodin' 

Abstract. A new species of Chelodina (Testudines: Pleurodira: Chelidae) is 
described from Roti Island, west of Timor, East Nusa Tenggara Province, in the 
southeastern Indonesian Archipelago. The species is endemic to Roti, a small and 
relatively xeric island. It is most similar and most closely related to Chelodina 
phtchardi from Papua New Guinea and C. longicollis from Australia, less closely 
related to C. novaeguineae and C reimanni from New Guinea. 

INTRODUCTION 

The side-necked turtles of the family Chelidae that inhabit the 
Australasian Archipelago of eastern Indonesia and Papua New 
Guinea remain one of the least well known turtle faunas of the 
world. Until recently only two species of the snake-necked genus 
Chelodina were known from the regions north of Australia: Chel- 
odina novaeguineae Boulenger, 1888 and Chelodina siebenrocki 
Werner, 1901. Since 1975, systematic studies have revealed an 
additional three species: Chelodina parkeri Rhodin and Mitter- 
meier, 1976, C/z^/o<a'//7ar£'/m(2««/Philippen and Grossman, 1990, 
and Chelodina pritchardi Rhodin, 1993. The last of these, from 
the Kemp Welch River of southeastern Papua New Guinea, was 
described in the first paper of this series on the chelid turtles of 
the Australasian Archipelago. In this second paper of the series, 
I describe another new species of Chelodina, this time from Roti 
Island, west of Timor in southeastern Indonesia. 

The first species of Chelodina to be described from anywhere 
in the Australasian Archipelago was C novaeguineae Boulenger, 



j^ 



' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts, and Chelonian Research Foundation, Lunenburg, Massachusetts. 



BREVIOR.A 



No. 498 




Figure 1 . Distribution of Chelodina subgeneric group "A" in northern Aus- 
tralia, New Guinea, and southeastern Indonesia. 1. Distribution of Chelodina 
mccordi on southwestern Roti Island, Indonesia. 2. Australian distribution of C. 
novaeguineae (sensu lato) in Northern Territory and Queensland. 3. Distribution 
of C. phtchardi in Papua New Guinea. 4. New Guinean distribution of C. no- 
vaeguineae in Irian Jaya and Papua New Guinea. 5. Approximate distribution of 
C reimanni in Irian Jaya, Indonesia. Major watershed limits in New Guinea and 
Australia indicated as thick lines; thin dotted lines mark the 200 meter water 
depth limits around the eastern Indonesian islands as well as the continental Sahul 
Shelf surrounding Australia and New Guinea. 



1888, which came from the southern New Guinea coastal region 
of what is now designated as the Western Province of Papua New 
Guinea. It is distributed throughout southern lowland New 
Guinea including extreme southeastern Indonesian Irian Jaya and 
southwestern Papua New Guinea. Similar forms of Chelodina 
novaeguineae (sensu lato) also occur in northern Queensland and 
northeastern Northern Territory in Australia (Fig. 1), although 
these forms probably represent separate and distinct taxa (J. Cann, 
personal communication to W. P. McCord). Soon after the orig- 
inal description of C. novaeguineae, Lidth de Jeude (1895) de- 



1994 NEW CHELODINA FROM ROTI 3 

scribed three specimens of this species from the island of Roti 
(originally spelled "Rotti"), west of Timor in southeastern pres- 
ent-day Indonesia. These specimens were collected on Roti in 
1891 by Dr. Herman F. C. Ten Kate and donated to the Leiden 
Museum. Though extremely isolated from the rest of the range 
of C. novaeguineae, this locality has subsequently been duly noted 
in the distribution of C novaeguineae without any further at- 
tempts at critical systematic comparison of its population to the 
populations in the New Guinean and Australian portions of the 
range. From the zoogeographic isolation of Roti, I long suspected 
that this population, if natural, must certainly represent a separate 
and distinct species, though probably closely related to C no- 
vaeguineae (sensu stricto). 

Unfortunately, Roti has become a difficult place to visit for 
field studies, due in part to the political upheaval on neighboring 
Timor, associated with the recent incorporation of Portuguese 
East Timor into Indonesia. For many years I attempted unsuc- 
cessfully to travel to Roti. A few years ago, I was fortunate to 
meet Dr. William P. McCord, who maintains an enormous col- 
lection of live turtles and whose primary interest is the diversity 
of species in the genus Chelodina. Dr. McCord, through his col- 
lection contacts in Indonesia (notably Mr. Frank Yowono), had 
recently succeeded in obtaining a series of 16 Chelodina from 
Roti. When I examined them, my original conviction that the 
Roti animals would be different from mainland New Guinean 
and Australian C. novaeguineae was confirmed. 

Subsequent to my analysis of the McCord collection of Roti 
animals I obtained on loan two of the original specimens collected 
in 1891 by Dr. Ten Kate on Roti and still present in the National 
Museum of Natural History in Leiden. A comparison of those 
specimens with the description by Lidth de Jeude (1895) confirms 
that they are the original specimens, and that they are also the 
same taxon as McCord's specimens obtained from Roti by Frank 
Yowono. Both Leiden specimens also bear tags with the manu- 
script name ""Chelodina rottiensis Brongersma." This name must 
now be considered a nomen nudum, as it was never published or 
described, but clearly Dr. Brongersma also deserves mention and 
credit for having previously recognized the distinctiveness of this 
new taxon from Roti. I thank him and Dr. Hoogmoed for releasing 



4 BREVIOHA No. 498 

the specimens and relinquishing claims to the formal description. 
These 1 8 specimens of Chelodina from Roti Island were then 
critically compared to a series of 43 Chelodina pritchardi from 
southeastern Papua New Guinea, 5 1 C. novaeguineae from south- 
western Papua New Guinea and adjacent Irian Jaya, 10 C. no- 
vaeguineae ^rom. northern Australia, 54 C longicoUis from eastern 
Australia, 12 C reimanni from southeastern Irian Jaya, Indo- 
nesia, and 7 C. steindachneri from western Australia, for a total 
study series of 195 specimens. Analysis of external morphology 
and cranial osteology demonstrated that this isolated Roti pop- 
ulation of Chelodina is indeed a new and distinct species of Chel- 
odina subgeneric group "A" (sensu Goode, 1967). It is more 
similar to C. pritchardi of Papua New Guinea than it is to either 
New Guinean or Australian populations of C novaeguineae. It 
now gives me great pleasure to formally describe this new species 
of Chelodina and to name it in honor of Dr. William P. McCord 
who succeeded in obtaining the series of animals that made con- 
firmation and description of the new species possible. 

TAXONOMY 

Chelodina mccordi, sp. nov. 
(Figs. 2-5) 

Chelodina novae guineae (panim) Ten Kate, 1894:689 

Chelodina novae-guineae (partim) Lidth de Jeude, 1 895: 1 1 9; De Rooij, 1915:315 

Holotype. MCZ 1 76730 (Fig. 2), alcohol-preserved adult female 
of 197.5 mm carapace length, purchased from native villagers by 
Frank Yowono in Kupang, western Timor, originally collected 
on Roti Island, East Nusa Tenggara Province, Indonesia; speci- 
men is formerly from the private live collection of William P. 
McCord, and also bears old tag AGJR 450 from the personal 
preserved collection of Rhodin. 



Figure 2. Holotype oi Chelodina mccordi (MCZ 176730), adult female mea- 
suring 197.5 mm carapace length, from Roti Island, Indonesia. 



1994 



NEW CHELODINA FROM ROTI 




6 BREMOR.A No. 498 

Paratypes. MCZ 176731, alcohol-preserved adult male of 153 
mm carapace length, purchased from native villagers by Frank 
Yowono in Kupang, western Timor, originally collected on Roti 
Island; specimen is formerly from the private live collection of 
William P. McCord, and also bears old tag AGJR 364 from the 
personal preserved collection of Rhodin; MCZ 176732 (Fig. 3), 
alcohol-preserved adult female of 1 94 mm carapace length, pur- 
chased from native villagers by Frank Yowono in Kupang, west- 
em Timor, originally collected on Roti Island; specimen is for- 
merly from the private collection of William P. McCord, and also 
bears old tag AGJR 368 from the private preserved collection of 
Rhodin; RMNH 10187 (Figs. 4-5), skeletal preparation of shell, 
skull, and limbs of adult of unknown sex (but probably female) 
of 179.5 mm carapace length, collected by Dr. Herman F. C. Ten 
Kate on "Rotti" [Roti Island, Indonesia] in 1891, originally de- 
scribed by Lidth de Jeude (1895) as a specimen of Chelodina 
novaeguineae'QovXQngQr, 1888. 

Referred Specimens. RMNH 4349, collected by Dr. Herman 
F. C. Ten Kate on "Rotti" [Roti Island, Indonesia] in 1891, 
originally described by Lidth de Jeude (1895) as a specimen of 
Chelodina novaeguineae^oultngtv, 1888; AGJR 365-7, 369, 448- 
9, 452-7, 460, purchased from native villagers by Frank Yowono 
in Kupang, western Timor, originally collected on Roti Island, 
Indonesia, and formerly from the private live collection of Wil- 
liam P. McCord. Personal specimens available on loan through 
the Chelonian Research Foundation, and eventually to be de- 
posited at the Museum of Comparative Zoology. 

Distribution. Known only from Roti Island, located about 20 
km southwest of the western end of Timor in the Lesser Sunda 
Islands in the Province of East Nusa Tenggara of the southeastern 
Indonesian Archipelago (Fig. 1). Not known to occur on Timor 
itself No current localities on Roti Island precisely documented 
as yet, but collectors indicate that the species is most readily found 
in rice paddies. Most rice paddies are located in the southwestern 
half of the island west and north of the village of Tudameda, and 
along the cross-island road between Tudameda and Ba'a. In ad- 
dition, there are some limited rice paddies and small lakes in the 
northeast peninsular region of the island where the species might 
also occur, but a large portion of the rest of the island is relatively 



1994 



NEW CHELODINA FROM ROTI 







3 
T3 
CO 

rT 

r- 

N 
U 

aj 
a 
>. 

C3 

Cu 



o 
g 

5 



X3 

u 

u 

H-1 



00 



BREVIOKA 



No. 498 






Figure 4. Dorsal, ventral, and lateral views of skull of Chelodina mccordi 
(Paratype RMNH 10187, adult of unknown sex measuring 179.5 mm carapace 
length) from Roti Island, Indonesia. 



xeric with little surface water. The total area of Roti Island is only 
about 1,214 km", while the highest elevation reaches 444 m. 

Type Locality. The collection locality of the holotype specimen 
obtained by Yowono from Roti Island cannot be stated precisely. 



Figure 5. Shell of Chelodina mccordi (Paratype RMNH 10187), originally 
collected in southwestern Roti in 1891 by Dr. Herman F. C. Ten Kate and de- 
scribed by Lidth de Jeude (1895). 



1994 



NEW CHELODINA FROM ROTI 




10 BREVIORA No. 498 

However, a review of the natural history and travel writings of 
Dr. Ten Kate, who collected three specimens in 1891, including 
one of the paratypes, and a careful study of older and modem 
maps of Roti can help further restrict the probable type locality. 
Dr. Ten Kate states in his travelogue (Ten Kate, 1894:688-689) 
that he observed a small specimen of Chelodina novae guineae 
[= Chelodina mccordi] that fishermen had just caught on 7 Sep- 
tember 1891 at the inland freshwater lake "Danau Naloek (Na- 
roek)" near lake "Danau Linggoe" [Danau = Lake in Bahasa 
Indonesian]. Danau Naloek was reached approximately three fifths 
of the way along a four and a half hour trip while Dr. Ten Kate 
traveled from "Ti-Kanaketoe" (also referred to as simply "Ti") 
[= modem Danaheo, or Danahoe, along the southwestem shore, 
4 km south of Tudameda] to "Baa" [= modern Ba'a along the 
northem shore]. The locality "Danau Naloek" does not appear 
on modem maps, but Dr. Ten Kate mentions that he followed 
Dr. Wichmann's earlier route through the area, and Wichmann 
(1892) shows the lake "Danu naluk" on his map about halfway 
between Ti and Baa, which he also indicates by their more modern 
names of Danoheoh and Namudale [= Namodale, a suburb of 
Ba'a]. Dr. Ten Kate further states that Danau Naloek was located 
in the middle of dry rolling grassland, and that shortly after leaving 
the lake he entered forested landscape at "boschrijke . . . Loa- 
meko" where he crossed the administrative border between Deng- 
ka and Lelain. This provincial boundary still occurs on modem 
maps with the villages of Busalangga and Longgo [probably = 
Danau Linggoe] just west of there, at approximately the three 
fifths point along the Danaheo-Ba'a road. It is therefore reason- 
able to state that the species occurred at Danau Naloek in the 
Busalangga region in 1891 and to assume that Dr. Ten Kate 
probably collected or obtained at least one of his paratype spec- 
imens there. Therefore, the type locality of Chelodina mccordi is 
hereby restricted to Danau Naloek, near Busalangga, ca. 1 1 km 
northeast of Tudameda and ca. 8 km southwest of Ba'a, elevation 
ca. 115 m, southwestem Roti Island (10°48'S, 123°00'E), East 
Nusa Tenggara Province, Indonesia. 

Diagnosis. A medium-sized isolated Rotinese snake-necked 
chelid turtle of Chelodina subgeneric group "A" (sensu Goode, 
1967 and Burbidge et ai, 1974) with relatively wide head, wide 



1994 



NEW CHELODINA FROM ROTI 



11 



Table 1 . Features distinguishing four species of Chelodina subgeneric 
GROUP "A". For meristic basis of distinguishing features, see Tables 3 and 
5 and Figures 4-6. Pluses and minuses for the intermediate categories 
indicate slight differentiation between the two species in that category. 



Feature 


longicollis 


pritchardi 


mccordi 


novaegidneae 


External 










Carapace width 


Interm. 


Wide 


Wide 


Narrow 


Head width 


Narrow 


Narrow 


Wide 


Wide 


Plastron width 


Wide 


Interm. 


Narrow 


Narrow 


Marginal 1 width 


Wide 


Wide 


Narrow 


Interm. 


Composite ratio 


Low 


Interm. (-) 


High 


Interm. ( + ) 


Osteology 










Triturating width 


Narrow 


Interm. (-) 


Interm. ( + ) 


Wide 


Skull depth 


Shallow 


Interm. 


Interm. 


Deep 


Skull width 


Narrow 


Narrow 


Wide 


Wide 


Robusticity 


Low 


Interm. (-) 


Interm. (+) 


High 



carapace, narrow plastron, narrow first marginal, wide skull, in- 
termediate triturating surface width, intermediate skull depth, and 
intermediate to mildly increased skull robusticity. Most similar 
to C pritchardi, but differentiated fi^om it by the much narrower 
first marginal, relatively wider second marginal, wider head, nar- 
rower plastron, wider skull, and very slightly increased skull ro- 
busticity. Also similar to C. novaeguineae, but differentiated fi-om 
that taxon by the wider carapace, slightly narrower first marginal, 
significantly narrower triturating surface width, shallower skull, 
and significantly decreased skull robusticity. 

In most morphological respects C mccordi is intermediate be- 
tween C. novaeguineae and C. longicollis, and most similar to C. 
pritchardi, especially in juvenile and subadult stages. See Table 1 
for a summary of differences among C. mccordi, C. pritchardi, C. 
longicollis, and C novaeguineae. 

Etymology. The specific epithet is a patronym honoring Dr. 
William P. McCord, a veterinarian with a deep interest in turtles, 
who was instrumental in securing the large study series demon- 
strating the distinctiveness of the species. 

Related taxa. Chelodina mccordi is most similar to the following 
five chelid taxa from New Guinea and Australia, all members of 
Chelodina subgeneric group "A": Chelodina longicollis (Shaw, 



12 BREVIOm No. 498 

1794), Chelodina novaeguineae Boulenger, 1888, Chelodina 
pritchardi Rhodin, 1993, Chelodina reimanni Philippen and 
Grossman, \990, din<X Chelodina steindachneri'^iQbtnroQk, 1914. 
Type localities and specimens for these species are listed in the 
first paper of this series (Rhodin, 1993). 

DESCRIPTION 
External Morphology 

Carapace. Carapace of C. mccordi moderately rugose and 
broadly oval, width averaging 77.9% of length [Fig. 6(1)A], mod- 
erately wider posteriorly at about marginals 6-7, with slight ex- 
pansion of marginals 6-8. Moderately prominent lateral marginal 
recurving from about marginal 4 through 7, often partially in- 
volving marginals 3 and 8 as well. Somewhat less prominent 
recurving in smaller specimens. Moderate supracaudal ridging 
with slight adjacent concavity of marginal 1 1 . No vertebral knobs, 
keel, or ridging. Slight vertebral flattening and shallow midline 
furrow in larger females, smoothly convex in smaller females and 
males. No supracaudal notch or marginal serrations. Dorsal nu- 
chal long and broad, not projecting anterior to carapace margin. 
Ventral underlap of nuchal also relatively long and broad. Ver- 
tebral 1 widest, then 2, 3, 5, and 4 in descending order of width 
in normal specimens. Basic external measurements of C. mccordi 
presented in Table 2, differences from C pritchardi, C novae- 
guineae, and C. reimanni presented in Table 3 and Figure 6 (parts 
1 and 2). 

Many individuals with altered vertebral and costal pattern with 
supernumerary or fused scutes. Five of 1 8 specimens have 6 ver- 
tebral scutes, three with a small intercalated supernumerary scute 
between V3 and V4, one between V2 and V3, and one between 
V4 and V5. Two specimens have three symmetrical costals on 
each side, one has three costals on one side, four on the other. 
This deformity creates a very wide fifth vertebral, indicating a 
fusion of the fourth costals with V5. The larger Leiden specimen 
(RMNH 10187) originally collected by Dr. Ten Kate has this 
deformity with a symmetrical pattern of three costals on each 
side and a very wide fifth vertebral (Fig. 5). Lidth de Jeude ( 1 895) 
interpreted this as a normal pattern. One of Yowono's specimens 



1994 



NEW CHELODINA FROM ROTI 



13 



Table 2. Basic external dimensions of Chelodina mccordi. all measure- 
ments IN MM. CL = CARAPACE LENGTH (STRAIGHT-LINE IN MIDLINE); CW = CARA- 
PACE WIDTH (greatest); CD = carapace DEPTH (GREATEST IN MIDLINE); PL-M = 
PLASTRON LENGTH (IN MIDLINE); PL-T = PLASTRON LENGTH (TOTAL, INCLUDING 
ANAL SPURS); PW = PLASTRON WIDTH (AT AXILLARY NOTCH); HW = HEAD WIDTH 

(tympanic); J = juvenile; f = female; m = male; u = unknown sex. 



Specimen 


















Number 


Sex 


CL 


CW 


CD 


PL-M 


PL-T 


PW 


HW 


RMNH 4349 


J 


99.5 


73.0 


31.0 


76.0 


81.0 


43.5 


18.6 


AGJR 460 


f 


150.0 


118.0 


49.0 


119.5 


126.0 


70.0 


25.1 


AGJR 455 


m 


150.5 


114.5 


49.5 


117.5 


124.5 


67.0 


26.3 


AGJR 454 


m 


151.0 


116.5 


47.0 


115.5 


123.5 


66.0 


25.0 


MCZ 176731 


m 


153.0 


120.5 


47.5 


116.5 


125.0 


67.0 


26.0 


AGJR 449 


m 


155.0 


123.5 


51.5 


121.5 


128.0 


73.0 


25.3 


AGJR 367 


f 


159.0 


121.0 


52.0 


124.0 


131.0 


71.0 


26.3 


AGJR 456 


m 


162.5 


127.5 


53.5 


120.5 


130.5 


70.0 


25.0 


AGJR 369 


u 


163.0 


125.0 


52.0 


123.0 


132.5 


71.0 


— 


RMNH 10187 


u 


179.5 


140.0 


60.0 


138.0 


147.0 


81.5 


28.7 


AGJR 457 


f 


180.0 


137.0 


60.0 


138.0 


147.0 


80.0 


28.7 


AGJR 448 


f 


181.5 


143.5 


63.5 


139.0 


146.5 


81.5 


27.8 


AGJR 366 


f 


182.0 


142.5 


60.0 


145.0 


153.0 


84.5 


27.9 


MCZ 176732 


f 


194.0 


152.0 


67.0 


153.0 


162.0 


85.0 


30.5 


MCZ 176730 


f 


197.5 


156.0 


67.5 


150.5 


159.5 


86.5 


29.6 


AGJR 452 


f 


202.0 


159.0 


73.0 


161.5 


171.0 


93.0 


31.2 


AGJR 365 


f 


206.0 


162.0 


72.0 


161.0 


171.0 


92.0 


31.8 


AGJR 453 


f 


213.0 


162.0 


77.5 


162.5 


172.0 


95.0 


32.8 



(AGJR 368) has this identical deformity, and another (AGJR 
365) has it on one side only. Five specimens have markedly 
reduced fourth costals, often on one side only, giving a slightly 
wider asymmetrical fifth vertebral. One of the specimens with 
this deformity is the smaller Leiden specimen (RMNH 4349) 
originally collected by Dr. Ten Kate. 

Width relationships of marginal scutes 1 and 2 distinctive, with 
Ml much narrower than M2 as measured along the VI -CI scute 
border. Ml typically half as wide as M2. In C phtchardi Ml is 
wider or subequal to M2, and in C. novaeguineae Ml averages 
about 80% of the width of M2 (Fig. 7). 

Carapace slightly less broad in C. mccordi than in C. pritchardi, 
but significantly broader than in C. novaeguineae. Lateral mar- 
ginal recurving typical for C mccordi, not present in C pritchardi. 



.875 

5 -85 

S 

"^ .825- 

a> 

U .8 

a 

W -i-ic 

« 

U 

...^ .75- 
^ .725- 



.7- 
Oi .675- 

u 

63 



.625 



O 



O 
O 



O 
O 



O 
O 



o o 
o o 



o 



o 



o 
o 



A A-^^ 
O O 

A A 

o ^ 



cf ■*  



» 4,»3, 



A 

A A 



O A 
O O 



o 



^ OA \ ^^ 



Broader 
Carapace 




OC 
«C 

AC 

 c 

AC 


longicollis 

mccordi 

novaeguineae 

prilchardi 

retmanni 



A aA ^ 

^aoA 

A 



AAA^ 



AO 
A A 



40 



60 



80 



100 



120 



140 



160 



180 



200 



220 



240 



S 

a. 
U 



-a 



.21 



19 



.18- 



.17 



.16- 



14 



.13 



.12 
40 



B 


A 

O 




A 




A 


A 




A 
A 




o 




A 

a 


^ 




A ^ 

A 




A 
A - 








Oq 


® 
o 


o 
o 

o ° 

°@ 

o 


o 


A 
A 

o 


A -^ A 
AA^ A 

°o°o *« 


A 

A 
A ^ 

A o 0<h<t 
f A » 

'^■- "a o 






Wider 
Head 


i k 










o C. longicollis 
<» C. mccordi 










O 


o 


o tf  o 


 




a C. novaeguineae 
 C. pritchardi 














o 






A C. reimanni 














o 









60 



80 



100 120 140 160 180 

Carapace Length (mm) 



200 



220 



240 



Figure 6. (Part 1). Shell morphometries. Graphs plotting morphometric vari- 
ation for four species of Chelodina subgeneric group "A", showing the relationships 
of: A. carapace width ratio (Carapace Width/Carapace Length); B. head width 
ratio (Head Width/CL). 



.575 

*^ 
OX) 

ID 

lU 525 

a 
a 

U .475 



-C .45 

."2 

^ .425 

C 

.35- 
























c 








o 


o 




o 








o 


o 
o 


o 

o©g 
o 


o 


% 


o 

Oo 

o 
o°°o 


o 

o 
o 
o 


o o 
o o 


o 




oo 


A 
A 




^/^3Va 4 


*  
A a a 




Broader 
Plastron 


i k 




s 




A A 


aa 

A 
A 


^a' 


9(^ 

A 










C. longjcollis 




3 C. mccordi 


















A C. novaeguineac 


















 C. pritchardi 












▲ 






A C. reimanni 





















40 



60 



80 



100 



120 



140 



160 



180 



200 



220 



240 












a 
Si 

0) 

o 
eu 

E 

o 



38 
.36 
.34 

.32 
.3 
28 
26 

.24 



18- 



D 



O^ A 



o 



o 



A 
O 






o o 



Broader Carapace 

Wider Head 
NaiTOwer Plastron 



^ 



o 

Or 



Ad 



A 



■'^ 



3 ^ 

A 



O C. longicoUis 
o C. mccordi 
A C. novaeguineae 
 C. pritchardi 
A C. reimanni 



o 



o o 
o 



cP 



40 



60 



80 



100 



120 



140 



160 



180 



200 



220 



240 



Carapace Length (mm) 



Figure 6. (Part 2). Shell morphometries. C. plastron width ratio (Plastron 
Width/CL); and D. composite ratio of (CW x HW x 1/PW)/CL expressed as a 
trivariate product; all versus Carapace Length. Note the position of C mccordi 
as most similar to C pritchardi in terms of carapace width (A), but most similar 
to C. novaeguineae in terms of head width (B), plastron width (C), and composite 
ratio (D). 



16 



BREV10R.A 



No. 498 



1.4 



'Si' 
-a 



s 

'OJD 
^ .4 













 














o C mccordi 






ti C. novaeguineae 




















 C pritchardi 


























  


 


 




 






Wider 










4 




Marginal 




i 


Number 1 




A 
A 


 
 

A 
A 


 
 
 
 




A 




 










Narrower 


 








A 


A^ 


A 




A 




A 




A 




A 

A 
^ 3 


A 


A 




A 








3 

3 
3 


3 
3 




3 
3 


3 

3 




3 




<b 


3 




3^ 
3 


33 ^ 
3 





80 100 120 140 160 180 200 

Carapace Length (mm) 



220 



240 



Figure 7. Shell morphometries. Graph plotting morphometric variation for 
M1/M2 ratio (marginal number 1 width/marginal number 2 width) versus cara- 
pace length for Chelodina mccordi, C. pritchardi. and C. novaeguineae. showing 
that C. mccordi has Ml about half as wide as M2. C. pritchardi has Ml slightly 
wider or subequal to M2, and C. novaeguineae has Ml about 0.8 times as wide 
as M2. 



not present or only very minimally developed in C. novaeguineae. 
Supracaudal ridging with adjacent mild marginal concavity also 
typical for C. mccordi, not present in C pritchardi or C novae- 
guineae. 

Carapace moderately deep in large specimens, relatively flatter 
in younger ones. No sexual dimorphism in carapace depth of 
specimens examined, with like-sized males and females with same 
carapace depth. Older females of larger body size all with deeper 
shells. Carapace depth in C mccordi slightly deeper than in either 
C. pritchardi or C. novaeguineae. 

Color variable, with most specimens having a distinctive light 



1994 NEW CHELODINA FROM ROTI 17 

grayish-brown carapace unusual for Chelodina subgeneric group 
"A". Some specimens darker chestnut brown, more typical of the 
color seen in C. novaegiuneae and C. pritchardi. Carapace scutes 
moderately rugose with retention of concentric growth lines only 
in small specimens. 

Plastron. Plastron broad, axillary width averaging 57.9% of 
midline plastral length. Anterior lobe moderately broad, similar 
to C. novaeguineae, but not as broad as C pritchardi [Fig. 6(2)- 
C]. Slight expansion of anterior plastral lobe at posterior edge of 
humeral scutes, similar to C. pritchardi. Anal notch moderately 
deep, no sexual dimorphism noted. Intergular broad, long, and 
without marginal contact. Plastral scute medial length formula Ig 
> An > Abd > Fem > Pec > Gul. C. pritchardi usually with 
Pec > Fem, C novaeguineae with either Pec > Fem or Fem > 
Pec. No axillary or inguinal scutes. 

Plastron color light yellowish-white with many specimens hav- 
ing thin irregular light-brown areas along the plastral sutures. 
Original specimens of Dr. Ten Kate with oxidized plastrons, all 
of Yowono's specimens without oxidation. Plastron most similar 
to C pritchardi, but generally with more pigment. Hatchlings with 
a beautiful orange and gray pattern covering plastron and ventral 
soft parts, gradually fading with growth (McCord, personal com- 
munication). 

Head and Soft Parts. Head with small irregular scales covering 
temporal skin, smooth over parietal and interorbital roof Neck 
with low soft tubercules, generally more similar to C. pritchardi 
than the slightly more prominent, firmer tubercules in C. novae- 
guineae. Soft parts light to moderate gray dorsally, whitish ven- 
trally, generally lighter in color than in either C. pritchardi or C 
novaeguineae. Hands and feet with four claws each. 

Head width moderately wide, similar to the relatively broad- 
headed C novaeguineae, significantly wider than the narrow- 
headed C pritchardi and C. longicollis [Fig. 6(1 )B], and narrower 
than the broad-headed C. reimanni. Head width not as wide as 
in the broad-headed members o{ Chelodina subgeneric group "B" 
(C. expansa, C. rugosa, C. siebenrocki, C. parkeri, and C. oblonga). 
Relative narrowing of the head ontogenetically. 

A few specimens of "C. novaeguineae'" said to come from Roti, 
but provided without reliable data, have broad heads most similar 



18 BREVIOR.4 No. 498 



Table 3. Means and standard deviations for shell measurement ratios 
OF three Chelodina species. Abbreviations as in Table 2. Data based only 

ON SPECIMENS OF CARAPACE LENGTH GREATER THAN 100 MM. C. NOVAEGUINEAE 

includes only New Guinean specimens, not Australian ones. Ml = width 
OF marginal number 1 , M2 = width of marginal number 2; for this measure- 
ment only, C. pritchardi n = 17 and C. novaeguineae n = 19. 





C. pritchardi 


C mccordi 


C novaeguineae 


Ratio 


(n = 43) 


(n= 17) 


(n = 51) 


CW/CL 


.785 ± .018 


.779 ± .012 


.723 ± .028 


PW/PL-M 


.610 ± .012 


.579 ± .010 


.576 ± .018 


HW/CL 


.150 ± .005 


.160 ± .007 


.167 ± .010 


CD/CL 


.328 ± .014 


.335 ± .015 


.327 ± .021 


M1/M2 


1.089 ± .129 


.465 ± .090 


.803 ± .105 



to C. reimanni and extremely deep robust shells of a dark black 
color. These distinct animals appear to represent a different taxon 
and have been excluded from this analysis of C mccordi, awaiting 
further confirmation of their probably disparate geographic origin. 
Size and Sexual Dimorphism. The largest specimen of C mccordi 
recorded is a female of 213 mm carapace length. The largest male 
examined has a carapace length of 1 62.5 mm, indicating probable 
sexual dimorphism, with females larger than males. Calculating 
the sexual dimorphism index according to the method of Gibbons 
and Lovich (1990) yields an SDI value of approximately 1.31 for 
C. mccordi. The SDI for C. pritchardi is 1 .22, that for New Guin- 
ean C. novaeguineae 1.37. The maximum size recorded for C 
pritchardi is 228 mm, and for New Guinean C. novaeguineae 2 1 8 
mm (Rhodin, 1 993). However, in general, and for most specimens 
representing typical mature adults, C. mccordi is significantly larg- 
er than New Guinean C. novaeguineae, slightly smaller than C. 
pritchardi, and significantly smaller than Australian C. novae- 
guineae, which reaches carapace lengths of 279 to 300 mm (Cann, 
1978; Rhodin, 1993). 

Osteology 

Skull. The description of skull osteology is based on the ex- 
amination of 36 skulls of Chelodina subgeneric group "A". Of 
these, 4 are C. mccordi, 6 C. pritchardi, 1 C. longicollis, 15 C. 
novaeguineae {\2 from New Guinea, 3 from Australia), 2 C stein- 



1994 



NEW CHELODINA FROM ROTI 



19 



Table 4. Basic skull measurements for Chelodina mccordi. SL = skull 

LENGTH (snout-occipital CONDYLE); SWT = SKULL WIDTH, TYMPANIC MAXIMUM; 
SWM = SKULL WIDTH, MAXILLARY MAXIMUM; SDM = SKULL DEPTH AT POSTERIOR 
EDGE OF maxillae; SD = SKULL DEPTH IN MIDLINE BETWEEN SUPRAOCCIPITAL SPINE 
AND BASISPHENOID; low = INTER-ORBITAL WIDTH. MINIMAL; OW = ORBITAL WIDTH, 
SHORT axis; PtW = PTERYGOID WIDTH, MINIMAL; TW = TRITURATING WIDTH, 
MAXILLARY (MEASURED IN MIDLINE FROM TOMIAL EDGE TO ANTERIOR CHOANAL 

BORDER). Refer to Table 5 and Figures 8(1) and 8(2) for analysis of skull 

MEASUREMENT RATIOS. 



Specimen 




















Number 


SL 


SWT 


SWM 


SDM 


SD 


lOW 


OW 


PtW 


TW 


AGJR 449 


35.5 


24.5 


20.2 


8.8 


10.0 


3.7 


8.4 


11.9 


4.7 


RMNH 10187 


40.1 


27.7 


22.9 


9.5 


11.0 


4.3 


8.9 


13.2 


5.7 


AGJR 452 


43.6 


30.1 


25.6 


10.1 


12.1 


4.9 


9.5 


14.2 


6.6 


AGJR 453 


45.5 


32.0 


25.9 


10.6 


13.4 


5.5 


9.6 


14.5 


7.1 



dachneri, and 2 C reimanni. Refer to Tables 4 and 5 and Figure 
8 (parts 1 and 2) for additional skull measurements and ratios, 
and Figure 3 for skull illustrations of C mccordi. Comparative 
figures of skulls of C. pritchardi, C. novaeguineae, C. reimanni, 
and C. longicollis are in Rhodin (1993). 

The skull of C. mccordi is a typical Chelodina subgeneric group 
"A" type skull, not overly elongate, flattened, or wide as in sub- 
generic group "B". It is strikingly similar to C pritchardi and 
differs markedly from C. novaeguineae. Like in C pritchardi, the 
skull of C mccordi is differentiated from C novaeguineae by its 
relative lack of robusticity. C. novaeguineae has wide and robust 



Table 5. Means and standard deviations for skull measurement ratios 
of three Chelodina species. Abbreviations as in Table 4. C. novaeguineae 
includes only New Guinean specimens, not Australian ones. 





C. pritchardi 


C. mccordi 


C. novaeguineae 


Ratio 


(n = 6) 


(n = 4) 


(n= 15) 


SWT/SL 


.648 ± .013 


.694 ± .006 


.687 ± .024 


SWM/L 


.553 ± .006 


.574 ± .009 


.532 ± .032 


SD/SL 


.290 ± .009 


.282 ± .009 


.320 ± .018 


TW/SL 


.144 ± .005 


.145 ± .010 


.194 ± .009 


PtW/SWT 


.522 ± .014 


.472 ± .014 


.461 ± .018 


lOW/OW 


.470 ± .037 


.503 ± .056 


.570 ± .055 



20 



BREVIORA 



No. 498 



OX) 

c 



3 
C/3 






3 

H 



ci. 



3 
C/3 



16 



M .14 



.12 



Broader 
Triturating 

Surface 



O C. longicoUis 
<* C. niccordi 
A C. novacguineae 
 C. pritcharcli 
* C. reimanni 



a 
o 



80 



100 



120 



140 



160 



180 



200 



220 



240 



.42 



J- .38 

DC 

C 

«^ .36 



3 ,34 
— 32 



.28 



.26 



B 



o C. longicollis 
o C. mccordi 
A C. novaeguineae 
 C. pritchardi 
* C. reimaniii 



Deeper 

Skull 



100 120 140 160 180 

Carapace Length (mm) 



200 



220 



240 



Figure 8. (Part 1). Skull morphometries. Graphs plotting morphometric vari- 
ation for four species ofChclodina subgeneric group "A", showing the relationships 
of: A. maxillary triturating width ratio (TW/Skull Length); B. skull depth ratio 
(SD/SL). 



.85 



825 



,8 



775' 



75 



725 



.675 



65 



.625 



Wider 
Skill I 



longjcollis 

mccordi 

novaeguineae 

pritcliardi 



^ 



A 
O 



Ch 



o 



80 



100 



120 140 



160 180 200 220 



240 



180 



160 



140 



120 



100 



80 



60 



40 



20 



D 














More 

Robust 
Skull 


' 










OC. longjcollis 
» C. mccord! 
A C. novaeguineae 
 C. pritchardi 
* C. reimaiini 



A A 



oo 



o 



o o 



80 



100 120 140 160 180 200 220 



240 



Carapace Length (mm) 



Figure 8. (Part 2). Skull morphometries. C. tympanic skull width (SWT/SL); 
and D. composite graph of Robusticity Index (TW x SWT x SD)/SL expressed 
as a trivariate product; all versus Skull Length. Note the position of C. mccordi 
as most similar to C. novaeguineae in terms of tympanic skull width (C), but most 
similar to C. pritchardi in terms of triturating width (A), skull depth (B) and 
composite Robusticity Index (D). 



22 BREVIORA No. 498 

maxillary and mandibular triturating surfaces, with correspond- 
ingly wide and robust homy rhamphothecae. C. longicollis has 
very thin triturating surfaces, whereas both C pritchardi and C. 
mccordi have surfaces that are intermediate and similar to each 
other, though C mccordi tends to have slightly wider surfaces in 
larger specimens [Fig. 8(1 )A]. Chelodina novaeguineae has a deep 
skull, C. longicollis a shallow skull, and C. mccordi and C pritch- 
ardi have skull depths that are intermediate and similar to each 
other [Fig. 8( 1 )B]. Skull width as compared to skull depth is greater 
in both C mccordi and C novaeguineae, while C longicollis and 
C. pritchardi have relatively narrower skulls [Fig. 8(2)C]. Chel- 
odina mccordi has an intermediate-sized parietal roof, like C. 
pritchardi, not as large as C longicollis or as reduced as C no- 
vaeguineae. The pterygoid trochlear processes are minimally di- 
vergent and unflared in C mccordi, as they are in C. pritchardi, 
lacking the extreme flaring and prominent divergence seen in C. 
novaeguineae. The skull depth and supraoccipital crest height are 
similar in C. pritchardi and C mccordi. 

The Robusticity Index (see Rhodin, 1993) for the skull of C. 
mccordi is very similar to C pritchardi, but shows a slight increase 
in skulls of larger specimens, related to slightly increased tritu- 
rating width and skull depth, and moderately increased skull width 
in the larger specimens. Both of these species have skulls that are 
much more robust than C. longicollis and much less robust than 
C. novaeguineae [Fig. 8(2)D], with C. mccordi generally slightly 
more robust than C. pritchardi. 

Overall, the skull of C. mccordi seems to represent a phylo- 
genetic intermediate step in a transformation series leading from 
C. pritchardi to C novaeguineae. Chelodina mccordi retains nar- 
row triturating surfaces, though they are slightly widened, and a 
shallow skull; but has developed significantly increased skull width 
with slightly increased skull robusticity. These features correlate 
with an increase in temporal muscle mass, intermediate between 
the relatively reduced mass in C pritchardi and the markedly 
increased mass in C novaeguineae. Evidently C. mccordi has 
developed the need for moderately increased mandibular adduc- 
tor force generation in its jaw closure mechanism, but has not 
reached the point of requiring massively enlarged opposing trit- 
urating crushing surfaces. 



1 994 NEW CHELODINA FROM ROTI 23 

From skull morphology, one would predict that C. mccordi is 
a generalized carnivore or omnivorous scavenger, intermediate 
between the presumed specialized molluscivorous C. novaegui- 
neaedind the more limited piscivorous or carnivorous C. pritchar- 
di. 

Cervical Spine. Central cervical articulation pattern is 
(2(3(4(5)6)7(8) in 4 specimens (2 by direct exam, 2 by radiographic 
investigation), the only known pattern for all Chelidae as de- 
scribed by Williams (1950). Atlanto-axial (C1-C2) cervical mor- 
phology in C. mccordi identical to the pattern in other Chelodina 
subgeneric group "A". 

Shell. No neural bones in 2 specimens, all pleurals meeting in 
the midline. Axillary buttress moderately robust, articulating with 
lateral first pleural and posterior third peripheral; inguinal buttress 
less robust, articulating with postero-lateral edge of fourth and 
antero-lateral edge of fifth pleurals, as well as anterior seventh 
peripheral. Suprapygal relatively wide, contacting tenth periph- 
eral. Broad contact between first peripherals and first pleurals. 

Ecology and General 

Reproduction. Radiographs or dissections were performed on 
the females in the series, with one female demonstrating multiple 
small ovarian follicles bilaterally, as well as enormous paired 
cloacal bursae (one on each side). No eggs were noted. Repro- 
ductive parameters have not yet been fully documented for the 
species, but McCord (personal communication) has hatched sev- 
eral clutches of eggs from captive individuals. Average clutch size 
is 8-9 eggs, with oval eggs similar in shape to C. longicollis and 
C pritchardi, but slightly larger than for either of those species, 
and slightly smaller than the eggs of C reimanni, which are larger 
and more rounded. The eggs have hatched in about 2 months 
when incubated at about 82°F. 

Growth. The smallest specimen of 99.5 mm carapace length 
shows three concentric growth rings. The rings are clearly visible 
on the costal scutes and allow for measurements of growth. The 
first ring encompasses the indistinguishable original scute and 
subsequent growth in the first season; the second ring, growth 
through the end of the second season; and the third ring, growth 
until capture. By measuring the corresponding costal-vertebral 



24 BREVIORA No. 498 

suture lengths for each of the rings it is possible to create a ratio 
of costal length to carapace length for each ring and thereby cal- 
culate the carapace length of the animal at the end of each growth 
season. By this method, this specimen (RMNH 4349), which is 
now 99.5 mm long at the end of its third and last growing season, 
was approximately 73.5 mm long at the end of the second season, 
and 5 1 .0 mm long at the end of the first season. The actual original 
hatchling scute is no longer visible, but extrapolation from the 
regression curve created by the first three values yields an expected 
hatchling size of about 32.0 mm (Fig. 9). This predicted hatchling 
size is within the range of hatchlings of other species ofChelodina 
I have examined: C. parkeri at 35.0 mm, C. siebenrocki at 35.0 
mm, C rugosa at 32.0 mm, C oblonga at 30.0 mm, and C. 
longicollis at 28.8 mm. It is of course not known whether the 
growth rings are reflective of an annual cycle, but Roti has well 
defined wet and dry seasons, and it appears likely that this spec- 
imen is therefore about three years old. 

Predation. Five large females display evidence of possible pre- 
vious crocodile encounters. Four animals have what appear to be 
typical healed tooth holes and bite striations on the carapace, one 
has the hind portions of the carapace missing with resultant de- 
formed regenerated scar tissue. The saltwater crocodile Croco- 
dylus powsus is the most likely predator, but freshwater crocodiles 
may also occur in the Roti area (Ross, 1986). Native collectors 
also indicate that many specimens receive carapacial damage from 
farmers' plow blades in the rice paddies where the species is 
known to occur (McCord, personal communication). 

Sympatry. No other freshwater turtles are known to occur on 
Roti, but the semi-aquatic emydid turtle Cuora amboinensis may 
well occur on the island, having previously been recorded on 
Timor (Iverson, 1986). In addition, the trionychid aquatic soft- 
shell XutXXq Amyda cartilaginea may occur on either Roti or Timor. 
Iverson (1986) records the nearest confirmed locality as Lombok 
Island just east of Bali, but Trionyx cartilagineus newtoni Ferreira, 
1897, was described as having been obtained on Timor, and may 
represent evidence for a population in this area. 

DISCUSSION 

The occurrence of a population of chelid turtles on Roti Island 
in Indonesia comes as a relative surprise because of the known 



1994 



NEW CHELODINA FROM ROTI 



25 



no 



100 



90 



JZ 




■^^ 


so 


0£ 




c 




0^ 


70 


0^ 




W 


60 


CQ 




a 




5^ 


SO 


Urn 




CO 




U 


40 



30 



Growth of Chelodina mccordi 




y = 32 + 17.25X + 1.75x2 



1.5 



2.5 



3.5 



Age (years) 



Figure 9. Graph showing probable growth of individual specimen of C mccor- 
di (RMNH 4349) as calculated by measurements of costal scute growth rings. 
Actual size of specimen recorded as last data point on graph; sizes at age 1 and 
2 calculated from growth rings; size at age extrapolated from the curve. 



zoogeography of the family. Other than in South America, no 
other natural populations of chelid turtles have been recorded 
outside of continental Australia and New Guinea and islands on 
their contiguous Sahul Shelf. Although Elseya novaeguineae has 
been recorded in the Palau Islands in the northwestern Pacific 
(Aoki, 1977), that record probably represents an introduction. In 
addition, I have seen photographs of a specimen of Emydura 
subglobosa purportedly from New Britain in northeastern oceanic 
Papua New Guinea, which also probably represents an introduc- 
tion. McCord (personal communication) has obtained specimens 
of this population o^ Emydura subglobosa collected in the vicinity 
of Rabaul, a major commercial center with a huge natural prod- 
ucts market where exotic species introductions would come as no 
major surprise. 

The collection of specimens of C mccordi on Roti by Dr. Ten 
Kate back in 1891 and now by Frank Yowono about 100 years 
later confirms the presence of an established viable breeding pop- 
ulation of this taxon. The demonstrated similarity in morphology 



26 BREVIORA No. 498 

of the original 1891 specimens and of the recently collected spec- 
imens confirms the identity and the source of the two series. 

The marked morphological differences in C. mccordi from geo- 
graphically proximate New Guinean and Australian C. novae- 
guineae argue strongly against recent introduction via human 
trade. In like manner, the significant similarities of C mccordi 
with the more geographically distant southeastern New Guinean 
C. pritchardi argues against a recent introduction. 

A more likely scenario to explain the presence of C mccordi 
on Roti is the possibility that both C pritchardi and C. mccordi 
represent relict populations of ancestral Chelodina subgeneric 
group "A" stock, living on the outlying periphery of the previously 
exposed margins of the continental Sahul Shelf during earlier 
periods of lower sea levels and shelf emergence (Jongsma, 1970; 
Doutch, 1972; Galloway and Loffler, 1972). During one of the 
periods that the Sahul Shelf was fully exposed to its 200 meter 
depth (Fig. 1), C. mccordi or its ancestor could potentially have 
reached Roti by rafting across the narrow deep oceanic channel 
that would have then separated the island from the northwestern 
shore of the exposed shelf In addition, C. pritchardi could have 
reached southeastern Papua New Guinea across what would then 
have been a more broadly exposed Torres Strait land-bridge. 
Subsequently, with the partial submergence of the shelf the two 
species were left as peripheral, isolated, relict populations while 
the continental Australo-New Guinean form evolved into what 
is now C novaeguineae. That large continental population then 
eventually became secondarily split by the much later appearance 
of Torres Strait separating New Guinea from Australia (occurring 
about 8,000 years ago), when sea levels rose to their present levels. 
This hypothesis is partially supported by the evidence found in 
skull morphologies, which suggests that both C. mccordi and C. 
pritchardi are intermediate between the primitive C. longicollis 
and the derived C novaeguineae. In addition, it suggests a long 
period of isolation of both C mccordi and C. pritchardi from 
"continental" C. novaeguineae stock. Further, it raises the pos- 
sibility that New Guinean and Australian forms of C. novaegui- 
neae may also be differentiating, as suggested by findings of slight 
differences in skull osteology between these two geographic iso- 
lates (Rhodin, 1993). 



1994 NEW C7/Z:"L6>D//V.4 FROM ROTI 27 



The time frame for this hypothesized phylogenetic scenario is 
hard to specify. The oldest known fossil of Chelodina is from the 
Early to Middle Miocene (ca. 28 million YBP) of northwestern 
Queensland, Australia (Gaffney et al, 1989), and is very similar 
to modern representatives of the genus. It is certainly conceivable 
that much of the dispersal suggested above could have taken place 
during Late Miocene and Early Pliocene times (12-28 million 
YBP) when large land-bridge connections were present between 
New Guinea and Australia (Doutch, 1972; Galloway and Loffler, 
1972). During this time there may even have existed some short- 
lived land-bridges between Australia and the southeastern In- 
donesian islands such as Timor and Roti (Doutch, 1972). Inter- 
estingly, one species of marsupial mammal from Australia (the 
cuscus, Phalanger orientalis) is found on Timor, also suggesting 
possible previous connections between the two areas (Cox, 1970), 
though Glover (1971) states that the cuscus probably represents 
an introduction by prehistoric man sometime later than 13,500 
YBP (earliest evidence of man on Timor). 

In addition, Jongsma (1970) has shown that the Sahul Shelf 
was fully exposed down to a depth of 200 meters as recently as 
the lUinoisan-Riss glaciation, about 1 70,000 years ago. Later, sea 
levels were again down to about 1 60 meters during the most recent 
Wisconsin-Wurm glaciation about 1 8,000 years ago. During these 
recent times, the Torres Strait land-bridge served as a continual 
connection between New Guinea and Australia between at least 
80,000 and 8,000 years ago (Chappell, 1976). 

It is therefore likely that C mccordi reached Roti during one 
of several distinct times: 1) Late Miocene to Early Pliocene times, 
ca. 12-28 million YBP; 2) Illinoisan-Riss glaciation, ca. 170,000 
YBP; 3) Wisconsin-Riss glaciation, ca. 18,000 YBP; or 4) intro- 
duced by prehistoric man sometime later than ca. 13,500 YBP. 
Other periods of potential dispersal probably also occurred be- 
tween the Pliocene and Recent periods. 

The phylogenetic relationships within Chelodina subgeneric 
group "A" have already been hypothesized and discussed by 
Rhodin (1993). Within the group, I regard C. steindachneri as the 
most primitive, with the group becoming more specialized and 
derived in a series that progresses through C longicoUis, C. pritch- 
ardi, and C novaeguineae to C reimanni, the most derived mem- 



28 



BREVIORA 



No. 498 



reimanni novaegu'meae mccordi pritchardi longicollis steindachneri 




Figure 10. Hypothesized phylogenetic relationships of the six currently rec- 
ognized species ofChelodina subgeneric group "A". The monophyly ofChelodina 
"A" follows Georges and Adams (1992). Characters supporting the intrageneric 
nodes are as follows: Node 1 : partial or complete loss of chelid foramen; Node 
2: wide parietal roof, narrow triturating surfaces, parallel pterygoids; Node 3: 
partial reduction in parietal roof width, slightly widened triturating surfaces; Node 
4: narrow parietal crest, flaring pterygoids, wide triturating surfaces, deep robust 
skull. 



ber of the group. Within this phylogeny, C. mccordi appears to 
be most closely related to C pritchardi (Fig. 10), sharing the 
derived features of lack of chelid foramina and partially narrowed 
parietal roof, as well as the plesiomorphic features of a shallow 
skull, decreased robusticity, narrow triturating surfaces, and par- 
allel pterygoid processes. The two species C reimanni and C 
novaeguineae share the derived features of a narrow parietal crest, 
flaring pterygoid processes, wide triturating surfaces, deep skulls, 
increased robusticity, and loss of chelid foramina. 

In view of the isolated occurrence ofChelodina mccordi on the 
very small island of Roti, where available habitat may be limited, 
and human utilization pressures are perhaps heavy, an investi- 
gation into the population and survival status of the species needs 
to be undertaken. Basic ecological and life history data on the 
species are also extremely limited and further investigation is 
needed. Finally, the application of modem methods of molecular 



1994 NEW CHELODINA FROM ROTI 29 

phylogenetic analysis to the species should be pursued to help 
confirm or falsify the hypothesized relationships presented here. 

ACKNOWLEDGMENTS 

I am grateful to William P. McCord who made this work pos- 
sible by obtaining the majority of the study specimens and do- 
nating them to me and to the Museum of Comparative Zoology 
for formal description. The collecting efforts and logistics help of 
Frank Yowono are also much appreciated. In addition, I thank 
both Marinus S. Hoogmoed and L. D. Brongersma of the Leiden 
Museum for making the original Dr. Ten Kate specimens avail- 
able and for relinquishing claims to the description. Curatorial 
assistance was gratefully obtained from Jose P. Rosado. Manu- 
script comments by John B. Iverson and John L. Carr are also 
appreciated. All illustrations were prepared by the author. 

APPENDIX 

Comparative material examined of Chelodina longicoUis, C. 
novaeguineae, C. pritchardi, C. reimanni, and C. steindachneri a\\ 
listed in Appendix in first paper of this series (Rhodin, 1993). See 
text for specimens of C. mccordi examined. Collection acronyms 
utilized in present paper are as follows: 

AGJR = personal collection of Rhodin; MCZ = Museum of 
Comparative Zoology; RMNH = National Museum of Natural 
History, Leiden. 

LITERATURE CITED 

AoKJ, R. 1977. The occurrence of a short-necked chelid in the Palau Islands. 

Japanese Journal of Herpetology, 7(2): 32-33. 
BouLENGER, G. A. 1888. On the chelydoid chelonians of New Guinea. Annali 

del Museo Civico di Storia Naturale de Genova, (2a)6: 449^52. 
. 1889. Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles 

in the British Museum (Natural History). London, Trustees of the Museum. 

3 1 1 pp. 
BuRBiDGE, A. A., J. A. W. KiRSCH, AND A. R. Main. 1974. Relationships within 

the Chelidae (Testudines: Pleurodira) of Australia and New Guinea. Copeia, 

1974: 392-409. 
Cann, J. 1978. Tortoises of Australia. Sydney, Angus and Robertson. 79 pp. 
Chappell, J. 1976. Aspects of late Quaternary palaeogeography of the Austra- 
lian-East Indonesian region, pp. 1 1-22. In R. L. Kirk and A. G. Thome (eds.), 



30 BRE\ lOm No. 498 



The Origin of the Australians. Canberra, AustraHan Institute of Aboriginal 
Studies. 

Cox, C. B. 1970. Migrating marsupials and drifting continents. Nature, 226: 
767-770. 

De Rooij. N. 1915. The Reptiles of the Indo-Australian Archipelago. I. Lac- 
ertilia, Chelonia, Emydosauria. Leiden, E. J. Brill. 334 pp. 

DouTCH, H. F. 1 972. The paleogeography of Northern Australia and New Guin- 
ea and its relevance to the Torres Strait area, pp. 1-10. In D. Walker (ed.), 
Bridge and Barrier: The Natural and Cultural History of Torres Strait. Can- 
berra, Australian National Univ. 

Ferreira, J. B. 1 897. Sobre alguns reptis ultimamente enviados a seccao zoologi- 
ca do Museu de Lisboa. Jomal de Sciencas Mathematicas Physicas e Naturaes, 
Lisboa, (2)5: 111-116. 

Gaffney, E. S., M. Archer, and A. White. 1989. Chelid turtles from the 
Miocene freshwater limestones of Riversleigh Station, northwestern Queens- 
land, Australia. American Museum Novitates, 2959: 1-10. 

Galloway, R. W., AND E. LoFFLER. 1972. Aspects of geomorphology and soils 
in the Torres Strait region, pp. 1 1-28. //; D. Walker (ed.). Bridge and Barrier: 
The Natural and Cultural History of Torres Strait. Canberra, Australian Na- 
tional Univ. 

Georges, A., and M. Adams. 1992. A phylogeny for Australian chelid turtles 
based on allozyme electrophoresis. Australian Journal of Zoology, 40: 453- 
476. 

Gibbons, J. W., and J. E. Lovich. 1990. Sexual dimorphism in turtles with 
emphasis on the slider turtle (Trachemys schpta). Herpetological Mono- 
graphs, 4: 1-29. 

Glover, I. C. 1971. Prehistoric research in Timor, pp. 158-181. In D. J. Mul- 
vaney and J. Golson (eds.). Aboriginal Man and Environment in Australia. 
Canberra, ANU Press. 389 pp. 

GooDE, J. 1967. Freshwater Tortoises of Australia and New Guinea (in the 
Family Chelidae). Melbourne, Lansdowne Press. 1 54 pp. 

IvERSON, J. B. 1986. A Checklist with Distribution Maps of the Turtles of the 
World. Richmond, Indiana, Privately Printed. 283 pp. 

Jongsma, D. 1970. Eustatic sea level changes in the Arafura Sea. Nature, 228: 
150-151. 

LiDTH DE Jeude, T. W. VAN. 1895. Reptiles from Timor and the neighbouring 
islands. Notes Leyden Museum, 16: 1 19-127. 

Philippen, H.-D., and P. Grossman. 1 990. Fine neue Schlangenhalsschildkrote 
von Neuguinea: Chelodina reimanni sp. n. (Reptilia, Testudines, Pleurodira: 
Chelidae). Zoologische Abhandlungen, Staatliches Museum fur Tierkunde, 
Dresden, 46(5): 95-102. 

Rhodin, a. G. J. 1 994. Chelid turtles of the Australasian Archipelago: I. A new 
species of Chelodina from southeastern Papua New Guinea. Breviora, 497: 
1-36. 

Rhodin, A. G. J., and R. A. Mittermeier. 1976. Chelodina parkeri. a new 
species of chelid turtle from New Guinea, with a discussion of Chelodina 



1994 NEW CHELODINA FROM ROTI 31 



siehenrocki Werner. 1901. Bulletin of the Museum of Comparative Zoology, 

147(11): 465-^88. 
Ross, C. A. 1986. Comments on Indopacific crocodile distributions, pp. 349- 

354. In Crocodiles. Proceedings of the 7th Working Meeting of the Crocodile 

Specialist Group of the Species Survival Commission of the lUCN convened 

at Caracas, Venezuela, 21 to 28 October 1984. lUCN Publ. NS. 
Shaw, G. 1794. Zoology of New Holland. Vol. 1. London, J. Davis. 33 pp. 
SiEBENROCK, F. 1914. Einc neue Chelodina Art aus Westaustralien. Anzeiger 

Akademischen Wissenschaften Wien. 17: 386-387. 
Ten Kate, H. F. C. 1894. Verslag eener reis in de Timorgroep en Polynesie. 

IV. Roti. — Savoe. Tijdschrift van het Koninklijk Nederlandsch Aardrijkskun- 

dig Genootschap, (2)1 1: 659-700. 
Werner, F. 1901. Ueber Reptilien und Batrachier aus Ecuador und Neu-Guinea. 

Verhandlungen der Zoologisch Botanischen Gesellschaft Wien, 51: 593-603. 
WiCHMANN, A. 1 892. Die Insel Rotti. Petermanns Geographischer Mitteilungen, 

51:97-103. 
Williams, E. E. 1 950. Variation and selection in the cervical central articulations 

of living turtles. Bulletin of the American Museum of Natural History, 94: 

510-561. 



B R E V I O R A 

I' r r s i 1 

Museiim of Comparative Zoology 

us ISSN 0006-9698 f -" A R^/ ^ P "^ 

Cambridge, Mass. 2 February 1994 Number 499 



AN ECOLOGICAL STUDY OF THE ENDEMIC 

HISPANIOLAN ANOLINE LIZARD, 

CHAMAELINOROPS BARBOURI 

(LACERTILIA: IGUANIDAE) 

Glenn Flores' -, John H. Lenzycki, 
AND Joseph Palumbo, Jr. 

Abstract. We studied the ecology and behavior of Chamaelinorops barbouri 
at two sites. C. barbouri has very specific habitat requirements: montane ravines 
with abundant leaf litter, well-shaded by intact forest canopy. It is an almost 
exclusively terrestrial lizard, preferring leaf litter in deep shade. Despite its non- 
basking, shade-loving habits, C. barbouri maintains its body temperature well 
above air temperature, and linear regression of body temperature and air tem- 
perature data yield a fairly low regression coefficient; this finding is surprising in 
comparison to the thermal biology of other forest-dwelling, non-basking anoles, 
and Greater Antillean anoles in general. We found Chamaelinorops barbouri to 
be cryptic, sedentary, and elusive, and thus difficult to study behaviorally. It is 
highly specialized ecologically, morphologically, and behaviorally for life in the 
leaf litter, much more so than any other anole. 

INTRODUCTION 

Since its discovery in 1 9 1 9 by K. P. Schmidt, the anohne hzard 
Chamaelinorops barbouri has remained in animal of enigma. Over 
half a century passed from the time of Schmidt's (1919) descrip- 
tion before the systematics and distribution of this endemic His- 
paniolan anoline were worked out satisfactorily, and yet the pre- 
cise type locality is still not known and probably never will be. 
This lizard has a unique vertebral column, not duplicated by any 
other vertebrate, of which the functional significance (if any) is 
still completely unknown (Forsgaard, 1983). Equally mystifying 



' Present Address: Robert Wood Johnson Clinical Scholars Program, Yale Uni- 
versity School of Medicine, IE-61 SHM, P.O. Box 3333, New Haven, Connecticut 
06510-8025. 
- To whom reprint requests should be addressed. 



BREVIORA No. 499 



are the relationships of Chamaelinowps; it has been argued that 
the genus is either very derived (Etheridge, 1960; Wyles and Gor- 
man, 1 980), having arisen from within Anolis, or is very primitive 
(Williams, 1977; Case and Williams, 1987), indeed, the most 
primitive of living anoles. 

Yet of all the enigmas of the biology of Chamaelinowps bar- 
bouri, we are most ignorant of its behavior and ecology. Our only 
information to date of C barbouri behavior is limited to a single 
study of display behavior in captive males (Jenssen and Feely, 
1991). No study has ever been conducted before on the ecology 
of C barbouri. The sum of our knowledge in this area is limited 
to a few sentences on the preferred habitat (Thomas, 1966) and 
some remarks on the conditions under which eleven specimens 
were collected (Schwartz and Inchaustegui, 1980; Franz and Cor- 
dier, 1986). Our limitation in knowledge is primarily attributable 
to the inability of a non-native collector to procure a series of 
specimens. Only local residents of an area where C barbouri is 
found are able to secure a series, usually with great ease. At 
the time of the most recent work on Chamaelinorops (Schwartz 
and Inchaustegui, 1980), a total of fifty specimens had been col- 
lected, of which the great majority (39) and the only large series 
(10) had been captured by natives. 

Herein we report the results of an ecological study intended to 
elucidate some of the mysteries of the ecology and behavior of 
Chamaelinorops barbouri. 

MATERIALS AND METHODS 

A large series of Chamaelinorops barbouri can be obtained at 
a small settlement known to herpetologists as The Haitian Village 
and to natives as " Ande Javi," located about 1 5 km SE of the 
town of Cabral and near the city of Barahona in the Provincia de 
Barahona, Dominican Republic. We received as many as 75 liz- 
ards collected by village residents in a "lizard market" in a two- 
hour period, and subsequently had to turn away additional spec- 
imens. Whereas no non-native had ever been able to collect more 
than a few Chamaelinorops, and then only serendipitously, and 
because it was unclear where to search for these lizards, we located 
a Haitian Village resident familiar with Chamaelinorops to act as 
a guide and to demonstrate how to collect them. 



1 994 ECOLOGY OF CHAMAELINOROPS BARBOVRI 



Study Site. There are two areas in the vicinity of The Haitian 
Village where Chamaelinorops is most abundant. Both are close 
to the village, located in ravines sloping to a dry stream bed. The 
slopes have been cleared of most of the understory and planted 
with food plants under an intact mesic primary forest canopy (and 
hence much shade), and are covered by abundant leaf litter often 
interspersed with small, crushed limestone rocks. 

The first site ( 1 ,000 m elevation) is called Caiia Segudinas by 
natives. A small portion of slope on one side of a ravine has been 
burned out and planted over with malanga", a tuberous crop with 
large leaves. Penetrating further into the forest, one encounters 
the dry stream bed and mostly uncultivated ravine slopes where 
Chamaelinorops is found. 

The second site ( 1 , 1 40 m elevation) is known by the villagers 
as Tejul. This site includes broadleaf forest, and an adjacent coffee 
plantation where the canopy is still intact and most of the un- 
derstory has been cleared and planted over with coffee trees. As 
with Caria Segudinas, the area is a ravine whose slopes descend 
to a dry stream bed. 

Methods. To collect Chamaelinorops, residents of The Haitian 
Village require only a short stick (about 2-3 ft long). The collector 
must walk slowly, brushing the leaf litter with the stick in mod- 
erately short, slow strokes while paying close attention to any 
movement. When encountered, the lizard betrays its presence by 
a short hop or run, followed by an abrupt stop or a dive beneath 
the litter. If not clearly sighted and kept track of during flight, 
Chamaelinorops is easily lost by dint of its superb camouflage 
and its ability to rapidly hide within the leaf litter. 

Our study was conducted from 28 July 1985 to 4 August 1985. 
We logged 1 5 person-days during the study period, collecting data 
on 70 Chamaelinorops barbouri. The optimal time for observing 
Chamaelinorops was throughout the morning and into the early 
hours of the afternoon, and so most of our study was conducted 
between 0900 and 1400 hrs. 

The study focused on three aspects of the biology of Chamae- 
linorops barbouri: (1) habitat preference; (2) thermoregulatory 
strategy; and (3) behavior. For habitat preference the "Rand cen- 
sus" was employed. The observer walks through the habitat, re- 
cording the height, diameter, and insolation of the perch where 



4 BREVIORA No. 499 

the lizard was first sighted (Rand, 1964). Perch height was re- 
corded as "underground," "ground," or estimated to the nearest 
foot. Because Chamaelinorops is almost completely terrestrial, 
one of five categories was recorded for every ground observation: 
on dead leaves, under dead leaves, on bare earth, on rocks, or 
under rocks. For perch diameter, seven categories were used: 
underground, on rocks, on ground covered with leaf litter, on bare 
ground, trunk (perch diameter greater than three inches), branch 
(perch diameter between one-half inch and three inches), and twig 
(perch diameter less than one-half inch). For insolation, the amount 
of shade at the perch was estimated as one of the following: full 
shade, moderate shade, light shade, or unshaded. For thermo- 
regulatory data, Schultheis quick-reading mercury thermometers 
were inserted into the cloaca as soon as possible after lizard cap- 
ture in order to minimize hand contact with the lizard's body. 
Once sighted, Chamaelinorops was easy to capture; all lizards 
were captured within five seconds, eliminating the possibility of 
false body temperature elevation due to a protracted chase. The 
air temperature 1 cm above the perch site was then immediately 
recorded with the thermometer bulb shaded, after the bulb was 
completely dry. Cloacal temperatures were recorded for 64 of the 
70 Chamaelinorops sighted. For behavioral data, individuals were 
often observed for several minutes before or after capture. Ad- 
ditional observations were also made on captive individuals 
maintained in a terrarium. 

Thermoregulation was assessed by the method of Huey and 
Slatkin (1976), using the regression coefficient from the linear 
regression of body temperature and air temperature: a regression 
coefficient near implies careful thermoregulation (body tem- 
perature independent of air temperature), whereas a regression 
coefficient near 1 implies thermoconformity (complete thermal 
passivity). A Spearman's coefficient of rank correlation of body 
and air temperature data was also calculated. There was no sig- 
nificant difference (/-test of the differences between two means, 
P » 0.05) between the body temperatures of adult males and 
adult females, and, although there was a significant difference 
between the body temperatures of adults and juveniles (0.05 > 
P > 0.02), the sample size of juveniles (3) was too small to be 



1 994 ECOLOGY OF CHAMAELINOROPS BARBOUR/ 



Table 1 . Perch height observations for Chamaelinorops barbouri. Perch 

HEIGHT categories: -G = UNDERGROUND; G = GROUND; ALL OTHER CATEGORIES 
ARE ESTIMATES TO THE NEAREST FOOT. (UnSEXED ADULTS = ADULTS OBSERVED BUT 

NOT CAPTURED.) 

-G G 1 2 3 4 5 



Males 




35 


1 


Females 


1 


25 


1 


Juveniles 




2 


1 


Unsexed adults 




2 




Totals 


1 


64 


3 



useful; consequently all sex and age classes were pooled in the 
presentation of thermoregulatory data. The standard criterion of 
statistical significance was utilized {P < 0.05). All statistical tests 
follow Sokal and Rohlf (1981). 

RESULTS 

Observations at the two study sites indicate that Chamaeli- 
norops has the following special habitat requirements: (1) An 
intact forest canopy providing abundant shade; (2) Abundant leaf 
litter; (3) Conditions (1) and (2) located in a ravine with slopes 
ending in a dry stream bed (the association with a dry stream bed 
in both of our study sites may have been a coincidental finding, 
but we never succeeded in finding Chamaelinorops in habitats 
meeting conditions (1) and (2) but not (3), and Chamaelinorops 
collected for us by residents from the Sierra de Neiba on the 
North Island of Hispaniola were always reported as having come 
from habitats exhibiting the above three conditions); (4) Montane 
elevations, usually about 1,000 m (but can range from 300 m to 
1,710 m [Schwartz and Inchaustegui, 1980]). 

Data on perch height (Table 1) indicate that Chamaelinorops 
is almost exclusively a terrestrial anole and not at all arboreal. 
Chamaelinorops was never observed in any type of situation that 
could be considered arboreal— trees, saplings, bushes, low vege- 
tation, etc.— despite intensive searches for lizards in these situ- 
ations. Although it is possible that Chamaelinorops was missed 
in arboreal situations due to its extremely cryptic appearance and 



BREVIORA 



No. 499 



Table 2. Types of terrestrial perches chosen by Chamaelinorops 



BARBOURI. 





On 


Under 


On dead 


Under dead 


Bare 




rocks 


rocks 


leaves 


leaves 


earth 


Males 


1 


1 


24 


5 


5 


Females 


1 


2 


15 


3 


4 


Juveniles 






1 


1 




Unsexed adults 




1 




1 




Totals 


2 


4 


40 


10 


9 



'Other terrestrial situations in which a single individual was observed: on a log, 
underground, on a dead banana leaf, within a pile of dead branches on dead leaves. 

habits, lizards could never be induced to climb, even when placed 
next to or on arboreal perches and provoked to a state of severe 
alarm or agitation. 

Chamaelinorops predominantly prefers ground covered with 
leaf litter (Tables 2 and 3). Chamaelinorops was observed on dead 
leaves more frequently than on all other terrestrial perches com- 
bined (Table 2). Similarly, the number of observations of Cha- 
maelinorops on leaf litter exceeded the number of observations 
on all other categories of perch diameter combined (Table 3). In 
addition, no observations for the trunk or twig categories were 
recorded. 

Chamaelinorops overwhelmingly prefers shady perches to sun- 



Table 3. Perch diameter observations for Chamaelinorops barbouri. Tr 
= trunk (>3" IN diameter); Br = branch ('/2"-3" in diameter); Tw = twig 
(<'/2" IN diameter). (See Materials and Methods Section in text for addi- 
tional explanation of perch diameter categories.) 









Ground 
















covered 












Under- 




by leaf 












ground 


Rock 


litter 


Ground 


Tr 


Br 


Tw 


Males 




1 


30 


6 








Females 


1 


1 


20 


6 








Juveniles 






2 






1 




Unsexed adults 






1 


1 








Totals 


1 


2 


53 


13 




1 





1994 



ECOLOGY OF CHAMAELINOROPS BARBOURI 




SHADE CATEGORY 

Figure 1 . Shade category observations for Chamaelinorops barbouri. FS = full 
shade; MS = moderate shade; LS = light shade; US = unshaded. 



ny ones. About half of our observations found Chamaelinorops 
in full shade, and lizards were seen in full and moderate shade 
(the two categories with the greatest shade) in over three-quarters 
of all observations (Fig. 1). Indeed, only three of 70 individuals 
were observed in unshaded conditions, over 1 1 times less fre- 
quently than in full shade. 

Seven lizard species are sympatric with Chamaelinorops at the 
study sites, including five species of anoles (Fig. 2). None of the 
other lizard species substantially overlaps with Chamaelinorops 
in its habitat preference. 

The endemic Hispaniolan anguid Wetmorena haitiana mylica 
can be found in the same forest situation as Chamaelinorops, but 
is encountered only under rocks and appears to be a burrower. It 
occurs not only in well-shaded forest habitats but also in dis- 
turbed, open habitat, as long as there are rocks for it to hide 
under. The second anguid lizard at the study sites, Celestus cos- 
tatus oreistes, is seen only in open, disturbed habitat and never 
is syntopic with Chamaelinorops. 

Five species ofAnolis are found at the study sites, and all occur 
primarily in ecotone habitat where the forest abruptly meets the 
heavily disturbed, open areas. Five of Williams's (1983) eco- 
morphs are represented. There are two trunk-crown ecomorph 
species, Anolis coelestinus (a trunk-crown I ecomorph species = 
large) and A. singularis (a trunk-crown II ecomorph species = 
small). Both occur on leaves and branches of the canopy and 
upper trunk of trees. Anolis distichus is a trunk ecomorph species, 
occurring primarily on tree trunks between the trunk-crown and 
trunk-ground species. Anolis cybotes is a trunk-ground species, 



BREVIORA 



No. 499 




1 994 ECOLOGY OF CHAMAELINOROPS BARBOVRI 



Table 4. Thermal biology data for Chamaelinorops barbouri. ' Tg = body 
temperature; T,, = air temperature; N = number of individuals observed; 

X ± SD = MEAN PLUS OR MINUS ONE STANDARD DEVIATION; Ry, = SpEARMAN'S 
coefficient of rank CORRELATION OF BODY AND AIR TEMPERATURES. 





N 


Th 


1 


T, 






A- ± SD 


Range 


A ± SD 


Range 


Males 


36 


25.6 ± 1.4 


22.0-31.0 


22.3 ± 1.5 


20.0-26.0 


Females 


23 


26.4 ± 2.0 


22.0-30.0 


22.8 ± 1.5 


21.0-27.0 


Juveniles 


3 


27.5 ± 0.5 


27.0-28.0 


23.8 ± 2.0 


22.0-26.0 


Unsexed adults 


2 


28.2 ± 1.8 


27.0-29.5 


24.0 ± 1.4 


23.0-25.0 


Total 


64 


26.0 ± 1.9 


22.0-31.0 


22.6 ± 1.5 


20.0-27.0 



'rs = 0.521 (P < 0.001). 

occurring on the lower trunks of trees and on the ground, usually 
in close proximity to a tree trunk. Anolis bahorucoensis is found 
primarily on bushes (bush ecomorph). Of the five Anolis, A. coe- 
lestinus and A. singidaris are rarely encountered at the study sites, 
and only A. bahorucoensis and A. distichus are common in or 
near forests inhabited by Chamaelinorops. Of all Anolis, A. ba- 
horucoensis penetrates the forest most deeply (although still pri- 
marily an ecotone species) and on several occasions was collected 
on bushes in Chamaelinorops habitat. 

Given that Chamaelinorops shows such a strong preference for 
well-shaded habitat, we were surprised to discover that its mean 
body temperature (MBT) is well above the mean air temperature 
(MAT) (Table 4 and Fig. 3). The MBT is 26.0°C, and ranges from 
22.0°C to 31.0°C; the MAT is 22.6°C and ranges from 20.0°C to 
27.0°C. The difference between the MBT and the MAT, Xtb - 
Xta, is about 3.5°C. Linear regression of body temperature and 
air temperature yields a regression coefficient of 0.69, suggesting 
Chamaelinorops is more of a thermoconformer than a thermo- 
regulator. 

The very patchy abundance oi Chamaelinorops, both spatially 



Figure 2. Perch and climatic preferences for the eight lizard species occurring 
at The Haitian Village study sites. Names beginning in lower case letters are species 
of Anolis. Hatched lines represent shaded habitat. 



10 



BREVIORA 



No. 499 



O 

LiJ 

or 

ID 

q: 

UJ 
Q. 

UJ 

I- 
>- 

Q 
O 

m 













31 


- 




• 


' .d' 


29 


_ 


















. 




• « • 


,' • A^/ 


27 


- 






/ 




- 


• 


*> fry # 


• 


25 


- 


/ 


^<^' « # # 


/ 




_ 


/ 


• • •^Z 






> 




• // 




23 


- 


1 


• / 

* y/ 
1/ , 1 


1 I 1 i 1 



19 21 23 25 27 

AIR TEMPERATURE (°C) 



29 



Figure 3. A plot of the body temperature and air temperature data for Cha- 
maelinorops barbouh. The solid line represents the isothermal line (body tem- 
perature = air temperature, Tg = T^). The dashed line is the linear regression of 
body temperature and air temperature, for which the equation is provided. The 
correlation coefficient (rj = 0.52. Multiple individuals with the same data are 
depicted by overlapping circles. 



and temporally, was impressive. Chamaelinorops only occurs in 
the selected areas around The Haitian Village where its stringent 
habitat requirements are met; elsewhere in the area it appears to 
not occur at all. It is abundant only during the morning hours 
and early afternoon; after 1400 hrs, Chamaelinorops completely 
disappears. Additionally, even when we visited ideal habitat at 
ideal hours where the previous day Chamaelinorops had been 
found in abundance, on some occasions, it was difficult or im- 
possible to find any lizards. 

Data on behavior is limited, primarily due to the sedentary 
habits and cryptic nature o{ Chamaelinorops. We often attempted 
to observe individuals in the field, but were continually rewarded 
with nothing but a prolonged view of an immobile lizard (even 
after up to 15 minutes of observation). Observations on captive 
individuals yielded similar results. We became most famiHar with 
Chamaelinorops escape behavior, which usually consists of a very 
brief scampering dash or series of hops (of no more than several 



1 994 ECOLOGY OF CHAMAELINOROPS BARBOURI 1 1 

inches) followed by an abrupt freeze. This tactic proves quite 
effective: the leaf litter, bare earth, and crushed limestone back- 
ground beautifully conceal a stationary Chamaelinorops with its 
color pattern of black markings on pale white, gray, and tan tones, 
and an outline that closely resembles a dead leaf. When pressed 
further, Chamaelinorops often dives into the leaf litter. If it does 
not dive into the leaf litter, Chamaelinorops continues its initial 
tactic of a brief dash or series of hops followed by an abrupt stop, 
starting and stopping until the threat abates or an object to hide 
behind is encountered. When subjected to a prolonged threat, 
Chamaelinorops seems to tire quickly, increasingly abbreviating 
the flight and extending the stationary portion of the escape. 

When captured, males can be quite aggressive, holding out the 
stark black dewlap, opening the mouth to expose the black mu- 
cosa, and even producing a weak bite if greatly agitated. Females 
and juveniles were never observed exhibiting such behavior when 
captured, and indeed one could reliably identify a male by such 
aggressive behavior alone. Chamaelinorops also moved abruptly 
and became quite agitated when one of us produced high-pitched 
whistles and bird-like clicking sounds (resembling the call of 
grackles). 

Although no behavioral interactions were observed in the field, 
on several occasions multiple individuals were found together in 
a small area, including one observation of five individuals in a 
three by three meter area. One question which still remains un- 
settled is whether Chamaelinorops burrows. One individual was 
found under several inches of sandy soil and stones, and several 
were collected under rocks and leaf litter (Table 2). Residents of 
The Haitian Village believe Chamaelinorops burrows, and we 
were shown holes in which lizards were believed to live. However, 
we never found lizards after excavating such holes, and in cap- 
tivity individuals were never observed to burrow despite being 
provided with ample soil in terraria. 

DISCUSSION 

General Ecology. Chamaelinorops barbouri is the only known 
West Indian anole specialized for life in leaf litter. Indeed, of all 
the anoles, only Anolis humilis from Central America approaches 
Chamaelinorops in its preference for leaf-litter habitat (Brattstrom 



12 



BREVIORA 



No. 499 



Table 5. Chamaelinorops barbouri and Anolis humilis compared. Data 
for a. humilis is ftiom fitch (1973, 1975). 



Feature 



Chamaelinorops barbouri 



Anolis humilis 



Snout-vent length 
Color 

Dewlap 

Modal perch 



Body proportions 



Scales 



Foraging behavior 



Defensive behavior 



Usually <50 mm 
Light brown/tan with 

variable pattern 
Small, black with yellow 

edge 
On ground, mostly leaf 

litter, marked shade 

preference 



Head relatively short, 
body compressed, long 
limbs, very long tail 

Middorsal scales greatly 
enlarged, keeled, 4-10 
rows 

Sit-and-wait predator (?) 
on ground 

Primarily crypsis, via im- 
mobility, start and stop 
flight; always flees on 
ground, never climbs 



<45 mm 

Dark brown/olive with 
variable pattern 

Large, red with yellow 
edge 

Often leaf litter, but most- 
ly above ground on but- 
tressed roots of large 
trees; marked shade 
preference 

Stout body, short limbs 
and tail 

Middorsal scales greatly 
enlarged, keeled, 8-12 
rows 

Active forager on ground 
and buttressed roots of 
large trees 

Primarily crypsis, via start 
and stop flight; always 
flees towards tree root 
buttresses on ground or 
climbs tree base 



and Howell, 1954; Fitch, 1973, 1975; Talbot, 1977). Anolis hu- 
milis exhibits several similarities to Chamaelinorops (Table 5), 
including small size, elements of the color pattern, yellow edging 
of the dewlap, preference for leaf litter and shade, greatly enlarged 
keeled middorsal scales, aspects of escape behavior, and absence 
ofbasking behavior (Fitch, 1973, 1975). However, the two anoles 
differ markedly in a number of other features which directly reflect 
the greater specialization oi Chamaelinorops for life in leaf litter. 
Chamaelinorops is exclusively terrestrial and predominantly found 
in leaf litter, whereas A. humilis spends much of its time above 
ground and, when in leaf litter, is almost always centered around 
buttress roots of large trees. Chamaelinorops relies more on stay- 
ing immobile or hiding, and flees only on the ground; A. humilis 



1994 ECOLOGY OF CHAMAELINOROPS BARBOURI 13 

is much more mobile and usually flees toward trees, occasionally 
"squirreling" around the tree base to the opposite side of the 
threat. In form, Chamaelinorops more closely resembles a dead 
leaf, with the "body extremely compressed, the sides vertical or 
concave . . ." (Schmidt, 1919) and an overall angular appearance, 
compared to the "stubby-bodied" (Fitch, 1975) A. humilis. So, 
in every way— ecologically, behaviorally, and morphologically— 
Chamaelinorops is more specialized for life in leaf litter than A. 
humilis. Indeed, Chamaelinorops is the only known true leaf-litter 
specialist among all of the anoles for which ecological information 
is available. 

Although no other West Indian anole is a leaf-litter specialist, 
three species are known to be as markedly terrestrial as Cha- 
maelinorops. Anolis armouri and A. shrevei of the Dominican 
Republic are commonly found under stones, and A. armouri also 
perches horizontally on fallen logs (E. E. Williams, in lit.). Ruibal 
(1964) provided a description of the habitat preferences of the 
Cuban A. ophiolepis, which is of interest in comparison to Cha- 
maelinorops: "This is not a rare species; it is merely rarely caught. 
This is the only truly terrestrial species of the Cuban anoline 
lizards. The species is found in pastures and savannas, on the 
ground and runs to take refuge in grass tussocks. I have observed 
the species sleeping on the leaves of small bushes." 

We tentatively suggest that the shared terrestrial habitat pref- 
erences and behaviors seen in Chamaelinorops and these three 
Anolis may represent a "weak" ecomorph, a "ground" category. 
Certain anole ecologies, such as a preference for ground habitat, 
may not select for strong, completely congruent behavior and/or 
morphology. This may reflect the great variability of ground hab- 
itats (in soil types, cover such as leaf litter versus grass, open 
surfaces versus dense undergrowth, to name a few) as compared 
to a more uniform surface such as a tree trunk. Thus, these four 
"ground" anoles, although alike in their terrestriality and partic- 
ular behaviors, are morphologically different, in contrast to the 
"standard sequence" ecomorphs (Williams, 1983) which show 
strong correlations among morhpology, ecology, and behavior. 

Data on perch height preference in Chamaelinorops show that 
it is an exclusively terrestrial anole. In situations where it was not 
clear whether an individual might have chosen a perch above 



14 BREVIORA No. 499 

ground level, we always decided in favor of the greatest perch 
height category possible. However, in all five such observations, 
stone or boulder perches were involved which closely resembled 
the ground in having leaf litter and/or moss cover; furthermore, 
these perches never arose abruptly but were in continuous and 
gradual contact with the ground. Hence, to Chamaelinorops, such 
perches are probably just another varied portion of the constantly 
changing ground surface. If Chamaelinorops ever climbs above 
the ground, it appears to be a rare exception. 

The observations of others (Schwartz and Inchaustegui, 1980; 
Franz and Cordier, 1986) are largely in agreement with our data, 
except some cases of use of low arboreal perches have been noted. 
Schwartz and Inchaustegui (1980) reported that all but two spec- 
imens for which they had information came from ground situa- 
tions; one individual was found "... at night sleeping totally 
exposed on the curving bare branch of a small woody legume 0.3 
m above the ground surface," and another was ". . . in a crevice 
in a large tree about 1.2 m above the ground in a field being 
actively cut and weeded by a number of native workmen." Franz 
and Cordier (1986) found all but three and their specimens in 
ground situations; three specimens were collected ". . . among 
twigs in dead shrubs," but no perch height was provided. 

Our data suggest that observations of diurnal arboreality in 
Chamaelinorops probably represent extreme circumstances or un- 
usual instances. Our experiences suggest that the individual found 
1.2 m above the ground in a large tree (Schwartz and Inchaustegui, 
1980) may have been driven there by severe, immediate habitat 
destruction and disturbance, a situation and response unlikely to 
be observed under more natural conditions. The three individuals 
found among twigs in dead shrubs (Franz and Cordier, 1986) may 
actually have been on the ground within the matted twigs, a 
situation we frequently encountered. However, a sleeping indi- 
vidual found on a low arboreal perch (Schwartz and Inchaustegui, 
1980) may represent either the true perch choice for sleeping 
Chamaelinorops, as is the case with most anoles, or unusual cir- 
cumstance; we failed to observe Chamaelinorops sleeping, and, 
to date, this single observation is the only published report avail- 
able. 

The habitat preferences oi Chamaelinorops appear to be rigidly 



1994 ECOLOGY OF CHAMAELINOROPS BARBOURl 15 



specific. We believe that the specific habitat requirements we 
observed will be closely adhered to wherever Chamaelinorops is 
encountered, perhaps varying in the presence of a small stream, 
or the absence of a dry stream bed (but under mesic conditions). 
Although our study sites were in montane broadleaf forest canopy, 
it is not surprising that Chamaelinorops has also been found in 
montane pine forest (Franz and Cordier, 1986), the other type of 
Hispaniolan forest at higher elevations. In Haitian pine forest, 
Franz and Cordier (1986) found Chamaelinorops in ground sit- 
uations, particularly in association with dead pine needles— the 
"leaf litter" of pine forests— and among their locality data ravines 
and basins are mentioned. Schwartz and Inchaustegui (1980) also 
noted Chamaelinorops in association with ravine habitat. 

The patchy abundance of Chamaelinorops, both spatially and 
temporally, is striking. Its activities are apparently limited by the 
time of day and microhabitat requirements, and vary from one 
day to the next. Given the elusive habits and cryptic nature of 
Chamaelinorops, it is no wonder that, for decades, only experi- 
enced residents could collect it in any abundance. 

Behavior. Although the sedentary nature, cryptic coloration, 
and elusive habits of Chamaelinorops prevented all but the most 
cursory portrait if this anole's behavior, some generalizations 
became apparent. Defensive behavior is based primarily on the 
use of camouflage and hiding: Chamaelinorops relies on immo- 
bility or short, abrupt starts and stops, along with leaf litter and 
other objects for hiding. Although foraging behavior was not ob- 
served, we suspect, as Schwartz and Inchaustegui (1980) have 
suggested, that Chamaelinorops is probably a "sit-and-wait" 
predator rather than an active pursuer, since its camouflage and 
sedentary tendencies well suit it for such a predation mode. 

Whether Chamaelinorops burrows, as is claimed by residents 
of The Haitian Village, is still unclear. Supporting such claims is 
the observation of an individual under several inches of soil, and 
the testimony of several local residents. Contradicting these claims 
was our failure to unearth Chamaelinorops from alleged burrows 
pointed out to us by residents, and a lack of evidence of burrowing 
activity in any of the many individuals observed in captivity. 

The possibility of burrowing lends an attractive potential func- 
tional explanation for the peculiar, extremely ossified vertebral 



16 BREVIORA No. 499 

column of Chamaelinowps, particularly in light of the resem- 
blance of its vertebral column to that of only one other vertebrate, 
the mole Scutisorex (see Allen, 1917). However, as discussed 
above, the data available on burrowing is far from conclusive and 
the issue clearly deserves further investigation. Bohme (1982) 
hypothesized that the bony dorsal "shield" found in Chamaeli- 
nowps and certain chameleons of the genus Brookesia serves to 
deter bird predation by maintaining immobility and rigidity after 
being struck by a bird's beak. However, we observed conspicuous 
agitation and alarm elicited in Chamaelinowps in response to 
bird-like whistles and clicks, suggesting Chamaelinowps most 
likely responds to threatened bird predation in a more active than 
passive fashion. Besides a number of bird species, other potential 
predators encountered at the study sites are the colubrid snake 
Antillophis parvifwns and very large centipedes common under 
rocks. 

It is clear from our discussion o{ Chamaelinowps behavior that 
it is quite difficult to obtain useful behavioral data on this cryptic, 
inactive, and highly elusive anole; ethological studies are patently 
needed but will demand the utmost in patient, careful observation 
and perseverance. 

Themoregulation. Chamaelinowps was never encountered 
basking during this study. The possibility might be raised that we 
failed to observe basking because the exceptional camouflage of 
these lizards caused us to overlook some individuals, or startle 
tactics necessary for locating Chamaelinowps resulted in indi- 
viduals moving out of sun patches too quickly for us to note 
basking. However, although we actively searched for Chamaeli- 
nowps in sun patches throughout the study, basking behavior was 
not observed (the three individuals observed in unshaded cir- 
cumstances did not exhibit "classic" lizard basking behavior, i.e., 
they had not oriented and positioned their body to receive max- 
imum solar radiation). Moreover, in the apparent preferred hab- 
itat of Chamaelinowps, sun patches are rare, small, and usually 
far apart even at midday, due to the thick forest canopy and 
frequent additional coffee trees, saplings or low bushes. Hence, 
we feel confident in stating that basking, if it occurs at all in 
Chamaelinowps, constitutes an insignificant proportion of this 
lizard's daily activities. 



1994 ECOLOGY OF CHAMAELINOROPS BARBOURI 17 

Given that Chamaelinowps was never observed basking and 
that it overwhelmingly prefers shaded deep forest habitat, the 
data on thermal biology is baffling (Table 4, Fig. 3). Huey and 
Slatkin (1976), in proposing a model of lizard thermoregulation, 
provided important predictions relevant to Chamaelinowps ther- 
mal biology: (1) Thermoregulation is beneficial only when asso- 
ciated costs are low. (2) The cost of raising body temperature 
should be proportional to the distance necessary for shuttling 
between sun and shade or hot and cold microenvironments. Thus, 
the cost of raising body temperature should be greater in closed 
forests than in more open habitats. (3) Lizards living in shaded 
forests (excluding the canopy), where costs of raising body tem- 
perature should be much higher than in open habitats (patches 
of sun for basking are more widely spaced in forests), tend not to 
bask and seemingly are relatively passive to ambient conditions. 
Chamaelinowps occurs exclusively in closed forests where patch- 
es of sun are few and far between; furthermore, Chamaelinowps 
shows a predominant preference for shade within such forest 
habitat (Fig. 1). Hence, the distance necessary for shuttling be- 
tween sun and shade in Chamaelinowps habitat is great, and so 
is the cost of thermoregulation, by prediction (2) above. Since the 
thermoregulation is beneficial only when associated costs are low, 
Chamaelinowps, like many other lizards in shaded forests (Huey 
and Slatkin, 1976), should tend not to bask and should be rela- 
tively passive to ambient conditions. As one might predict, no 
basking behavior was observed, and Chamaelinowps was ex- 
pected to be quite a thermoconformer, maintaining a body tem- 
perature varying little from ambient temperature. 

However, as the data graphically and surprisingly depict (Table 
4, Fig. 3), Chamaelinowps clearly maintains a body temperature 
well above air temperature, with the value of .Vtb - -tjA approx- 
imating 3.5°C. These data are even more impressive when com- 
pared with similar thermal data on other anoles (Table 6). Note- 
worthy is that the value of Xtb - Xta of Chamaelinowps greatly 
exceeds that of all the mainland shade-loving forest anoles for 
which data are available, and also well exceeds that of Anolis 
allogus, A. gundlachi, and A. lucius, the only purely shade-loving 
Greater Antillean forest anoles for which such data are available. 
Indeed, the Xj^ - Xj^ of Chamaelinowps exceeds that of all but 



18 



BREVIORA 



No. 499 






Z 


>■ 


w 


z 
< 


UJ 
UJ 


Q 


^ 


z 


u 

oa 





Q 

UJ 

H 

C/3 


-1 


UJ 


z 




J 


u 





U 


i 


z 




2 


< 


u 

u 
u 

5 


2 


S 


OS 

u 


2 


0. 




II 


on 

< 


< 


PS 


< 


UJ 


■^ 





>^ 


5 


§ 


X 

H 


1 


UJ 

X 
H 
u. 
o 

u 

Oh 

2 

C/5 




5 


ui 


u 




oa 


u. 




§ 


< 
> 


II 


z 




a; 


< 


h 
Z 

UJ 


E 




b£ 




o 


i 


< 

< 


1. 

UJ 

o 
u 


X 

UJ 


s< 


H 


u 


5 


< 


X 


§ 


Q 


H 




UJ 

< 


i 


I 


i 


S 


So 


^ ^ 


3 


Cu 


w 


O 


,* 





0^ 


uj S; 


^ 


Oi 





_J ■T 


u. 
O 


0. 

< 


s 


a 


>■ 


I 
o 

X 


ill 

Bi 
O 


> 
< 


O 


^ 


^ 


f- 




s 





UJ 

CL 


z 


J 


u. 


s 


II 


< 


75 


u 




s 


UJ 


H 


< 


OS 





oi 


Z 


I 


z 


< 




<; 




Uj' 


1- 




Z 


B^ 


UJ 

H 


eh: 


< 

UJ 


D 
1- 
< 





Q 
Z 
< 


Bi 


u. 
O 


u. 

O 




z 


z 


UJ 


UJ 





^ 


X 


H 


2 
< 

1 


-1 

UJ 


< 


< 




oi 

UJ 
0. 


Q 
Z 


P 


Q 


5 


< 


O 


< 


UJ 


UJ 

Bi 

H 


vd 


X 
in 

Q 


>■ 
D 


UJ 


Z 





< 


J 


^ 


a 


Ot! 


ffl 




UJ 


< 


Z 


Z 


0. 


h- 


< 


< 

UJ 


UJ 




5 


S 


H 






X) 
C3 



c 


.^ 


o 


c 




o 


'c/5 




C/5 


'C 




iE 


0£ 


(U 


OJ 


o 


Qi 


CJ 









o 




































__ 








_ 






NO 
















Ov 


00 








00 






ON 




00 


ON 








^ 


vO 


ON 








ON 






"" 




ON 


NO 








L- 


ON 
















»■ 






ON 








U 


















13 












































75 


. 


>^ 








>1 






X) 




>. 


. 








X> 


^^ 


<u 








lU 






■-H 




<u 


. ; 








<U 


a 


3 








3 






3 




3 


a 




VO 


VO 


^ 


_4> 


X 




^^ 


S 


X 


NO 


TT 


OJ 


'a- 


X 


^ 
u 


T3 — 


O^ 


T3 


•a 


o 


00 
On 


ON 


•o 


ON 


On 


o 


ON 


T3 


3 


_^ 


_. 


C 
03 


"o 


c 


00 
On 




_- 


c 


, 




00 

ON 




C 


"o 


CO 
C/5 




03 






N 

c 


^' 


N 






1 




J 

o 






re 


.E 


'3 


'3 


3 


aj 


<u 


m 


'B 


OJ 


3 


o 


OJ 


O 


u 


H 


Di 


QC 


X 


I 


X 


X 


a: 


X 


Qi 


X 


J 


X 


X 


X 





*-» 






re 






c 






CO 






c^ 






re 






2m 






re 






■c 












•c 




x> 


re 
a 

75 -r- 

re •- 


c 
_re 


ID lA ID ^ 

re y « S 


u 
•o 
re 

75 


■o 

re 


75 


X 

o 

75 

c 


■V 

re 


75 

T3 

C 

re 


ed are; 
d and 


75 
75 

2 


75 " un re 


•o 


JZ 


c 


o 


j= 


•a 


^ c 


u 


7) 


Q 




75 


c 


o re 


c 


p-forest 
n sunny 
p-forest 
n, xeric 


n areas 
n areas 
itly close 


♦-< 
75 

4J 
u 

1 

a 


<-• 

re 

3 

't! 


'-£ 

o 
a 

OJ 

■o 
re 
s: 


75 

<2 
a 


_re 

75 
75 

2 


n and cl 
ic grassl 


1 monta 
n park 


QOQOOO^Q 


OX) 

■a 

UJ 


4j c a «J •- a 
Q < O S X O 




NO 


rn 








ON 


NO m 






t^ 


— 








00 


ON 00 




S < < < 

d z z z 


0.71 
1-0. 
0.63 


T 

00 


< 
Z 


00 

d 


< 
Z 


r- — Tj- 


0.35 
0.77 




T 


q 








d 


NO r- 






d 


— 










d d 







o 




u 


75 


a 


 H 


75 






'o 


c 


w 


re 


D. 




C/5 


'•5 




c 



ON 

NO d 

•^ 00 00 

d d 
I 



So 

I 

O 

SI 



00 r\ <^ 

— — Tt 





o 










lA) 


_. d 


— • 


in 


in 




d 


1 1 


fi 
1 


—  


ri 

1 


— ON 


1 1 


1 


00 


1 


r-' d 


d 


d d 


H 


d 


H 





5 



<<1 

3 



^ 

a 



to 



.to -S 



-P ^ :; ;^ s: '-o 
C' -5 S: -« -2 -2 



1^ s t. 

>) Co >5 



1994 



ECOLOGY OF CHAMAELINOROPS BARBOURI 



19 



w 

3 
C 
^-» 
C 

o 

U 



LU 
-1 
PQ 
< 






X) 



c — 
o c 



I 



u 
a 

(/5 



VO 




vO 






VO 


r- 




t^ 






r- 


o 




Ov 






o\ 


— ^ 




^— 






^^ 


. 


rn 


. 


^ 


00 


. 


"a 


ON 


"5 


ON 


ON 


~5 












•«^ 


^ 




^ 


, 




<i> 


x: 


X 


j= 


N 

c 


x:' 


sz 


CJ 


o 


o 


o 


o 




i~t 




CJ 


4-* 


4-» 














■pu 


U- 


U- 


K 


E 


E 



x: 



-o 



!« —; !« 

<U « lU 

fe H 4J 

T3 ' O 

n *^ O 

O Q O 



< < < s < < 

Z Z Z r^ Z Z 



Si 




o 
o 


<o 


o 


? 




0= 


lU 


a 


c 


(rt 


OJ 


c 


u 


'c5 


<u 


Q 


Qi 


Q 



^ O NO 00 — r\l 

d — d — — — 



• *-< 

u 

u 
o. 

CO 

1/5 



T3 

C 

c 



a 

to 
O 



i>5 

to s: 



6o-« s: ci. i? S 



-Si -s: 



20 BREVIORA No. 499 

one Greater Antillean Anolis, A. shrevei, which is known to bask 
frequently (Hertz and Huey, 1981), in contradistinction to our 
observations on Chamaelinorops. Similarly, the regression coef- 
ficient of the linear regression of body temperature and air tem- 
perature for Chamaelinorops, 0.69, puts it distant to non-basking, 
shade-loving anoles like the Costa Rican A. polylepis (regression 
coefficient = 0.93; Hertz, 1974) or the Puerto Rican A. gundlachi 
(regression coefficient = 1.08-1.13; Hertz, 1981), but close to 
average for an open habitat, basking Greater Antillean anole, such 
as A. cristatellus or A. cybotes (Table 6). Hence, given that a 
regression coefficient of represents perfect thermoregulation and 
1 perfect thermoconformity (Huey and Slatkin, 1976), Chamae- 
linorops appears to be thermoregulating much more than it 
"should" be. 

Chamaelinorops appears to be a shade-loving, non-basking 
anoline lizard that inhabits deep montane forest where sun patch- 
es are rare and costly to get to. How can it, then, maintain such 
a high value for Xjq - Xja and such a low regression coefficient 
for the linear regression of body temperature and air temperature? 
As an explanation for this enigma, three possibilities surfaced: 
(1) Chamaelinorops hides and/or sleeps beneath substrates that 
are warmer than air temperature. Bustard (1967) reported that 
geckos active at night can and may thermoregulate under bark or 
rock flakes during the day and achieve body temperatures near 
preferred levels. In order to utilize such a strategy, Chamaeli- 
norops would have to use the limestone stones abundant in its 
habitat, in order to thermoregulate during the day. However, such 
stones are likely to heat up only in open habitat, not in the closed 
shady forest Chamaelinorops inhabits. (2) Chamaelinorops utiliz- 
es leaf litter, while both active and sleeping, to raise its body 
temperature. Leaf litter is a natural thermal insulator, and its 
decomposition generates some heat. Chamaelinorops could uti- 
lize the warmth retained or generated by leaf litter to raise its 
body temperature without necessarily having to bask. Though 
plausible, this explanation seems unlikely, given that leaf litter 
probably cannot generate adequate heat for a lizard. (3) Cha- 
maelinorops may actually bask but was missed engaging in such 
activity during our study. This is always a distinct possibility. 



1994 ECOLOGY OF CHAMAELINOROPS BARBOURI 21 



especially given that some deep-forest lizards, such as Kentropyx 
calcaratus (Rand and Humphrey, 1968), follow sun flecks on the 
forest floor. However, given that our insolation data do not in- 
dicate even the weakest correlation between Chamaelinorops perch 
selection and sun patches (Fig. 1) and that we never observed 
individuals sunning, we believe that this explanation may also be 
unlikely. 

In summing up the paradoxical thermal biology of Chamae- 
linorops we can state only that resolution is not currently possible 
and must await more detailed study. 

ACKNOWLEDGMENTS 

We thank E. E. Williams for several useful discussions of Cha- 
maelinorops biology, information on Scutisorex, and for alerting 
us to the Bohme paper. J. B. Losos, G. C. Mayer, and E. E. 
Williams provided helpful comments on the manuscript. We also 
wish to especially thank F. X. Geraldes, S. J. Inchaustegui, and 
R. Rimoli of the Museo Nacional de Historia Natural, Santo 
Domingo, Dominican Republic, for help in obtaining permits, 
maps, and a vehicle, and for general assistance. A. Schwartz also 
provided needed assistance and his unparalleled expertise on West 
Indian herpetofauna while we were in the field. We wish to ac- 
knowledge the invaluable editorial suggestions on F. Boisse-Kilgo. 
The senior author wishes to thank A. A. Arian for assistance, 
typing, and moral support, C. D. Bain for logistical support, and 
H. E. Flores, H. Rosenberg, I. Flores, and E. E. Williams for 
financial support during the planning and writing of this study. 
The senior author is also grateful to C. H. Suh for assistance in 
the bibliography, German translation, and moral support. The 
third author (JP) would like to thank J. Palumbo Sr., L. D. Palum- 
bo, and C. Arpee for assistance and moral support. P. Alberch, 
J. L. Knight, J. P. Rosado, and F. D. Ross provided workspace, 
access to facilities, use of field equipment, and general invaluable 
assistance at the Museum of Comparative Zoology, Harvard Uni- 
versity. Field work was supported by two grants from the Youth 
Fund of The Explorers Club, a Grant-in-Aid of Research from 
Sigma Xi, and a grant from the Barbour Fund of the Museum of 
Comparative Zoology, Harvard University, all to the first author. 



22 BREVIORA No. 499 



LITERATURE CITED 

Allen, J. A. 1917. The skeletal characters of Scutisorex Thomas. Bulletin of 

the American Museum of Natural History, 37: 769-783. 
BoHME, W. 1982. Ein neues Chamaleon aus Tanzania, mit Bemerkungen uber 

Mimese bei Echsen (Reptilia: Sauria). Bonn Zoologische Beitrage, 33: 349- 

361. 
Brattstrom, B. H., and T. R. Howell. 1954. Notes on some collections of 

reptiles and amphibians from Nicaragua. Herpetologica, 10: 1 14-123. 
Bustard, H. R. 1967. Activity cycle and thermoregulation in the Australian 

gecko Gehyra variegata. Copeia, 1967: 753-758. 
Case, S. M., and E. E. Williams. 1987. The cybotoid anoles and Chamaeli- 

norops: Evidence of mosaic evolution. Zoological Journal of the Linnean 

Society, 91: 325-341. 
Etheridge, R. E. 1 960. The relationships of the anoles (Reptilia: Sauria: Iguani- 

dae): An interpretation based on skeletal morphology. University Microfilms. 

xiii + 236 pp. 
Fitch, H. S. 1968. Temperature and behavior of some equatorial lizards. Her- 
petologica, 24: 35-38. 
. 1973. A field study of Costa Rican lizards. University of Kansas Science 

Bulletin, 50: 39-126. 

1975. Sympatry and interrelationships in Costa Rican anoles. Occasional 



Papers of the Museum of Natural History, University of Kansas, 40: 1-60. 

Fitch, H. S., A. F. Echelle, and A. A. Echelle. 1976. Field observations on 
rare or little known mainland anoles. University of Kansas Science Bulletin, 
51: 91-128. 

FoRSGAARD, K. 1983. The axial skeleton of Chamaelinorops, pp. 284-295. In 
Anders G. J. Rhodin and Kenneth Miyata (eds.). Advances in Herpetology 
and Evolutionary Biology. Essays in Honor of Ernest E. Williams. Cambridge, 
Massachusetts, Museum of Comparative Zoology, Harvard University, xix 
+ 725 pp. 

Franz, R., and D. Cordier. 1986. Herpetofaunas of the national parks of Haiti. 
USAID/Haiti Publication, 1-73. 

Heatwole, H., T-H. Lin, E. ViLLALON, A. MuNiz, AND A. Matta. 1969. Some 
aspects of the thermal ecology of Puerto Rican anoline lizards. Journal of 
Herpetology, 3: 65-77. 

Hertz, P. E. 1974. Thermal passivity of the sympatric grass anoles (Anolis 
semilineatus and A. olssoni) in lowland Hispaniola (Reptilia, Lacertilia, Igua- 
nidae). Journal of Herpetology, 13: 329-333. 

. 1981. Adaptation to altitude in two West Indian anoles (Reptilia: Igua- 

nidae): Field thermal biology and physiological ecology. Journal of the Zoo- 
logical Society of London, 195: 25-37. 

Hertz, P. E., and R. B. Huey. 1981. Compensation for altitudinal changes in 
the thermal environment by some Anolis lizards on Hispaniola. Ecology, 62: 
515-521. 

Huey, R. B., and M. Slatkin. 1976. Cost and benefits of lizard thermoregu- 
lation. Quarterly Review of Biology. 51: 363-384. 



1994 ECOLOGY OF CHAMAELINOROPS BARBOURl 23 



HuEY, R. B.. AND T. p. Webster. 1976. Thermal biology of Anolis lizards in a 

complex fauna: The cristatelliis group on Puerto Rico. Ecology, 57: 985-994. 
Jenssen, T. a., and p. C. Feely. 1991. Social behavior of the male anoline 

lizard Chamaelinorops barbouri, with a comparison to Anolis. Journal of 

Herpetology, 25: 454-462. 
Lee, J. C. 1980. Comparative thermal ecology of two lizards. Oecologia, 44: 

171-176. 
Rand, A. S. 1964. Ecological distribution in anoline lizards of Puerto Rico. 

Ecology, 45: 745-752. 
Rand, A. S., and S. S. Humphrey. 1 968. Interspecific competition in the tropical 

rain forest: Ecological distribution among lizards at Belem, Para. Proceedings 

of the National Museum, 125: 1-17. 
Ruibal, R. 1961. Thermal relations of five species of tropical lizards. Evolution, 

15: 98-111. 
. 1964. An annotated checklist and key to the anoline lizards of Cuba. 

Bulletin of the Museum of Comparative Zoology, 130: 473-520. 
Schmidt, K. P. 1919. Descriptions of new amphibians and reptiles from Santa 

Domingo and Navassa. Bulletin of the American Museum of Natural History, 

41: 519-525. 
Schwartz, A., and S. J. Inchaustegui, 1980. The endemic Hispaniolan lizard 

genus Chamaelinorops. Journal of Herpetology, 14: 51-56. 
SoKAL, R. R., and F. J. RoHLF. 1981. Biometry. San Francisco, California, W. 

H. Freeman, xiii + 776 pp. 
Talbot, J. J. 1977. Habitat selection in two tropical anoline lizards. Herpeto- 

logica, 33: 114-123. 
Thomas, R. 1 966. A reassessment of the herpetofauna of Navassa Island. Journal 

of the Ohio Herpetological Society, 5: 73-89. 
Williams, E. E. 1977. The macrosystematics of the anoles, pp. 122-131. In 

Ernest E. Williams (ed.). The Third Anolis Newsletter. 131 pp. 
. 1983. Ecomorphs, faunas, island size, and diverse end points in island 

radiations of Anolis, pp. 326-370. In Raymond B. Huey, Eric R. Pianka, and 

Thomas W. Schoener (eds.). Lizard Ecology. Studies of a Model Organism. 

Cambridge, Massachusetts, Harvard University Press, vi + 501 pp. 
Wyles, J. S., AND G. C. Gorman. 1980. The classification of Anolis: Conflict 

between genetic and osteological interpretation as exemplified by Anolis cy- 

botes. Journal of Herpetology, 14: 149-153. 



B R E V i,ja,P A 

Museum of Compa^tjj^Ci Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 10 January 1995 ERSITNumber 500 

THE PENNSYLVANIAN TEMNOSPONDYL 

COCHLEOSAURUS FLORENSIS RIEPPEL, FROM THE 

LYCOPSID STUMP FAUNA AT 

FLORENCE, NOVA SCOTIA 



2,3 



Stephen J. Godfrey' and Robert Holmes 



Abstract. A previously undescribed skull of the Pennsylvanian cochleosaurid 
Cochleosaurusflorensis Rieppel, from Florence, Nova Scotia, Canada, contributes 
the following characters to the generic diagnosis: extreme elongation of both the 
vomers and choanae, exclusion of the ectopterygoids and maxillae from the sub- 
temporal fossae by a broad lateral ala of the pterygoid, and triangular tabular 
"horns." The transverse width of the skull through the midorbital region is less 
than the antorbital length. The configuration of the vomers and the general pro- 
portions of the palate resemble those of the Permian cochleosaurid Chenoprosopus 
milleh Mehl. The two genera are united in a monophyletic family— the Cochleo- 
sauridae— by the following autapomorphies: absence of the parietal foramen and 
lateral line sulci, subdued sculpturing on depressed areas of the dermal skull roof, 
an elongate antorbital region of the skull that is reflected in the palate by the 
greatly elongate vomers, ectopterygoid and maxilla excluded from the rim of the 
subtemporal fossa by a lateral ala of the pterygoid that makes contact with the 
jugal, and very long, triangular choanae. 



INTRODUCTION 

Members of the order Temnospondyli comprise the most nu- 
merous and diverse of all amphibian groups, enjoying a strati- 
graphic range extending from the Lower Carboniferous to the 
Lower Cretaceous. The superfamily Edopoidea has traditionally 



' Paleo-Skullpture Studio, Box 692, Rosedale. Alberta TOJ 2V0, Canada. 

^ Redpath Museum, McGill University, 859 Sherbrooke Street West, Montreal, 

Quebec H3A 2K6, Canada. 

^ To whom reprint requests should be addressed. 



2 BREVIOIL-i No. 500 

been identified as the stem group for the order (Carroll, 1988). 
Recent phylogenetic analyses (Milner, 1 990a; Trueb and Cloutier, 
1991) tend to confirm a relatively plesiomorphic status for those 
genera normally assigned to the group, but because their anatomy 
is incompletely known, this phylogenetic position, and even the 
integrity of the superfamily, is supported by few characters. Con- 
sequently, it is possible that the Edopoidea as presently consti- 
tuted represents a structural grade within primitive tetrapods, and 
not a monophyletic group (Trueb and Cloutier, 1991). 

In 1956, four bone-bearing lycopsid stumps of the genus Sig- 
illaria were excavated from the wall of an abandoned strip coal 
mine about 2 km north of Florence, Cape Breton County, Nova 
Scotia, Canada, by a field party from the Museum of Comparative 
Zoology, Harvard (MCZ). These stumps, all from a single horizon, 
were rooted about 4 m above the Lloyd Cove (Lower Bonar) Coal 
Seam of the Morien Group (Late Pennsylvanian), which probably 
corresponds in age to the Westphalian D of Europe (Carroll et 
al, 1972). Reptilian remains from the stumps include the early 
protorothyridid Palaeothyris acadiana (Carroll, 1969) and the 
ophiacodontid synapsids Archaeothyris florensis and Echinerpe- 
ton intermedium (Reisz, 1972). Non-amniote tetrapods are rep- 
resented by the limnoscelid Limnostygis relictus (Carroll, 1967a), 
the embolomerous anthracosaur Carbonoherpeton carrolli (Klem- 
bara, 1985), and the primitive temnospondyl Cochleosaurus flo- 
rensis (Rieppd, 1980). 

On the basis of two crushed but reasonably complete skulls of 
Cochleosaurus florensis (MCZ 4343 and 4344), Rieppel (1980) 
noted several similarities to Chenoprosopus milleri from the Low- 
er Permian of New Mexico (Langston, 1953) and endorsed its 
assignment to the Edopoidea. Imperfect preservation, however, 
precluded detailed comparison of many anatomical features. Iso- 
lated postcranial remains subsequently attributed to C florensis 
(Klembara, 1985) did little to clarify its taxonomic position. A 
third, more complete skull (MCZ 4342) from the same locality 
permits a more precise diagnosis of C florensis, expands our 
knowledge of the family Cochleosauridae, and will contribute to 
a better understanding of the interrelationships of primitive tem- 
nospondyls. 



1 994 COCHLEOSA UR US FLORENSIS FROM NOVA ^CJ^J/^p^ p Y^ 

SYSTEMATIC PALEONTOLOGY .gg^ 

Order Temnospondyli Zittel, 188/ 

Superfamily Edopoidea Romer, 1945^ i->\ / a i-»p^ 

\KVAKD 
Family Cochleosauridae Broili (in Zittel), I'^^RSITY 

Diagnosis. Temnospondyl amphibians with a maximum known 
snout-postparietal length of about 290 mm (Chenoprosopus). Pa- 
rietal foramen and lateral line sulci absent. Extremely large pre- 
maxillae and vomers resulting in elongate antorbital region of the 
skull that exceeds skull width in midorbital region. Both external 
nares and elongate, anteriorly broadened choanae positioned well 
posterior to the tip of the snout. Ectopterygoid and maxilla ex- 
cluded from the subtemporal fossa by a lateral ala of the pterygoid 
that forms a pterygoid-jugal contact. 

Cochleosaurus Frixsch, 1885 

Diagnosis. Temnospondyl amphibian with a maximum known 
skull length of 124 mm (snout-postparietal lappets). Sculpturing 
muted on depression on anterior part of lacrimal and on a median, 
depressed strip on the skull roof extending from the posterior 
margin of the postparietal to a shallow expanded depression on 
the snout. Lacrimal bears an anteroventrally directed ridge on its 
posterior half. Sculptured postparietal lappets. The choanae form 
a right-angled triangle with an anteriorly positioned base. 

Cochleosaurus florensis Rieppel, 1980 

Horizon and Locality. Morien Group, Westphalian D, Domin- 
ion Coal Co., strip mine no. 7 (abandoned), about 2 km north of 
Florence, Cape Breton County, Nova Scotia, Canada. 

Diagnosis. Pterygoid with a broad ala that forms an extensive 
contact with the jugal to exclude the wedge-shaped posterior mar- 
gin of the ectopterygoid and maxilla from the margin of the sub- 
temporal fenestra. Shorter postparietal lappets and relatively wid- 
er skull table than C. bohemicus. Triangular tabular "horns" 
variably developed. At comparable skull lengths, dermal sculp- 
turing in C. florensis comprises predominantly rounded pits rather 



BREVIORA 



No. 500 




Table 1 . Cranial dimensions of the skulls of Cochleosaurus florensis {in 

millimeters). 







MCZ 


MCZ 


MCZ 






4342 


4343 


4344 


A. 


Skull length (snout-tabular) 


109 


124 


87* 


B. 


Snout-quadrate length 


108 


123* 


84* 


C. 


Antorbital length 


58 


62 


49 


D. 


Skull table length 


37 


44 


28 


E. 


Orbital length 


14* 


18 


12 


F. 


Interorbital width 


21* 


27* 


-t 


G. 


Skull table width 


47* 


49* 


30 


H. 


Interpterygoid vacuity length 


27 


— 


— 


I. 


Interpterygoid vacuity width 


29* 


31 


— 


J. 


Snout-interpterygoid vacuity length 


50 


— 


— 


K. 


Snout-vomerine fang distance 


29 


26* 


— 



* Measurement estimated. 

t — = Dimension not preserved or exposed. 



1994 COCHLEOSAURUS FLORENSIS FROM NOVA SCOTIA 5 

than elongate grooves like those of C bohemicus, indicating a 
smaller maximum (adult) size. 

SPECIMENS EXAMINED 

The following, comprising all known material attributed to C 
florensis, were examined. 

MCZ 4342: The most complete and least distorted skull of C. 
florensis. Both dorsal and ventral surfaces are almost completely 
free of matrix. This skull is slightly smaller than that of the ho- 
lotype (Table 1). 

MCZ 4343 (holotype): A severely crushed but articulated, large- 
ly complete skull; associated girdle and limb elements include a 
clavicle, a femur, and a tibia (Rieppel, 1980, text-fig. 1). 

MCZ 4344: A small, dorsoventrally compressed skull originally 
prepared to expose the palate (Rieppel, 1 980, text-fig. 3), but since 
then embedded in Carbowax (polyethylene glycol) and prepared 
to expose the dorsal aspect. 

MCZ 2773: Isolated postcranial remains (Klembara, 1985, 
text-figs. 8, 9). 

ABBREVIATIONS 

a. cor— anterior coronoid pm — premaxilla 

ang— angular po— postorbital 

art— articular po. cor— posterior coronoid 

c. pr— cultriform process pof— postfrontal 

d— dentary posp— postsplenial 

ect— ectopterygoid pp— postparietal 

f— frontal prf— prefrontal 

it— intertemporal ps— parasymphyseal tusk 

j— jugal pt— pterygoid 

1— lacrimal q— quadrate 

m— maxilla qj— quadratojugal 

m. cor— middle coronoid sa— surangular 

Mech — Mechelian fenestra sm — septomaxilla 

n — nasal sp— splenial 

p— parietal sq — squamosal 

pa— prearticular st— supratemporal 

pal — palatine t— tabular 

para— parasphenoid v— vomer 



6 BREVIORA No. 500 

DESCRIPTION 
Skull Roof 

The skull of Cochleosaurus florensis (Fig. 1) resembles both 
Cochleosaurus bohemicus (Steen, 1938) and Chenopwsopus mil- 
leri (Langston, 1953) in having crocodiloid proportions with a 
broad, spatulate snout and flared cheeks, although the snout is 
somewhat broader than that of Chenopwsopus. Other propor- 
tional similarities include an antorbital region that exceeds its 
transverse width at the level of the orbits (the reverse of the 
condition seen in other Paleozoic temnospondyls such as Caero- 
rhachis Holmes and Carroll, 1977; Dendrerpeton Carroll, 1967b, 
and Godfrey et al, 1987; Greererpeton Smithson, 1982; Nelda- 
5awrw5 Chase, 1 965; and £'<^o/75Romer and Witter, 1942, although 
shared by the eryopoid Archegosaurus Milner, 1978) and small 
dorsolaterally facing external nares located well back from the 
anterior margin of the snout. The orbits are variable in size (Table 
1). The suspensorium projects only slightly posterior to the back 
edge of the skull table, resulting in a relatively anterior placement 
of the quadrate and shallower squamosal embayment as com- 
pared to the much larger Chenopwsopus. Similar intra- and in- 
terspecific, size-dependent variations, however, have been ob- 
served in other labyrinthodonts (Romer, 1939), rendering the 
taxonomic significance of such proportional differences to the 
Cochleosauridae ambiguous. 

The parietal foramen and lateral line sulci are absent. Dermal 
ornamentation, as in other temnospondyls, consists primarily of 
course, rounded pits. However, unlike the type (MCZ 4343), some 
elongate grooves indicative of zones of rapid growth in immature 
individuals (Bystrow, 1935) are evident on some bones of both 
MCZ 4342 and 4344, especially the jugal and quadratojugal, al- 
though these are not as well developed as in C. bohemicus. The 
skull roof bears two conspicuous, broadly rounded, and coarsely 
sculptured longitudinal ridges, each passing anteriorly from the 
tabular along the margin of the skull table to the posterior rim of 
the orbit. Each wraps around the dorsal and anterior orbital bor- 
ders, crossing the pre- and postfrontals as well as the lateral margin 
of the frontal, and then continues anteriorly along the lateral 
margin of the nasal, finally subsiding on the dorsolateral surface 



1 994 COCHLEOSA UR US FLORE NS IS FROM NOVA SCOTIA 




Figure 1 . Cochleosaurusflorensis Rieppel. Reconstniction of skull based large- 
ly on MCZ 4342 in A) dorsal view and B) ventral view. 



of the premaxilla. Similar ridges are seen in the eryopoid tem- 
nospondyls Zatrachys (Langston, 1953), but unlike in the latter 
a deep longitudinal trough, nearly devoid of sculpturing, runs up 
the middle of the central depression, gradually becoming broader 
and more shallow as it passes anteriorly between the orbits onto 
the snout, where it terminates as an expansive, shallow depression 
between and anterior to the external nares. The lacrimal bears a 
low but distinct rounded ridge that passes anteroventrally across 
its surface. Anteromedial to the ridge, its concave surface bears 
only muted ornamentation, as in Chenoprosopus but in contrast 
to C bohemicus, where no difference in texture occurs (personal 
observation). Chenoprosopus shows similar, sparsely ornamented 
depressions on the snout, but apparently not in the postorbital 
region (Langston, 1953). 

The well-developed nasals equal the frontals in midsagittal 
length, but as in Chenoprosopus, the unusually long antorbital 
region is primarily formed by the remarkably large premaxillae, 



BREVIORA 



No. 500 



posp 



Mech 




1994 COCHLEOSAVRUS FLORE NS IS FROM NOVA SCOTIA 9 

which measure about 65% of the nasal length. Posterolaterally 
oriented nasopremaxillary sutures position the external nares well 
back from the tip of the snout. A less pronounced, but otherwise 
similar structure occurs in Edops. Large premaxillae and poste- 
riorly positioned external nares also occur in the eryopoid Za- 
trachys, but the snout morphology is otherwise very different, and 
further comparisons are of doubtful value. The morphology of 
Cochleosaurus contrasts to that of long-snouted embolomeres such 
as Archeria (Holmes, 1989) in which the premaxillae are small 
and external nares are located anteriorly. Although the premax- 
illae are somewhat larger in Archegosaurus (Milner, 1978), it is 
primarily elongation of the frontals and in particular the nasals 
that account for the snout elongation and, unlike Cochleosaurus, 
the anterior margins of the elongate external nares are close to 
the anterior end of the snout. The posterior 6 of the approximately 
20 premaxillary teeth gradually increase in size to form a pseu- 
docanine peak just anterior to the premaxillary-maxillary suture. 
Large premaxillary teeth also occur in colosteids (Smathson, 1 982) 
but comprise a tusk pair distinctly larger than the other teeth in 
a much smaller premaxilla. There is no marked size variation in 
the approximately 35 maxillary teeth except posteriorly, where 
they gradually become smaller. 

The septomaxilla forms a posteriorly directed, apparently unor- 
namented wedge on the dorsal surface of the snout. It nearly (right 
side of MCZ 4342) or completely (left side of MCZ 4342 and 
4344) excludes the lacrimal from the margin of the external naris. 
In contrast to the condition in Chenoprosopus, a nasomaxillary 
contact is absent. 

Variation in size and shape of some dermal elements occurs. 
The left postfrontal of MCZ 4342 is much wider than the right 
(Fig. 2). The postorbitals, although elongate and triangular in 
outline in most specimens (e.g., MCZ 4343 [Rieppel, 1980], MCZ 
4344 [Fig. 3]), are significantly narrower posteriorly in MCZ 4342 
(Fig. 2). The significance of this variation is unknown. 



Figure 2. Cochleosaurus florensis Rieppel. Specimen drawing of MCZ 4342 
in dorsal view, showing right lower jaw in medial view. 



10 



BREVIORA 



No. 500 




Figure 3. Cochleosaurus florensis Rieppel. Specimen drawing of MCZ 4344 
in dorsal view, showing right lower jaw in ventral view. 



The skull table in MCZ 4342, as in the type, is relatively wider 
than in C bohemicus. A separate intertemporal is clearly present 
in all specimens. As in Chenoprosopus, the anterodorsal notch of 
the squamosal embayment reaches far forward along the edge 
of the table to the midpoint of the supratemporal. The latter bone 
thus forms a significant portion of the dorsal border of the em- 
bayment. This is distinctly different from the condition in other 
temnospondyls possessing embayments in which a posterodorsal 
process of the squamosal forms much of its dorsal border, severely 
limiting participation of the supratemporal or completely ex- 
cluding it from the rim. The posterolateral comer of the right 
tabular in MCZ 4342 extends to form a blunt, triangular tabular 
"horn" that projects posteriorly and slightly ventrally. In contrast 
to the tabular horn of embolomeres (Panchen, 1970), it is not 
biramous and bears ornamentation on both dorsal and ventral 
surfaces. Prominent tabular projections, rare in temnospondyls, 
also occur in Zatrachys\ however, they differ from those in Cochle- 
osaurus in being posterodorsally curved, attenuated spikes that 
probably developed in association with the unique spiny orna- 
mentation festooning the skull. Homologies with the tabular pro- 



1 994 COCHLEOSA VR US FLORENSIS FROM NOVA SCOTIA 1 1 

jections of other labyrinthodonts are doubtful. The "horn" is not 
as well developed in either MCZ 4343 (Rieppel, 1 980), MCZ 4344 
(Fig. 3), or any specimens of C. bohemiciis. Such structures are 
often subject to positive allometry, but the horns are not partic- 
ularly prominent in the largest specimen (MCZ 4343). The size 
of these horns is quite variable in C. bohemiciis (Sequeira, per- 
sonal communication), and this may also be the case in C. flo- 
rensis. The postparietal lappets, although well developed in both 
specimens, are noticeably smaller than in equivalently sized C. 
bohemicus. They project straight posteriorly with their dorsal sur- 
faces contiguous with that of the table rather than occupying a 
more ventral position on the occipital surface of the postparietal 
as in Caerorhachis or Dendrerpeton. Their ornamented dorsal 
surfaces are considered to be diagnostic for the genus (Sequeira 
andMilner, 1993). 

Palate 

The excellently preserved palate in MCZ 4342 (Fig. 4) bears 
denticles over much of the lateral portions of the vomers, the 
entire palatines, and ectopterygoids and most of the pterygoids 
except the posterodorsal part of the quadrate ramus. The snout 
is distinctly longer than described by Rieppel (1980), who based 
his reconstructions on the less well preserved palates of MCZ 
4343 and 4344. Other than being broader, it is proportioned like 
that of Chenoprosopus. This elongation is produced by massive 
vomers, which account for about 45% of the snout-quadrate length 
in contrast to embolomeres (Panchen, 1970), other temnospon- 
dyls such as Caerorhachis, Dendrerpeton (Godfrey et a!., 1987), 
Greererpeton, Edops, and even the long-snouted Archegosaurus 
(Whittard, 1928), in which the vomers are relatively much short- 
er. Among temnospondyls, only Neldasauriis and the aberrant 
Zatrachys have vomers approaching these proportions. The tri- 
angular choanae, with their bases positioned anteriorly, occupy 
positions well posterior to the tip of the snout. Each vomer bears 
a small tusk and replacement pit at the apex of a pronounced 
triangular thickening located anteromedial to the choana. As in 
Chenoprosopus, a broad depression occupies the central portion 
of the plate between the anteriorly diverging medial margins of 
the thickenings. 



12 



BREVIOR.4 



No. 500 



1 CM 




1994 COCHLEOSA UR US FLORE NS IS FROM NOVA SCOTIA 1 3 

Table 2. Proportions of the palate in some early temnospondyls: quad- 
rate-anterior MARGIN of the INTERPTERYGOID VACUITIES/SNOUT- ANTERIOR 



Dendrerpeton acadianum {Godfrey et al., 1987) 0.34 

Greererpelon burkemorani (Smhhson. 1982) 0.44 

Edops craigi (Romer and Witter, 1942) 0.53 

Coch leosaurus bohemicus (Sieen, 1938) 0.59 

Cochleosaurus florensis (Rieppd, 1980) 0.78 

Chenoprosopus milleri (LangsXon, 1953) 0.84 

Cochleosaurus florensis (this paper) 0.96 

Caerorhachis bairdi {HoXmes and. CarroW, 1977) 0.61 

Neldasaurus wrightae (Chase. 1965) 0.65 

* 1 .00 would indicate that the anterior margin of the interpterygoid vacuities lay 
equidistant between the snout and quadrate. These ratios are based on the res- 
torations contained in the original publications. 



The anterior borders of the modest interpterygoid vacuities are 
equidistant between the tip of the snout and quadrate condyles. 
In other early tetrapods, this margin is more anteriorly placed 
(Table 2). 

The sutural outline of the palatine is difficult to follow, but its 
lateral extent is indicated by the presence of a palatine tusk and 
replacement pit clearly visible on the left side of MCZ 4342 im- 
mediately medial to the maxillary tooth row and lateral to the 
posterior corner of the choana. 

In MCZ 4342, the left quadratojugal and posterior portion of 
the jugal have folded under the skull, exposing the lateral portion 
of the attached ectopterygoid and posterior portion of the maxilla. 
(Fig. 4). The concave dorsal surface of the displaced left ectoptery- 
goid faces laterally, and the tusk pair— the anterior one complete, 
the posterior one missing the tip and overlaid by two small max- 
illary teeth— project medially. The region of the right ectoptery- 
goid, where dentition might be expected, is obscured by an un- 
identified bone, possibly a hyoid element or limb bone. 



Figure 4. Cochleosaurus florensis Rieppel. Specimen drawing of MCZ 4342 
in palatal view, showing left lower jaw in ventral view. 



14 BREVIORA No. 500 

Although the narrow palatal rami of the pterygoids almost cer- 
tainly do not meet anteriorly, they contact the anterior end of the 
cultriform process, thus excluding the vomers from the rim of 
the moderate-sized interpterygoid vacuity. This morphology is 
here considered to be intermediate between the primitive tem- 
nospondyl condition in which the vacuities are completely bor- 
dered by pterygoids that meet anteriorly (Caerorhachis, Greer- 
erpeton, and Edops) and the more derived temnospondyl condition 
in which the vomers enter the margins of the vacuities (Milner, 
1990a). This inferred evolutionary sequence is apparently cor- 
related with an increase in the size of the vacuities. More pos- 
teriorly, the pterygoid projects laterally into the anterior portion 
of the opening of the subtemporal fossa, but no ventral deflection 
like that reported in Chenoprosopus is apparent. 

A unique feature of the palate is the configuration of the pter- 
ygoid-ectopterygoid suture. In MCZ 4342, each pterygoid wraps 
around the roughly triangular posterior end of the ectopterygoid 
and extends a broad ala anterolaterally to form a relatively ex- 
tensive contact with the inner surface of the jugal, thereby ex- 
cluding the ectopterygoids and maxilla from the rim of the sub- 
temporal fossa. In Chenoprosopus, a more modest lateral projection 
of the pterygoid forms a limited contact with the jugal. A point 
contact appears to occur in Caerorhachis, but indistinct sutures 
make this equivocal (Holmes and Carroll, 1977). A pterygoid- 
jugal contact also occurs in anthracosaurs (Panchen, 1970) and 
the saurerpetontid temnospondyl Acroploiis (Foreman, 1990) but 
is accomplished by a medial process of the jugal (alary process) 
rather than by a lateral ala of the pterygoid. A pterygoid-jugal 
suture does not occur in other primitive amphibians such as 
Greererpeton, Dendrerpeton, Edops, loxommatids (Beaumont, 
1977), and Ichthyostega (Jarvik, 1980). 

In MCZ 4343, denticles extend beyond the basicranial articu- 
lation onto the quadrate ramus of the pterygoid (contra Rieppel, 
1980, text-figs. 1, 2). The quadratojugal wraps around the pos- 
terolateral corner of the quadrate and appears to have made a 
modest contribution to the lateral portion of the quadrate condyle, 
although crushing makes it impossible to be certain. 



1 994 COCHLEOSA UR US FLORENSIS FROM NOVA SCOTIA 1 5 

Braincase 

The braincase is much more completely preserved in MCZ 
4342 (Fig. 4) than in either MCZ 4343 or 4344. The stout basipter- 
ygoid process inserts into a simple rectangular depression on the 
posteromesial margin of the remarkably stout basal process (Fig. 
4, right side). Unlike Greererpeton, no evidence indicates a basal 
socket. The posterior lip of the cup-shaped distal end of the basi- 
pterygoid process wraps around the posterior margin of the basal 
process. 

Between the basipterygoid processes is a conspicuously raised, 
triangular tubercle bearing a patch of denticles. The shape of this 
tubercle, consistent in both MCZ 4342 and 4343, distinguishes 
C florensis from Greererpeton and Caerorhachis, where the tu- 
bercle is oval in outline. Lateral to the tubercle, an anteromedially 
directed groove, probably marking the course of the internal ca- 
rotid and palatine arteries and nerves, crosses the base of each 
basipterygoid process. 

The basal plate of the parasphenoid bears two V-shaped de- 
pressions, the "tubera basisphenoidales" (Romer, 1930; Smith- 
son, 1982), the lateral margins of which are bordered by prom- 
inent crests. The smooth periosteal floor of each pocket and the 
low median ridge that separates them extend to the posterior 
margin of the parasphenoid. 

The cultriform process in MCZ 4342 becomes progressively 
broader anterior to the midpoint of the interpterygoid vacuities 
(contra Rieppel, 1980). In many early temnospondyls such as 
Caerorhachis, Greererpeton, and Edops, the process remains ap- 
proximately parallel-sided or may even narrow anteriorly. How- 
ever, an anteriorly expanded process is present in a range of 
temnospondyls such as Dendrerpeton (Godfrey et al, 1987), Nel- 
dasaurus, and Zatrachys, making the significance of this feature 
unclear. Anteriorly, the process forms a broad wedge between the 
pterygoids and appears to contact the vomers, but lack of well- 
defined sutures makes this difficult to confirm (Figs. IB, 4). 

A well-ossified sphenethmoid is exposed through the left in- 
terpterygoid vacuity of MCZ 4342 (Fig. 4). During dorsoventral 
compression of the skull, it was rotated to expose its featureless 



16 BREVIORA No. 500 

left lateral surface. It extends anteriorly almost to the front of the 
interpterygoid vacuity. Its rear margin is obscured by the basal 
process. 

The otico-occipital portion of the braincase of MCZ 4342 has 
been crushed against the ventral surface of the skull, obscuring 
its morphology. 

Lower Jaw 

The mandible, which bears no lateral line sulci, is particularly 
slender anteriorly (Fig. 5). The right half, exposed in a medial 
view (Fig. 4), is crushed, but complete. A parasymphyseal tusk 
is present. There is no canine peak, although the teeth in the 
anterior half of the tooth row are somewhat larger. Each dentary 
could have held at least 55 labyrinthine-infolded teeth. 

Three denticle-covered coronoids form most of the dorsome- 
dial surface of the jaw anterior to the adductor fossa. MCZ 4342 
preserves what appears to be a coronoid-surangular suture on the 
medial surface of the outer wall of the adductor fossa. A coronoid 
contribution to the surangular crest, present in some derived tem- 
nospondyls like Tersomius (Carroll, 1964) and Phonerpeton 
(Dilkes, 1990), would represent a derived condition relative to 
primitive tetrapods such as Caerorhachis, Greererpeton, and Edops 
in which the posterior coronoid does not contribute to the sur- 
angular crest, but its level of apomorphy remains uncertain. In 
MCZ 4342, however, the usually straight course of the proposed 
"suture" suggests that it may be a break, and a distinct change in 
texture of this element at the anterior end of the adductor fossa 
provides an alternate position for the coronoid-surangular suture. 
Medially, the posterior coronoid forms the anterodorsal rim of 
the adductor fossa and then extends to a rounded termination 
anteriorly. The long, narrow anterior coronoid can be distin- 
guished from the dentary dorsally and presplenial ventrally, but 
the sutures become obscured toward the symphysis. The limits 
of the middle coronoid are problematic. Although it is easily 
distinguished from the dentary dorsally and prearticular ventrally, 
its anterior suture is obscured by broken bone surface. Posteriorly, 
it appears to pass ventral to the posterior coronoid to form part 
of the ventrolateral margin of the adductor fossa (Fig. 4), a unique 



1994 



COCHLEOSAURUS FLORENSIS FROM NOVA SCOTIA 



17 




Figure 5. Cochleosaurus florensis Rieppel. Reconstruction of lower jaw in 
medial view, based primarily on MCZ 4342 with additional information from 
MCZ 4344. 



condition for early tetrapods. However, the bone surface of this 
apparent posteroventral extension bears a much closer resem- 
blance to that of the adjacent prearticular, suggesting that the 
latter has been broken into two longitudinal splints. A conser- 
vative reconstruction of the coronoids is presented here (Fig. 5), 
although the alternative interpretations already discussed cannot 
be dismissed unequivocally. 

A broadly rounded ridge, visible on the dorsal portion of the 
medial surface of the surangular, extends from its suture with the 
posterior coronoid to the mandibular condyle. 

There is a relatively large Meckelian fenestra at the common 
junction of the prearticular, postsplenial, and angular. Crushing 
makes it impossible to confirm the presence of more anterior 
fenestrae. 

The right ramus of MCZ 4344, preserved in ventral view, allows 
the mutual relationships of the angular, articular, surangular, and 
splenials to be determined (Fig. 3). The large, coarsely sculptured 
angular, occupying the posterior one-third of the ramus, forms a 
suture posteriorly and dorsally with the surangular. The latter is 
not exposed on the medial surface of the ramus, which is formed 
in this region by a lamina of the articular (Fig. 3). A reexamination 
of MCZ 4343 confirms this (contra Rieppel, 1980, text-figs. 1, 2). 
A 1-cm section is missing from the middle of the ramus of MCZ 
4344, which otherwise is undisturbed. The dentary forms a broad 
contact posteriorly with a large postsplenial (splenial) that has an 
extensive, coarsely sculptured lateral exposure. A much smaller, 
splint-like splenial (presplenial) with only ventral and medial ex- 
posure and muted sculpturing occupies a position between these 



18 BREVIORA No. 500 

two elements. Reexamination of MCZ 4343 indicates that the 
"angular" as described by Rieppel (1980) is actually the angular 
and postsplenial separated by a cryptic suture. Consequently, the 
suture pattern in this area conforms to the primitive temnospon- 
dyl pattern (e.g., Dendrerpeton, Greererpeton, Edops, Neldasau- 
rus), and an angular-splenial (presplenial) contact does not occur. 
This putative character was thought to be shared with Cheno- 
prosopus (Rieppel, 1980). Langston (1953:369) clearly indicated 
that the ". . . intersplenial sutures are not clear . . . ," raising the 
possibility that his "angular" (Langston, 1953, text-fig. 6) is, like 
the angular as described by Rieppel, a compound element. If this 
is the case, then Chenoprosopus would also conform to the prim- 
itive pattern. 

Denticles are absent on the prearticular. The articular lacks a 
retroarticular process. 

DISCUSSION 

The information provided by MCZ 4342 and 4344 allows a 
more precise diagnosis of Cochleosaurus florensis and the family 
Cochleosauridae. An initial attempt is made to establish the po- 
larity of characters discussed below by accepting the reasonable 
hypothesis (Milner, 1990b) that the primitive character states for 
the Temnospondyli are exhibited by dendrerpetontids, edopsids, 
and trimerorhachoids. Dendrerpeton, Edops, and Neldasaurus are 
used for this purpose. Where data are missing or inconsistent, 
interpretation is equivocal, or relationships above the family level 
are discussed, the more distantly related Caerorhachis, Greerer- 
peton, loxommatids, embolomeres, and Ichthyostega are used as 
outgroups. 

Rieppel (1980) used the following characters to distinguish 
Cochleosaurus florensis from C bohemicus: 1) smaller adult size, 
2) shorter postparietal lappets, 3) wider skull table, 4) smaller 
orbits, and 5) shorter snout. To support the character of smaller 
size, he argued that in contrast with C. bohemicus the uniformly 
rounded pitting on the skull roof of the type of C florensis in- 
dicated that maximum adult size had been reached (Bystrow, 
1935). Although some elements of MCZ 4342 and 4344 bear 
elongate pits toward their edges, indicating that rapid growth was 



1 994 COCHLEOSA UR US FLORE NS IS FROM NOVA SCOTIA 1 9 

Still occurring at the time of death, they are far less developed 
than in larger specimens of C. bohemicus and so tend to support 
his original hypothesis, albeit not as strongly. Both MCZ 4342 
and 4344 confirm the presence of smaller postparietal lappets in 
C.florensis. The skull table in MCZ 4342, as in the type, is wider 
than that of C. bohemicus, and this condition probably represents 
a valid distinction. The much smaller size of MCZ 4344 makes 
the reliability of such comparisons dubious. The status of the 
remaining characters is equivocal. The orbits of MCZ 4342 are 
distinctly smaller than an equivalent-sized C. bohemicus, but a 
reexamination of the type reveals an orbital diameter of about 
1 8 mm after correcting for crushing (contra Rieppel, 1 980), within 
the range expected for C. bohemicus. The well-preserved vomers 
of MCZ 4342 indicate that the snout of C.florensis is longer than 
estimated on the basis of the distorted type, exhibiting proportions 
similar to those of C. bohemicus. The unusually well-developed 
lateral ala of the pterygoid that wraps around the V-shaped pos- 
terior extremity of the ectopterygoid to form an extensive contact 
with the jugal may constitute an additional diagnostic feature, 
but the detailed morphology of this region has not been described 
in C. bohemicus. Although it appears probable that the Florence 
material represents a distinct species, more detailed comparisons 
await publication of a description of C. bohemicus presently being 
prepared by Sandra Sequeira at Birkbeck College, London. 

The Cochleosauridae 

Four genera are presently included in the family Cochleosaur- 
idae: Cochleosaurus, Chenoprosopus, and the poorly known 
Gaudrya and Macrerpeton (Carroll, 1977, 1988). The type of 
Gaudrya (Fritsch, 1885), comprising the anterior end of a snout, 
has been synonymized with Cochleosaurus bohemicus (Sequeira 
and Milner, 1993), making Gaudrya its junior synonym. Two 
specimens from Linton, Ohio (AMNH 2933 and 6954), originally 
assigned to Leptophractus by Romer (1930) but later transferred 
to Gaudrya on the basis of close similarities in palatal structure 
to the type (D. Baird, personal communication) should therefore 
also be reassigned to Cochleosaurus. Other specimens referred to 
Gaudrya (Romer, 1947) are not cochleosaurids (A. R. Milner, 



20 BREVIORA No. 500 

personal communication). The poorly known Macrerpeton, pres- 
ently being studied by Robert Hook, appears to be a derived 
cochleosaurid, but until a thorough review of its anatomy is com- 
pleted more specific relationships cannot be established. 

Members of the family Cochleosauridae share the following 
derived features: 

1 . Absence of parietal foramen and lateral line sulci. 

2. Elongate, triangular choanae wider anteriorly than posteriorly. 

3. Squamosal lacking posterodorsal process, leaving much of the 
lateral edge of the supratemporal exposed along the antero- 
dorsal margin to the squamosal embayment. 

4. Lateral ala of the pterygoid contacts the jugal to exclude the 
ectopterygoid and maxilla from the rim of the subtemporal 
fossa. 

In addition, all cochleosaurids share at least two features that, 
although probably homologous, are problematic: 

5. Large premaxillae with posterolaterally directed nasal sutures, 
resulting in posteriorly positioned external nares. Although not 
developed to the same degree, this snout configuration also 
occurs in Edops, and may diagnose the more inclusive super- 
family Edopoidea (see later). 

6. Extreme elongation of the premaxillae and vomers, producing 
an elongate preorbital region and posterior position of both 
external and internal nares. Enlarged vomers also occur in 
Archegosaurus and Zatrachys. In the former, however, the 
anterior borders of the choanae are much closer to the front 
of the snout, and there is no comparable development of the 
premaxillae. In the latter, the circular choanae are posterior 
in position, but the highly derived palatal structure of this 
aberrant eryopoid makes further comparisons difficult. 

7. System of prominent, rounded ridges on the skull roof sepa- 
rating depressed areas exhibiting muted sculpturing. Although 
this striking set of features allows one to immediately distin- 
guish cochleosaurs from other Carboniferous tetrapods, it is 
not unique to the family. The distantly related Permian ery- 
opoid Zatrachys bears a similar system of ridges and depres- 
sions (Langston, 1953). However, direct comparison is difficult 



1 994 COCHLEOSA UR US FLORE NS IS FROM NOVA SCOTIA 2 1 

because in Zatrachys a large fenestra occupies the region of 
the snout bearing the bowl-shaped depression in cochleosaurs, 
and in contrast to cochleosaurs the other depressions on the 
dermatocranium appear to bear well-developed ornamenta- 
tion. Although this is probably a convergent feature, its oc- 
currence in at least one other temnospondyl family renders it 
unreliable for establishing relationships. 

Differences between Cochleosaurus and Chenoprosopus 

Some of the differences between Cochleosaurus and Chenopro- 
sopus are essentially proportional and arguably correlated with 
the smaller size of the former (with a skull of one-half to one- 
third the length of the latter). Although they serve to distinguish 
the known specimens of these two taxa, their status as diagnostic 
characters is uncertain. These include the following: 

1 . A longer snout in Chenoprosopus relative to that of Cochleo- 
saurus, with a ratio of antorbital length (measured from the 
midpoint of the orbit) to postorbital length (measured to the 
posterior edge of the postparietal lappet) of 2.0 in the former 
and 1.5 in the latter. 

2. A more narrowly parabolic skull outline in Chenoprosopus 
with a maximum skull length-to-width ratio of 1.9 as com- 
pared to 1.6 in Cochleosaurus. 

3. A relatively longer, more posteriorly projecting suspensorium 
in Chenoprosopus. 

4. Contact between the septomaxilla and lacrimal reduced or 
absent in Chenoprosopus, resulting in a nasomaxillary contact. 
This also occurs in the long-snouted Archegosaurus (Milner, 
1978) and the trimerorhachoid Neldasaurus. 

Other differences, not obviously size-related, are considered as 
derived features of Chenoprosopus. These include the following: 

5. Prominent, denticle-bearing ridges present on the vomers, 
pterygoids, palatines, and ectopterygoids. 

6. Basicranial articulation apparently sutured and immobile in 
adults. This also occurs in most eryopoid temnospondyls but 
it not considered characteristic of edopoids (Carroll, 1988). 



22 BREVIORA No. 500 

7. Ventral surface of the cultriform process bearing a string of 
denticles. 

8. Vomerine pits (Sequeira and Milner, 1993). 

The Edopoidea 

The generally plesiomorphic status of the Cochleosauridae has 
prompted most authors to place the family near the base of tem- 
nospondyl phylogeny, traditionally comprising, with the Edopi- 
dae, the superfamily Edopoidea (e.g., Carroll, 1988). However, 
no undisputed synapomorphies have been identified. This has 
resulted in different interpretations of relationship between the 
Edopidae and Cochleosauridae, including sister-group relation- 
ship within a monophyletic Edopoidea (Milner, 1990a), structural 
grade (Trueb and Cloutier, 1991), and distant relationship, with 
each family included within distinct groups of temnospondyls 
(Boy, 1990). 

A reassessment of the Edopoidea is beyond the scope of this 
paper, but a few comments are appropriate. Boy (1990), in a 
phylogenetic analysis of European Lower Permian temnospon- 
dyls, argued against a close relationship between Edops and Che- 
noprosopus, instead hypothesizing a sister-group relationship be- 
tween the latter and the clade Archegosaurus + Sclewcephalus 
based on the following characters: 1) nasomaxillary suture, 2) 
elongate, anteriorly constricted prefrontal, and 3) pterygoid (?read 
parasphenoid)-vomer contact. However, all three characters are 
correlated with the elongation and narrowing of the snout. A 
nasomaxillary suture also occurs in the relatively long-snouted 
Neldasaurus. Although not an inevitable correlate of snout elon- 
gation, it could be expected to occur in any skull exhibiting rapid 
anteroposterior growth of the nasal and/or premaxilla relative to 
the lacrimal. A comparable prefrontal morphology is seen in the 
similarly proportioned but clearly unrelated Archeria (Holmes, 
1989), and, as Boy pointed out, the pterygoid (?read parasphe- 
noid)-vomer contact arises independently in most other genera 
included in his analysis (Boy, 1990, fig. 9). 

Sequeira and Milner (1993) hypothesized a monophyletic Edo- 
poidea based on the presence of enlarged premaxillae that form 
a long common medial suture and extend far posteriorly along 



1994 COCHLEOSAURUS FLORENSIS FROM NOW A SCOTIA 23 

the jaw margin behind this median suture, bordering small inset 
external nares. Unlike the characters used by Boy (1990), this 
represents a unique form of snout elongation in which the pre- 
maxillae account for most of the increased preorbital length and 
is here considered more reliable for establishing relationships. 
The anatomy of Cochleosaurus florensis supports this hypothesis. 
The occurrence of the prefrontal-jugal contact excluding the lac- 
rimal from the orbit, although probably derived within the group, 
is correlated with snout elongation and found in other groups 
(e.g., embolomeres) and is of less certain value. 

ACKNOWLEDGMENTS 

We would like to thank Mr. A. Lewis, then of the Museum of 
Comparative Zoology, for expert preparation of the specimens, 
Dr. F. Jenkins, Jr. and Mr. C. Schaff, both of the same institution 
for permission to borrow and study specimens of Cochleosaurus 
florensis, and Dr. R. Reisz for bringing the existence of MCZ 4342 
to our attention and suggesting this project be undertaken. Com- 
ments by Dr. R. Reisz, Mr. D. Dilkes (University of Toronto), 
and Drs. R. L. Carroll, Donald Baird, and Robert Hook improved 
an earlier draft. Special thanks to Dr. Andrew Milner and Sandra 
Sequeira for aid and advice in interpretation of a few difficult 
features and sharing their views on the anatomy and relationships 
of cochleosaurids. Meticulous criticism of two anonymous re- 
viewers saved us from many errors and omissions both major 
and minor. This research was supported by the Natural Sciences 
and Engineering Research Council of Canada, the Fonds pour la 
Formation de Chercheurs et L'aide a la Recherche (FCAR) of the 
Government of Quebec, the Gakken Corporation of Japan, and 
the Royal Tyrrell Museum of Palaeontology (Department of Com- 
munity Development, Alberta). 

LITERATURE CITED 

Beaumont, E. H. 1977. Cranial morphology of the Loxommatidae (Amphibia: 
Labyrinthodontia). Philosophical Transactions of the Royal Society of Lon- 
don (B), 280: 29-101. 

Boy, J. A. 1 990. Uber einige Vertreter der Eryopoidea (Amphibia: Temnospon- 
dyli) aus dem europaischen Rotliegend (?h6chstes Karbon-Perm). 3. Onchio- 
don. Palaontologische Zeitschrift, 64: 287-312. 



24 BREVIORA No. 500 

Bystrow, a. p. 1935. Morphologische Untersuchungen der Deckknochen des 
Schadels der Wirbeltiere. 1 . Mitt. Schadel der Stegocephalen. Acta Zoologica, 
16:65-141. 

Carroll, R. L. 1964. Early evolution of the dissorophid amphibians. Bulletin 
of the Museum of Comparative Zoology, 131(7): 161-250. 

. 1967a. A limnoscelid reptile from the Middle Pennsylvanian. Journal 

of Paleontology, 41: 1256-1261. 

. 1967b. Labyrinthodonts from the Joggins Formation. Journal of Pale- 
ontology, 41: 111-142. 

. 1969. A Middle Pennsylvanian captorhinomorph, and the interrela- 
tionships of primitive reptiles. Journal of Paleontology, 43: 151-170. 

. 1977. Patterns of amphibian evolution: an extended example of the 

incompleteness of the fossil record, pp. 405-437. In A. Hallam (ed.). Patterns 
of Evolution. Amsterdam, Elsevier Science Publishing Co. 

1988. Vertebrate Paleontology and Evolution. New York, W. H. Free- 



man. 698 pp. 
Carroll, R. L., D. S. Belt, D. Di>fELEY, and C. McGregor. 1 972. Guidebook, 

Vertebrate Paleontology of Eastern Canada. 24th International Geological 

Congress, Canada. Field Excursion A-59, Montreal. 1 13 pp. 
Chase, J. N. 1965. Neldasaurus wrightae, a new rhachitomous labyrinthodont 

from the Texas Lower Permian. Bulletin of the Museum of Comparative 

Zoology, 133: 153-225. 
DiLKES, D. W. 1990. A new trematopsid amphibian (Temnospondyli: Disso- 

rophoidea) from the Lower Permian of Texas. Journal of Vertebrate Pale- 
ontology, 10(2): 222-243. 
Foreman, B. 1990. A revision of the cranial morphology of the Lower Permian 

temnospondyl amphibian Acroplous vorax Hotton. Journal of Vertebrate Pa- 
leontology, 10(3): 390-397. 
Fritsch, A. 1885. Fauna der Gaskohle und der Kalksteine der Permformation 

Bohmens. Vol. 2. Prague. 1 14 pp. 
Godfrey, S. J., A. R. FiORJLLO, AND R. L. Carroll. 1987. A newly discovered 

skull of the temnospondyl amphibian Dendrerpeton acadianum. Canadian 

Journal of Earth Sciences, 24: 796-805. 
Holmes, R. 1989. The skull and axial skeleton of the Lower Permian anthra- 

cosauroid amphibian Archeria crassidisca Cope. Palaeontographica, Abtei- 

lungA, 207: 161-206. 
Holmes, R., and R. L. Carroll. 1977. A temnospondyl amphibian from the 

Mississippian of Scotland. Bulletin of the Museum of Comparative Zoology, 

147:489-511. 
Jarvik, E. 1980. Basic Structure and Evolution of Vertebrates. Vol. I. London, 

Academic Press. 575 pp. 
Klembara, J. 1985. A new embolomerous amphibian (Anthracosauria) from 

the Upper Carboniferous of Florence, Nova Scotia. Journal of Vertebrate 

Paleontology, 5(4): 293-302. 
Langston, W. 1953. Permian amphibians from New Mexico. University of 

California Publications in Geological Sciences, 29(7): 349^16. 



1994 COCHLEOSAURUSFLORENSIS FROM NOW A SCOTIA 25 

MiLNER, A. R. 1978. A reappraisal of the early Permian amphibian Memono- 
menus dyscriton and Cricotillus brachydens. Palaeontology, 21(3): 667-686. 

. 1990a. The radiation of temnospondyl amphibians, pp. 321-349. /« P. 

D. Taylor and G. P. Larwood (eds.). Major Evolutionary Radiations. Sys- 
tematics Association Special Volume 42. Oxford, Clarendon Press. 

. 1990b. The relationships of the eryopoid-grade temnospondyl amphib- 



ian from the Permian of Europe. Acta Musei Reginaehradecensis S.A.: Scien- 

tiae Naturales, 22: 131-137. 
Panchen, a. L. 1970. Teil 5/A. Anthracosauria. Handbuch der Palaoherpeto- 

logie. Stuttgart, Fischer. 84 pp. 
Reisz, R. 1972. Pelycosaurian reptiles from the Middle Pennsylvanian of North 

America. Bulletin of the Museum of Comparative Zoology, 144: 27-62. 
Rieppel. O. 1 980. The edopoid amphibian Cochleosaurus from the Middle Penn- 
sylvanian of Nova Scotia. Palaeontology, 23(1): 143-149. 
Romer, a. S. 1930. The Pennsylvanian tetrapods of Linton, Ohio. Bulletin of 

the American Museum of Natural History, 59: 77-147. 
. 1939. Notes of branchiosaurs. American Journal of Science, 237: 748- 

761. 
. 1 947. Review of the Labyrinthodontia. Bulletin of the Museum of Com- 



parative Zoology, 99: 1-366. 

Romer, A. S., AND R. V. Witter. 1942. £"^0^5, a primitive rhachitomous am- 
phibian from the Texas red beds. Journal of Geology, 50(8): 925-960. 

Sequeira, S. E. K. and A. R. Milner. 1993. The temnospondyl amphibian 
Capetus from the Upper Carboniferous of Nyrany, Czechoslovakia. Palaeon- 
tology, 36(3): 657-680. 

Smithson, T. R. 1982. The cranial morphology of Greererpeton birkemorani 
Romer (Amphibia: Temnospondyli). Zoological Journal of the Linnean So- 
ciety of London, 76: 29-90. 

Steen, M. C. 1938. On fossil Amphibia from the Gas Coal of Nyrany and other 
deposits in Czechoslovakia. Proceedings of the Zoological Society of London 
(B), 108: 205-283. 

Trueb, L., and R. Cloutier. 1991. A phylogenetic interpretation of the inter- 
and intrarelationships of the lissamphibia (Amphibia: Temnospondyli), pp. 
223-313. In H. P. Schultze and L. Trueb (eds.). Origins of the Higher Groups 
of Tetrapods: Controversies and Consensus. Ithaca, New York, Cornell Uni- 
versity Press. 

Whittard. W. 1928. On the structure of the palate and mandible of .4 /r/zf^o- 
saurus decheni. Goldfuss. Annals and Magazine of Natural History, Series 10 
i: 225-264. 



B R E V I O R A 

MCZ 

useiim of Comparative Zoology 

JAN 2 3 1995 



us ISSN 0006-9698 



Cambridge, Mass. 10 jA>ajARY 1995 Number 501 

I iMiv/rp 

A NEW LIZARD OF THE GENUS MACROPHOLIDUS 

(TEIIDAE) FROM A RELICTUAL HUMID FOREST OF 

NORTHWESTERN PERU, AND NOTES ON 

MACROPHOLIDUS RUTHVENI NOBLE 

John E. Cadle'^ and Pablo Chuna M.- 

Abstract. Macropholidus ataktolepis. new species, is a microteiid lizard known 
only from the type locality, Bosque Cachil, in the western Andes of extreme 
southwestern Cajamarca Department, Peru. It differs from the only other species 
of the genus, Macropholidus ruthveni Noble, in having prefrontal scales and in 
having the paired series of enlarged dorsal scales disrupted at or before midbody, 
rather than continuing to the tail. In addition, taxonomic data and natural history 
observations for M. ruthveni are summarized, including data for a large sample 
from the Rio Zaria valley of northwestern Peru (Cajamarca Department). 

Resumen. Macropholidus ataktolepis, nueva especie, es un microteido cono- 
cido solamente en la localidad tipica, Bosque Cachil, en los Andes occidentals 
del extremo suroeste del departamento de Cajamarca, Peru. La nueva especie 
difiere de Macropholidus ruthveni Noble, la linica otra especie del genero, por 
tener escamas prefrontales y tener la serie pareada de escamas dorsales agrandadas 
disruptidas hasta o antes de la mitad del cuerpo, antes que continuar hasta la cola. 
Ademas, se resumen datos taxonomicos y observaciones de la historia natural 
para M. ruthveni. incluyendo datos para una muestra grande de esa especie pro- 
cedente del valle del Rio Zana al noroeste del Peru (Departamento de Cajamarca). 

INTRODUCTION 

As a result of recent explorations, the Andean slopes of north- 
western Peru continue to yield many new species of amphibians 
and reptiles (Cadle, 1989, 1991; Cadle and McDiarmid, 1990; 
Duellman and Wild, 1993). In addition, distributions of species 



' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 

- Universidad Antenor Orrego, Apartado 1075, Trujillo, Peru. 
' To whom reprint requests should be addressed. 



2 BREVIORA No. 501 

previously known from few localities in this region are being 
refined. Most of the new discoveries have come from remnants 
of mesic to humid forests that occur in scattered patches on the 
western slope of the Andes from the Ecuadorian border to central 
Peru (H. W. Koepcke, 1957, 1961; H. W. Koepcke and M. 
Koepcke, 1958; M. Koepcke, 1954). These forests are thus islands 
in the sea of arid mountainous terrain characteristic of this portion 
of western South America. This paper describes a small lizard 
recently discovered in one such forest remnant and provides ad- 
ditional data on its presumed closest relative, Macropholidus ruth- 
veni Noble. Macropholidus ruthveni proved to be common at 
Bosque Monte Seco, another forest isolate in the Rio Zaiia valley 
just north of the type locality of the new species and from which 
other species of frogs, lizards, and snakes have been recently 
described (Cadle, 1989, 1991; Cadle and McDiarmid, 1990). 

Noble (1921a) erected the genus Macropholidus for a species 
of microteiid lizard (type species, M. ruthveni) from the "cordil- 
lera forming the boundary between the Departments of Piura and 
Cajamarca [Peru]." He compared Macropholidus only to Pholi- 
dobolus, a genus of the Ecuadorian Andes, from which Macro- 
pholidus was distinguished by ( 1 ) its possession of two enlarged, 
smooth, hexagonal rows of medial dorsal scales, the character 
that provided the etymology for the generic name, and (2) the 
lack of reduced scales laterally on the body. Macropholidus ruth- 
veni has, until now, been known from only the four specimens in 
the type series. Subsequently, Parker (1930) described Macro- 
pholidus annectens from the vicinity of Loja City, Ecuador, and 
noted that this species shared scutellational characters with both 
Macropholidus and Pholidobolus. In particular, M. annectens 
lacked the enlarged dorsals characteristic of M ruthveni but, in- 
stead, had dorsal scales similar to some species of Pholidobolus. 
Montanucci (1973) transferred annectens to Pholidobolus but re- 
tained Macropholidus as a monotypic genus characterized by the 
enlarged dorsals, a feature not seen in Pholidobolus (Montanucci, 
1973:5). Other than Montanucci's brief discussion, Macropholi- 
dus has rarely been mentioned in the literature subsequent to 
Parker's description of annectens. Some workers (e.g., Presch, 
1980) treated the two genera as synonymous based on a consid- 
eration o{ annectens, which is much better known than ruthveni, 



MCZ 
LIBRARY 

1994 NEW TEIID LIZARD FROM PERU JAM p? |hQi: 

the type species of Macropholidus. The new species described 
herein has the enlarged dorsals characteristic of Macropholidus,^]^ 
but they do not form as extensive a series as in the type species,! TY 
AI. ruthveni. 

MATERIALS AND METHODS 

The type series of the new species consists of nine specimens. 
The type series (A^ = 4) of Macropholidus ruthveni was also ex- 
amined, as well as a series (A^ = 24) of that species recently col- 
lected by the senior author from the Rio Zaiia valley, southwestern 
Cajamarca Department, Peru, and one other specimen referred 
to ruthveni from a locality (Lima Department) far south of the 
other known localities for that species. Details on these samples 
are given in the section on ruthveni, later. 

Snout-vent length (SVL; the straight line distance from the tip 
of the snout to the vent) and tail length (TL; vent to tip of tail, 
regenerated portion separated by a + sign) were measured to the 
nearest 0.5 mm with a metric ruler. All other measurements were 
taken to the nearest 0. 1 mm with dial calipers: head length (HL; 
tip of snout to posterior margin of ear), maximum head width 
(HW) and depth (HD), and body length (BL; posterior margin of 
arm to anterior margin of leg). 

The terminology of scales in lizards generally is difficult to 
standardize; we used Peters (1964) and Smith (1946:17-30) as 
guides herein. Most of the potentially confusing scale terminology 
concerns the circumorbital series and the series of scales on the 
ventral surface of the head. Definitions used herein are the fol- 
lowing: Superciliaries include scales contacting the supraorbitals 
and at least one-half of whose area is dorsal to the orbit. Genials 
are large paired scales posterior to the postmental, in contact on 
the midline and contacting the infralabials laterally. Postgenials 
are enlarged scales posterior to the genials, in contact laterally 
with the infralabials and not in contact medially. Gulars are scales 
enclosed by the genial-postgenial series anteriorly and the gular 
fold posteriorly; in Macropholidus, the gular series includes a 
paired series of enlarged medial gular scales extending anteriorly 
from the gular fold as well as smaller scales between the enlarged 
series and the genial-postgenial series. Dorsals are considered to 
be all scales on the trunk except for the squarish ventral plates; 



4 BREVIORA No. 501 

this definition includes "laterals" as defined by Smith (1946:27). 
In Macropholidus, the middorsal pair of dorsal scales is consid- 
erably enlarged {medial dorsals). Paradorsals are a pair of scale 
rows, somewhat larger than other dorsals, that border the medial 
dorsal rows. 

Transverse dorsal scale rows were counted from the occipitals 
to the posterior margin of the hindlimb. Transverse ventrals were 
counted between the limbs (axilla to groin). Counts of subdigital 
lamellae included the terminal claw sheath. 

Museum abbreviations for specimens referred to are Academy 
of Natural Sciences of Philadelphia (ANSP), American Museum 
of Natural History, New York (AMNH), Field Museum of Nat- 
ural History, Chicago (FMNH), University of Kansas Museum 
of Natural History (KU), and Museum of Comparative Zoology, 
Harvard University (MCZ). Specimens o{ Macropholidus ruthveni 
referred to only by J. E. Cadle field numbers (JEC) will be de- 
posited in the Museo de Historia Natural de San Marcos, Lima. 

DESCRIPTION 

Macropholidus ataktolepis, new species 
Figures 1, 3, 4 

Holotype(Figs. 1, 3, 4). MCZ 1 78050 (field number JEC 10320), 
an adult female collected by Pablo Chuna Mogollon, 28 Septem- 
ber 1991, at Bosque Cachil, approximately 3 km (airline) SE 
Contumaza, 2,400 m, Cajamarca Department, Peru (07°23'S, 
78°47'W; Fig. 2). 

The type locality (Fig. 2) is the site of ongoing biological surveys 
by Abundio Sagastegui, Pablo Chuna, and their colleagues of the 
Universidad Antenor Orrego, Trujillo, Peru. It lies in a small 
montane valley near the main road between Cascas and Contu- 
maza in extreme southwestern Cajamarca Department. 

Paratopotypes. The following eight specimens, all collected at 
the type locality: MCZ 178038-39 collected 27 July 1993 by P. 
Chuna M., P. Lezana, and S. Leiva; MCZ 178045-46 collected 
17 May 1993 by P. Chuna M. and P. Lezana; and MCZ 178264- 
67 collected 12 December 1993 by P. Chuna M. 

Etymology'. The species name, a noun in apposition derived 
from the Greek ataktos (disordered, irregular, not arranged) + 



1994 



NEW TEIID LIZARD FROM PERU 




Ci '"% .^ r% 



Figure 1. The female holotype of Macropholidus ataktolepis (MCZ 178050) 
in dorsal view. 



lepis (scale), alludes to the disruption of the orderly array of en- 
larged dorsal rows in this species, as compared to the completely 
regular series in the type species of the genus. 

Diagnosis. Macropholidus ataktolepis differs from the only oth- 
er species of the genus, M. ruthveni, in having (1) a pair of pre- 
frontal scales (absent in ruthveni); (2) the paired series of enlarged 
middorsal scale rows continuous only on the anterior part of the 
body (continuous to, or nearly to, the tail base in ruthveni); and 
(3) a regular arrangement of four enlarged temporal scales (irreg- 
ular in number [1-7] and arrangement in ruthveni). Macropholidus 
ataktolepis differs from Pholidobolus (formerly Macropholidus) 
annectens (Parker) in having prefrontal scales and a double row 
of enlarged medial dorsal scales on the anterior part of the body. 
No species of Pholidobolus, as currently defined (Montanucci, 
1973), has a double row of enlarged medial dorsals (see additional 
comments later). 

Description (Type Series). The type series comprises the female 
holotype (38.5 mm SVL) and the following paratypes: four males 
(MCZ 178038-39, 178265-66; 29-35 mm SVL); two females 
(MCZ 1 78045, 1 78264; 39-43 mm SVL); and two juveniles (MCZ 



BREVIORA 



No. 501 




Figure 2. The Andes of northwestern Peru showing distributions of species of 
Macropholidns. place names, and physical features. Stippled area is above 1 ,000 
m; hatched area is above 3,000 m. Star marks the type locality of M. ruthveni. 
Star within circle is the other known locality for M. ruthveni, Bosque Monte Seco 
(Cajamarca Department). Dot marks the type locality for M. ataktolepis, Bosque 
Cachil (Cajamarca Department). The question mark in northern Peru denotes the 
possible approximate locality for the "Chongollapi" paratypes of ruthveni (see 
text). The arrow within the inset map shows the location of Chaclacayo (Lima 
Department), from which comes an enigmatic specimen provisionally referred to 
M. ruthveni (KU 220845; see text). 



178046, 178267; 19.5-22 mm SVL). Thus, adult females attain 
a larger size than adult males. Measurements and scale counts of 
the holotype are given in Table 1 , and meristics and proportional 
data are given for the series in Table 2. 

HL 21-24% SVL in adults (28% in juveniles), 1.5-1.9 times 
longer than wide, 1.3-1.7 times wider than high. Head slightly 



1994 



NEW TEIID LIZARD FROM PERU 



Table 1 . Measurementts (in millimeters) and scale counts for the holotypes 
OF Macrophoudus ataktolepis and Macropholidus ruthveni. 



ataklolepis 


ruthveni 


Holotype, 


Holotype, 


MCZ 178050, 


MCZ 14041, 


Female 


Female 



Snout-vent length 

Tail length 

Head length 

Head width 

Head depth 

Body length 

Scales around midbody 

Subdigital lamellae, finger IV 

Subdigital lamellae, toe IV 

Total transverse dorsal rows 

Total enlarged medial dorsal pairs 

Transverse ventrals between limbs 



38.5 


45.5 


2 + 24 


34 + 37 


8.2 


9.6 


4.8 


5.5 


3.2 


3.4 


20.3 


26.1 


19 


19 


13, 14 


16, 16 


16, 16 


19, 18 


31 


33 


15 


30 


21 


21 



wider than neck, which is as wide as anterior body. Body cylin- 
drical, slightly depressed. Complete tail in adults greater than 
twice SVL (2.1 times SVL in MCZ 178045 with tail tip missing, 
2.4 times SVL in MCZ 178038 with complete tail); 68% and 70% 
of total length in these two specimens, respectively. Tail squarish 
to oval in cross section at base, tapering toward tip. Limbs pen- 
tadactyl, with well-developed digits; all digits with terminal claws. 
Forelimb extended forward along neck and head reaches posterior 
border of eye, or slightly anterior to this. Tongue (examined in 
MCZ 178045) lanceolate, covered with thin, scale-Hke papillae 
arranged in oblique rows, tip bifid; 8/8 heavily pigmented in- 
fralingual plicae. Anterior teeth conical, posterior teeth laterally 
compressed, tricuspid. 

Head. Head short, depressed (depth 58-79% of width); snout 
blunt (Figs. 1, 3). Rostral wider than deep, visible from above, 
laterally in contact with first supralabial and anterior nasal, dor- 
sally in contact with frontonasal. Frontonasal pentagonal, with 
slightly curved anterior border and obtusely pointed posterior 
border, separating nasals; posterolaterally narrowly contacting lo- 
real(MCZ 178039, 178045-46, 178050, 178264-67) or narrowly 
separated from it by prefrontal-posterior nasal contact (MCZ 



BREVIORA 



No. 501 



+1 



s ^ 



^ 1/3 



(/) 

w 

Z 

> 

Q 
w 
X 
w 

Z 
D 



^ 



"1 

~2 . - 



lO 

S 
■Q 



§•1 






a 

— (^1 
§) II 
o < 

U 



o 
E 

3 

o 

3 
O 

U 



o 
o 



"^ l^ ^~* 



§■1 

I 



u 
3 
O" 
(/I 

o 

03 



>3 _- 

"I -a -g 
2 &o ^ 



 

00 



00 m 
oo 



< 



— m (N u-1 O ^-. 





m 




r<) 


__, 


+1 


1 


+1 


_l +1 


1 




(N 






o 


W-1 


m 


O 


m iri 











d 
+1 

d 



d 
+1 

00 



2:0 
oi:: 



^^ d 

:2 +1 

00 



I 

00 



O ^_^ m ^_^ O ^^ in 
— I — d -^ — < <^ — ^ 



R q 



«-) ^^ o ^^ 

—  Co — ' On" 



O 



(N ro fN 

+1 Jl +1 -I +1 J +1 _L <:y^ - +1 I +1 I 



O 
<N 



00 






00 



m 



00 



O <N — c t^ (N «^ — 



+1 



rs| ro r^-i 

I +1 I +1 I 

0\ <N >/^ 

0\ — . — I r<^ 00 (N OS 

Tf ^ d "" '^. O^ 

m r^ r^ — I 



OS 







00 


so 


r^ 


r- 


,_^ 


00 


^^ 00 ^^ 


-^ 


^ d 





d 


^ d 




t^ 


fN 




— (^) 


+1 


1 +1 


1 


+1 


1 +1 1 







t^ 




— vo 



■^ — OS — . Tt — 



o 



O 01 Tf — . r~- rn 

SO ^__^ Os r-- ". r-- so — I 
cjrsi — ir>(-io'^(Ndod?— oT 

+1 I +1 I +1 I +1 I +1 I +1 I +1 I 

O OS fN_^(N 00 <N SO 

OsfNOOojm-^^. — .(^-H(N — SO— • 



o 






ir> 



OS 



m 







a 


00 










^^ 


1- 




;*. 






OJ 


c^ 




T3 






y5 


"c 







«0 

U-i 

c 
<u 
> 


C/3 






■3 


2 


U1 


X3 


■T3 


u 


_ 





C 


Si 

1- 


in 




-0 


C3 
(A) 
U 



3 


> 

c 


> 
t/) 
C 




T3 

15 


15 

1/1 

u 






2 


"c 





■0 


15 



— 1 


H 


w 


H 




t/5 



m 


t-^ 


00 

c 




<-• 


aJ 


aJ 


iH 


2. 










4J 


u 


s 


s 


w 


C3 



■0 

3 
C/5 



3 



1994 



NEW TEIID LIZARD FROM PERU 



Q 



O 

U 



U 



l>5 






a 

QO II 

o < 



U 



=0 

S^ "S y > 

b s 3 < 

r* "^ O 

I ^ 



O 

Co W 

:s . - i> - 

;; s « < 
a "^ 3 ^ 
^ cr 

m 



a 









v2 a 



3 

o 



00 








oo 




00 




00 




00 


m 


r<1 


CTv 


CO 


d 


X 

z 


IT) 

d 


X 

Z 




:s 




S 




< 




< 



00 

00 
00 







o <^ o 


m 


■Ai 


± 0. 
-0.2 
± 0. 
.68) 


rf 


^ 


m 


'^ 


0.22 
(0.21 
0.68 
(0 



O 

d 
+1 

in 



in — 



moo 



>n r<^ Tt 

m ■rj- 04 

I I I 

"^ P P 

— ' ^ 00 

m rn — ' 



o 
d 

+1 



m 

— m 

^^ o 



O 



O 

+1 
: d 



O 

d 



o 
d 
+1 

00 



m <^) — ' 

^ o ^ 



0\ O v^ 

:? +1 :^ 
?-? 

t^ m oo 

^ d '^. 

O O 



s £ S 

S" 9 o 
■n m ,^i 

q Ln u^ 

o^ 00 (j; 

<N rn X. 



o^^o^..^o^^o,.^ 
d> <^ d "^ 



° ^ ^ 



ON 



+ 1 



^ 



+1 d 



+1 



? 



+1 d ^ 
OoOd<^d<^d 



<u 

N 



_1> 

"0. 






oo 

c 

-J 
> 






> 

1/5 






00 

c 

a 

4= 






c 
at 
> 

3 



00 

c 




<u 


X) 


T5 


T3 


« 


« 


OJ 


w 


X 


X 



c c 

JJ O 

05 r:: > 

o 2 c^ 

^ ^ S 

•5 S 00 

c B- w 

."3 o T3 

W ^^ O 
H 02 



X 






00 
00 

X 

z 
< 



c 
o 



03 
C 



X> 
O 

(U 
u 
C3 

s 

<4j 



^ 






03 



a 

i2 

"o 
00 

c 
o 

s: 
U 






e 

o 



10 BREVIORA No. 501 

1 78038). Prefrontals hexagonal, in narrow medial contact. Frontal 
hexagonal, longer than wide, broader anteriorly. Frontoparietals 
hexagonal with long medial suture, each individually much longer 
than wide, collectively nearly as wide as long. Interparietal hep- 
tagonal, longer than wide. Parietals irregularly polygonal, about 
as wide as long; equal to or shorter than interparietal. Parietal 
contacts the upper postorbital on each side (MCZ 1 78039, 1 78046, 
178050, 178264-67), or parietal and postorbital separated by 
contact between posterior supraocular and upper anterior tem- 
poral (MCZ 178045), or there is narrow parietal-postorbital con- 
tact on the right side, which is reduced to a point on the left (MCZ 
178038). Three postparietals (occipitals), two lateralmost scales 
large, hexagonal; medial scale small, pentagonal. First pair of 
medial dorsal scales on neck (nuchals) distinctly enlarged (broader 
and wider than following dorsals). Head scales smooth, with scat- 
tered pores mostly located around the periphery of dorsal head 
plates, temporals, and supralabials; a few pits on other head scales. 

Lower eyelid with transparent disk. Two subequal supraoculars 
in direct contact with superciliaries (posterior one in contact with 
upper postocular; anterior supraocular irregularly hexagonal, pos- 
terior one squarish or pentagonal. Four superciliaries (three on 
one side in MCZ 1 78039), anterior scale more than twice as large 
as any other, and overlapping onto top of head. (Noble [1921a: 
138] considered there to be five superciliaries in M. ruthveni, with 
the last being the scale here considered the upper postocular. We 
consider the latter scale part of the postorbital series because 
essentially none of its area is above the orbit. Noble used neither 
pre- nor postocular for any of the circumorbital series. Both atak- 
tolepis and ruthveni typically have only four superciliaries under 
the present scheme.) 

Nostril in extreme posterior part of anterior nasal scale, bulging 
into anterior part of posterior nasal; anterior nasal larger than 
posterior nasal (this condition is the same as in M. ruthveni, for 
which Noble [192 la: 137] stated the condition as "[n]ostril be- 
tween the nasals"). Loreal generally large, higher than wide, con- 
tacting the posterior nasal, prefrontal, anterior superciliary, preoc- 
ular, second supralabial, and (except MCZ 178038) also narrowly 
contacting the frontonasal (see later for exceptions to this pattern). 



1994 



NEW TEIID LIZARD FROM PERU 



11 






Figure 3. Head of Macropholidus ataktolepis in dorsal, ventral, and lateral 
views (MCZ 178050, female holotype). Bar = 1 mm. 



Three unusual loreal conditions were observed: (1) loreal di- 
vided transversely into dorsal and ventral portions (MCZ 1 78039); 
(2) posterior nasal unusually small and with bilateral dorsal con- 
tact between loreal and anterior nasal (MCZ 178266); and (3) 
bilateral fusion of posterior nasal with ventral portion of loreal, 



12 BREVIORA No. 501 

and contact of this enlarged scale with preocular (MCZ 178264); 
thus, loreal in this specimen does not contact supralabials. 

Preocular triangular, small, in contact with loreal, second and 
third supralabials, and anterior subocular (contact with second 
supralabial reduced to a point in MCZ 178038 and 178050). 
Suboculars 3 (MCZ 178045, 178267) or 4 (all others); anterior 
and posterior scales largest. Postoculars 3, dorsal scale largest, 
ventral smallest (ventral scale in series with the suboculars, but 
more than twice as large as any subocular). Supralabials 7 (uni- 
lateral conditions of 8 and 6 in MCZ 178039 and 178264, re- 
spectively), 4th under eye and also longest. 

Temporal region covered by four large polygonal juxtaposed 
scales (five on one side in MCZ 178046). plus a series (6-1 1) of 
smaller polygonal scales located generally anterior and ventral to 
the enlarged temporals (Fig. 3). Anterior dorsal enlarged temporal 
contacts upper postocular, parietal, and lateral postparietal (in 
MCZ 178045 also narrowly contacting posterior supraocular). 
Posterior dorsal enlarged temporal contacts lateral postparietal 
and the first transversely enlarged dorsal scale. Posterior ventral 
enlarged temporal separated from ear by one row of denticles. 
Ear opening round to vertically oval, bordered by small denticles; 
tympanum deeply recessed. 

Mental with straight posterior margin (Fig. 3). Postmental large, 
obtusely pointed posteriorly, in lateral contact with infralabials 
1-2. Two pairs of genials in contact on midline; anterior pair 
large, squarish, in contact with infralabials 2-3. Posterior pair 
large, pentagonal, in contact with infralabials 3-4. Two pairs of 
enlarged postgenials, anterior pair much larger than posterior pair, 
separated medially by two (anterior postgenials) to six (posterior 
postgenials) gular scales; anterior postgenials in contact with in- 
fralabials 4 or 4-5; posterior postgenials in contact with infrala- 
bials 4 (narrowly) and 5, or 5 only. Infralabials 5 (6 on one side 
in MCZ 178039 and 178267), 4th longest. 

Neck and Body. Anterior gular region (between posterior pair 
of genials and enlarged gulars; see Fig. 3) filled with small polyg- 
onal gular scales in roughly six to seven irregular rows. Posterior 
gular region (level of posterior margin of ear opening to the gular 
fold) covered by four to six pairs (5'/2 pairs in MCZ 178038) of 
enlarged, rhomboid gular scales, each wider than long. Gular fold 
weak, ill defined, and without hidden scale rows. 



1994 



NEW TEIID LIZARD FROM PERU 



13 





1 



I 




Figure 4. Comparison of dorsal body scalation of Macropholidus ataktolepis 
and M. ruthveni. Left: Holotype of M. ataktolepis (MCZ 178050), showing the 
break-up of the enlarged medial dorsals shortly behind the shoulder region (at 
level indicated by arrow). Right: M. ruthveni (ANSP 31765, from Bosque Monte 
Seco, Rio Zana, Cajamarca Department), showing array of enlarged medial dorsals 
continuing to the tail (compare also Fig. 8). Note also the more squarish shape 
of the enlarged dorsals on the anterior part of the body in M. ataktolepis and their 
more hexagonal form in M. ruthveni. 



Side of neck anterior to arm covered with medium-sized, 
rounded, juxtaposed to weakly overlapping scales. Axillary scales 
small, rounded, juxtaposed. 

Dorsal scales of neck in two enlarged smooth rows (medial 
dorsals); anteriorly each medial dorsal >1 times as wide as long, 
gradually becoming more squarish by the shoulder region, con- 
tinuing in two parallel series, becoming gradually smaller. Medial 
dorsals bordered on each side by a parallel series of somewhat 
enlarged paradorsals. Near midbody medial dorsal scales ap- 
proximately equal in size to lateral body scales, no longer obvi- 



14 BREVIORA No. 501 

ously in parallel series (Fig. 4); number of pairs of enlarged medial 
dorsals varies from 1 2 to 20: 1 2 (MCZ 1 78266), 1 3 (MCZ 1 78045), 
14 (MCZ 178046, 178265), 15 (holotype, MCZ 178038), 17 (MCZ 
178267), 19 (MCZ 178264), and 20 (MCZ 178039). Posterior 
dorsal scales smooth, squarish, slightly imbricate, in irregular 
transverse series with lateral body scales (Fig. 4). Usually slight 
misalignment between lateral and dorsal body scales over mid- 
dorsal region, caused by differing shapes and slightly different 
sizes of the two sets of scales. Posterior middorsal scales often 
irregular in shape (quadrangular to obtusely cycloid) and size. 
Scales around midbody 18-20. 

Lateral body scales smooth, bluntly pointed, imbricate, slightly 
smaller than posterior dorsal scales. No lateral rows reduced in 
size, although a few scattered small, imbricate scales about one- 
half the size of other lateral scales are present. Lateral fold absent. 

Ventralmost dorsals (i.e., the scales immediately bordering the 
ventral plates) somewhat larger than the other dorsal rows. Ven- 
trals smooth, larger than ventralmost dorsals, squarish to rect- 
angular, in four longitudinal rows, in 20-22 transverse rows be- 
tween limbs. (Noble [1921a] included the ventralmost dorsal rows 
in his count of six transverse abdominal plates for M. ruthveni. 
Since the scales in these rows have the shape typical of the other 
dorsals, although slightly larger, they are considered part of the 
dorsal series here. Both ataktolepis and ruthveni have four lon- 
gitudinal rows oi quadrangular ventrals.) One pair of anal scales; 
one pair of enlarged preanals. Femoral and preanal pores absent. 

Tail and Limbs. Caudal scales at base of tail dorsally and lat- 
erally hexagonal, imbricate, weakly striated to very weakly keeled 
dorsally. Ventral surface of tail with paired series of somewhat 
enlarged, squarish, smooth, weakly imbricate scales. 

Upper surface of arm and hand covered with large, smooth, 
polygonal, imbricate scales that gradually decrease in size distally. 
Ventral surface of arm covered with smaller imbricate scales, 
somewhat conical and nonoverlapping proximally. Anterior, dor- 
sal, and ventral surface of thigh with large, smooth, imbricate, 
plate-like polygonal scales. Posterior surface of thigh with small 
conical or pavement-like scales. Lower leg dorsally and ventrally 
with weakly imbricate scales half the size of those on anterodorsal 
surface of thigh. Top of foot with large imbricate scales twice or 



1994 NEW TEIID LIZARD FROM PERU 15 

more the size of those on lower leg. Palms and soles covered with 
small conical to pavement-like juxtaposed scales. 

Subdigital lamellae as follows (roman numerals = digits; arabic 
numbers = range for subdigital lamellae in type series counted 
on one of each pair of feet for each specimen): forefoot, I 5-7, II 
8-10, III 12-14, IV 12-14, V 9; hindfoot, I 6-8, II 9-12, III 13- 
17, IV 16-19, V 11-12. 

Coloration in Life (Holotype). Dorsum medium brown. Top of 
head grayish brown. Dorsal and ventral surfaces of tail and dorsal 
surface of hindlimbs dark charcoal gray. Tan dorsolateral stripes 
from temporal region, fading into dorsal color just behind scap- 
ular region; bordered dorsally by dark gray/blackish thin line. 
Whitish supralabial/neck stripe present. Loreal, temporal, lateral 
neck regions and flanks dark charcoal gray, paling somewhat on 
flanks. Anterior gular region dull whitish with grayish wash. Ven- 
ter similar, but with dull orangish wash in pectoral region and 
laterally. Edge between belly and flanks and pelvic area speckled 
with dark blackish flecks. Ventral surface of forelimbs with dull 
orangish wash and tiny dark flecks. 

Coloration in Preservative (Holotype). Top of head and dorsal 
surface of body grayish brown, becoming greenish gray posteriorly 
on body, and slate gray on tail. Dorsal head scales and enlarged 
scales on top of neck heavily and finely speckled with black (speck- 
ling on neck concentrated on medial edges of light dorsolateral 
stripes); the speckling continues onto body but gradually decreases 
in intensity. Coloration of lateral surface of body and dorsal sur- 
face of hindlimbs dark grayish brown, sharply set off'from dorsal 
coloration, heavily speckled with black. Lateral surface of neck 
and temporal region slate gray, with fine lighter flecks (visible 
only under microscope). Loreal region and supralabials yellowish 
brown, heavily suffused with black. Top of forelimbs yellowish 
brown, heavily speckled with black concentrated proximally. Pale 
(yellowish) dorsolateral stripe beginning as thin line on antero- 
lateral edge of parietals, widening on enlarged paradorsals of neck 
(occupying about V^ of these scales), gradually fading posterior to 
shoulder region. Very thin white supralabial stripe beginning at 
posterior edge of second supralabial, continuing along middle of 
supralabial row, then dropping to labial border on last two su- 
pralabials and continuing to anteroventral border of ear; several 



16 BREVIORA No. 501 

scales with light centers forming line behind the ear, but not 
forming distinct stripe. Ventral surface of head, neck, and body 
grayish white, finely (on head and neck) to heavily (posteriorly 
and laterally on body) speckled with black. Ventral surface of 
forelimbs whitish, with only a few black speckles. Ventral surface 
of hindlimbs whitish, heavily speckled with black. Palms dusky. 
Soles dark gray brown. Ventral surface of tail dark slate gray with 
fine lighter speckling. 

The holotype retains more details of coloration and pattern 
than any paratype, probably as a result of differential preservation. 
Six paratypes (MCZ 178045-46, 178264-67) are very dark, al- 
most black. The other two (MCZ 178038-39) are essentially as 
described for the holotype, but the dark ventral pigmentation is 
more evident, and there is no sharp distinction between the dorsal 
and lateral body pigment. The dorsolateral stripes are visible in 
all paratypes. 

Comparison oi MacrophoUdus ataktolepis 
with M. ruthveni 

In general form, body proportions, and coloration, M. atak- 
tolepis and M. ruthveni are virtually indistinguishable (preserved 
specimens of both species can differ markedly in color and pattern, 
but we attribute these differences to the effects of preservation 
rather than to substantive color differences in life; see the follow- 
ing description of coloration in ruthveni for a potential pattern 
difference between the species). Because scale fusions and some 
intraspecific variation are characteristic of many species of mi- 
croteiids generally (the so-called "normal" fusions or divisions 
that give rise to variation in, for example, the number of supra- 
labial scales), the three scutellational differences between ruthveni 
and ataktolepis noted in the diagnosis are commented upon briefly 
here. 

Prefrontal Scales and Other Head Plates (Figs. 3, 5). The pres- 
ence of paired prefrontal scales between the frontonasal and fron- 
tal scale in ataktolepis creates differences between ataktolepis and 
ruthveni in the shapes of these scales. In ataktolepis, both the 
frontonasal and frontal are hexagonal, with oblique angles pos- 
teriorly and anteriorly, respectively. In ruthveni, the frontonasal 
is squarish and the frontal pentagonal, and the two scales meet 



1994 



NEW TEIID LIZARD FROM PERU 



17 






Figure 5. Head of Macropholidus ruthveni in dorsal, ventral, and lateral views 
(MCZ 14041, female holotype). Lateral view is right-side reversed. Bar = 1 mm. 



in a straight border. In some individuals of ruthveni, the frontal 
shape might be interpreted as marginally heptagonal, produced 
by more than point contact between the frontal and first super- 
ciliary; in these cases, the frontal still retains a straight anterior 



18 BREVIORA No. 501 

border, and the anterolateral sides of the "heptagon" are very 
short. 

The prefrontal scales in ataktolepis are well defined and con- 
sistent in size, shape, and position in the available specimens (Fig. 
3). In the geographically heterogeneous sample of 28 specimens 
of ruthveni, three specimens show significant variation in the 
region of the frontal/frontonasals. With the exception of MCZ 
1 78036, these cases result in highly irregular and asymmetric scale 
patterns. MCZ 178036 has a pair of small triangular prefrontals 
at the lateral juncture of the frontal, frontonasals, and first su- 
perciliaries, which appear to have formed from fused portions of 
each of those scales; they are widely separated on the midline by 
a broad frontal-frontonasal contact, as is normal in ruthveni. The 
other two ruthveni specimens are more aberrant. FMNH 232606 
is unusual in the form of the frontal-frontonasal suture on the 
left side (curved posteriorly rather than straight), and it has two 
partial sutures within the frontonasal, which partially delimit a 
large irregular azygous scale on the left side between the frontal 
and frontonasal. A similar condition is seen in FMNH 232599, 
except that the supernumerary sutures are complete and the azy- 
gous scale itself is longitudinally divided into a medial and smaller 
lateral portion (the right side of this specimen is, as in FMNH 
232606, "normal"). The scales so formed are highly irregular in 
shape and do not approach the regular prefrontal shapes seen in 
all specimens of ataktolepis. 

The only unusual conditions of head plates in the series of 
ataktolepis are (1) the nearly complete fusion of the left fronto- 
parietal with the posterior supraocular in MCZ 178045, a fusion 
also seen in a paratype o^ ruthveni (MCZ 147313; Noble [1921a: 
139] stated that this was on the left side of one of the paratypes, 
but it is on the right side of MCZ 147313 and none of the other 
paratypes has this condition); and (2) the fusion of each posterior 
nasal scale with the corresponding ventral portion of the loreal 
(MCZ 178264), as already described. 

Prefrontal scales vary in their presence/absence within and 
among species of the presumably (see later) closely related genus 
Pholidobolus. Other than Pholidoholus macbrydei, however, in 
which the condition is apparently variable (Montanucci, 1973: 
1 6), prefrontals are typically present or absent in the other species. 



1994 NEW TEIID LIZARD FROM PERU 19 

Montanucci (1973:37) stated that there was a "high frequency" 
of prefrontal scales in P. macbrydei but did not give a specific 
proportion, nor state whether the variation was intra- or inter- 
populational; in a sample of six macbrydei from widely separated 
geographic areas (Cotopaxi and Azuay Provinces, Ecuador; MCZ 
154631-33, 163958-59), all individuals lacked prefrontals. It is 
the characteristic shape and population-specific nature of the pre- 
frontals in Macropholidus ataktolepis, their absence in a geograph- 
ically heterogeneous sample of ruthveni, and the coincidence of 
this character with unusual dorsal scale pattern (described in greater 
detail later) that lead us to interpret the nature of the prefrontals 
as characteristic of M. ataktolepis. 

A high proportion of scale aberrancies in ruthveni involving the 
region of frontal-frontoparietal contact is perhaps significant, in 
that this region gives rise to one of the diagnostic differences 
between M. ruthveni and M. ataktolepis. Such an association be- 
tween intraspecific variants and interspecific differences has been 
inferred for interspecific scale differences in one other group of 
microteiid lizards (Donnelly et al., 1992). 

Dorsal Scales of the Body (Fig. 4). The paired series of vertebral 
scales in both Macropholidus ruthveni and M. ataktolepis begin 
on the neck as transversely elongate hexagonal or rectangular 
scales. In both species (but seemingly more so in ataktolepis than 
in ruthveni; see Fig. 4), they become gradually more squarish 
posteriorly by extension in the longitudinal dimension, usually 
noticeably so just behind the shoulder region. In ruthveni, the 
enlarged scales continue virtually to the tail base (Figs. 4, 8). In 
ataktolepis, however, there is a generally rather abrupt transition 
to small dorsals by midbody (Fig. 4). The posterior dorsal scales 
behind this transition zone in ataktolepis are slightly larger than 
the lateral body scales at the same level and usually have straight 
posterior borders, in contrast to the bluntly pointed borders of 
the lateral scales. The transition to smaller scales in ataktolepis 
is sometimes accompanied by slight irregularities in the arrange- 
ment of dorsal scales, caused by varying scale sizes in the tran- 
sition zone. 

A comparison of the number of pairs of enlarged vertebral scales 
relative to the total number of transverse rows of dorsal scales 
(occiput to posterior margin of the leg) demonstrates this differ- 



20 



BREVIORA 



No. 501 





Figure 6. Top: Bosque Cachil as seen from the slope on the opposite side of 
the valley. Arrows mark the approximate upper extent of forest, above which is 
short bushy vegetation. The forest extends to the left and right off both sides of 
the photograph, but most of its extent is encompassed within the photograph. 
Bottom: General view of the terrain immediately down the valley from Bosque 
Cachil. Most slopes are denuded of vegetation. The trees in the lower left are 
Eucalyptus. Both photographs were taken on 28 September 1 99 1 . 



1994 NEW TEIID LIZARD FROM PERU 21 

ence between nithveni and ataktolepis well. Both species have 
comparable total numbers of scales in this area (32-37 in ruthveni\ 
29-35 in ataktolepis), but the number of pairs of enlarged dorsals 
in ruthveni is 25-34 (average of 3.3 for the difference between 
total transverse dorsals and enlarged dorsals), whereas in atak- 
tolepis it is only 12-20 (average of 17.4 scales difference) (see 
Table 2). 

Temporal Scales (Figs. 3, 5). All specimens of M ataktolepis 
have a regular arrangement of four enlarged temporal scales (Fig. 
3; MCZ 178046 has five on one side), and the total number of 
temporal scales (the region bounded by the postoculars, suprala- 
bials, parietals-postparietals, and the anterior margin of the ear) 
is 10-14 on each side. On the other hand, the arrangement of 
temporal scales in ruthveni is more irregular (Fig. 5). The frag- 
mentation of scales in the temporal region in ruthveni makes the 
delineation of "enlarged" from "normal" temporals a somewhat 
arbitrary distinction, as there is often a continuous gradation in 
scale sizes. The total number of temporal scales in ruthveni is 1 1- 
21, with the number that might be considered "enlarged" ranging 
from 1 to 7 (mode = 4, with other strong modes at 2 and 5). 
Some individuals of ruthveni are highly asymmetrical in the num- 
ber and size of temporals on each side; this was not observed in 
any ataktolepis. 

Distribution, Habitat, and Natural History 

Macropholidus ataktolepis is known only from the type locality, 
"Bosque Cachil." The following observations are largely extracted 
from Cadle's field notes made during a visit to Bosque Cachil on 
28 September 1991 (see Figs. 6, 7). The site presently has a small 
remnant of humid forest lying in a montane valley (Fig. 6), at the 
bottom of which is a small stream, known locally as Quebrada 
Cachil. Most of the forest is between 2,400 and 2,500 m, with 
some riparian forest extending somewhat lower. Dillon et al. ( 1 994) 
estimated the area of Bosque Cachil at about 100 ha. Surrounding 
the forest, the general habitat is scrubby chaparral-like bushland 
and disturbed terrain and obviously much drier (Fig. 6); however, 
the original extent of forest prior to human intervention is un- 
known and possibly was not much greater than at present. Mon- 
tane forests this far south in the western Andes of Peru are usually 



22 



BREVIORA 



No. 501 




Figure 7. Understory vegetation at Bosque Cachil, showing generally small, 
short-statured trees, but with abundant mosses and, in this view, bromeliads. A 
relatively long exposure due to low ambient light level resulted in slight fuzziness 
in the photograph. Photographed 28 September 1991. 



quite localized because local features of climate and aspect limit 
the extent of forest development (see H. W. Koepcke, 1957, 1 96 1 ; 
H. W. Koepcke and M. Koepcke, 1958). 

The end of September, when these observations were made, is 
well into the dry season (approximately May to December in this 



1994 NEW TEIID LIZARD FROM PERU 23 

portion of the Andes in Peru). At that time, the general aspect of 
Bosque Cachil was quite dry, although clear evidence of seasonal 
humidity was present. The forest is dominated by Podocarpus, 
with lesser amounts of Cliisia and Guarea. Some large bromeliads 
were present (Fig. 7), but nowhere were these dense. Orchids were 
present but not abundant, but no tree ferns {Cyathea) were seen. 
Mosses festooned most trees, but at this time these were all dry, 
crackly, and brown. Another indication of the dryness was the 
fact that soil under even large boulders was dry. The lower part 
of the valley below Bosque Cachil is scrubby chaparral-like bush- 
land and secondary growth, including introduced Eucalyptus (Fig. 
6). According to local inhabitants, the area is quite wet and cold 
from about January to April, sometimes with dense fog. The 
characteristics of the streambed at the bottom of the valley give 
another indication of the seasonal abundance of water at Bosque 
Cachil. In September 1991, stream flow was reduced to approx- 
imately 1-2 m wide at most points. However, the channel is deep 
and scoured in places, with many moss-covered boulders, cas- 
cades, and deep pools. This suggests much greater water flow 
during parts of the year. During a visit to Cachil on 1 7 May 1 993, 
M. O. Dillon (personal communication) reported a constant rain 
of about 2 hr, with high humidity lasting through the afternoon 
and evening. 

The forest of Cachil was apparently not studied by Hans and 
Maria Koepcke during their extensive surveys of the western An- 
dean forest remnants of Peru (H. W. Koepcke and M. Koepcke, 
1958; H. W. Koepcke, 1957, 1961; M. Koepcke, 1954, 1961). 
They did, however, study several other forest isolates on the south 
side of the Rio Chicama valley. They characterized the forests of 
Hacienda Llaguen (07°40'S, 78°40'W), directly south of Bosque 
Cachil across the Rio Chicama valley, and spanning comparable 
elevations (1,700-2,900 m), as "[c]actus and bushsteppe; tran- 
sition to riparian forest and mesothermic rainforest; light ever- 
green mountain forest to primary rainforest of the oligothermic 
zone in some places . . . between 2400 and 2900 meters" (H. W. 
Koepcke, 1961:31). The forest of Cachil, in both elevational zone 
and dominant vegetation, appears to correspond well to the "oli- 
gothermic rainforest zone," characterized by the presence of 
Podocarpus and trees locally densely covered with epiphytes (H. 
W. Koepcke, 1961:164). 



24 BREVIORA No. 501 

In general respects, the climatic regime (rainy season roughly 
January to April) appears similar to that at another forest isolate 
just north of Bosque Cachil, Bosque Monte Seco on the Rio Zafia 
(see Cadle, 1989, 1991; Cadle and McDiarmid, 1990; Sagastegui 
and Dillon, 1991). However, despite the presence of abundant 
mosses on the trees and a similar floral composition, Bosque 
Cachil has a drier aspect and is at a slightly higher elevation 
(2,400-2,500 m) than much of the humid forest at Monte Seco 
(1 ,500-2,500 m). Indicators of the greater aridity at Cachil include 
fewer streams, and those present with reduced water flow; shorter- 
stature forest in general; reports by locals of only occasional dense 
fog during the rainy season (ubiquitous during the rainy season 
at Bosque Monte Seco); and the absence of tree ferns (although 
possibly these were removed by earlier inhabitants of the region, 
as they are used medicinally; tree ferns, however, are common at 
Bosque Monte Seco; see Sagastegui and Dillon, 1991; Dillon et 
ai, 1994). 

The flora of Bosque Cachil appears to be a small subset of that 
at Bosque Monte Seco (Sagastegui and Dillon, 1 99 1 ; M. O. Dillon, 
personal communication), although with different dominant el- 
ements. Whereas the forest of Cachil appears to correspond to 
Koepcke's (1961) "oligothermic" rainforest, the more humid for- 
est of Bosque Monte Seco corresponds more with the "mesother- 
mic rainforest" and "cloud forest" habitats discussed by Koepcke 
(1961), which in general occur at slightly lower elevations in west- 
ern Peru than the "oligothermic rainforests" (H. W. Koepcke, 
1961). Other than an enigmatic species of Dipsas (Colubridae), 
no species of amphibians or reptiles presently known from Bosque 
Cachil occur also at Bosque Monte Seco, but much more extensive 
sampling of the herpetofauna needs to be done at Cachil. 

All specimens of Macropholidus ataktolepis were collected un- 
der rocks during the day. This species is presumably diurnal, as 
is its close relative M. ruthveni (see later). One female (MCZ 
178045) collected on 17 May contained one developing follicle, 
whereas the female holotype, collected at the end of September, 
and MCZ 178264, collected in December, contain no yolking 
follicles. The only other amphibians or reptiles now known from 
Bosque Cachil are undescribed species of Stenocercus (Tropidur- 
idae), Dipsas (Colubridae), and Eleuthewdactylus (Leptodactyli- 
dae) presently under study and a species of Gastrotheca (Hylidae). 



1994 NEW TEIID LIZARD FROM PERU 25 

NOTES ON MACROPHOLIDUS RUTHVENI NOBLE 

Types, Type Locality, and a New Series 
from the Rio Zaiia 

Noble (192 la: 138) described Macropholidus ruthveni on the 
basis of four specimens: the holotype and one paratype in MCZ 
and two additional paratypes in AMNH. The holotype, MCZ 
14041 (Figs. 5, 8), is from "Coucumayo, a half-way station be- 
tween the towns of Huancabamba and Tabaconas" (Noble (1921a: 
139), and MCZ 147313 (originally MCZ 14043) is from "Chon- 
gollapi" (=Chongoyape; see additional comments on localities 
later). The two AMNH paratypes, AMNH 38817-18, were both 
originally cataloged with "Coucumayo" as the locality. However, 
Noble (192 la: 139) stated that two specimens were obtained at 
each of the two localities mentioned in the type description; hence, 
one of the AMNH specimens is from Coucumayo and the other 
from Chongoyape. Noble gave sufficient detail in his description 
to identify AMNH 38818 as the "Chongollapi" specimen (21 
ventral scales between the collar [=gular fold] and the anal plates, 
and presence of a third supraocular in one of the Chongoyape 
specimens, as is the case in the holotype; AMNH 38818 is the 
only paratype that satisfies these criteria). AMNH 38817, then, 
is the other specimen from the type locality, Coucumayo. 

We have not found Coucumayo (stated by Noble to consist of 
a single house) listed in gazetteers or on maps, but Noble's de- 
scription places it on the eastern spur of the Cordillera Huanca- 
bamba on the border between the departments of Piura and Ca- 
jamarca (Fig. 2). According to Noble (192 la: 139), Coucumayo is 
"one of the highest points of the trail but is probably not over 
8,000 feet [=2,440 m] in altitude." Most parts of the cordillera 
between Huancabamba and Tabaconas rise well over 3,000 m, 
but a break with maximum elevations between 2,000 and 3,000 
m occurs 18-20 km southeast of Huancabamba at approximately 
05°20'S, 79°20'W (1:480,000 maps for the departments of Piura 
and Cajamarca, Instituto Geografico Nacional, Lima). This is the 
present location of the main route between Huancabamba and 
Tabaconas. Based on the elevation stated in the description, the 
type locality o{ ruthveni is most likely along this part of the ridge. 
Thus, the type locality is east of the continental divide in upper 
reaches of the Rio Huancabamba/Chamaya drainage. 



26 



BREVIORA 



No. 501 







Figure 8. Holotype of Macropholidus ruthveni, MCZ 14041, an adult female. 
Noble (1921a:139) stated that the light borders of the scales visible in the type 
were "not so distinct" in life. He also did not mention the dorsolateral stripe, 
visible faintly in the photograph. 



Noble did not give information about the general environment 
of Coucumayo, stating only that the two specimens of Macro- 
pholidus collected there were from a pasture. Huancabamba lies 
in a dry rain-shadow valley (Noble, 1921b; T. A. Parker et al, 
1 985); however, humid forest persisted even recently at elevations 
above 2,100 m on the slopes north and east of Huancabamba (T. 
A. Parker et al., 1985) and possibly lower in 1916 when Noble 




31 



1^.^ 



Figure 9. Dorsal view of Macropholidus ruthveni (ANSP 31764), an adult 
female from Bosque Monte Seco, Rio Zafia valley, Cajamarca Department, Peru. 



1994 NEW TEIID LIZARD FROM PERU 27 

visited the area. Tabaconas itself lies in a valley characterized by 
humid montane forest (Barbour and Noble, 1920; T. A. Parker 
et al., 1985). 

Noble purchased two of the paratypes (MCZ 147313, AMNH 
38818) in Chongollapi (=Chongoyape) from a restaurant keeper, 
who had preserved them in aguardiente. Chongoyape is at 200 
m elevation on the Rio Reque in the coastal desert region, Lam- 
bayeque Department, 06°39'S, 79°24'W (Fig. 2), but Noble sug- 
gested that the specimens probably came from "some of the high 
altitudes fifty or a hundred miles inland." This assessment makes 
sense with respect to natural history observations for ruthveni 
made by Cadle on a population at the Rio Zana (see later). It 
seems unlikely, therefore, that ruthveni would be found in low 
coastal desert (but see discussion of KU 220845, later). Noble's 
estimate, however, of "fifty or a hundred miles inland" is probably 
a substantial overestimate, whether considered in airline or trail 
distances; the nearest humid forests are about 50 km airline dis- 
tance and 100 km by the existing road east of Chongoyape (Cadle, 
personal observations). 

Presently, the type specimens of M. ruthveni are in fair con- 
dition and somewhat soft (the MCZ paratype is a small juvenile, 
which has had its head removed, critical-point-dried, and mount- 
ed for scanning electron microscopy; the mounted head still re- 
tains the outline of the head scutellation). The holotype (Fig. 8) 
is an aduh female, AMNH 38817 is an aduh male, and AMNH 
38818 is probably a male based on swelling of the tail base. 

In addition to the type series, we studied a series of ruthveni 
collected by Cadle from the upper Rio Zana (Bosque Monte Seco), 
Cajamarca Department (Figs. 2, 9; for details concerning the area, 
see Cadle, 1989, 1991; Cadle and McDiarmid, 1990; Sagastegui 
and Dillon, 1991). These specimens include ANSP 31764-69; 
FMNH 231771, 232599-608; MCZ 178036-37; and JEC 7202, 
7211, 7463, 7528, 7798-99, 8062. All of these specimens are 
from within a 3-km airline radius north to east of Hacienda Monte 
Seco, Rio Zana, Cajamarca Department, Peru (79°07'W, 06°51'S; 
Fig. 2). KU 220845, which we refer provisionally to M. ruthveni, 
is from Chaclacayo, Lima Department, a locality far south of the 
other known localities for ruthveni (Fig. 2). Details concerning 
this specimen are given later, but it was not included in the sum- 



28 BREVIORA No. 501 

maries of variation for nithveni immediately following. Data on 
the type series and the Rio Zaha sample ofAIacropholidus nithveni 
are summarized here, as are observations on the natural history 
of this species at the Rio Zaiia study site. 

Descriptive and Variational Comments 

The series referred to Macropholidus ruthveni from the Rio 
Zaiia (Bosque Monte Seco; see Fig. 9) is similar to the type series 
in all scutellational features, except for what might be considered 
normal (and, in this case, minor) intraspecific variation. Table 1 
gives measurements and scale counts of the holotype of M. ruth- 
veni, and Table 2 summarizes scutellation and mensural features 
in all samples of the species. The following comments augment 
the characterization given by Noble (1921a). 

Noble (1921a) noted several minor variations in head scalation 
in the type series, and the larger series examined by us does not 
significantly alter his characterization. Some of the variation was 
already discussed in the context of comparing ruthveni and atak- 
tolepis. Other variations are noted here. Virtually all specimens 
have two large supraocular scales on each side. The holotype 
differs in having three supraoculars on each side, the third being 
a small quadrangular scale formed by a short suture across the 
posterolateral portion of the second supraocular (Fig. 5). One 
other specimen, AMNH 38818 from Chongoyape, has three su- 
praoculars on the right side, similar to the pattern in the holotype, 
and the usual condition of two supraoculars on the left side. No 
specimens from the Rio Zaiia showed this variation. 

Proportional measurements and meristic counts for the spec- 
imens of ruthveni we examined are as follows. Complete tail 2.03- 
2.26 times SVL in three adults, 1.61-1.87 times SVL in five 
juveniles, and 0.62-0.69 times TL (adults and juveniles com- 
bined). The holotype, a female (SVL 45.5 mm), is the largest 
specimen. Range of SVL for other females was 36-43 mm, greater 
than the range for males (29-35.5 mm). The forelimb extended 
forward reaches the posterior border or middle of the eye. 

Superciliaries usually four, occasionally three or five (Noble 
included the scale we consider the upper postocular in his super- 
ciliary series, and therefore stated the condition in the type series 
as five superciliaries); postoculars usually three, occasionally two. 



1994 NEW TEIID LIZARD FROM PERU 29 

Supralabials seven, with the fourth under the midpoint of the eye 
(eight and six on two sides each). Infralabials five (six on one side, 
four on two sides). Scales around the middle of the body 1 7-20 
(19.3 ± 0.73). A distinct loreal, higher than wide, separating the 
posterior nasal from the preoculars. 

No lateral body fold or lateral scales of reduced size. Four 
transverse rows of quadrangular ventrals (Noble's count of six 
rows included the ventralmost rows of dorsals; see the comments 
in the comparison oi ataktolepis with ruthveni). 

Subdigital lamellae as follows (roman numerals = digits; arabic 
numbers = range for subdigital lamellae counted on one of each 
pair of feet for each specimen): forefoot, I 5-7, II 8-11, III 11- 
14, IV ll-16,V7-9;hindfoot, 16-8, II 10-12,111 12-15, IV 16- 
20, V 10-12. 

Distribution 

The new series o{ Macropholidus ruthveni from the Rio Zana 
is from about 175 km south of the type locality and in the south- 
erly adjacent river valley to the Rio Reque system, from which 
the two "Chongollapi" paratypes presumably came (Fig. 2). Other 
species of amphibians and reptiles known from the Rio Zana 
study site, including Stenocercus percultus, S. eunetopsis, Tel- 
matobius latiwstris, and Philodryas simonsi (Cadle, 1991, and 
personal observations), are also known from montane forests of 
the upper Rio Reque. To the extent that the distribution of M. 
ruthveni is representative of the same pattern, Noble's suspicion 
that the two "Chongollapi" paratypes of ruthveni were from higher 
elevations east of Chongoyape seems likely. This species thus 
appears to be another example of a taxon occurring in humid 
montane forest isolates on the western slope of the Andes in 
northern Peru and might be expected in other similar forests of 
this area (see Cadle, 1991:85-89, for discussion and other ex- 
amples). As is the case for Telmatobius latirostris and several 
other elements of the west slope herpetofauna of northern Peru 
(Cadle, unpublished data for the Bosque Monte Seco herpeto- 
fauna), Macropholidus ruthveni also occurs east of the continental 
divide (the type locality is in the upper Rio Huancabamba drain- 
age; Fig. 2). 

If the population represented by KU 220845 proves to be re- 



30 BREVIORA No. 501 

ferable to Macwpholidus ruthveni (see discussion later), then it is 
widely disjunct from the nearest known more northern locality 
(Fig. 2). Moreover, this would imply that the range of ruthveni 
encompasses that of ataktolepis. However, given the fragmented 
nature of habitats for ruthveni in northwestern Peru, any areas of 
sympatry are likely to be quite narrow or restricted to single forest 
isolates (see Montanucci, 1973:20-24, for a similar pattern for 
montane Pholidobolus in Ecuador). 

At the Rio Zana study site, MacrophoUdus ruthveni was col- 
lected between 1,440 and 2,210 m elevation. The only other 
elevational data for the species is Noble's statement (192 la: 139) 
that the type locality was "probably not over 8000 feet [=2,440 
m]." 

Coloration in Life and Preservative 

The following color notes are based on a detailed field descrip- 
tion of FMNH 232602 (female), with supplemental notes from 
35-mm Kodachrome transparencies. The coloration is nearly 
identical to that already described for M. ataktolepis. Dorsum 
medium brown with fine black specks, becoming grayish brown 
toward head, grayish on tail. Lateral surface of body darker brown 
to grayish, more or less abruptly set off from dorsal coloration; 
becoming darker on neck and temporal region. Dorsolateral gold- 
en stripe begins behind eye and fades rather abruptly at approx- 
imately midbody (see Fig. 9). Cream-colored labial stripe begins 
on upper labials and extends to base of forearm. Tail dark gray 
with obscure small dark spots dorsally. Chin white. Belly yellow- 
ish white, golden toward sides. Ventral surface of tail gray with 
darker markings. 

The coloration of MacrophoUdus ruthveni in preservative is 
essentially like that of A/, ataktolepis. The amount of dark ventral 
pigment varies enormously, from almost none to essentially the 
entire venter very dark, in the series available. The chin and 
throat, however, are usually paler than the rest of the venter. The 
dorsolateral stripe in ruthveni occasionally (e.g., ANSP 31764, 
FMNH 232605) continues as a vague discontinuous line extend- 
ing anteriorly along the superciliary scales and canthal region. 
Both ruthveni and ataktolepis have a supralabial stripe extending 
to the anterior margin of the ear. In the Rio Zana specimens of 



1994 NEW TEIID LIZARD FROM PERU 31 

ruthveni, this stripe continues posterior to the ear for some dis- 
tance, and in most specimens extends to the base of the forehmb. 
In the small series of ataktolepis, at most only a few scales pos- 
terior to the ear have whitish pigment, resulting in a line of small 
white dots; in no case do these form a continuous line or extend 
farther than the midpoint between the posterior margin of the ear 
and the base of the forelimb. This pattern difference may prove 
to be a consistent difference between the two species, although 
we hesitate to conclude this given the small samples presently 
available for ataktolepis and differences in preservation between 
the ruthveni and ataktolepis samples. 

Noble (1921a) did not describe the colors in life of the types 
of M. ruthveni in any detail, noting only that the ground color 
was "browner" than the blackish coloration in preservative (see 
Fig. 8). Curiously, he mentioned neither the dorsolateral light 
stripes nor the light supralabial-ear stripe characteristic of the Rio 
Zafla population o{ ruthveni. The state of preservation of the type 
series is such that details of pattern are difficult to discern. How- 
ever, the dorsolateral light stripes are clearly visible in all speci- 
mens of the series (see Fig. 8); they extend from the supraoculars 
and fade on the body behind the shoulder. The supralabial-ear 
stripe is less discernible. It is visible in the holotype as a vague 
light line below the eye; the posterior and anterior extent cannot 
be discerned. In AMNH 38817 it is visible as a broken line 
extending from below the eye to the anterior margin of the ear. 

Habitats and Activity 

Habitats in which Macropholidus ruthveni occurred at the Rio 
Zaiia study site spanned a range: relatively pristine montane cloud 
forest, secondary and disturbed forests, overgrown to relatively 
open cafetals, and open brushy hillsides. All active specimens 
were encountered during the day from midmoming (0945 hours) 
to late afternoon (1700 hours), usually in areas with much leaf 
litter (although they were also found crossing open trails or dirt 
roads). Many specimens also were found inactive during the day 
under rocks (occasionally logs) or, in one instance, under moss 
1.5 m above the ground on a large boulder. Noble (192 la: 139) 
stated that two specimens of the type series were collected under 
flat rocks in a pasture. One specimen from the Rio Zaiia (FMNH 



32 BREVIORA No. 501 

232605) was disgorged by a specimen of the colubrid snake, Coni- 
ophanes longinquus (Cadle, 1989). 

Reproduction and Communal Nesting 

Females of Macwpholidus ruthveni with eggs were collected at 
the Rio Zaiia study site on 1 5-1 8 January and 1 7 June; in addition, 
egg clutches were discovered on 1 7 June (see the following). Five 
females (FMNH 232603, 232608, collected in June; ANSP 3 1 764, 
31767, 31769, collected in January) contained two eggs each; 
ANSP 31765, collected in January, contained one. Hatchlings 
(FMNH 232599-600, JEC 7528; SVL 18-19.5 mm) were col- 
lected on 14 May and 18 June. These observations span the early 
rainy season (January) and early middle dry season (May to June), 
and suggest either a lengthy or multimodal period of reproduction 
in this population. 

Communal nesting is known in a variety of lizards and snakes 
and is probably more widespread than has been reported. The 
only published records for teiids are for the macroteiid Kentropyx 
calcaratus (Magnusson and Lima, 1 984) and the microteiid Neus- 
ticurus ecpleopus (Uzzell, 1966). In addition to observations re- 
ported here for Macropholidus ruthveni, Cadle has observed one 
instance of communal nesting in another microteiid, Proctoporus 
boliviamis, in southern Peru (Upper Rio Santa Maria, Cuzco De- 
partment). The communal nesting habits oi Macropholidus ruth- 
veni were observed at Bosque Monte Seco in the Rio Zafia valley, 
Cajamarca Department, Peru (for discussion of this locality, see 
Cadle, 1989, 1991; Cadle and McDiarmid, 1990). In this case, 
the communal nest of M. ruthveni was coincident with a com- 
munal nest of the colubrid snake Dipsas oreas^ 

These observations were made by Cadle on 17 June 1987 at 
1,490 m elevation at the Rio Zafia study site. On that date, he 



•* This species has been taxonomically confused (see Orces and Almendariz, 1 987; 
Kofron, 1 982) and has not been previously reported from Peru. Specimens recently 
collected by Cadle at several localities in northern Peru, and additional museum 
specimens to be reported elsewhere, are tentatively identified as Dipsas oreas 
pending further study of geographic variation in this complex. These collections 
confirm the species' presence in humid montane forests of the western Andes of 
Peru south to at least the Rio Zafia. 



1994 NEW TEIID LIZARD FROM PERU 33 

observed a Macwpholidiis nithveni crossing a dirt road and seek- 
ing refuge in a hole close to the base of the adjacent road em- 
bankment, which was a mixture of clay and small rocks. Upon 
digging to extract the lizard, a communal nest of this species was 
discovered, and with little further digging several snake eggs and 
egg shells were discovered. The eggs were laid in a crevice (1-3 
cm wide and 15-20 cm in vertical dimension) beneath a loose 
conglomerate of flaky rocks and clay. The crevice was lined mostly 
with moist clay mixed with some gravel. Lizard eggs were found 
between 5 and 30 cm from the entrance to the crevice. The snake 
eggs were found between 20 and 30 cm from the entrance and 
were intermingled with the lizard eggs. Although eggs were found 
up to 30 cm from the opening, since the crevice was oriented 
roughly parallel to the face of the roadcut, the deepest part of the 
crevice was only about 1 5 cm from the surface of the soil. Because 
the road embankment faced roughly eastward and was not cov- 
ered or overshadowed by dense vegetation, the soil at this site 
probably would have been warmed daily by the morning sun. 

Both the microteiid and the snake eggs were apparently of vary- 
ing ages (minimally, two viable snake clutches were present, plus 
one recently hatched clutch). The remains included some micro- 
teiid eggs that were already well decomposed, other more recent 
shell remains, and several unhatched eggs. A total of 220 micro- 
teiid shell remains were found (this is a minimum count), plus 
eight viable eggs, one of which hatched the next day (FMNH 
232599; SVL 18 mm). The first snake eggs encountered included 
five empty shells together in a group; a bit farther in were three 
additional shells that looked roughly the same age as the first five 
and might have been part of the same clutch. A second clutch 
included seven viable eggs. The deepest clutch included five viable 
eggs plus one fungus-ridden egg. These two latter clutches had 
embryos of slightly different ages when one egg of each was opened 
on 18 June (the deepest clutch had a smaller embryo).^ Clearly 



' Five of the snake eggs were transported to Lima on 27 June, where they were 
kept in moist paper towels in a plastic bag while fieldwork was completed elsewhere 
in Peru. They were then transported back to the United States, where they hatched 
between 23 September and 1 October. Egg-laying in subterranean retreats may 
seem an unusual behavior for "arboreal" snakes such as Dipsas oreas. Although 



34 BREVIORA No. 501 

the microteiid nest represents a true "communal" nest, with many 
females contributing eggs. The same is probably true for the snake 
clutches, although improbably they could represent successive 
clutches of a single female. 

Notes on KU 220845, Tentatively Referred to 

Macropholidus ruthveni 

KU 220845 was collected at Chaclacayo, Lima Department, 
Peru, 11°59'S, 76°46'W, by M. Urbina, date unknown. This lo- 
cality is approximately 620 km south of the nearest known locality 
for M. ruthveni (Rio Zaiia valley; see Fig. 2), but a series of 
specimens from Chaclacayo is apparently now available and will 
be reported on elsewhere by Antonio W. Salas and his colleagues 
(personal communication). Measurements of the KU specimen 
are as follows: SVL 31 mm, HL 7.21 mm, HW 4.5 mm, HD 3.25 
mm, BL 15.9 mm, and TL 38 + 2 mm. Meristic counts (e.g., 
Table 2), the arrangement of head plates and body scales, and 
aspects of pattern of KU 220845 are within the range of those 
already reported for the types and Rio Zaiia samples of M ruth- 
veni. However, the specimen is unusual in the following features: 

(1) the loreal scale is completely fused with the posterior nasal 
scale on both sides in KU 220845, whereas in all other specimens 
o{ ruthveni examined, the loreal is present as a discrete scale; and 

(2) the paired series of enlarged medial dorsal scales are incom- 
plete and irregular on the posterior part of the body. There is a 
suggestion that scales of the temporal region in KU 220845 are 
broken up more than in the other specimens o{ ruthveni, but this 
difference is subtle and not reliable on the basis of the single 
Chaclacayo specimen we examined. 



the species of Dipsas are usually regarded as highly arboreal, Dipsas oreas at the 
Rio Zaiia site and other sites in northwestern Peru (personal observations; see 
also Orces and Almendariz, 1987) spends its inactive diurnal period in retreats 
within or under surface objects on the ground, or in holes in the ground, and 
ascends vegetation at night. Observations on Dipsas peruana at several sites in 
southern Peru suggest a similar behavior pattern. Orces and Almendariz (1987) 
reported a clutch of seven eggs of D. oreas beneath decomposing humid logs in 
Ecuador. 



1994 NEW TEIID LIZARD FROM PERU 35 

The second characteristic requires further comment, as it ap- 
proaches the condition in M. ataktolepis. The specimen has a 
total of 34 medial dorsal scales between the occipitals and the 
posterior margin of the hindlimb, which is typical for the other 
ruthveni samples (Table 2). However, each of the enlarged rows 
is disrupted by the intercalation of small scales. That is, neither 
of these rows is continuously enlarged from the occipitals to the 
tail base. There are 14-16 enlarged medial dorsals anteriorly, 
followed by additional medial dorsals of varying sizes, some ap- 
proximately the same size as the "enlarged" anterior scales, others 
much smaller. Furthermore, the larger and smaller of these pos- 
terior scales are intermixed within each medial dorsal row and 
have no consistent arrangement between the two rows. This con- 
dition is similar to that in ataktolepis, but in available specimens 
of that species the size reduction occurs only once, and small 
scales then continue to the tail base. 

Given the unusual features of KU 220845 relative to other 
Macwpholidus ruthveni we have examined, we defer an opinion 
about the taxonomic status of this population, which is currently 
under study by Antonio W. Salas and his colleagues. However, 
in addition to being widely disjunct from the other localities known 
for ruthveni, it is worth noting that Chaclacayo is about 500 m 
lower in elevation (889 m; Stiglich, 1922) than any other ruthveni 
localities, and the region is presently much more arid than the 
habitats at the other known localities for ruthveni.^ Details mi- 
crohabitat information and study of larger samples from this pop- 
ulation should help clarify its systematic status. 



* In this context, the uncertain origin of the "Chongollapi" paratypes of M. ruthveni 
may be significant. Chongollape is a town of the coastal desert, and Noble only 
suspected that the paratypes came from higher, more mesic environments farther 
inland. We agree with this inference. Nevertheless, isolated populations o{ ruthveni 
or related species may exist in pockets of mesic or riparian habitats in the Peruvian 
coastal region. Well-documented specimens will be necessary to verify this pos- 
sibility, which, in any event, we consider unlikely. However, species of the pre- 
sumed closest relative of Macropholidus, Pholidoholus, inhabit mesic to xeric 
montane habitats in Ecuador (Montanucci, 1973:21); given the few documented 
localities for M. ruthveni, it may be premature to draw general conclusions about 
habitat specificity in this species. 



36 BREVIORA No. 501 

STATUS OF MACROPHOLIDUS VIS-A-VIS 
PHOLIDOBOLUS 

Noble (192 la: 137) noted one point of similarity (lack of pre- 
frontal scales) and one point of difference (presence of lateral scales 
of reduced size in Pholidobohis\ uninterrupted lateral scales in 
Macwpholidus) between Macropholidus and Pholidobohis. Spe- 
cies of Pholidobolus described subsequent to Noble's paper (Mon- 
tanucci, 1973) showed that both characters are variable within 
Pholidobolus. With the discovery of Macropholidus ataktolepis, 
prefrontal scales are now known to be variably present in Macro- 
pholidus. Of the 16 nonosteological/nonhemipenial features used 
by Montanucci (1973:31) to define Pholidobolus, the two species 
of Macropholidus share all but the weakly keeled to striated dorsal 
scales. Dorsal scales of the tail in both species of Macropholidus 
may, however, be weakly striated. We have seen neither osteo- 
logical nor hemipenial material of Macropholidus. 

Macropholidus is further distinguished from all species of Phol- 
idobolus by the two parallel series of enlarged dorsal scales, which 
are foreshortened in M. ataktolepis, and by having a translucent 
disk in the lower eyelid (but this feature is present in most spec- 
imens of Pholidobolus annectens\ Montanucci, 1973:5). The two 
species of Macropholidus are smaller than any of the described 
species of Pholidobolus (maximum SVL for the largest specimens 
of M. ruthxeni and M. ataktolepis, both females, 45.5 mm and 
39 mm, respectively; maximum SVL for females of Pholidobolus 
>56 mm for all species [Montanucci, 1973]). 

Macropholidus and Pholidobolus share one apparently derived 
scutellational feature: the presence of two median rows of enlarged 
gular scales (Kizirian and Coloma, 1991:420; this character is 
shared also with Prionodactylus). Thus, if this feature proves in- 
dicative of a close relationship between Pholidobolus and Macro- 
pholidus, as seems likely given their shared character suites and 
geographical distributions (see Cadle, 1991:85-89), then the en- 
larged dorsal rows characteristic of Macropholidus may be simply 
autapomorphic for these two species, and Pholidobolus may be 
paraphyletic with respect to Macropholidus (a similar interpre- 
tation of the lower eyelid disk might be possible, pending its 
eventual interpretation in P. annectens). Phylogenetic studies of 



1994 NEW TEIID LIZARD FROM PERU 37 

the broader relationships among microteiids should clarify this 
issue. 

ACKNOWLEDGMENTS 

Our work at Bosque Cachil has been facilitated and supported 
by Abundio Sagastegui of the Universidad Antenor Orrego de 
Trujillo, whose help in so many other ways is also appreciated. 
Several great field companions have helped us at Cachil and at 
Monte Seco: Jose Guevara, Segundo Leiva, Pedro Lezana, Raul 
Quiroz, Jose Santisteban, and Helena Siesniegas. Cadle's field- 
work at Cachil was supported by the American Philosophical 
Society and at the Rio Zana by the Field Museum of Natural 
History; the fieldwork was also partially supported by NSF BSR 
84-00166. The support of the Ministerio de Agricultura, Direc- 
cion General Forestal y de Fauna, the Museo de Historia Natural 
de San Marcos (Lima), and the people of Monte Seco is greatly 
appreciated. M. O. Dillon (FMNH) encouraged our work, shared 
his botanical knowledge of western Andean forests and an un- 
published manuscript, and clarified some botanical information. 
Antonio W. Salas provided information about the Chaclacayo 
specimen of M ruthveni. During most of the 1991 fieldwork, 
Cadle was accompanied by Rosa Ortiz, Camilo Diaz, and Alwyn 
Gentry, whose tragic early death deprived us of one of the world's 
great tropical biologists. E. E. Williams and M. Henzl assisted 
with German translations, and R. F. Inger (FMNH), C. W. Myers 
(AMNH), and J. E. Simmons (KU) loaned us specimens. Laszlo 
Meszoly prepared Figures 3 and 5. David Kizirian and an anon- 
ymous reviewer provided many useful comments that helped us 
clarify the manuscript. 

LITERATURE CITED 

Barbour, T., and G. K. Noble. 1920. Some amphibians from northwestern 
Peru, with a revision of the genera Phyllohates and Telmatobius. Bulletin of 
the Museum of Comparative Zoology, 63(8): 395-427. 

Cadle, J. E. 1989. A new species of Coniophanes (Serpentes: Colubridae) from 
northwestern Peru. Herpetologica, 45: 41 1-424. 

. 1 99 1 . Systematics of lizards of the genus Stenocercus (Iguania: Tropi- 

duridae) from northern Peru: New species and comments on relationships 
and distribution patterns. Proceedings of the Academy of Natural Sciences 
of Philadelphia, 143: 1-96. 



38 BREVIORA No. 501 

Cadle, J. E., AND R. W. McDiARMiD. 1990. Two new species of Ce«fro/^/7W/<3 

(Anura, Centrolenidae) from the western slope of the Andes in northern Peru. 

Proceedings of the Biological Society of Washington, 103: 746-768. 
Dillon, M. O., A. Sagastegui, I. Sanchez, S. Llatas, and N. C. Hensold. 

1994. Floristic inventory and biogeographic analysis of montane forests in 

northern Peru. Memoires of the New York Botanical Garden, in press. 
Donnelly, M. A., R. W. McDl\rmid, and C. W. Myers. 1992. A new lizard 

of the genus Arthrosaura (Teiidae) from southern Venezuela. Proceedings of 

the Biological Society of Washington, 105(4): 821-833. 
Duellman, W. E., and E. R. Wild. 1993. Anuran amphibians from the Cor- 
dillera de Huancabamba, northern Peru: Systematics, ecology, and biogeog- 

raphy. Occasional Papers of the Museum of Natural History, University of 

Kansas, 157: 1-53. 
KiziRiAN, D. A., and L. a. Coloma. 1 99 1 . A new species of Proctoporus (Squa- 

mata: Gymnophthalmidae) from Ecuador. Herpetologica, 47(4): 420^29. 
Koepcke, H. W. 1957. Uberdie Walderanderwestseitederperuanischen Anden 

und ihre tiergeographischen beziehungen. Verhandlungen Deutschen Zool- 

ogische Gesellschaft, 1957: 108-119. 
. 1961. Synokologische studien an der Westseite der peruanischen Anden. 

Bonner Geographische Abhandlungen, 29: 1-320. 
Koepcke, H. W., and M. Koepcke. 1958. Los restos de bosques en las vertientes 

occidentals de los andes peruanos. Boletin del Comite Nacional para la 

Proteccion de la Naturaleza, Lima, 16: 22-30. 
Koepcke, M. 1954. Corte ecologico transversal en los Andes del Peru central 

con especial consideracion de las aves. Parte I: Costa, vertientes occidentales 

y region altoandina. Memorias del Museo de Historia Natural "Javier Prado," 

3: 1-119. 
. 1961. Birdsofthewestemslopeofthe Andes of Peru. American Museum 

Novitates, 2028: 1-31. 
KoFRON, C. P. 1982. The identities of some dipsadine snakes: Dipsas elegans. 

D. ellipsifera and Leptognathus andrei. Copeia, 1982(1): 46-51. 
Magnusson, W. E., AND A. P. Lima. 1984. Perennial communal nesting by 

Kentropyx calcaratus. Journal of Herpetology, 18: 13-75. 
MoNTANUcci, R. R. 1973. Systematics and evolution of the Andean lizard genus 

Pholidobolus (Sauria: Teiidae). Miscellaneous Publications, Museum of Nat- 
ural History, University of Kansas, 59: 1-52. 
Noble, G. K. 1921a. Some new lizards from northwestern Peru. Annals of the 

New York Academy of Sciences, 29: 133-139. 

. 1921b. A search for the marsupial frog. Natural History, 21: 474-493. 

Orces, G., AND A. Almendariz. 1987. Sistematica y distribucion de las ser- 

pientes dipsadinae del grupo oreas. Politecnica (Revista de Informacion Tec- 

nico-Cientifica, Quito), 12(4): 135-144. 
Parker, H. W. 1930. Two new reptiles from southern Ecuador. Annals and 

Magazine of Natural History, ser. 10, 5: 568-571. 
Parker III, T. A., T. S. Schulenberg, G. R. Graves, and M. J. Braun. 1985. 

The avifauna of the Huancabamba region, northern Peru, pp. 169-187. //; 



1994 NEW TEIID LIZARD FROM PERU 39 

P. A. Buckley, M. S. Foster, E. S. Morton, R. S. Ridgely, and F. G. Buckley 

(eds.). Neotropical Ornithology. Washington, D.C., American Ornithologists' 

Union. 
Peters, J. A. 1964. Dictionary of Herpetology. New York, Hafner Publishing 

Company, ix + 392 pp. 
Presch, W. 1 980. Evolutionary history of the South American microteiid lizards 

(Teiidae: Gymnophthalminae). Copeia, 1980: 36-56. 
Sagastegui, a., and M. O. Dillon. 1 99 1 . Inventario preliminar de la flora del 

Bosque Monteseco. Amaldoa, 1(1): 35-52. 
Smith, H. M. 1946. Handbook of Lizards, Lizards of the United States and 

Canada. Ithaca, New York, Comstock Publishing Co., xxi + 557 pp. 
Stiglich, G. 1 922. Diccionario Geografico del Peru, 2 vols. Lima, Torres Aguirre, 

1,193 pp. 
UzzELL, T. M. 1966. Teiid lizards of the genus Neusticwus (Reptilia, Sauria). 

Bulletin of the American Museum of Natural History, 132: 277-328. 



NOTES ADDED IN PROOF 

1 . Antonio W. Salas (personal communication) provided additional details con- 
cerning the Chaclacayo (Lima Department) population of Macropholidus mthveni 
(p. 34). Specimens are known only from an artificial forest created in association 
with a resort establishment by plants imported from many other places and main- 
tained by aggressive cultivation in this otherwise arid locale. Salas believes, and 
we concur, that the mthveni population at Chaclacayo very likely resulted from 
introduction with these imported plants. If so, this would remove the distributional 
and microenvironmental anomaly produced by the record. 

2. We recently collected specimens of A/, mthveni (to be deposited in MCZ and 
in the Museo de Historia Natural de la Universidad Antenor Orrego) from the 
vicinity of Sangal, Cajamarca Dept., Peru, 2000 m [07°08' S. 78°50' W], approx- 
imately 48 km SE (airline distance) of the Monte Seco population of this species; 
the new locality represents the southernmost population known for mthveni. Spec- 
imens were collected under rocks in disturbed habitats (pastures, agricultural land, 
and brushy hillsides). Significantly, the Sangal population is only about 30 km 
north across the broad, dry valley of the Rio Chilete from the only known pop- 
ulation of ataktolepis at Bosque Cachil. Thus, the distributions of the two species 
may be parapatric at the boundary formed by the Rio Chilete, or the ranges of 
the two species may approximate one another at appropriate elevations around 
the headwaters of this river (assuming adequate habitats remain in this highly 
populous region). Conceivably, the two species could even be sympatric in as yet 
undiscovered populations in this area. 



8 



B R E V I O E A 



n/mT 



oseum of Comparative Zoology 

lAM ^ 1995 



us ISSN 0006-9698 



Cambridge, Mass. 10 January 1995 , Number 502 
Hai"< v/ 



UNIVER: 



>i 1 I 



A COMPUTER APPROACH TO THE COMPARISON AND 

IDENTIFICATION OF SPECIES IN DIFFICULT 

TAXONOMIC GROUPS* 

Ernest E. Williams,'^ Hugh Rand,^ A. Stanley Rand,^ 
AND Robert J. CHara"* 

Abstract. A computer program for the identification of unknown taxa in 
"difficult groups" based on matching rather than sequential exclusion is proposed 
as a substitute for both the conventional dichotomous key and for the random 
entry matrix that has been suggested as a replacement for the dichotomous key. 
The matching program is modeled after the steps that a practicing taxonomist 
would employ in the identification of an unknown specimen: 

1. Data for the unknown are compiled. 

2. Data for the unknown are compared with those of relevant known taxa. 

3. On the basis of the comparison, certain of the named taxa are considered 
possible matches with the unknown. 

4. Final choice of the named taxon best matching the unknown is made and 
confirmed from additional data. If there is no match, the possibility of an 
undescribed species must be confronted. 



* Apple, HyperCard, and Macintosh are registered trademarks of Apple Computer, 
Inc. Turbo BASIC is a registered trademark of Borland International, Inc. IBM 
is a registered trademark of International Business Machines Corporation. MS- 
DOS is a registered trademark of Microsoft Corporation. 

' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 

- Department of Applied Mathematics, University of Washington, Seattle, Wash- 
ington 98195. 

' Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa, Republic of 
Panama. 

^ Center for Critical Inquiry in the Liberal Arts, University of North Carolina at 
Greensboro, Greensboro, North Carolina 21412-5001. 
' To whom reprint requests should be addressed. 



2 BREl lORA No. 502 

The program enables a computer to follow these steps: 

1 . It first enters the states of selected characters for the unknown individual or 
series. For each character of the unknown individual or series, two numbers 
are recorded, a minimum and a maximum; in the case of the series allowing 
for variation within the sample, and in the case of the individual allowing for 
differences between its two sides andaho obscurities and ambiguities in counts 
or coding. 

2. Once these data are entered, the program compares them against the ranges 
recorded in a reference matrix for selected known species. The user specifies 
the maximum number of characters in which a taxon in the reference matrix 
is allowed to differ from the unknown before being accepted as a "match." 

3. The resulting report lists all the matching taxa and, for each, the number of 
characters not matched, the specific characters not matched, and by how much. 

4. Included in the report for each matching taxon is a "descriptor" that cites 
characters that are not coded for the computer as well as characters regarded 
as "diagnostic" of the taxon. The descriptors assist in the final choice of the 
most plausible identification for the unknown. In certain cases (e.g., a new 
taxon), evaluation of the descriptors may require the user to reject all matches. 

While the program was inspired by problems encountered during exploration 
of the systematics of anoline lizards, it does not deal with phylogeny at all. It is 
only— in our eyes— a better substitute for the dichotomous key. It aids in the 
identification of animals. As such, it has been customized for the anoles. The 
reference matrices, character descriptions, and "descriptors" provided as examples 
in the second and succeeding sections of this paper are for anoline lizards only. 
The concept of the matching program is, however, applicable to taxa of any sort. 



I. HISTORICAL INTRODUCTION 
Ernest E. Williams 

Some years ago, while visiting Stanley Rand in Panama, I won- 
dered aloud whether and to what extent computers could be used 
in species identification. I expressed dissatisfaction with the com- 
puter keys I knew about and with the usual dichotomous keys 
employed in taxonomic works in general. The dichotomous key 
did not at all resemble the process by which the working taxon- 
omist, engaged, for example, in revising a large and complex 
genus, would actually employ to distinguish the species. 

In my own work on anoline lizards, I had grown into the habit 
of using a standard data sheet for almost every anole specimen 
that I encountered, whether I recognized the species on sight or 
not, and very definitely for any specimen that I did not recognize 



MCZ 
LIBRARY 

1994 COMPUTER IDENTIFICATION OF SPECIES JAN 2 33 I9V5 

with complete certainty. I routinely recorded museum number 
and locality and then a set of about 25 characters, some of them '^^ 
counts, sometimes brief descriptions, and, last of all, some very '' ' ' 
brief remarks on color as preserved. I used these data sheets to 
compel myself to always take the same data on all specimens and 
to standardize my information with regard to species that I did 
not know well. In the case of West Indian species, which ordinarily 

I knew in life, color, shape, and some scale differences that seemed 
diagnostic were the way that I distinguished species, but with 
mainland species that I knew only as museum specimens— and 
furthermore as specimens preserved, as the older specimens usu- 
ally were, in ways that minimized color distinctions, and having 
scale characters that I had little acquaintance with— my data sheets 
seemed the only way to go. 

My method of comparison for these unfamiliar species and 
characters was, however, primitive: a random, intuitive search 
for matching characters and populations. It seemed to me that 
there must be a way to automate the comparisons— the match- 
ing—that I was already doing by hand and that computers must 
be the best way to do this. 

Not all of this do I know to have been said at one time, or even 
in one place, but it is certain that at one supper hour Stan Rand 
agreed with my general philosophy and, with his son Hugh's 
concurrence, voiced the opinion that a computer matching pro- 
gram of the sort that I imagined was feasible. By my visit the 
next year, Hugh Rand had written a Basic program for the Apple 

II computer that was the first version of the system described 
here. I provided the first reference matrix, one for Puerto Rican 
anoles. Stanley Rand emended the Apple II version and then 
created an IBM PC version, for which I have added matrices for 
Panamanian, Costa Rican, and Ecuadorian anoles to the one for 
Puerto Rico. This version is used at the Smithsonian Tropical 
Research Institute. 

Much later, when Robert O'Hara was my assistant and es- 
pecially my advisor-expert for the Macintosh computer, he in- 
troduced me to HyperCard, and using that very special application 
he created for me the Macintosh version that is also the database 
on which I work and from which I furnish the reference matrices 
used by the IBM PC. 



4 BREVIOK4 No. 502 

II. A COMPUTER MATCHING PROGRAM FOR 
USE AS AN IDENTIFICATION KEY 

Ernest E. Williams, Hugh Rand, and A. Stanley Rand 

Difficult taxa are those for which none but the speciahst can 
identify species with any confidence, and the speciahst not without 
tremors of unease. They are typically large genera in which species 
distinctions are subtle or ambiguous. They may even be relatively 
easy to identify in one sex or under special conditions. Typically 
the species list is still very incomplete, and the possibility of 
encountering a new species is still uncomfortably high. These are 
taxa for which the dichotomous key may be as often a trap as an 
aid. It is our thought that a computer key of a novel type will be 
able to help where conventional dichotomous keys fail. 

It was recognition of the difficulties associated with certain taxa 
that led Peters and Orejas Miranda (1970) and Peters and Donoso 
Barros (1970), in the two- volume Catalogue of the Neotropical 
Squamata, to present character matrices instead of dichotomous 
keys in three cases (the snake genera At r actus and Bothrops in 
Volume 1, and the lizard genus Anolis in Volume 2) and to ad- 
vocate computer sorting. It is interesting to quote in full their 
reasons for doing so (pp. vi-vii, identically paginated in the snake 
and lizard volumes): 

"Most of the keys presented here are the standard dichot- 
omous type .... In the case of very large genera, however, 
we have introduced a different concept. Any attempt at writ- 
ing keys for poorly known large genera is likely to be futile, 
and we have avoided this by presenting as much data as 
possible in the form of a matrix. This permits 'random entry' 
identification, for the user, in the matrix that he wishes to 
check and eliminates all taxa that do not possess that char- 
acter, finally arriving at a considerably reduced number of 
taxa (hopefully only one) after checking a series of characters. 
This concept has formed the basis of computer identification, 
since the machine can do such sorting more rapidly and 
efficiently than the human, and the random entry matrices 
presented here are organized in such a way that they can be 
incorporated in the computer programs now available for 
such machine sorting. It is our assumption that this method 



1994 COMPUTER IDENTIFICATION OF SPECIES 5 

of identification will be used more and more in the future, 
and we hope that presenting these matrices will encourage 
others to begin to organize their data similarly, thus antici- 
pating the day when computer keys are available to all users." 

The matrices provided by Peters, Orejas Miranda, and Donoso 
Barros partly parallel the procedure we shall advocate below, but 
to our knowledge they have received little use. The sampling for 
the contained species was inadequate, a major problem for genera 
in which high individual variability is a characteristic. The data 
for our own matrices were collected from 20 specimens whenever 
possible and as many as are available in all other cases. 

More fundamental, however, is a conceptual difference in meth- 
od between the Peters, Orejas Miranda, and Donoso Barros ap- 
proach and our own. Their method (and Peters advocated similar 
techniques in other papers; see Peters and Collette, 1968; Morse 
et al, 1971) was to sort by sequential exclusion: i.e., by the fun- 
damental technique of the dichotomous key, but here pursued by 
"random entry" with the aid of a computer. The goal of such a 
method is elimination of all but one of the possible choices, and 
no individual of a species is allowed to have the alternative of 
the successively chosen character. Our technique is the inverse 
of this: matching, instead of exclusion. Variation and overlap are 
expected, and the goal is maximum congruence, rather than elim- 
ination. 

We have called our system a "key." It is certainly not a con- 
ventional "key," and, although it will assist in the identification 
of species, it is not at all comparable to a conventional dichot- 
omous key. Instead, it is a computer-assisted procedure delib- 
erately modeled after the steps that a taxonomist would employ 
in attempting the assignment of new specimens to the recognized 
taxa for which no dichotomous key existed. No taxonomist re- 
viewing a large snake genus, for example, would begin by relying 
on the dichotomous keys prepared by his or her predecessors. 
The taxonomist would begin by collecting data— data on the num- 
ber and, perhaps, shape of head scales; on the number of dorsal 
rows and ventrals; on the pattern on the head, body, and the tail; 
on sexual dimorphism, locality, and habitat; etc. Thereafter, the 
procedure would be to associate populations of phenetically sim- 



6 BREVIORA No. 502 

ilar animals, animals similar in detail, even if not identical, since 
nothing biological is exactly similar. The taxonomist would, in 
fact, be matching individuals and populations, determining in 
what regards and by how much the members of selected species 
might differ. Judgment would necessarily be involved. The ex- 
pertise for which the experienced taxonomist is known is neces- 
sarily a familiarity gained over time with the chosen taxon. During 
this learning period, there grows an appreciation of which char- 
acters are most meaningful for species distinction or, at the least, 
the most readily determinable without major error. There comes 
also an appreciation of the kind and extent of the variation that 
seems to accord with the gaps that separate some underlying 
biological realities that are called "species." 

The steps that the practicing taxonomist would use in the iden- 
tification of a series of some unknown taxon for which there 
happened to be no conventional key would certainly be the fol- 
lowing: 

1 . On the basis of prior experience, preliminary investigation, 
and/or literature search, characters are selected to be routinely 
checked in all specimens. 

2. A table or data matrix is prepared recording the ranges of the 
states of the chosen characters in the named taxa to be com- 
pared. 

3. The new specimens are examined for as many as possible of 
the chosen characters. 

4. The table or data matrix is searched for matches and mis- 
matches with the characters of the new specimens. 

5. Tentative assignment of specimens to recognized taxa is made 
on the basis of closeness of match. 

6. The tentative taxonomic assignment is confirmed or rejected 
with the aid of additional characters, whether from the liter- 
ature or from previously determined specimens. 

Our computer "key" breaks this procedure into two parts. 

The first part, corresponding to steps 1 through 5, depends on 
reference matrices that contain codings for qualitative character 
states or standard counts, such as those that are (or should be) 
taken routinely on any specimen of the relevant taxa. These counts 



1994 COMPUTER IDENTIFICATION OF SPECIES 7 

or character states are stored and compared by the computer with 
the data entered for specimens or series to be identified. 

The second part, corresponding to step 6, is a supplementary 
set of species descriptors (automatically provided in the computer 
search report) that highlights the most valuable diagnostic features 
of each species and reports its known geographic range, ecology, 
behavior, or other significant features such as color in life. 

Together these two data sets are intended to permit the "keying 
out" of all species presently known for any taxonomic group from 
any studied area. Importantly the program has three especially 
useful capabilities: 

1 . The program will report not only those species in the reference 
matrices that match the specimens or series under examination 
but also those that are similar. The computer will, in fact, ask 
the user of the key by how many characters the unknown may 
be allowed to differ. 

2. The program will report how many and which of the characters 
do not match the coded data for species in the reference ma- 
trices and by how much they differ. 

3. The program can be used to discover shared characters— 
whether plesiomorphic or synapomorphic— or combinations 
of shared characters of either kind simply by employing as the 
unknown the relevant character state or states and searching 
for those taxa that match. 

More importantly, however, our computer key contains pro- 
visions for its own improvement. Built into the program are pro- 
cedures such that both the reference matrices and the descriptors 
may be modified or expanded to include new or more useful 
characters or newly discovered species or species of other regions. 

When new species are discovered, a dichotomous key can be 
very hard to change in ways that accommodate the new data. 
Often old dichotomies no longer work. The addition of the new 
species may require that the key be rewritten entirely, and this 
may be a job that compels reexamination of all the taxa involved. 
In contrast, in our program data matrices for the new species can 
easily be constructed and added to the reference set with no changes 
to the program itself or to the other reference matrices. 



8 BREVIORA No. 502 

It is to be emphasized that this system routinely provides for 
variation and ambiguity. The expectation of variation is, in fact, 
built into the system. All character states or counts must be spec- 
ified by two numbers. If the character state or count is mono- 
morphic for any species, this fact is entered in the matrix by simple 
repetition of the coding number (e.g., 1,1 or 3,3). If there is vari- 
ation between the two sides of an animal or within a population 
or if there is intermediacy or ambiguity in the assessment of a 
character state or count, then the two appropriate extreme num- 
bers should be entered (e.g., 1,3 for a character state or 16,19 for 
a count). 

There are caveats that need mention. Several negative features 
of our "key" are intrinsic to its concept and therefore irremediable 
without discarding the concept. These are as follows. 

1 . The computer accepts as a match the minimum or the max- 
imum recorded in the matrix or any number that falls between 
these extremes. This results in three problems. 

First, in the three characters that involve enlarged scales (char- 
acters 7, 14, and 21), coding has to deal with conditions ranging 
from (no scales enlarged) to a condition in which there is a 
series of gradually enlarging scales that must be coded arbitrarily. 
We have chosen to code such gradually enlarging scales as 30,30 
or 50,50. This means that any species or series of an unknown 
where some individuals have no enlarged scales and others have 
many very slightly enlarged scales, or any unknown in which the 
choice between these two is ambiguous might be coded 0,30. Any 
species with this coding in the reference matrix will be matched 
by any unknown with an intermediate coding for this character, 
and any unknown with this coding will match all species in the 
reference matrix on this character; i.e., there will be many false 
matches. The descriptors should resolve this difficulty. We advise 
the user to treat matches that involve these enlarged scale char- 
acters with caution. 

Second, many species also have overlapping ranges for indi- 
vidual characters but, although the means may differ greatly, the 
computer will, of course, accept as a match to both species any 
value that falls within the zone of overlap. Because of this aspect 
of the program, individuals will often be unambiguously identi- 
fied, whereas series, even when subjectively the same species, may 



1994 COMPUTER IDENTIFICATION OF SPECIES 9 

not be. It must also be stressed emphatically that a report of zero 
difference by our program is not a statement of species identity. 
It is only a statement of zero difference in the chosen characters. 

Third, in some cases zero differences appear to indicate true 
relationship, but many times species that are judged to be remote 
from one another on the characters not in the reference matrix 
are reported by our program as close or as exhibiting zero differ- 
ences. 

Remedies: In all cases of ambiguity on the initial run-through, 
recourse must be had— special consideration must be given to the 
descriptors, the second feature of our program, which cite such 
features as locality or color that are not readily computer-codable 
and that highlight special morphological, behavioral, or ecological 
traits. 

Often the descriptors will eliminate taxa matched by the com- 
puter from further consideration. The excluded species may differ 
radically from the unknown in color or shape or in some other 
way not coded in the reference matrix. Alternatively, the descrip- 
tors may afford clear support for an identification by calling at- 
tention to or emphasizing very evident characters of the un- 
known—such as color, shape, or some feature special to the species. 
In any case, both tentative mismatches or matches should always 
be corroborated by examination of previously identified speci- 
mens and/or a check of the pertinent literature. 

2. Instead of matching too many species, the unknown will 
sometimes not match any closely. If among the reference matrices 
there are unique types or small series, there will ordinarily be a 
failure to perfectly match a specimen or small series that, in the 
judgment of the inquiring systematist, should be conspecific. This 
is expectable. No unique biological specimen is ever exactly 
matched by another. In consequence, if any species is described 
from a single specimen, the identification of the next specimen 
is always a matter of extrapolation. A type or any unique specimen 
may set the universe of discourse, but it does not bound it. A 
type is an example only; it necessarily does not and cannot exhibit 
the range of variation that is contained in the population of which 
it is a member. 

Remedies: In cases of failure to match any descriptor, the very 
real possibility of a new species will have to be confronted. Here, 



10 BREVIORA No. 502 

as in the case of too many matches, a conservative approach is 
appropriate. 

Special consideration should be given in both the cases of too 
many matches and of no match at all to unique features of the 
unknown, whether it is an individual or a series. Decision, in any 
case, depends on differences between populations. A unique type 
or potential type needs to be compared with populations, opti- 
mally with relevant sympatric or adjacent populations. If the 
characters of any unique type or potential type fall within the set 
that is characteristic of a well-sampled population, and the de- 
scriptors do not exclude it, synonymy is the first and most prob- 
able decision. We say only probable decision, because, before any 
decision, every kind of error must be ruled out. In all cases, the 
characters reported in the descriptors need to be carefully scru- 
tinized—in particular color, pattern, and geography, including 
sympatry or near sympatry, and habitat. 

3. It will never be possible to code all characters of possible 
taxonomic significance. Some, like color, pattern, or body shape, 
are difficult to code for computer use unless broken down into 
fine details. Every taxon will have its special features, which may 
often be more readily expressed verbally rather than in computer 
code. Some of these features may be unique and make the species 
instantly identifiable. (This is, in fact, a danger, because unique 
and easily recognizable characters may often conceal even species 
differences among populations so easily recognized by one char- 
acter that no further scrutiny is given them.) 

The flexibility of our program should be strongly emphasized. 
It permits the distribution of single character states or of special 
combinations of character states to be tracked through entire taxa 
or subsets of them, making possible tests of the validity of old 
taxa or providing evidence for the erection of new taxa. In general, 
the system here presented will prove readily adaptable internally 
to individual needs. Certainly the system itself can be modified 
or superseded. 

Some final statements should be made. First, our program pro- 
vides no escape from matters of judgment. In the characters em- 
ployed as well as the species recognized, human judgment has 
entered in. The program can only mechanically sort and match 
what human judgment has given it. The program will not, in any 



1994 COMPUTER IDENTIFICATION OF SPECIES II 

genuine sense, solve any taxonomic problems; it can, however, 
serve the purpose of calling attention to and exploring problems. 
One of us (EEW) now routinely uses it for the exploration of 
problems in the systematics of anoline lizards. 

Second, it is important to emphasize that our program does 
not deal with phylogeny; it is a phenetic program, dealing with 
overall similarity. Homoplasy may often be a reason that species 
estimated to be remote phyletically may be reported with zero 
difference by our program. In such cases, rather than discovering 
true relationships, it discovers ecomorphs (Rand and Williams, 
1969; Williams, 1972, 1983). However, it may also reveal the 
often subtle phyletic constraints that require differences between 
ecomorphs. 

Finally, in the subsequent sections of this paper, the single 
example provided is one herpetological taxon, the anoline lizards. 
This is only an historical accident. The idea of a matching key 
arose, as recounted in Section I, among students of these peculiar 
and special animals. No feature of the fundamental concept of 
our system restricts it at all to this specific taxon. Other taxa and 
other environments may use our system. 

For example, Knowlton (1993) has emphasized the abundance 
of sibUng species (^"difficult" genera) in marine environments. 
She cited this (p. 189) as due to "both inadequate study of mor- 
phological features of living organisms ('pseudo-sibling species') 
and divergence in habitat, life history, and chemical recognition 
systems without parallel divergence in morphology." 

There is no reason not to extend this generalization to terrestrial 
systems. One of us (EEW) has been very aware in anoline lizards 
of "inadequate morphological study," including very inadequate 
and very incomplete descriptions of morphology. (The section in 
this paper that deals explicitly with anoline morphology is an 
attempt to describe some of the characters that should routinely 
be considered in every description of an anoline lizard. More can 
and should be added.) EEW has also been aware of habitat, life 
history, and visual (color and behavioral) recognition systems in 
anole lizards. 

In the actual anoline matrices, only external morphological 
characters have been utilized. The ecological and color characters 
have been relegated to the descriptors. This is not at all a necessary 



12 BREVIORA No. 502 

action. It may not even have been a prudent one. Any codable 
feature, ecological, behavioral, skeletal, or biochemical, may be 
utilized by our system. We sincerely hope that our matching key 
will be utilized by students of marine organisms as well as by 
students of other terrestrial biota. 



III. ANOLINE LIZARDS AS A 
TEST CASE FOR THE MATCHING KEY 

Ernest E. Williams 

The group first explored with the aid of our system is the anoline 
lizards, a monophyletic group that may include more than 400 
species. We are able at this time to provide data for only limited 
segments of this chosen group. These small segments are provided 
only as examples and samples: examples of the method and sam- 
ples simultaneously of some general problems and, of course, of 
the special features that must be tailored to each individual case. 

The anoline lizards have inspired the development of our sys- 
tem because they fit so well into the category of difficult taxa. 
They have, indeed, traditionally been so regarded. In part— the 
most trivial part— this is a result of the high number of species. 
But, especially, the difficulty in identifying the anoles is a result 
of overlapping variability in scale characters and the absence or 
near absence of the invariant characters so necessary for the classic 
dichotomous key. The dearth of invariant characters is especially 
egregious for females and juveniles, and this by itself tends to 
make a dichotomous key inoperable. 

In addition, the anoline lizards are highly visually oriented, and 
color vision is important in social interactions. Unfortunately, 
the importance of color in life to the animals themselves does not 
at all help in sorting out preserved specimens. Species often differ 
very little in any external feature except color, yet the usual meth- 
ods of preservation obliterate or obscure characteristic colors and 
patterns or, alternatively, reveal patterns unusual in the live an- 
imal. The colors of the living animal are always more or less 
altered and may be darkened in poorly preserved specimens of 
many species to a uniform muddy brown. The dewlap, it is true, 
is usually much less altered, and especially less darkened, than 



1994 COMPUTER IDENTIFICATION OF SPECIES 13 

Other portions of the body, but it is much more often than not 
quite absent in females and, unless preserved fully extended, is 
of little use even in males. Furthermore, even live animals may 
differ greatly in color or pattern according to physiological and 
psychological states. It is therefore small occasion for surprise 
that keys that rely heavily on color characters in anoline lizards 
are less than universally useful. 

This is true, unhappily, for our matching program as much as 
for dichotomous keys. For species distinguished primarily on col- 
or characters, many formaldehyde-preserved specimens are use- 
less. Our matching program may, indeed, be able to do more than 
the conventional dichotomous key: the 37 external characters that 
are used in the key may sometimes achieve identification when 
color, because of artifactual uniformity, fails entirely. But even 
scales may be poorly visible on badly preserved specimens, and 
it should be emphasized that old specimens of anoline lizards 
that are uniformly brown or black as preserved, without infor- 
mation on colors in life, or dry, damaged, or inadequately pre- 
served are, for many or most purposes, worthless. 

So species-rich and widely distributed a taxon as the anoline 
lizards is for many purposes unmanageable as a unit. Since Bou- 
lenger's key of 1885 — never very useful and now, in the most 
genuine sense, hopelessly out of date— no dichotomous key for 
the anoline lizards has ever existed, nor is one ever likely to exist. 
It is the plausible practice of the taxonomist to break such a large 
unit into smaller segments, at least geographically, perhaps by 
political boundaries, or perhaps by presumed or provisional "nat- 
ural" units. Even the computer system that we propose will work 
most usefully and easily on smaller units than the 400 or more 
species that is the expected total for the whole taxon. 

As a first— but, we feel, a fair— test of our system, we have tried 
it on the 1 2 species occurring on the Puerto Rican bank: the land 
area exposed at various times during the Pleistocene of which 
Puerto Rico and the Virgin Islands and a few other small islands 
are the present emergent parts. Eleven of the species are unques- 
tionably valid, ten of them— all those of mainland Puerto Rico— 
widely overlapping geographically or intimately interdigitating. 
The eleventh (A. roosevelti Grant) is rare or extinct, known only 
from six specimens. It was, indeed, known from only two indi- 



14 BREVIORA No. 502 

viduals, until the recent reexamination of old collections. It is, 
however, distinctive enough that its species status has never been 
questioned. A twelfth species, A. ernestwilliamsi, recently de- 
scribed (Lazell, 1983), has a very small range, a single cay, an 
enclave within the range of its very widely distributed closest 
relative; its status is still disputed. Indeed, only the latter is a 
problem species in the sense that its species status can be ques- 
tioned; all other species are somewhere sympatric with their clos- 
est relatives, A. cuvieri and A. roosevelti, which are allopatric and 
are distinguished by strong and consistent morphological features. 

The twelve species include several species groups and range 
from siblings to taxa, the relationships of which, at least within 
Puerto Rico, are quite unclear. Small as the anole fauna of the 
Puerto Rican bank is, it will exemplify many of the problems of 
"difficult" taxa. 

The anoles as a group cannot be specified by any single character 
nor by any unique combination of characters. The group is spec- 
ified by the combination of two or three characters, any one of 
which may be missing. Again, it is usual, or at least not infrequent, 
that subgroups within the anoles present the same problem of 
definition. We suggest that this is the sort of difficulty that makes 
"difficult" taxonomic groups difficult. The special case of Puerto 
Rican anoles is in some regards an atypically favorable case. There 
are no known hybrids and no equivocal species except the one 
that has an extraordinarily restricted geographic range. Using both 
data sets, there should be in the case of the Puerto Rico anoles 
no specimens for which the determination is doubtful. There is 
negligible probability of the discovery of new species. This will 
not be true for other geographic segments of the anoline lizards. 
For these, there will be other problems to confront case by case. 
For example, in the case of the Panama-Costa Rica matrix, we 
know of at least nine Panamanian species that are undescribed 
and one or two others that are possibly valid; we would also add 
to species recognized by Savage for Costa Rica some that he has 
synonymized or left undescribed. The existence of undescribed 
or disputed species is a very general problem in all the Central 
American and South American areas of anole distribution. Only 
Cuba presents many similar issues in the West Indies. 



1994 COMPUTER IDENTIFICATION OF SPECIES 15 

IV. THE MORPHOLOGICAL CHARACTERS USED 

IN THE ANOLEKEY DESCRIBED, 

DEFINED, AND ILLUSTRATED 

Ernest E. Williams 

The 37 counts and character descriptions provided in the ma- 
trices used by the IBM and Macintosh ANOLEKEYs have been 
chosen by myself as those beheved, after long trial, to be the most 
useful examined and readily codable characters (size, proportions, 
scale character states, and counts) for anoline lizards as a whole. 
This is a personal judgment and perhaps a very temporary one. 
It can be justified only by some measure of empirical success, 
and it remains to be widely tested. 

The following counts and character states are presented as a 
coded sequence of alternatives (Table 1). It will always be possible, 
and often probable, that counts a little higher than or states ad- 
jacent to those presented for any species in the coded matrix will 
be found in a specimen presented for identification. In considering 
whether to accept a match or a failure to match in an ANOLEKEY 
Report, allowance should always be made for this possibility. 
However, counts or conditions numerically farther away from 
those predicted by the matrix for the species found closest to the 
examined material on first tnal should be regarded as evidence 
for preferring the identification of some other species. Certainly 
this is grounds for reexamining, perhaps recoding, some of the 
characters of the unknown anole and, as well, very carefully con- 
sidering the alternative possibilities suggested by the supplemen- 
tary descriptor provided for each matched species in the report. 

It may well be that many current species matrices will require 
modification with an increase in sample size. (Those matrices 
based on samples below 20 will certainly need modification.) But 
such modifications should, of course, only be done // there is 
strong evidence from the congruence of the other characters and 
the details provided by the descriptor that the specimen with the 
discrepant character really belongs to the same species. 

In such cases of failure to match the likelihood of a new species 
is especially real for the anoles of mainland South and Central 
America. Both areas are inadequately known, and local new spe- 



16 BREVIORA No. 502 

Table 1. 

1. Head Scales. Smooth: 1,1. Rugose: 2,2. Unicarinate: 3,3. Multicarinate: 4,4. 
Striate: 5,5. 

2. Scales between Second Canthals (Fig. 1). 1-30. 

3. Postrostrals (Figs. 1, 2, and especially 3). 2-15. 

4. Nasal (Figs. 1, 2, and especially 3). Circumnasal: 0,0. Anterior nasal: 1,1. 
Divided anterior nasal: 2,2. Inferior nasal 3,3. 

5. Scales between Nasal and Rostral (Figs. 1, 2, and especially 3). 0-5. 

6. Scales between Supraorbital Semicircles (Fig. 1). 0-10. 

7. Enlarged Scales in Supraocular Disk (Fig. 1). 0-30.* 

8. Elongate superciliaries (Figs. 1, 2, and 4). 0-7. 

9. Superciliary Series (Figs. 1, 2, and 4). Granules: 1,1. Small scales: 2,2. Larger 
square or swollen scales: 3,3. 

10. Loreal Rows (Fig. 2). 1-15. 

1 1. Loreal Number (Fig. 2). 2-tO.* 

12. Interparietal Relative to Ear (Figs. 1, 2, and 5). Much smaller: 0,0. Smaller: 
1,1. Equal to: 2,2. Larger: 3,3. Much larger: 4,4. Interparietal absent: 5,5. 

13. Scales between Interparietal and Semicircles (Fig. 1). 0-15. In the absence of 
an interparietal: 888,888 in the IBM version, NA in the Macintosh version. 

14. Scales between Interparietal and Nape Scales (Fig. 1). In the absence of an 
interparietal: 888,888 in the IBM version, NA in the Macintosh version. 
Count of enlarged scales behind the interparietal distinctly larger than nape 
scales: 0-15. Scales behind interparietal grading into nape scales: 50,50.* 

15. Scale Rows between Suboculars and Supralabials (Fig. 2). 0-3. 

16. Supralabials to below Center of Eye (Fig. 2). 4-15. 

17. Postmentals (Fig. 6). 1-15. 

18. Sublabials (Fig. 6). 0-2. 

19. Sublabials in Contact with Infralabials (Fig. 6). 0-10. 

20. Dorsals (Figs. 7 A, B). Rat, smooth: 1,1. Swollen: 2,2. Unicarinate: 3,3. Mul- 
ticarinate: 4,4. Triangular or conical crest scales: 5,5. 

21. Enlarged Middorsal Rows (Figs. 7 A, B). 0-30.* 

22. Middorsal Crests (Fig. 7A). None: 0,0. Low crest: 1,1. High crest: 2,2. 

23. Rank Scales (Figs. 7A. B). More or less widely separated: 0,0. Juxtaposed: 
1,1. Imbricate: 2,2. Heterogeneous: 3,3. 

24. Size of Ventrals Relative to Dorsals (Figs. 7 A, B). Larger: 1,1. Equal: 2,2. 
Smaller: 3,3. Much smaller: 4,4. 

25. Smooth/Keeled Ventrals (Figs. 7A, B). Smooth: 1,1. Weakly keeled: 2,2. 
Strongly keeled: 3,3. 

26. Ventrals (Figs. 7 A, B). Separated: 0,0. Juxtaposed: 1,1. Subimbricate: 2,2. 
Imbricate: 3,3. 

27. Toe Pads (Figs. 8 A, B). Pad overlapping first phalanx: 1,1. Pad not distinct 
from first phalanx: 2,2. No pads: 0,0. 

28. Lamellar Number (Figs. 8A, B). 0-50. 

29. Supradigitals. Smooth: 1,1. With indistinct or single keels: 2,2. Multicarinate: 
3,3. 



1994 COMPUTER IDENTIFICATION OF SPECIES 17 

Table 1. Continued. 



30. Tail (Fig. 9). Round: 1,1. Weakly compressed: 2,2. Strongly compressed: 3,3. 

31. Tail Crest (Fig. 9). None: 0,0. Serrate: 1,1. Distinct crest: 2,2. High crest: 3,3. 

32. Postanals. Present: 1,1. Obscure: 2,2. Absent: 3,3. 

33. Dewlap (Male) (Fig. 10). Large: 1,1. Intermediate: 2,2. Small: 3,3. Absent: 
4,4. 

34. Dewlap (Female) (Fig. 10). Large: 1,1. Intermediate: 2,2. Small: 3,3. Absent: 
4,4. 

35. Snout-Vent Maximum (Male): 0-300. 

36. Snout-Vent Maximum (Female): 0-300. 

37. Tail Length/Body Length: Ratios between 0.8 and 1.2: 1,1. Ratios between 
1.3 and 1.7: 2,2. Ratios between 1.8 and 2.2: 3,3. Ratios between 2.3 and 
2.7: 4,4. Ratios between 2.8 and 3.0 or more: 5,5. 

*The last number is an arbitrary number (see text). 



cies are rather to be expected than not. Only widespread common 
species are well known, and these only relatively so. This is not 
true for the anoles of the West Indies, which have been extensively 
studied, although Cuba, which has been relatively neglected, may 
be expected to have some efflorescence of novelties. 

It is important to realize that the samples for even the best 
represented species have been selected opportunistically. When 
readily available, males, females, and juveniles have all been 
counted and coded, but no attempt has been made to secure a 
"fair" representation of age and sex classes. Sampling of geo- 
graphic variation also has been opportunistic: no provision has 
been made for "adequate" sampling of described subspecies. 

It is a matter of course that for poorly known species the samples 
are always biased and inadequate. There is always, as we have 
stated in Section II of this paper, extrapolation— judgment— in 
the association of a second specimen with a unique type or in the 
association of a population from a new locality with a species 
previously known from a small, geographically limited range. 

It is important to reemphasize that there is no escape from such 
matters of judgment. Indeed, some of the species in our ANO- 
LEKEY matrices may be composite. Named subspecies have been 
consciously lumped in the preparation of the reference matrices 
for the ANOLEKEY. Some subspecies will, rather surely, be rec- 
ognized as full, valid species. Others will turn out to be biologically 



18 BREVIORA No. 502 

meaningless. We do not attempt to solve such taxonomic prob- 
lems. We do provide a computer method that will assist the 
needed comparisons. 

The ranges for counts reported in Table 1 as the permissible 
limits for counts or measurements may deliberately extend be- 
yond the limits actually known for any anole species. Thus, the 
lowest known count of total number of loreals (character 1 1, later) 
is three (in two species of Phenacosaurus). I have coded the min- 
imum number for total loreals as two, because this would be a 
readily expectable variation. Similarly, although the maximum 
count for lamellae under phalanges ii and iii of the fourth toe 
(character 28) that I have actually counted is 44, I have coded 
the maximum for this count as 50. Coding of this kind has been 
done in several characters in the interest of allowing for easy 
modification of matrices when known counts are exceeded in 
either direction. 

In the case of character states. Table 1 reports the known states 
for each character, or, as in the preceding case of certain quan- 
titative characters, extrapolated, to take care of variation that 
may become known in the future. Any state other than those 
listed must be coded 888,888 for the IBM version of ANOLE- 
KEY, or NA (nonapplicable) for the Macintosh version. Any 
known character state not reported in some individual species 
matrix but found in some specimen judged to be conspecific with 
that species may be added to the appropriate matrix by use of 
the Change Menu in the IBM ANOLEKEY or by simple insertion 
of the missing coding in the relevant field in the Macintosh Anolis 
Handlist. 

Certain counts are more repeatable than others, and some char- 
acter states may not be readily interpretable. There should be, for 
example, except for obvious pathological conditions, no equivocal 
counts for the number of postrostral scales or for the number of 
scales between the supraorbital semicircles. In other cases, it is 
very necessary to be aware of arbitrariness and subjectivity in the 
evaluation of a character. Whenever this is true, advantage should 
be taken of the program's explicit recognition of the possibility 
of intermediacy or ambiguity in counts or character states by 
coding a range even for an individual. In all difficult cases, no 
time should be wasted by attempting false precision. Code inter- 



1994 COMPUTER IDENTIFICATION OF SPECIES 19 

mediacy, if that seems appropriate, or code the extremes allowed 
by differing interpretations. 

A special case exists in the several instances of graded series of 
enlarged scales. It is always difficult to decide when to count a 
scale as enlarged except in the cases of truly abrupt enlargement. 
In such cases, high counts often reflect a condition much closer 
to zero than to low counts. I have in such cases used arbitrary 
high numbers as signals that gradation in size has made counts 
subjective enough to be meaningless except as indicating gradual 
change in size. 

In the attempt to facilitate the examination of specimens, counts 
and characters are listed in an order in which a specimen might 
readily and naturally be examined, beginning with the head and 
proceeding to the body, limbs, and tail. 

Remember that all characters must be recorded by two numbers 
and that there must be a comma between the numbers. Missing 
characters should be coded 999,999 in the IBM version, or UA 
(=unavailable) in the Macintosh version. Nonapplicable charac- 
ters should be coded 888,888 in the IBM version, or NA (=not 
applicable) in the Macintosh version. I explicitly reinforce the 
preceding admonitions by repeating in the coding for character 
states below double numbers separated by commas. 

All terms used in Character Descriptions are defined in Peters' 
(1964) Dictionary of Herpetology. 

Character Descriptions 

/. HEAD SCALES. Smooth: 1,1. Rugose: 2,2. Unicarinate: 3,3. 
Multicarinate: 4,4. Striate: 5,5. Recognition of smooth, more or 
less parallel ribbed (striate) or rugose head scales or those with 
strong single ridges (unicarinate) or strong multiple ridges (mul- 
ticarinate) should not be difficult. The coding for any individual 
specimen should reflect the most extreme condition in terms of 
keeling. (The supraoculars may be the most useful scales to look 
at.) 

A very useful character. Striate is the rarest condition. 

2. SCALES BETWEEN SECOND CANTHALS (Fig. 1). 
Range: 1-30. Because they are better defined posteriorly, the can- 
thals are counted from the eye forward. The first canthal always 



20 



BREVIORA 



No. 502 



rostral 



postrostrols (3) 
[coding 8,8] 



inferior nasol (4) 
[coding 3, 33 , ond 
(5) [coding 0,0] 



head scales 
smooth (I) 
[coding I, l] 



second conthcl 

first ccnthol 
supraorbito 

elongate 
superciliary (8! 
[coding 1,1] 

superciliary 
series (9) 
[coding I, [] 



scoles between 
Interponetal and 
nape scales (14) 
[coding 6,6] 




scales between 
second conthal (2) 
[coding 8,B] 



scales between 
suproorbital 
semicircles (6) 
[coding 2, 2) 



supraocular disk (7) 
[codings, 9] 

scales between 
nterporietol and 
semicircles (13) 
[coding 2, 2) 



interparietal 



nape 



Figure 1. Dorsal view of head of "typical" anole (characters 2-9, 13, and 14). 
Boldface numbers in parentheses indicate relevant characters; plain numbers in 
brackets display codings for the illustrated character states. 



extends from the canthal ridge backward over the orbit; the second 
usually does not have such an extension. 

This is not necessarily an easy count to make. Variation ac- 
counts for some of the problem. At the lower range of counts, 
variation within a species should not exceed 1 or 2, but it may 
be 4 or more at the higher extremes. Because two or more scales 
may be in contact with the second (or the third) canthal on each 
side, counts within one individual by different observers or by 



1994 



COMPUTER IDENTIFICATION OF SPECIES 



21 




■£,8,«n 







u 






t-) 






s 






■•6 






B 






•^ 






U9 






S 






a> 






6 






fi 






p 






H 






a 






a 






• fH 






tn 






ii 






U 






JO 






g 






3 






C 






lU . 






O l« 






,C^ IL> 






l« *j 


^ 




•a <a 


o 




1-^ ■«-» 


3 




o «» 


^ <-) 




09 k. 


^ o 




<u 


r -^ 






Z) 




l/l 






"O 




C u 


ween 
Is an 
(15) 


^ 




^^ c3 


^5 o 


o" 






en 

c 

■a 

o 

-^1 


8-1 
illu 


O Q. -O 
U Z3 3 
to CO LO 


in ^ 


^.-^ 




o <0 




^' ^ 






. o 






f <i; 


^ ^^IFT 




(U (A 


o <u „ 




■*-» 00 


5 ^ N- 




charac 
codin 


supral 
center 
[cod in 






^-^ >. 




4> W 












"o a 






c .22 






« T3 






^ 2 












ca aj 






y ^ 






■&2 










r -^ 






C c 






o •- 












rt aj 






u X> 






^1 






?> c 






u .3 






•r' ca 






> —3 






— a 






rt . - 






C. w) 






(U t^ 






is (U 






ca -^ 






>— 1 CO 






u 






« 






cm" -C 






igure 
vant 






Uh 'U 



22 



BREVIORA 



No. 502 



postrostra 




C 



-/- 



circumnasQl 

(4) = 0,0/(5) = 0,0 



rostral / 

1st supralabial 








circumnasal separated 
(4) = 0,0/(5)=IJ 




single anterior nasal 
(4) = l,l/(5)=0,0 








divided anterior nasal 
(4)=2,2/(5) = 0,0 




inferior nasal 
(4)-3,3/(5)=0,0 



postrostral anterior nasal 




Q, \ inferior nasal 



Figure 3. Characters 4 and 5 (nasal scales of anoles). 



1994 COMPUTER IDENTIFICATION OF SPECIES 23 

one observer at different times may differ by 1 or more scales. 
For matrices based on few specimens, this may not be a useful 
character; it is primarily useful for the low counts, 2-5, but every 
species has a characteristic range of counts. 

3. POSTROSTR.4LS (Figs. 1, 2, and especially 3). Range: 2- 
15. Those scales behind the rostral, in contact with it, and between 
the supralabials are easily counted and will usually not vary within 
a species by more than 2 to 3. This count may include the cir- 
cumnasal and any other of the differentiated nasals discussed next. 

4. NASAL (Figs. 1, 2, and especially 3). Circumnasal: 0,0. An- 
terior nasal: 1,1. Divided anterior nasal: 2,2. Inferior nasal: 3,3. 
The nasal in anoles may be a single oval scale that contains the 
nostril. It is then coded as "circumnasal": 0,0. More often, the 
scales around the nasal are differentiated, overlap, and at least 
partially obscure it. Most common is a condition in which a scale 
anterior to the nasal becomes large and subtriangular and overlaps 
part of the nasal scale; it is then coded as "anterior nasal": 1,1. 
In some species, the anterior nasal is divided transversely; it is 
then reported as a "divided anterior nasal": 2,2. The other scales 
overlapping the nasal are not considered, except that the inferior 
nasal— a scale obscuring the lower surface of the nasal— if it comes 
to overlie the sulcus between the rostral and the first supralabial, 
is then, and only then, coded "inferior nasal": 3,3. Occasional 
anomalies (e.g., an anterior nasal replaced by small granules) or 
conditions due to injury should always be reported as 888,888 in 
the IBM version, or NA in the Macintosh version. 

This and the next (character 5) are important and very useful 
characters. In some cases both may be difficult to score. It is then 
best to code the alternatives; these may have been already in- 
cluded in the relevant matrices. 

Note that the sulci bounding the scales surrounding and over- 
lapping the nasal are sometimes obscure and that, therefore, the 
existence, for example, of the anterior nasal must be inferred. 
Most often, the plausible coding will be 1,1. These difficulties 
should, in any event, exist only in single specimens or one side 
of a single individual. Series should obviate or alleviate the prob- 
lem. 

5. SCALES BETWEEN NASAL AND ROSTRAL (Figs. 1, 2, 
and especially 3). Range: 0-5. In anoline lizards with the condition 



24 



BREVIORA 



No. 502 



(8) = 3,3 (9) = 2,2 




(8) = l,l (9)=l,l 




(8) = 0,0 (9) = 1,1 




(8) = 0,0 (9) = 3,3 




Figure 4. Characters 8 and 9 (superciliary scales in anoles). 



"circumnasal"— the simple unmodified nasal scale— that scale may 
be in contact with the rostral, but one or more scales usually 
intervene. In the latter case, the minimum number is coded. In 
the condition "anterior nasal," the nasal itself is obscured, but 
the anterior nasal is usually in contact with the sulcus between 
the rostral and first supralabial. This condition is coded 0,0. If 
the anterior nasal is wholly in contact with the rostral, and the 
inferior nasal has moved into a position above the sulcus between 
the rostral and first supralabial, the coding is still 0,0. If, however, 
a recognizable circumnasal or anterior nasal is separated from the 
rostral by one or more scales, then the relevant minimum count 
is recorded. 

Often more than one interpretation of any individual condition 
may be possible. It is, for example, difficult in some species to 
decide whether the nasal scales are properly regarded as the "an- 



1994 COMPUTER IDENTIFICATION OF SPECIES 25 

terior nasal" separated by one scale from the rostral (i.e., character 
4: 1,1; and character 5: 1,1), or whether the better interpretation 
is "anterior nasal" divided and in contact with the rostral (i.e., 
character 4: 2,2; and character 5: 0,0). Ordinarily the ambiguity 
will already be recorded in the matrix for the species in question, 
and either of the options will be accepted by the computer. 

6. SCALES BETWEEN SUPRAORBITAL SEMICIRCLES 
(Fig. 1). Range: 0-10. This is a minimum count. Any contact, 
even a point contact, between the supraorbital semicircles is coded 
as 0,0. 

This is an important count, but in some species there is excep- 
tional variation. 

7. ENLARGED SCALES IN SUPRAOCULAR DISK (Fig. 1). 
Range: 0-30. Enlarged scales arranged as a supraocular disk may 
be conspicuously larger than any surrounding scales; they are then 
easily countable. In many species, however, there is gradation 
such that all counts are subjective; in such counts, a range should 
be reported. In still other species, the enlargement is so gradual 
and so limited that there can be no pretense of an accurate count. 
These cases should be coded arbitrarily as 30,30. If there is no 
indication of a disk or of significant enlargement of any of the 
supraocular scales, the coding should then be 0,0. 

In many or most species of anoles, this is not the most useful 
of characters. It is useful in those cases that are unambiguous, 
i.e., those in which the disk consists of a few large scales that are 
sharply distinct from surrounding scales. High counts, i.e., smaller 
scales, are ordinarily subjective. 

It is possible to be undecided about whether the coding should 
be 30,30 or 0,0. Specimens or species in which this occurs should 
be coded 30,30. The alternative, 0,30, is unacceptable, because 
the computer will then assume that all counts between and 30 
are valid, although no species is known in which so highly variable 
a condition is true. This coding will be closer to reality. 

If you have before you any series containing low counts (large 
scales; e.g., 3-7) in the supraocular disk and also high counts 
(small scales; e.g., 20-30), that series should be suspect as com- 
posite, and low- and high-count specimens treated separately. 

8. ELONGATE SUPERCILIARIES (Figs. 1, 2, and 4). Range: 
0-7. The superciliaries are distinguished from the canthals by not 



26 BREVIORA No. 502 

extending anteriorly beyond the orbit at all. The first superciliary 
in anoles is usually distinguishable from more posterior super- 
ciliaries by being much longer anteroposteriorly. If there is more 
than one elongate superciliary (there may be as many as six), they 
will be strongly overlapping and grade in length posteriorly. In a 
few species, there is no elongate superciliary, and all the super- 
ciliary scales are granular. If so the coding is 0,0. 

9. SUPERCILIARY SERIES (Figs. 1, 2, and 4). Granules: 
1,1. Small scales: 2,2. Larger square or swollen scales: 3,3. The 
scales posterior to the elongate superciliaries— if any are present- 
may be granular, like the smaller scales of the supraocular area, 
and are then coded 1,1. Alternatively, there may be one or two 
rows of distinctly larger but still small scales following the elongate 
superciliary or superciliaries. If there is difficulty in deciding be- 
tween these two conditions, or if there is a mixture of granules 
and small scales, the code should be 1,2. If there is no elongate 
superciliary and all superciliaries are large and squarish or large 
and swollen, the conditions should be coded 3,3. The large and 
squarish and the large and swollen superciliaries are rare condi- 
tions, known only in two species in Colombia, Ecuador, and 
Panama. 

10. LOREAL ROWS (Fig. 2). Range: 1-15. Loreal rows are 
counted down from the second canthal or from the first canthal, 
if it clearly overlaps the loreal area (those scales that continue the 
subocular arc in front of the eye are preoculars, not loreals, and 
should never be counted). Sometimes, however, it is difficult to 
distinguish the preoculars from the loreals; this count is then 
subjective, by one or two. 

This is a count that is often subjective. The next character (total 
loreal number) is then more useful. 

77. LOREAL NUMBER (Fig. 2). Range: 2-40. Total loreal 
number is easily counted when the loreals are few and the subocu- 
lars broadly in contact with the supralabials. Difficulty occurs 
when the loreals are confluent with a scale row or rows separating 
the subocular and supralabials. Whenever such a row is complete, 
the number of loreals will, in most cases, be relatively high (sig- 
nificantly >30); it is then convenient to use the arbitrary coding 
40,40. In some cases the preoculars also may be difficult to dis- 
tinguish. Where the total count would in any case be less than 









1994 COMPUTER IDENTIFICATION OF SPECIES 27 



interparietal 



ear ^ 

(12) coding = 4,4 3,3 2,2 

Figure 5. Size of interparietal compared with that of ear (character 12). 



40, the possible interpretations may be coded as a range. If the 
ambiguous situation involves a high count, the arbitrary number 
40,40 may be the best solution. Low counts are clearly diagnostic 
for certain species; even errors of one or two do not matter. High 
counts do not appear ever to be useful for species discrimination. 

12. INTERPARIETAL RELATIVE TO EAR (Figs. 1, 2, and 
5). Much smaller: 0,0. Smaller: 1,1. Equal to: 2,2. Larger: 3,3. 
Much larger: 4,4. Interparietal absent: 5,5. The size of the inter- 
parietal relative to the size of the ear is ordinarily unambiguous. 
If there is ambiguity or variation within a species, code as a range 
(e.g., 1,3). Again extreme conditions tend to be diagnostic, but in 
many species codings of 1, 2, or 3 will be appropriate within a 
single population. 

The absence of an interparietal is a rare and apparently derived 
condition; it is confined to certain species in northwestern South 
America. 

13. SCALES BETWEEN INTERPARIETAL AND SEMI- 
CIRCLES (Fig. 1). Range: 0-15. In the absence of an interparietal: 
888,888 in the IBM version, NA in the Macintosh version. If there 
is no interparietal, the coding of this character must be 888,888 
for the IBM version, or NA in the Macintosh version. This is 
again a minimum count taken on each side from the interparietal 
to the nearest scales of the supraorbital semicircles. If the two 
sides differ, code them as, for example, 2,3. Ranges of 3 or 4 or 
more are not unusual in certain populations. 

14. SCALES BETWEEN INTERPARIETAL AND NAPE 
SCALES (Fig. 1). In the absence of an interparietal: 888,888 in 



28 



BREVIORA 



No. 502 



mentol 

media 
gular 



sublabial (18) 
[coding 2,2] 



sublabiols in contact 
with infralabiais (19) 
[coding 2,2] 



latere 
gular 



postmentols (17) 
[coding 8,8] 



infralabiais 




central 
gular 



Figure 6. Ventral view of head of "typical" anole (characters 17-19). Boldface 
numbers in parentheses indicate relevant characters; plain numbers in brackets 
display codings for the illustrated character states. 



the IBM version, NA in the Macintosh version. Count of enlarged 
scales behind the interparietal distinctly larger than nape scales: 
0-15. Scales behind interparietal grading into nape scales: 50,50. 
This count is made in the approximate midline behind the in- 
terparietal, and the scales must be appreciably larger than the 
nape scales immediately behind them. If there is gradation, and 
this frequently is the case, the count is surely subjective. If so, 
code as 50,50. 

Discrepancy compared with a matrix is not to be taken seriously 
in the case of high and subjective counts. However, may have 
a taxonomic meaning, i.e., an interparietal followed by small 
scales not significantly different from nape scales. This differs 
importantly from cases in which scales behind the interparietal 



1994 COMPUTER IDENTIFICATION OF SPECIES 29 

differ abruptly in size from nape scales. Thus, both a low count 
and an unambiguous count of 0,0 may be very useful species 
characters. 

75. SCALE ROWS BETWEEN SUBOCULARS AND SU- 
PK4LABIALS (Fig. 2). Range: 0-3. This is also a minimum count. 
Anv contact between suboculars and supralabials is coded 0,0. 

16. SUPRALABIALS TO BELOW CENTER OF EYE (Fig. 
2). Range: 4-15. Because the posterior termination of the supra- 
labial series may sometimes be difficult to determine, the su- 
pralabials are counted from the rostral posteriorly. The sulcus 
between two supralabials may lie below the center of the eye. The 
coding is then a range (e.g., 6,7). 

17. POSTMENTALS (Fig. 6). Range: 1-15. This is a count of 
all the scales in contact with the mental between the infralabials. 
It includes the anteriormost sublabials, if these are differentiated. 

18. SUBLABIALS (Fig. 6). Range: 0-2. A sublabial series is 
an abruptly enlarged series of scales on each side paralleling or 
radiating from the infralabials. Abruptly enlarged is here inter- 
preted to require that the first sublabial of each side be at least 
four to five times the size of the postmental (^^medial gular) medial 
to it. If no such abruptly enlarged scales adjacent to the infralabials 
exist (i.e., if the postmentals are subequal or grade evenly from 
larger laterally to smaller medially), or if the enlarged scales are 
only twice to three times any medial gular, the condition is to be 
reported as the absence of sublabials, coded 0,0. If both first 
sublabials are present, the coding is 2,2. If, as sometimes happens, 
a first sublabial is present on one side only, the coding is 1,1. If 
there is ambiguity (i.e., if you cannot decide whether or not the 
putative sublabials are as much as four times the other postmen- 
tals), code 0,2. 

This character again is a very useful one. If the sublabials are 
recognized only when the lateral postmentals are four to five times 
larger than any medial gular, the condition tends to be invariant. 
Species in which the lateral postmentals are only two to three 
times larger than the medial gulars tend to be variable in this 
regard and often have the lateral postmentals subequal to the 
medial gulars or grading into them. 

19. SUBLABIALS IN CONTACT WITH INFRALABIALS 
(Fig. 6). Range: 0-10. In anoles, the first sublabials, when present, 



30 BREVIORA No. 502 

are almost always in contact with the first infralabials. If the first 
sublabial is, as rarely may be the case, separated from the first 
infralabial by a lateral gular, the coding is, of course, 0,0. Any 
more posterior sublabials (=abruptly enlarged scales in sequence 
with the first) may also be in contact with the infralabials, or some 
or all of them may be separated by a row of smaller scales. If only 
the first sublabials are in contact with infralabials, or there are 
no posterior sublabials, the coding is 1,1. In some species, there 
may be as many as 9 in contact, and the number often differs on 
the two sides (e.g., 1,2 or 6,7). If there are no sublabials at all the 
coding is, of course, 0,0. 

20. DORSALS (Figs. 7 A, B). Flat, smooth: 1,1. Swollen: 2,2. 
Unicarinate: 3,3. Multicarinate: 4,4. Triangular or conical crest 
scales: 5,5. These conditions should be readily recognizable, but 
if there is any ambiguity, code a range. 

21. ENLARGED MIDDORSAL ROWS (Figs. 7 A, B). Range: 
0-30. There may be subjectivity in counts of enlarged middorsal 
rows. In all such cases, code not single numbers but an appropriate 
range of values. If the enlarged middorsal rows are not countable 
because of too gradual a transition to the flank scales, code ar- 
bitrarily as 30,30. 

As in certain other counts, low counts may be more meaningful 
and repeatable than high counts. 

22. MIDDORSAL CRESTS (Fig. 7 A). None: 0,0. Low crest: 
1,1. High crest: 2,2. In anoles, "crest" is applied only to one or 
two rows of sharply enlarged middorsal crests, characteristically 
triangular or conical, of varying height, but always projecting 
conspicuously above the paravertebral scales. Most anoles lack 
such crests. 

This is not usually a very useful character. A few anoles have 
such high crests that I would be remiss if I did not call attention 
to them. 

23. FLANK SCALES (Figs. 7 A, B). More or less widely sepa- 
rated: 0,0. Juxtaposed: 1,1. Imbricate: 2,2. Heterogeneous: 3,3. In 
some, usually giant species, the flank scales are relatively large 
and may be separated by more or less minute granules. This 
infrequent condition is coded 0,0. Much more frequently, the 
flank scales are smaller, and some or all of them are narrowly 
separated by naked skin or by minute granules, which may allow 



1994 



COMPUTER IDENTIFICATION OF SPECIES 



31 





CO 



I 

o 



o 






o 
(/) 

c 
> 

c 




> 
o 

03 



03 
o 



o 






3 

00 



32 BREVIORA No. 502 

partial contact. This condition is coded 0, 1 . Many species have 
the flank scales juxtaposed; this is coded 1,1. If any of the flank 
scales clearly overlap, the coding is 2,2. In a few species, the larger 
scales of the flank may be of very unequal size; these are then 
scored as heterogeneous: 3,3. The minute granules, which may 
be present in some cases, are not considered in this definition of 
heterogeneity. 

24. SIZE OF VENTRALS RELATIVE TO DORSALS (Figs. 
7 A, B). Larger: 1,1. Equal: 2,2. Smaller: 3,3. Much smaller: 4,4. 
Most anole species have the ventrals larger than the largest dor- 
sals. This may, however, be untrue for species with a distinct zone 
of enlarged dorsals or those species with crest scales. There may 
even be noticeable variation in this regard within species. As 
usual, such variation is coded as a range. The condition — ventrals 
much smaller than dorsals— is known only in Chamaeleolis. 

This is often a very useful character. 

25. SMOOTH/KEELED VENTRALS (Figs. 7 A, B). Smooth: 

1.1. Weakly keeled: 2,2. Strongly keeled: 3,3. Some species are 
truly variable, with ventrals in some individuals keeled (always 
unicarinate) and in others smooth. Series from such species should 
be coded 1,2 or 1,3, as appropriate, and individuals either 1,1, 

2.2, or 3,3. Other species have keeling so weak that it is a sub- 
jective judgment whether or not it exists at all. Such species may 
be coded 1,2. 

There are just a few species in which ventrals vary from smooth 
to keeled. In most species this is a strong character. 

26. VENTRALS (Figs. 7 A, B). Separated: 0,0. Juxtaposed: 1,1. 
Suhimbricate: 2,2. Imbricate: 3,3. Ventrals are sometimes sepa- 
rated by naked skin or granules and often are juxtaposed or sub- 
imbricate (intermediate) or distinctly imbricate (distinctly over- 
lapping). More than one condition can occur within a species or 
even within an individual. In contrast to the last, this is not a 
strong character, but some species do have strong separation or 
strong imbrication. 

27. TOE PADS (Figs. 8 A, B). Pad overlapping first phalanx: 
1,1. Pad not distinct from first phalanx: 2,2. No pads: 0,0. In most 
species of anoles, the adhesive pad under phalanges ii and iii of 
all the toes projects distally under the proximal infradigital scales 
of the first (claw-bearing) phalanx. This is coded 1 , 1 . In a number 



1994 



COMPUTER IDENTIFICATION OF SPECIES 



33 



Ph I 



>Ph 2 



>Ph3 



Ph4 



>Ph2 




Ph3 



Ph4 



Figures 8A, B. Two conditions of the digital pads (characters 27 and 28). 
Phalanges i-iv are labeled, and appropriate counts of lamellae under the pad are 
illustrated. 



of species, not closely related, the projection is minimal or absent, 
and the proximal termination of the pad is therefore indistinct 
(coded 2,2). In the adults of one species only (Fig. 8B), a pad is 
completely lacking (coded 0,0). 

28. LAMELLAR NUMBER (Eigs. 8A, B). Range: 0-50. La- 
mellae are the widened distally overlapping scales characteristic 



34 BREVIORA No. 502 

crest 







(30) = !, I =2,2 =3,3 =3,3 

(31) = 0.0 =0.0 =0.0 =3,3 

Figure 9. Tail character states (character 30 and 3 1 ). 



of the adhesive toe pads of anoles. Counts are made only on 
phalanges ii and iii of the fourth toe of the hind foot. A given 
range of lamellar counts is species-specific. 

Counting lamellar number, as understood here, involves finding 
the bend between phalanges iii and iv; this is arbitrarily considered 
the proximal termination of the adhesive pad. A bend also occurs 
within the pad between phalanges ii and iii, but the distal ter- 
mination is defined as the joint between phalanges i and ii. Counts 
are therefore restricted to the lamellae under phalanges ii and iii. 
At the joint between phalanges iii and iv, the scale at or within 
the bend is counted. Distally the small terminal lamella of the 
pad projection is always counted or, in default of such a scale, 
the scale that lies across the joint between phalanges i and ii. 

The intention here is to count only scales of the pad proper. 
For most anole species, the procedure outlined above probably 
closely approximates reality. However, scanning electron mi- 
croscopy has revealed that, in fact, the adhesive lamellae with the 
functionally adhesive hairs are not necessarily confined within 
the boundaries of phalanges ii and iii. However, these boundaries 
provide a convenient macrostructural definition on which to base 
reproducible counts. Ultimately, however, the justification for 
this count, rather than counts of all lamellae under the fourth toe, 
is historical: Boulenger (1885), who first provided standard and 
recognizable descriptions of anole species, used the count of la- 



1994 



COMPUTER IDENTIFICATION OF SPECIES 



35 




(33) ^1,1 

(34) ^i.l 






4,4 
4,4 



Figure 10. Dewlap extent (characters 33 and 34). Arrow points to posterior 
termination of dewlap. 



mellae under phalanges ii and iii. Stejneger, Cochran, Schwartz, 
and Williams have routinely used this count. 

It is a source of some confusion that workers on Mexican and 
Central American anoles have rather consistently used other 
counts, especially the total number of subdigitals under the fourth 
hind toe. The attempt here is to standardize lamellar counts with 
those conventionally used for West Indian and South American 
anoles. 

29. SUPRADIGITALS. Smooth: 1,1. With indistinct or single 
keels: 2,2. Multicarinate: 3,3. Most anoles have multicarinate su- 
pradigital scales. The alternative conditions— smooth (1,1) or with 
indistinct, usually single keels (2,2)— are relatively uncommon 
and, therefore, when they do occur, diagnostic. 

30. TAIL (Fig. 9). Round: 1,1. Weakly compressed: 2,2. Strong- 
ly compressed: 3,3. Strongly compressed tails are readily recog- 
nized. Weakly compressed or round tails are common. 

31. TAIL CREST (Fig. 9). None: 0,0. Serrate: 1,1. Distinct 
crest: 2,2. High crest: 3,3. Round tails have no crests. Weakly 
compressed tails may or may not be serrate above. Strongly com- 
pressed tails may or may not have a crest, and may not even be 
serrate above, and, if present, the crest may be high or low. Fe- 



36 BREVIORA No. 502 

males rarely have high crests and variation from population to 
population within a species and within a sex is not unusual. 

32. POSTANALS. Present: 1,1. Obscure: 2,2. Absent: 3,3. The 
characteristic enlarged postanal scales of male anoles are usually 
laterally expanded ovals, somewhat resembling airplane propel- 
lers, a variable distance behind the vent. They may also be round, 
large or small, obscure, or, as in males of not a few species, and 
always in females, absent. 

33. DEWLAP (Male) (Fig. 10). Large: 1,1. Intermediate: 2,2. 
Small: 3,3. Absent: 4,4. Dewlaps should be scored as large if they 
extend onto the belly past the insertion of the arms, intermediate 
if they reach just the level of the axilla, and small if they are 
shorter than this. If the animal is female, code 888,888 in the 
IBM version, or NA in the Macintosh version. 

Large ( 1 , 1 ), as defined here, is a very inclusive term, and species 
so coded will differ conspicuously in the dewlap area. The present 
definition has been adopted to avoid the problem of quantifying 
dewlap area and to permit the scoring of dewlaps that are not 
extended in preserved animals. 

34. DE WLAP (Female) (Fig. 10). Large: 1,1. Intermediate: 2,2. 
Small: 3,3. Absent: 4,4. Females may possess a dewlap, which is 
then scored just as in males. However, both an indication— a 
mere zone of differentiated scales— or a mere fold (i.e., any struc- 
ture that could not be substantially extended) and a complete 
absence should be scored 4,4. If the animal examined is a male, 
code 888,888 in the IBM version, or NA in the Macintosh version. 

35. SNOUT-VENT MAXIMUM (Male). Range: 0-300. Snout- 
vent length should be measured on every specimen. The number 
with which this length is compared in the coded matrix is the 
maximum known for the relevant sex in each species. Only if the 
matrix number is exceeded by a substantial margin should a ten- 
tative identification be rejected. 

A is used as the minimum for size only because hatchling 
size is unknown for most anoles. 

A female should be coded 888,888 in the IBM version, or NA 
in the Macintosh version. 

36. SNO UT- VENT MA XI MUM (Female). Range: 0-300. The 
principles suggested for the interpretation of the comparable entry 
in males apply here also. For a male, code this entry 888,888 in 



1994 COMPUTER IDENTIFICATION OF SPECIES 37 

the IBM version, or NA in the Macintosh version. As in males, 
is treated as the minimum only because hatchling size is nearly 
unknown or unreported in anoles. 

37. TAIL LENGTH/BOD Y LENGTH. Ratios between 0.8 and 
L2: 1,L Ratios between 1.3 and L7: 2,2. Ratios between L8 and 
2.2: 3,3. Ratios between 2.3 and 2.7: 4,4. Ratios between 2.8 and 
3 or more: 5,5. Tail length should be measured whenever the tail 
is neither broken nor regenerated. (The regenerated portion of a 
lizard tail always differs in scale form from the unregenerated 
portion; such difference in scale form is never seen in an unre- 
generated tail.) 

The measured tail length must then be compared with the snout- 
vent length, and the ratio of tail length over body (snout-vent) 
length obtained. Ratios between 0.8 and 1.2 should be coded 1,1. 
Ratios between 1.3 and 1.7 may be coded 2,2; those between 1.8 
and 2.2 coded 3,3; those between 2.3 and 2.7 coded 4,4; and those 
between 2.8 and 3 or more 5,5. All intermediates should be so 
coded (e.g., individuals with ratios 2.75 as 4.5; series with ratios 
ranging from 1.5 to 1.9 as 2,3). 

V. ANOLEKEY- OPERATIONAL INSTRUCTIONS 
Ernest E. Williams 

The ANOLEKEY is menu-driven and is available in two ver- 
sions, one in Basic for the IBM PC, written by Hugh and Stanley 
Rand, and one in HyperCard for the Macintosh, written by Robert 
O'Hara. Both versions do essentially the same job but differ in 
one aspect. The HyperCard version for the Macintosh is essen- 
tially a database for all anoles with an ANOLEKEY as one of the 
options and the reference matrices as part of the database. 

At this time, most reference matrices are not finished and lack 
the added feature of descriptors. Only Puerto Rico, Jamaica, Ec- 
uador, and Costa Rica-Panama subsets of the total database are 
available at this time. However, it is hoped to add other subsets 
in the near future. 

Because the ANOLEKEY itself is functional, and we welcome 
suggestions and improvements, we have decided to go public at 
this time. A copy of ANOLEKEY and associated documentation 
(Hugh and Stanley Rand, 1994, for the IBM PC, and Robert 



38 BREVIORA No. 502 

O'Hara and Ernest Williams, 1 994, for the Macintosh) is available 
to anyone interested in trying the ANOLEKEY and willing to 
report bugs and/or suggest improvements. Each copy will include 
a description of the 37 characters and how to count them (Wil- 
liams, this paper). (For the IBM version, write to A. S. Rand 
[include formatted disk]; for the Macintosh version, write to E. 
E. Williams [include formatted disk].) 

In addition to the ANOLEKEY and its documentation, the 
distribution disk for the IBM includes a version of the Basic 
Program that is not customized for use with the anoles but that 
can be used with any group. For a comparable bare-bones 
HyperCard for the Macintosh version, write to E. E. Williams. 

VI. ANOLEKEY IN BASIC FOR THE IBM PC 
A. Stanley Rand 

The IBM version of ANOLEKEY consists of a Basic Program, 
a "Readme" file that describes the use of the program, a text file 
describing the 37 anole characters, and several reference matrices, 
all stored on a single floppy disk. The program was created in 
MS-DOS, is stored in ASCII, and may be loaded and run using 
BASICA, Turbo BASIC, or Quick Basic. The program should run 
on any IBM PC or clone. 

The disk contains two versions of the program adapted to iden- 
tify anoles, one that prints out a hard copy of results and the other 
that does not. There is also a stripped down version that lacks 
the modifications for use with anoles so that it can be used for 
any group. 

The reference matrices each contain information on all the 
recognized species of anoles from a specific geographic area. Four 
areas are currently available: Puerto Rico, Jamaica, Panama- 
Costa Rica, and Ecuador. Each matrix contains the ranges for the 
37 standard characters for each species and the short list of di- 
agnostic characteristics, the "descriptor," for each species. 

The program allows you to load a reference matrix (e.g., Pan- 
ama-Costa Rica) and then enter the information for the 37 stan- 
dard characters for an individual or series to be identified. Most 
of these characters are scale characters, two involve size, and one 
involves proportions. A list of these characters and their codings 
is given in Appendix 2 of the "Readme" file. Some of these 



1994 COMPUTER IDENTIFICATION OF SPECIES 39 

characters are counts that are entered directly; the others must 
be coded. You are prompted for each character and informed of 
the possible codings. For each character, the program will accept 
only an appropriate range of values. You must enter for each 
character two numbers separated by a comma, a minimum and 
a maximum. This allows you to enter information from a series 
of animals, to code the two sides of an animal if they are different, 
and to code an ambiguous situation. The data for the unknown 
can be corrected and stored in a file. 

Once the data are entered and checked, the program asks you 
to enter the number of characters where a species in the reference 
matrix with which the unknown is being compared must differ 
before being eliminated as a possible match. The larger this num- 
ber, the more species are matched and the more different they 
may be from the unknown. 

The program then compares the unknown with the reference 
matrix and produces a report that lists the species that match, 
giving for each the total number of characters in which it differs, 
what the differing characters are, and how great the differences, 
positive or negative, are. For each matching species the program 
supplies a short description of characters such as color, distri- 
bution, and any peculiarities that could help in identification. 

The program allows you to create or change a reference matrix, 
although creating a reference matrix is probably more easily done 
with a word-processing program outside of ANOLEKEY. The 
details of the format for a reference matrix are given in the 
"Readme" file. 

As a shortcut, there is also the option of comparing an unknown 
with the reference matrix using only two characters, instead of 
all 37. This will be useful if the unknown has among its 37 stan- 
dard characters one or two states that are so rare or unusual that 
only one, two, or at most three species can possibly be matches. 
The descriptor or descriptors will confirm the identification. 

VII. THE ANOLIS HANDLIST: 
A HYPERCARD VERSION OF THE ANOLEKEY 

Robert J. O'Hara and Ernest E. Williams 

The Macintosh version of the ANOLEKEY is much more than 
a computer key, and to better indicate its multiple functions we 



40 



BREVIORA 



No. 502 




^'^^■t^Kffffvrw^mm^i^^^ 



\fywwwwwf.>.-'.^!:i:v:/^^^^^^^ 



Anolis frenatus COPE 



1899 Anolis frenatus COPE, Phila Mus. Bull,, 1:6, Plate 2, Fig 2 - TYPE 
LOCALITY: "Colonribia, near Bogota" (Holotype: lost, fide BARBOUR 1934). 



CA, SA: COSTA RICA, PANAMA, COLOMBIA. Giant Long hind limbs. Four 
or five oblique dark bars on flanks, each bar split by a light line A dark 
inter orbital bar One very elongate superciliary. Thigh scales unicarinate, 
Dev/lap large, cream white. Arboreal, 



O 



O 



QmmTonv Qbehruior ®chhrhcters O color Qecologv Qli terhture 



hsC O: 1,3 

snsc(2): 9, 15 

pr<3): 6, 10 

n<4): 0,3 

r,-r<5>: 0, 1 

ssc(6): 2,5 

sd(7>: 5,30 
esc(8): 1,4 
scsCQ:*: 1,2 
lorr< 10> 6, 10 



lornC 1 1 ) 

ip/e< 12) 

ip/sc< 13) 

ip/n< 14) 

so/sKIS) 

sK16) 

pm(17) 

sK18) 

sl/l K 19) 

cK20) 



40,40 

0,3 

3,6 

2.7 

0,2 

8, 12 

6, 11 

0,0 

0,0 

1, 4 



mlddr<:21 ) 

middc<:22) 

fs(23) 

v/d(:24) 

ws/k<25) 

wj/i<26) 

R/H<:27) 

K28) 

sd(29) 



0,30 t(30): 1,3 

0,0 tcOl): 0,0 

0, 1 pa(32): 1,3 

1, 1 mdeuj<33): 1,2 
1, 1 fdeuj<34): 2,3 
0,3 msize(35): 0, 143 
1, 1 fsize(36): 0, 118 
21,28 tl /bl <37): 2,4 
3, 3 samp Is: 20 



Click HELP to see character def i n i t i ons , 



OhELP OkEV O subsets (*) species 140 OF 404 



F IHD. 



I ^ ^» ^ I 



Figure 11. Species card for Anolis frenatus from the Anolis Handlist. The 
buttons at the lower left, present on every card in the stack, allow the user to 
choose the Help, Key, Subset Editor, or Species cards. The diagnostic characters 
are now being shown on this card; information on anatomy, behavior, color, 
ecology, or literature may be displayed by clicking on the appropriate button. 



refer to it as the Anolis Handlist. In addition to serving as a key, 
the Anolis Handlist, when completed, is intended to be an index 
and checklist for the possibly more than 400 species oi Anolis, 
Chamaeleolis, Chamaelinorops, and Phenacosaurus residing in 
North, Central, and South America. 

Although the Anolis Handlist is still incomplete, as already 
mentioned, we describe it here as though it were complete. While 
waiting for EEW to complete additional reference matrices along 
with the several subsidiary COLOR, etc., fields, described below, 
the ANOLEKEY is fully functional. 

The Anolis Handlist was created with Apple Computer's 
HyperCard, an object-oriented software development environ- 
ment that makes use of all of the standard elements of the Mac- 
intosh interface, including pull-down menus, windows, buttons, 
and scroll bars (Apple Computer, 1987). HyperCard documents 
may be thought of as groups of interactive electronic file cards: 



1994 COMPUTER IDENTIFICATION OF SPECIES 41 



SELECT KEY OR COHPHR I SOM REPORT: ® HtlOLE KEV Q COHPfiR I SOM REPORT 



THE ANOLE KEY • Type the name of an unknown specimen or sample m 

the top field below, and minimum and maximum values (for example '4,6' 

or '2,2 0') for the characters of the unknown mto the 37 character fields 

For invariant characters enter, for example, '8,8'. Put 'UA' and 'NA' for 

unavailable and not applicable characters. Click 'Help' for character 

information When done, choose 'Compare Unknown,..' from the Key menu. 

MCZ $6$10-§6$15. PANAMA^ Panama. Juan Diaz 

hsU ) : .3j.4 I orn( 11 > : .40, 40 m i ddr<2 1) : .9, .1.1 tOO ) ; ...L..2. 

snsc < 2 > : .7.,...!..! i p /e ( 1 2 ) : ..3 ,.,3 m i ddc ( 22 ) ; .,0.^.0 tc < 3 O : .0,...1 

pr < 3 > : ..6.,..5 i p /sc ( 1 3 ) : ...1..„,2 f s ( 23 ) : .,.L.2 pa ( 32 ) : ...L..3 

n < 4 > : ..3.,..3 i p /n < 1 4 > : .2 ^.4 y /d ( 24 ) : . .1.^ J. mdeiu C 33 ) : ...^..,1 

n-r < 5 > : ..0^.,0 so /s l< 1 5 > : „,!.,...1. ys /k < 25 > : ..3..3 f deiu C 34 ) : ..3.,..4. 

ssc<6 ) : ..OJ s l< 16 ) : ..5,..7 uj / i <25 ) : ..3..3 ms i zeC35 ) : .43.„.5g. 

sd(7):i.,j'III pm< 17):..6.,.5. fl /M< 27 ) : ..2.,..2 fs i ze(36): .4.4,47 

esc(8 ) : luilll'I s K 18 ) : ,,g,.0 K28 ) : ...1.3.,..1.5 1 1 /b I <37 ) : ..5,.,5 

scsCQ ) : .,2,..2. s I / i l< 19 > : „g,,,0 sd<29 ) : ..3.,,3 

Iorr<10>: 6 7 d(20): 2,2 Click HELP to see character definitions. 



OhELP (5) KEV O SUBSETS Q SPECIES 140 OF 404 \ i-ifili 



Figure 1 2. The Key card, with data from a series to be compared entered into 
the character fields. 



each HyperCard document is known as a "stack," and each stack 
is made up of one or more individual screens, or "cards" (Apple 
Computer, 1 989). The Anolis Handlist is a single HyperCard stack, 
and it consists of more than 400 Species cards (some of the species 
are undescribed or not well understood), a Key card, a Subset 
Editor card for use with the Key, and a Help card that provides 
general information as well as definitions of diagnostic characters 
(see Section V, earlier). 

Each Species card displays in the upper two fields the name, 
author, original citation, and type locality and in the bottom two 
fields two kinds of characters: (1) the "descriptors," those that 
are especially diagnostic, i.e., special for individual species, and 
(2) the characters that are routinely recorded for all species (Fig. 
1 1 ). Also on each Species card, provision is made for information 
on anatomy, behavior, color, ecology, and literature, and this 
information may be recorded and read on fields called up by 
choosing the corresponding button the middle row between the 
upper and lower fields. The Species cards may be browsed one 
by one using the navigational buttons in the lower right comer 



42 



BREVIORA 



No. 502 



Comparison Subset Editor for The nnolis Handlist 



flno 1 i s 


ach i 1 1 es TflVLOR 


rv 


Hnol is 


acutus HflLLOUELL 




flno 1 i s 


aeneus GRflV 




flno 1 i s 


aequo tor iai is UERNER 




Rno 1 1 s 


ogassizi STEJNEGER 




flno 1 1 s 


ainii BflRBOUR 




flno 1 i s 


olbi BflRBOUR 




flno 1 1 s 


albimaculatus HENLE 




flno 1 1 s 


alforoi GflRRIDO & HE 




flno 1 i s 


al iniger MERTENS 




flnol is 


alii oceus COPE 




flnol is 


al 1 isoni BflRBOUR 




flno 1 i s 


a 1 1 ogus BflRBOUR 8, Rfl 




flno 1 i s 


altae DUNN 


O 



[ >> €<ipy >> ] 



flnol i s cooki GRflNT 


i"^ 


flnol is cristate II us DUMERIL 




flnol is cuuieri flERREM 




flnol is desechensis HEflTUOLE 




flnol is ernes tali 1 1 i ams i LfiZE 




flnol is euermanni STEJNEGER 




flnol is gundlachi PETERS 




flnol i s krug i PETERS 




flnol is monensis STEJNEGER 




flnol is occultus WILLIflMS 8. 




flnol is poncensis STEJNEGER 




flnol is pulchellus DUMSRIL & 




flnol is scriptus GflRMflN 




flnol is stratulus COPE 


O 



Rll species 



[close Rll Species] 



Puerto Rico 
[ Saue Subset ] 



[ Clo 



se Subset 



OHELP OKEV ® subsets O species 388 OF 404 



r- i t 



W/' 



Figure 1 3. The Subset Editor card. The scroUing field on the left lists all the 
species in the Anolis Handlist, and the field on the right contains one of the 
comparison subsets. These subsets may be edited and saved for future use with 
the Key. 



of the cards, and they may be sorted according to species name, 
year of description, or any chosen diagnostic character by making 
the appropriate selection from a pull-down menu. Species cards 
may be added, deleted, or edited at the discretion of the user. 
Every scrolling field on each Species card can hold up to 30,000 
text characters; thus, the field for original citation could in fact 
contain a full synonymy, or the literature field an extensive bib- 
liography. All the data in the Species cards may be searched at 
will by selecting the FIND button (Fig. 1 1). 

The Key card (Fig. 12) permits the user to enter values for the 
diagnostic characters of an unknown specimen or series and to 
compare this unknown to all the Species cards or to a subset of 
the Species cards. When going to the Key card from a Species 
card, the option is provided to automatically read the data from 
that Species card into the Key as if it were an unknown, thus 
permitting comparisons among known species as well as among 
unknowns and knowns. Once the characters of a specimen or 



1994 COMPUTER IDENTIFICATION OF SPECIES 43 

species to be compared have been entered on the Key card, the 
user selects "Compare Unknown" from a pull-down menu and 
responds to a series of prompts asking, for example, by how much 
the unknown may differ from the reference data on the Species 
cards and still be considered a match. 

The Subset Editor (Fig. 13) is an adjunct to the Key card. 
Comparison of an unknown to all of the Species cards in the 



A report from The Anolis Handlist by E. E. Williams 
Sunday, April 10, 1994, 4:48 PM 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' compared with the 
subset ' 'PANAMA. ' ' 

• Anolis auratus DAUDIN 

CA, SA: eastern and central PANAMA, COLOMBIA, ECUADOR, VENEZUELA, 
BRAZIL. Small. Toepads not overlapping first phalanx. Light lat- 
eral line. Dewlap large, blue or black. Sharply enlarged keeled 
middorsals. In grass. 

' 'MCZ 86810-86815, PANAMA, Panama, Juna Diaz' ' misses by out 
of 37 key characters. 

• Anolis biporcatus (WIEGMANN) 

CA, SA: MEXICO, GUATEMALA, EL SALVADOR, NICARAGUA, COSTA RICA, 
PANAMA, COLOMBIA. Large. Heavy bodied. Green changing to dark 
brown. Dewlap moderate in male, white basally, mostly powder blue 
with red-orange free margin, in female smaller, sometimes with 
black flecks. In canopy. 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 5 out 
of 37 key characters : 

n(4) by 1, n-r(5) by -1, A/N(27) by 1, 1(28) by -1, tl/bl(37) by 

1 

• Anolis humilis PETERS 

CA: NICARAGUA, COSTARICA, PANAMA. Small, Brown. Dewlap large, red 
with bright yellow margin. Axillary pits. About 10 middorsal rows 
enlarged, larger than ventrals, the two median rows smallest. On 
or near ground. 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 5 out 
of 37 key characters : 

lorn(ll) by 1, d(20) by -1, v/d(24) by -2, A/N(27) by 1, tl/ 
bl(37) by 3 

• Anolis intermedius PETERS 

CA: COSTARICA, PANAMA. Small, Brown or greyish . Dewlap bone white. 
Arboreal . 



44 BREVIOR.4 No. 502 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 4 out 
of 37 key characters: 

lorn (11) by 13, so/sl(15) by 1, A/N(27) by 1 , tl/bl(37) by 2 

• Anolis kemptoni DUNN 

CA: PANAMA. Small . Greyish brown. White line under the eye . Dewlap 
skin red with orange anterior spot, scales whitish . Arboreal . 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz ' ' misses by 3 out 
of 37 key characters: 

so/sl(15) by 1, A/N(27) by 1, tl/bl(37) by 2 

• Anolis lemurinus COPE 

CA: MEXICO, GUATEMALA, HONDURAS, NICARAGUA, COSTARICA, PANAMA. 
Moderate size. Olive brown with dark dorsal blotches or ( females 
diamond- shaped middorsal markings or a black- edged middorsal 
stripe. Dewlapdark redwithblackscales . InPanamaonly in western 
and central regions. 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 5 out 
of 37 key characters: 

n(4) by 1, d(20) by -1, A/N(27) by 1, 1(28) by -1, tl/bl(37) by 

2 

• Anolis lionotus COPE 

CA: PANAMA. Moderate size. Light lateral line. A dorsal zone of 
about 10 rows of enlarged smooth scales much larger than ventrals. 
Dewlap large, orange. Semiaquatic. Only central Panama. 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 5 out 
of 37 key characters : 

n(4) by 1, d(20) by 1, v/d(24) by -2, A/N(27) by 1, tl/bl(37) by 

2 

• Anolis poecilopus COPE 

CA, SA: PANAMA, COLOMBIA. Moderate size. Light lateral line. Dew- 
lap large , orange . Head scales small . A dorsal zone of about 20 rows 
of enlarged keeled scales about as large as ventrals. Semiaquatic . 
In Panama only in eastern region, in Colombia only in western re- 
gion. 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 4 out 
of 37 key characters : 

snsc(2) by -3, n(4) by 1, A/N(27) by 1, tl/bl(37) by 2 

• Anolis tropidogaster HALLOWELL 

CA, SA: PANAMA, COLOMBIA. Small. Often an indication of a light 
middorsal stripe in both sexes. Dewlap yellow, orange or reddish. 
Trees and bushes. 



1994 COMPUTER IDENTIFICATION OF SPECIES 45 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 5 out 
of 37 key characters: 

n(4) by 2, ssc(6) by -1, d (20) by -1, A/N(27) by 1. tl/bl(37) by 
2 

• Anolis vittigerus COPE 

CA, SA: PANAMA, COLOMBIA. Moderate size . Variable and complex pat- 
ternonnape. Dewlap with central dark spot . In Panama only in east- 
ern region. 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 5 out 
of 37 key characters: 

middr(21) by -1, A/N(27) by 1, 1(28) by -1, mdew(33) by -1, tl/ 

bl(37) by 1 

• Anolis woodi DUNN 

CA: COSTARICA, PANAMA, Cordillera Talamanca . Large. Olive with 
indistinct rusty spots, Dewlap pink orange at edge, amber yellow 
in middle, bluish white at base. 

' 'MCZ 86810-86815, PANAMA, Panama, Juan Diaz' ' misses by 4 out 
of 37 key characters: 

n(4) by 1, ip/e(12) by 2 , A/N(27) by 1, tl/bl(37) by 1 



Figure 14. A Comparison Report from the Anolis Handlist. Reports may be 
edited and printed with the standard HyperCard facilities or copied into any word 
processor for editing and printing. 



Anolis Handlist may be time-consuming, and in most cases some 
additional information about the unknown (such as its collecting 
locality) or its species group will permit a comparison to be re- 
stricted to some subset of the total collection of species. The 
Subset Editor allows the user to assemble comparison subsets, 
either manually or automatically, to save these subsets for future 
use and to edit them as necessary. When making a comparison 
with the Key, the user may specify that the comparison be made 
against one of the existing subsets. 

The result of a comparison made with the Key is a Comparison 
Report (Fig. 14). This report specifies the name of the unknown 
that was compared, the name of the subset (if any) to which it 
was compared, the number of characters that differ between the 
unknowns and the knowns, and by how much these characters 
differ. The Comparison Report may be printed using the PRINT 



46 BREVIOR.4 No. 502 

FIELD option under FILE. (PRINT CARD will print only the 
portion of any field visible initially.) 

The Anolis Handlist has been designed specifically for the Mac- 
intosh computer. The principles it embodies are general ones, 
however, and these principles could be implemented on a variety 
of computer platforms. 

ACKNOWLEDGMENTS 

We acknowledge with special gratitude the struggles of two 
anonymous reviewers to catch errors, typographical and other, 
revise the logic of our discussions, and, with all good will, to 
generally emend and improve our paper. Nancy Knowlton and 
John Cadle saw a much later version. We also greatly appreciate 
their helpful comments. 

LITERATURE CITED 

Apple Computer. 1987. Human Interface Guidelines: The Apple Desktop In- 
terface. Reading, Massachusetts, Addison-Wesley Publishing Company. 

. 1989. HyperCard Stack Design Guidelines. Reading, Massachusetts, 

Addison-Wesley Publishing Company. 

BouLENGER, G. A. 1 885. Catalogue of the lizards in the British Museum (Natural 
History). London, Trustees of the British Museum, 2nd ed. 2: xiii + 497 pp. 

Knowlton, N. 1993. Sibling species in the sea. Annual Review of Ecology and 
Systematics, 24: 189-216. 

Lazell, J. D., Jr. 1983. Biogeography of the herpetofauna of the British Virgin 
Islands, with description of a new anole (Sauria: Iguanidae), pp. 99-1 17. In 
A. G. J. Rhodin and K. Miyata (eds.). Advances in Herpetology and Evo- 
lutionary Biology. Cambridge, Museum of Comparative Zoology. 

Morse, L. E., J. A. Peters, and P. B. Hamel. 1971. A general data format for 
summarizing taxonomic information. BioScience, 21: 174-181. 

Peters, J. A. 1964. Dictionary of Herpetology. New York, Hafner Publishing 
Company, ix + 392 pp. 

Peters, J. A., and B. B. Collette. 1968. The role of time-share computing in 
museum research. Curator, 11: 65-75. 

Peters, J. A., and R. Donoso-Barros. 1970. Catalogue of the Neotropical 
Squamata: Part II. Lizards and amphisbacnians. United States National Mu- 
seum Bulletm, 297: 1-293. 

Peters, J. A., and B. Orejas-Miranda. 1970. United States Museum, Part I. 
Snakes. United States National Museum Bulletin, 297: 1-347. 

Rand, A. S., and E. E. Williams. 1969. The anoles of La Palma: Aspects of 
their ecological relationships. Breviora, Museum of Comparative Zoology, 
327: 1-19. 



1994 COMPUTER IDENTIFICATION OF SPECIES 47 

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. Papeis Avulsos de 

Zoologia, Sao Paulo, 29(26): 259-268. 
— — . 1983. Ecomorphs, faunas, island size and diverse end points in island 

radiations of Anolis, pp. 326-370. In R. B. Huey, E. R. Pianka, and T. W. 

Schoener (eds.). Lizard ecology, studies of a model organism. Cambridge, 

Harvard University Press. 



B R E V I O R A 

LIBHARY 

Museiiim of ComparatiyjQ ZjOalpgy 

us ISSN 0006-9698 

Cambridge, Mass. 18 April 1996 UNlVt Number 503 

CYEMATID LARVAE OF THE LEPTOCEPHALUS HOLTI 

GROUP IN THE ATLANTIC AND PACIFIC OCEANS 

(PISCES: SACCOPHARYNGIFORMES) 

David G. Smith' and Michael J. Miller^ 

Abstract. Cyematid larvae of the Leptocephalus holti group consist of three 
distinct species or species groups, each found in the Atlantic and Pacific oceans. 
Species 1 has four gut loops and lacks pigment along the lateral midline. Species 
2 also has four gut loops, but it has lateral pigment. Species 3 has three gut loops 
and lacks lateral pigment. The name Leptocephalus holti is used as a convenient 
group name to refer to a complex of related species, none of which has been 
conclusively identified with an adult. Larvae of the Leptocephalus holti group may 
belong to Neocyema Castle, but this identification cannot yet be confirmed. 

INTRODUCTION 

The family Cyematidae is among the strangest and most highly 
modified of the deep-sea eels. Only the gulpers (Saccopharyngidae, 
Eurypharyngidae, and Monognathidae) exceed it in the degree of 
skeletal reduction. For nearly a century, the family was known 
from a single species, Cyema atrum Giinther, 1878, found in all 
oceans at depths of 1 ,500-3,000 m (Bertin, 1937:25). Castle ( 1 977) 
described a second genus and species, Neocyema erythrosoma, 
from two specimens collected in the South Atlantic Ocean. Evi- 
dence of a second cyematid species, however, had existed long 
before Castle's discovery in the form of an unidentified lepto- 
cephalus. The leptocephalus of Cyema atrum was first collected 
by the Michael Sars North Atlantic Expedition in 1910 and il- 
lustrated (but not named) by Murray and Hjort (1912, fig. 79). It 
was identified as Cyema atrum by Lea (1913:19), largely on the 



' Division of Fishes, National Museum of Natural History, Washington, D.C. 

20560. 

^ Department of Oceanography, University of Maine, Orono, Maine 04469-01 14. 



2 BREVIORA No. 503 

basis of the unusually low number of myomeres, and confirmed 
by Roule and Berlin (1929:108) through the discovery of meta- 
morphic specimens. Even before this, however, Schmidt (1909: 
6) described Leptocephalus holti from material collected by the 
Danish vessel Thor in the northeastern Atlantic. He made no 
attempt to identify it beyond speculating that it and some other 
leptocephali might represent "southern warm-water forms which 
have been taken at their northern limits in the 'Thor's' investi- 
gation." Larvae of the L. holti type were not reported again until 
Raju (1974:559) found a similar specimen in the South Pacific. 
Raju pointed out its resemblances to the larva of Cyema at rum 
and felt "compelled to relate it to an unknown species of the 
Cyemidae [sic]." Tabeta (1988:29) described two L. hoIti-\ikQ 
forms as "Cyematidae sp. 1" and "Cyematidae sp. 2"; species 1 
differed from species 2 and from Schmidt's and Raju's specimens 
in lacking the conspicuous midlateral pigment spots. Fortuno and 
Olivar ( 1986; also Olivar and Fortuno, 1991) reported a specimen 
collected in the South Atlantic off Namibia. They noted that their 
specimen lacked lateral pigment and speculated that this character 
might appear later in development. Smith (1989b:945) reported 
three additional specimens from the Sargasso Sea and the equa- 
torial Atlantic and agreed with Raju that they probably belonged 
to the Cyematidae. Smith's specimens also lacked midlateral pig- 
ment spots, and they had slightly fewer myomeres than Schmidt's 
holotype of L. holti. Based on the limited material available, he 
was unable to assess the significance of these differences. 

In this paper, we report on 47 additional specimens from both 
the Atlantic and Pacific oceans. These have revealed previously 
unsuspected diversity in several characters and allow us to give 
a more complete account of these distinctive larvae than has 
heretofore been possible. 

MATERIAL AND METHODS 

Most of our material (30 specimens) was collected during five 
cruises in the subtropical convergence zone of the Sargasso Sea 
between 1981 and 1989 (Kleckner et al, 1983; Kleckner and 
McCleave, 1988; Miller, 1993). These cruises were designed to 
study the spawning and larval distribution of the eel Anguilla 
rostrata. The other new Atlantic specimen was collected near 



1 996 LARVAE OF THE LEPTOCEPHALUS HOLTI GROUP 



Bermuda. Including the five previously recorded specimens 
(Schmidt, 1909; Fortufio and OHvar, 1986; Smith, 1989b), the 
total number of specimens known from the Atlantic is now 36. 
Of the 16 new Pacific specimens, 4 were found in collections at 
the Natural History Museum of Los Angeles County, 9 at Scripps 
Institution of Oceanography, and 3 at the National Marine Fish- 
eries Service Honolulu laboratory. With the nine previously re- 
corded specimens (Raju, 1974; Tabeta, 1988), 25 specimens are 
now know from the Pacific. Specimens examined are deposited 
in the Academy of Natural Sciences of Philadelphia (ANSP); Mu- 
seum of Comparative Zoology, Harvard University (MCZ); Nat- 
ural History Museum of Los Angeles County (LACM); National 
Museum of Natural History, Washington, D.C. (USNM); and 
Scripps Institution of Oceanography, La Jolla, California (SIO). 

Counts and measurements follow the methods of Smith ( 1 989a: 
665). Near the tip of the tail, myomeres become difficult to count, 
and in most cases only approximate counts were possible. The 
small size of most of our specimens made it difficult to obtain 
precise numerical values for any of the characters. The position 
of the last vertical blood vessel (LVBV) could not be seen clearly 
at the point where it entered the dorsal aorta in any of the spec- 
imens. We estimate this point to be on the average some six to 
eight myomeres anterior to a vertical line through the anus. Num- 
bers in parentheses following meristic values represent the num- 
ber of specimens on which the count is based. 

We use the term "" Leptocephalus hoUr in the sense of Orton 
(1964a; 199, 1964b:438) as a convenient group name to refer to 
what is apparently a complex of closely related species. In referring 
to the three distinct types (whether each represents a single species 
or a complex within the larger holti complex), we follow Tabeta 
(1988) in calling them species 1, species 2, and (newly described 
here) species 3. 

GENERAL DESCRIPTION OF LEPTOCEPHALUS HOLTI 
(AFTER SMITH, 1989b:946) 

Body moderately deep, depth about one-sixth to one-third stan- 
dard length (SL); body deepens gradually behind head. Gut with 
a distinct swelling at hepatogastric region and two or three loops 
or arches behind this; a compact liver lobe near 1 7th myomere, 



BREVIORA 



No. 503 






Figure 1. The Leptocephalus holti group. Top, species 1, MCZ 101007, 30 
mm SL; Middle, species 2, MCZ 101003, 26 mm SL; Bottom, species 3, MCZ 
101023, 25 mm SL. Drawn by L. Meszoly. 



contributing to swelling of gut; pancreas compact, located just 
posterior to liver and gall bladder; dorsal aorta sending several 
conspicuous vertical blood vessels that enter a parallel ventral 
vessel that lies distinctly above the gut. Dorsal fin begins ap- 
proximately 20 myomeres anterior to anus. Head and snout long; 
eye located posteriorly, close to anteriormost myomeres; snout 
long and pointed, profile relatively flat; nasal capsule small. Sev- 
eral expanded melanophores sometimes present on lateral mid- 
line. Moderately large melanophores on gut. One to four mela- 
nophores sometimes present near dorsal margin of body, in clear 
area above myomeres. Pigment usually present at anterior tip of 



1 996 LARVAE OF THE LEPTOCEPHALUS HOLTI GROUP 



•40 



20  



• 



20 . 




40 



Figure 2. Distribution of Leptocephalus holti and Neocyema erythrosoma in 
the Atlantic. Square = species 1; circle = species 2; triangle = species 3; cross = 
Neocyema erythrosoma. 



snout and lower jaw. Maximum size unknown, though probably 
not large. Largest specimen known 43 mm SL; all specimens 
premetamorphic. 

Species 1 
Figures 1 (top), 2, 3 

Diagnosis. Four gut loops, including hepatogastric swelling. No 
pigment on side of body along lateral midline. One to three me- 
lanophores near dorsal margin of body above myomeres. Paired 
melanophores laterally on gut adjacent to pectoral fin and pos- 
teriorly between third and fourth gut loops; a single or complex 
melanophore dorsal to each gut loop. Pigment at tip of snout and 



BREVIORA 



No. 503 




Figure 3. Distribution of Leptocephalus holti in the Pacific. Square = species 
1; circle = species 2; triangle = species 3. 



lower jaw. Myomeres: total ca. 99-117 (15 specimens), preanai 
45-65 (20). 

Size. Ca. 10-39 mm SL, all premetamorphic. 

Variation. All but two of the Atlantic specimens came from 
the Sargasso Sea, the others from off the west coast of Africa (Fig. 
2). The latter had approximately 99-105 total myomeres com- 
pared to ca. 108-1 17 for the western Atlantic specimens. There 
seem to be no other differences between the eastern Atlantic and 
western Atlantic specimens. Tabeta (1988:29) gave a range of 97- 
100 total, 51-62 preanai, and 46-49 LVBV myomeres for his 
seven western Pacific specimens, 16-31 mm in length. The single 
central Pacific specimen examined, USNM 324871, had signifi- 



1996 LARVAE OF THE L£^/'rOC£'/'///lLC/5//OLr/ GROUP 7 

cantly fewer preanal myomeres (ca. 45) than either the Atlantic 
specimens (ca. 49-65) or the western Pacific specimens (51-62). 
Material Examined. Atlantic (25, ca. 9-39 mm SL): MCZ 64484 
(1, 31), 34°27.0'N, 71°18.5'W, 250-0 m, 13 Apr 1977. 65647 (1, 
20), 4°05.2'N, 17°20.8'W, 75 m, 15 Nov 1978. 101005(1, <10), 
24°19.5'N, 70°24.5'W, 280 m, 27 Feb 1981. 101006 (1, 34), 
25°10.3'N, 71°33.0'W, 318 m, 13 Feb 1983. 101007 (1, 30, il- 
lustrated), 26°25.rN, 71°17.4'W, 280 m, 14 Feb 1983. 101008 
(2, ca. 12-ca. 22), 26°20.3'N, 71°18.0'W, 232 m, 14 Feb 1983. 
101009 (1, <15), 25°41.6'N, 71°31.0'W, 132 m, 15 Feb 1983. 
101010(1), 24°47.1'N, 70°27.0'W, 356 m, 17 Feb 1983. 101011 
(1, ca. 12), 24°11.4'N, 70°25.2'W, 303 m, 18 Feb 1983. 101012 
(1, ca. 22), 26°20.2'N, 74°12.5'W, 112 m, 26 Feb 1983. 101013 
(1, 19), 27°52.0'N, 66°45.7'W, 261 m, 3 Apr 1983. 101014 (1, 
39), 26°44.9'N, 66°38.8'W, 260 m, 4 Apr 1983. 101015 (1, ca. 
11), 29°56.4'N, 68°58.2'W, 298 m, 16 Mar 1985. 101016 (2, ca. 
9-ca. 25), 27°04.7'N, 70°03.4'W, 134 m, 13 Feb 1989. 101017 
(1, 11), 27°21.6'N, 70°12.3'W, 299 m, 14 Feb 1989. 101018 (5, 
13-15), 27°02.1'N, 73°59.7'W, 304 m, 16 Feb 1989. 101019 (1, 
13), 26°33.6'N, 73°53.9'W, 318 m, 19 Feb 1989. 101020 (1, < 10), 
26°42.7'N, 73°59.4'W, 302 m, 20 Feb 1989. 101021 (1, 19), 
26°14.3'N, 73M9.3'W, 300 m, 21 Feb 1989. Note: Another spec- 
imen, MCZ 101026 (<15 mm), probably belongs here, but it is 
badly damaged and we cannot determine the number of gut loops. 
Pacific (1, 9 mm SL): USNM 324871 (1, 9), 29°48'00"N, 
179°03'54"E, 50-100 m, 9 Feb 1985. 

Species 2 
Figures 1 (middle), 2, 3 

Diagnosis. Atlantic specimens (including data from holotype, 
Schmidt, 1909): Four gut loops. Five expanded melanophores 
along lateral midline at myomeres 14-16 (4 specimens), 29-31 
(4), 44_48 (4), 57-65 (4), 71-78 (4), centered below surface and 
often extending onto body wall on one side or other; two to four 
melanophores near dorsal margin of body, in clear area above 
myomeres. Myomeres: total ca. 108-ca. 130 (4), preanal 65-75 
(4). 

Pacific specimens: Four gut loops. Four or five expanded lateral 
melanophores, at myomeres 12-19 (14), 25-38 (14), 42-53 (14), 



BREVIORA No. 503 



53-68 (13), 61-75 (7); one or two dorsal melanophores; other 
pigment as in Atlantic specimens. Myomeres: total ca. 100-1 10 
(9), preanal 57-70(11). 

Size. Atlantic specimens 23-35 mm SL, Pacific specimens ca. 
19-43 mm; all premetamorphic. The specimen reported by Raju 
(1974) was given as 40 mm; we remeasured it as 37 mm. 

Variation. Three of the four Atlantic specimens came from the 
Sargasso Sea, the other (the holotype of Leptocephalus hold) from 
the northeastern Atlantic south of Ireland (Fig. 2). Despite its 
geographic separation from the others, the holotype shows no 
obvious differences from the three western Atlantic specimens. 
The holotype and MCZ 101003 have fewer total myomeres (ca. 
108-112) than MCZ 101002 and 101004 (ca. 120-130 and ca. 
128). The former pair also has fewer preanal myomeres (65-67 
vs. 74-75). In one specimen (MCZ 101003), the last vertical blood 
vessel enters the kidney slightly more anteriorly than in the others, 
i.e., in the trough between the third and fourth gut loops instead 
of near the top of the fourth loop. Another specimen (MCZ 1 1 002) 
has extra ventral melanophores, between the first-second and 
second-third gut loops. With the limited material available and 
the difficulty of obtaining precise myomere counts, we are unable 
to assess the significance of these differences. 

Thirteen of the 1 5 Pacific specimens came from an area north 
to northeast of the Hawaiian Islands, one came from Southeast 
Hancock Seamount in the central North Pacific, and one from 
the South Pacific, southwest of the Austral Islands (Fig. 3). The 
South Pacific specimen is at the low end of the range of a few 
meristic characters (preanal myomeres, position of some lateral 
melanophores), but the only character that is clearly outside the 
range of the other specimens is the position of the fifth lateral 
melanophore (61-62 vs. 64-75). Seven specimens have four lat- 
eral melanophores, seven others have five, and one has three. 
Tabeta's (1988) specimen has five lateral melanophores, and its 
total and preanal myomere counts (99 and 59, respectively) fall 
within the range of our specimens. The Pacific specimens appear 
to have fewer total myomeres (99-110) and preanal myomeres 
(57-70) than the Atlantic specimens (ca. 108-130 and 65-75, 
respectively). The position of the first four lateral melanophores 
coincides in the Atlantic and Pacific specimens. Only the fifth 



1 996 LARVAE OF THE LEPTOCEPHALUS HOLTI GROUP 



appears to differ, at myomere 61-75 in the Pacific vs. 71-78 in 
the Atlantic specimens. All four Atlantic specimens have five 
lateral melanophores, whereas more than half of the Pacific spec- 
imens examined by us have only three or four. 

Material Examined. Atlantic (3, 23-26 mm SL): MCZ 101003 
(1, 26 illustrated), 28°31.4'N, 69'02.1'W, 475 m, 4 Mar 1981. 
101002 (1, 23), 26°59.7'N, 68°52.0'W, 150 m, 23 Mar 1985. 
101004 (1, 23), 31°27.0'N, 64°21.0'W, 9 Apr 1990. Pacific (15, 
19-45 mm SL): LACM 36437-1 (1, 28), 26°32'N, 147°13'W, 0- 
160 m, 10 Apr 1966. 36438-2 (1, ca. 21), 26°32'N, 147°13'W, 
surface, 10 Apr 1966. 36447-4 (1, ca. 24), 27°55'N, 144°10'W, 
11 Apr 1966. 36454-3 (1, 22), 28°48'N, 141°59'W, surface, 12 
Apr 1966. SIO 70-1 18 (1, 37, specimen described by Raju, 1974), 
24°30.5'S, 154°54'W, 0-175 m, 4 Oct 1969. 89-57 (2, 42-43), 
31°N, 159°W, 200-0 m, 13-14 Apr 1989. 89-63 (4, 19-39), 31°N, 
159°W, 200-0 m, 18 Apr 1989. 89-65 (2, 42-42), 31°N, 159°W, 
400-0 m, 19 Apr 1989. 89-68 (1, 34), 31°N, 159°W, 0-900 m, 
22 Apr 1989. USNM 324872 (1, 26), 29°49'46"N, 179°07'54"E, 
0-100 m, 20 Apr 1987. 

Species 3 
Figures 1 (bottom), 2, 3 

Diagnosis. Three gut loops, including hepatogastric swelling. 
Lateral and dorsal pigment absent; paired melanophores on lateral 
surface of gut adjacent to pectoral fin; a melanophore dorsally 
and one on each side of hepatogastric swelling; a complex me- 
lanophore dorsally on the two posterior gut loops, extending lat- 
erally on each side of gut; no melanophore between second and 
third gut loops; pigment present at tip of snout and lower jaw. 
Myomeres: total ca. 104-1 15 (4), preanal ca. 54-57 (6). 

Size. Largest specimen ca. 25 mm; all specimens premetamor- 
phic. 

Variation. Five specimens came from the Sargasso Sea and one 
from the central North Pacific. Significant variation is not evident. 

Material Examined. Atlantic (5, 16-ca. 25 mm SL): ANSP 
153490 (1, 19), 21°03'N, 57°54'W, 0-150 m, 30 Mar 1979. MCZ 
101022(l,23),26°17.1'N,66M4.6'W,253m,8Apr 1983. 101023 
(1, 25, illustrated), 26°17.0'N, 66°45.0 W, 150 m, 9 Apr 1983. 
101024 (1, 23), 28°31.4'N, 69°02.1'W, 302 m, 17 Mar 1985. 



10 



BREVIORA 



No. 503 




Figure 4. Leptocephalus of Cyema atrum (after Smith, 1989b). 



101025 (1, 16), 27°02.1'N, 73°59.7'W, 304 m, 16 Feb 1989. Pa- 
cific (1, 10 mm SL): USNM 324783 (1, 10), 29°47'36"N, 
179°03'54"E, 50-100 m, 9 Feb 1985. 

IDENTIFICATION AND RELATIONSHIPS 

Leptocephalus holti and the larva o{ Cyema atrum (Fig. 4) share 
the following characters: a long, peg-like snout with a straight 
profile; a posteriorly placed eye; a gut with an anterior swelling 
at the hepatogastric region followed by two to four arches or loops; 
pigment dorsally on each gut loop; pigment near the dorsal margin 
of the body; an acute tail without distinct hypural elements; a 
large ventral blood vessel conspicuously separated from the gut 
tube; and V-shaped myomeres with a highly obtuse angle at the 
midlateral line. These characters distinguish Cyema atrum and 
L. holti from all other leptocephali and support the hypothesis 
that they belong to the same family. Cyema atrum (Fig. 4) has a 
deeper body than L. holti with a steeper anterior profile, it has 
an expanded mass of pancreatic tissue that fills much of the space 
between the dorsal margin of the intestine and the ventral margin 
of the myomeres, and its lateral pigment is scattered over the side 
of the body instead of being restricted to the midline. 

If L. holti is accepted as a cyematid, which cyematid is it? Both 
Castle (1977:75) and Smith (1989b:947) have made the obvious 
suggestion that L. holti is the larva of Neocyema, but they con- 



1996 LARVAE OF THE LEPTOCEPHALUS HOLTI GROUP 1 1 



sidered such an identification inconclusive. Particularly trouble- 
some was the absence of any trace of the conspicuous lateral 
melanophores in the specimens of Neocyema, despite the semi- 
leptocephaloid appearance of the latter. Bertin (1937:17) showed 
that the lateral pigment of the leptocephalus was retained in a 
1 1 5 -mm juvenile Cyema at mm. Both the holotype of L. holti and 
Raju's Pacific specimen had conspicuous lateral pigment. Castle 
in particular felt that the pigment character made an identification 
of L. holti with Neocyema unlikely. Smith agreed but pointed out 
that the age of the Neocyema specimens was unknown and that 
at least some specimens of L. holti lacked lateral pigment. 

The present material further reduces the objections to identi- 
fying L. holti as the larva of Neocyema. It is now clear that the 
majority of Atlantic L. holti lack lateral pigment, so the main 
obstacle has been removed, at least in theory. The number of 
myomeres is in the same range (Castle [1977] reported that the 
one intact specimen of Neocyema erythrosoma had 108 total 
myomeres). Although the identification cannot be disproved, it 
cannot be confirmed, either, especially in the absence of meta- 
morphic specimens. The only known specimens of Neocyema 
were taken far from the known range of L. holti, but lack of 
adequate collecting weakens this objection. After all, for more 
than 60 years L. holti was known from a single specimen. 

Larval fishes can provide important and valuable information 
that is not available from the study of adults alone. Regardless 
of whether L. holti is the larva of Neocyema or another still 
unknown genus, these larvae enable us to say without question 
that such a genus exists, it contains at least three species, and it 
is found in all oceans. 

ACKNOWLEDGMENTS 

We thank Dr. J. D. McCleave for access to the specimens from 
the University of Maine's Sargasso Sea collections. The following 
provided the opportunity to examine specimens in their collec- 
tions: R. J. Lavenberg (LACM), R. H. Rosenblatt (SIO), and B. 
C. Mundy (National Marine Fisheries Service, Honolulu). The 
collection and identification of the University of Maine specimens 
were made possible with financial support from the National 



12 BREVIORA No. 503 

Science Foundation (OCE-77 19440, OCE-8208394, and OCE- 
88 1 1005). Publication costs were covered in part by a grant from 
the Wetmore-Colles Fund. 

LITERATURE CITED 

Bertin, L. 1937. Les poissons abyssaux du genre Cyema Giinther (anatomic, 

embryologie, bionomie). Dana Report, 10: 1-30. 
Castle, P. H. J. 1977. A new genus and species of bobtail eel (Anguilliformes, 

Cyemidae) from the South Atlantic. Archiv fur Fischereiwissenschaft, 28(2/ 

3): 69-76. 
FoRTUNO, J.-M., AND M.-P. Olfvar. 1 986. Larvas de anguilliformes capturadas 

in el Atlantico sudoriental. Miscellanea Zoologica, 10: 223-231 (not seen). 
Kleckner, R. C, and J. D. McCleave. 1988. The northern hmit of spawning 

by Atlantic eels {Anguilla spp.) in the Sargasso Sea in relation to thermal 

fronts and surface water masses. Journal of Marine Research, 46: 647-667. 
Kleckner, R. C, J. D. McCleave, and G. S. Wippelhauser. 1983. Spawning 

of American eel, Anguilla rostrata, relative to thermal fronts in the Sargasso 

Sea. Environmental Biology of Fishes, 9: 289-293. 
Lea, E. 1913. Muraenoid larvae from the "Michael Sars" North Atlantic Deep- 

Sea Expedition, 1910. Report on the Scientific Results of the "Michael Sars" 

North Atlantic Deep-Sea Expedition, 3(1): 1-59. 
Miller, M. J. 1993. Species assemblages of leptocephali in the Sargasso Sea and 

Florida Current. Ph.D. Dissertation, University of Maine, Orono. 
Murray, J., andJ. Hjort. 1912. The Depths of the Ocean. London; Macmillan. 

XX + 821 pp. 
Olivar, M.-P., and J.-M. Fortuno. 1991. Guide to the ichthyoplankton of the 

southeast Atlantic (Benguela Current region). Scientia Marina, 55(1): 1-383. 
Orton, G. L. 1964a. Identification oi Leptocephalus acuticeps as the larva of 

the eel genus Avocettina. Pacific Science, 18(2): 186-201. 
. 1 964b. New information on a rare eel larva. Leptocephalus hyoproroides 

Stromman. Copeia, 1964(2): 434-444. 
Raju, S. N. 1974. Three new species of the genus Monognathus and the lep- 
tocephali of the order Saccopharyngiformes. Fishery Bulletin, 72(2): 547-562. 
RouLE, L., AND L. Bertin. 1 929. Les poissons apodes appartenant au sous-ordre 

des Nemichthyidiformes. Oceanographical Reports of the Danish "Dana" 

Expeditions 1920-22, 4: 1-1 13. 
Schmidt, J. 1 909. On the occurrence of leptocephali (larval muraenoids) in the 

Atlantic W. of Europe. Meddelelser fra Kommissionen for Havundersogelser, 

Ser. Fiskeri, 3(6): 1-19. 
Smith, D. G. 1989a. Introduction to leptocephali. In E. B. Bohlke (ed.). Fishes 

of the Western North Atlantic. Memoirs of the Sears Foundation for Marine 

Research, 1(9): 657-668. 
. 1989b. Family Cyematidae: leptocephali. In E. B. Bohlke (ed.). Fishes 

of the Western North Atlantic. Memoirs of the Sears Foundation for Marine 

Research, 1(9): 944-947. 
Tabeta, O. 1988. Anguilliformes. In M. Okiyama (ed.). An Atlas of the Early 

Stage Fishes in Japan. Tokyo, Tokai University Press. 1 154 pp (in Japanese). 



B R E V I O R A 

LIBRARY 

usenm of Comparative Zoology 

^ APR 3 1996 

us ISSN 0006-9698 



Cambridge, Mass. 18 April 1996 :'_^' Number 504 

i Vi:LF^...Jl j 1 

THE GENUS PHENACOSAURUS 
(SQUAMATA: IGUANIA) IN WESTERN VENEZUELA: 

PHENACOSAURUS TETARII, NEW SPECIES, 

PHENACOSAURUS EUSKALERRIARI, NEW SPECIES, 

AND PHENACOSAURUS NICEFORI DUNN, 1944 

Tito R. Barros,' Ernest E. Williams,- and Angel Viloria' 

Abstract. Two new, possibly parapatric species of Phenacosaurus are de- 
scribed from the Sierra de Perija, Estado Zulia, Venezuela: P. tetarii, larger, at 
least 85 mm snout-vent length, with heterogeneous squamation on the flanks, 
closest to P. nicefori but larger in adult size, and P. euskalerriari, small, about 56 
mm snout-vent length, with uniform flank squamation, closest to P. orcesi, but 
with larger flank scales and shorter interparietal. 

Phenacosaurus nicefori Dunn is redescribed on the basis of material from Be- 
tania, Estado Tachira, Venezuela, and from the Paramo de Tamo, overlapping 
the borders of both Colombia and Venezuela, as well as from topotypic material 
from Norte de Santander in Colombia. 

INTRODUCTION 

The members of the genus Phenacosaurus are anoline lizards 
endemic to the subparamo and paramo of northwestern South 
America and are characterized by a casqued head with converging 
lateral parietal crests with a smaller or larger notch at their oc- 
cipital ends, a variable vertebral crest, short limbs, and a probably 
prehensile tail. Initially they were believed to be present only in 
Colombia, from where the type species, Phenacosaurus heteroder- 
mus, was described by Dumeril and Dumeril in 1851. Dunn 
(1944), studying material in Colombian collections, added two 
new species to the genus, P. nicefori from the Department of Norte 
de Santander and P. richteri from the Department of Cundina- 



' Museo de Biologia, Facultad de Ciencias, Universidad del Zulia, Apartado 526, 
Maracaibo 4033, Zulia, Venezuela. 

^ Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 



2 BREVIORA No. 504 

marca. Hellmich ( 1 949) described as P. paramoensis a single spec- 
imen from the paramo overlapping the borders of the Depart- 
ments of Cundimamarca, Huila, and Meta. Lazell (1969), in a 
revision using most of the known specimens of the genus, syn- 
onymized P. richteri and P. paramoensis with P. heterodermus 
but added the species P. orcesi on the basis of two specimens 
from Ecuador. The first giant species of the genus from the Cor- 
dillera Oriental of Colombia was named P. inderenae by Rueda 
and Hemandez-Camacho (1988). Williams and Mittermeier (1991) 
cited a juvenile from Venceremos in the Department of San Mar- 
tin in Peru as a possible third specimen of P. orcesi. 

At the present time, several additional new species of the genus 
have been described or are in the process of description. From 
Venezuela, Myers et al. (1993) described a small series of one 
new species {P. neblininus) from the Cerro La Neblina in the 
extreme south, and Williams et al. (1996b) described an addi- 
tional new species based on a single specimen collected by S. 
Gorzula and A. Farrera in the Macizo del Chimanta Tepui in 
Estado Bolivar. A second giant species very similar to P. inderenae 
has been collected in La Alegria and adjacent localities in the 
Provincia de Sucumbios in Ecuador and was described by Wil- 
liams et al. (1996a). 

As the result of three expeditions to the Sierra de Perija in 
Estado Zulia, Venezuela, in 1989 and 1991 by the Museo de 
Biologia of the Universidad del Zulia (MBLUZ), four specimens 
of Phenacosaurus (MBLUZ R-2 1 5 and R-308, MCZ 1 76474, and 
176475) represent two new species. One species is represented by 
a male from the Paramo de Tetari and a female from the Cerro 
Pintado. A specimen (MHNLS 664) of this new species had al- 
ready been collected by Ramon Urbano in 1952 but was referred 
to P. nicefori by Aleman (1953) and Lazell (1969). The other is 
a smaller species, again a male and a female, related to P. orcesi. 

In addition, with the intention of completing a synopsis of the 
species of Phenacosaurus in western Venezuela, the material of 
P. nicefori from Betania, Estado Tachira, and Paramo del Tama, 
which overlaps both the Department of Norte de Santander of 
Colombia and Estado Tachira, Venezuela, and also topotypic 
specimens from the Department of Norte de Santander, Colom- 
bia, are reported and described. 



1996 PHENACOSAURUS m y^ESTERN VENEZUELA 3 

The descriptions of the new species herein are a slightly mod- 
ified version of the format utilized by Williams for lizards of the 
genus Anolis (for the nomenclature of the scales, see Williams et 
a!., 1996a,b). 

DESCRIPTION 

Phenacosaurus tetarii, new species 

Holotype. MBLUZ R-215, adult male, collected by Angel Vi- 
loria, October 24, 1989, on the roads that lead to the Paramo del 
Tetari, Sierra de Perija, Estado Zulia, Venezuela (10°06'34"N, 
72°53'00"W), 2,790 m elevation. 

Paratypes. MHNLS 664, adult male, collected by Ramon Ur- 
bano, 1952, at the base of Pico Tetari, Sierra de Perija, Estado 
Zulia, Venezuela, 2,900 m elevation; MCZ 176474, adult female, 
collected by Angel Viloria, March 24, 1989, at the base of Cerro 
Pintado, Sierra de Perija, Estado Zulia, Venezuela, 2,400 m el- 
evation. 

Etymology. Tetari is the name that the Yukpa Indians have 
given to the second highest peak in the Serrania de Perija (3,575 
m elevation). 

Diagnosis. A phenacosaur closest to P. nicefori but differing in 
the greater snout-vent length (SVL of tetarii: holotype male 80 
mm male, paratype 86 mm; female 70 mm, rather than a max- 
imum SVL of males 57 mm and females 58 mm in nicefori), with 
the parietal area converging to a very narrow notch at its occipital 
margin, and differing also in the dewlap color {tetarii: yellow 
rather than white with pale orange stripes in nicefori). 

Description. The description is based primarily on the male 
holotype and the male paratype; the differences of the female 
paratype are mentioned whenever visible. 

Head: A casque well developed. The head scales anteriorly 
mostly smooth, but strongly pustular posteriorly in the region of 
the parietal table. 

Dorsal head scales (Fig. 7^1— Antorbital area: Canthals 4 on both 
sides in both males, 4 on the left side in the female paratype, the 
other side obscure. In the male holotype, the anteriormost canthal 
separated from the circumnasal by 3 small scales on the left side, 
by 2 scales on the right side. In the male paratype in this area at 



BREVIORA 



No. 504 




Figure 1 . Map of the localities mentioned in this paper. 



least 6 small scales on the left, 5 on the right. In the female 
paratype 2 scales on the left, 1 on the right. Four scales between 
the second canthals in all specimens. Rostral wider than high, 
bordered posteriorly by 4 or 6 postrostrals, 2 of which are in 
contact with the first supralabials. The circumnasal on each side 
in contact with the first supralabial and separated from the rostral 
by the postrostral that overlies the sulcus between the first su- 
pralabial and rostral. In the male holotype dorsally between the 
circumnasals, a single large scale in a central position posterior 
to the postrostrals, flanked by 2 smaller scales. In the female 
paratype 2 subequal scales in this position. In the male paratype 
6 scales in 2 rows. Between the second canthals 2 (male holotype) 



1996 P//£-iV/lC05'/lt//?C/5'IN WESTERN VENEZUELA 5 

the largest scales of the head and, indeed of the entire animal, 3 
(male and female paratypes). 

Orbital area: The supraorbital semicircles separated by 1 row 
of moderately sized subrectangular scales in the holotype male. 
In the male paratype 1 anterior row in contact. In the female 
paratype 2 posterior pairs in contact. In all specimens both the 
scales of the semicircles and those of the median row being bluntly 
tubercular or ridges that may be coalesced tubercles. All the scales 
of the supraocular area much smaller than those of the semicircles, 
quite smooth, and differing much in size, larger medially, smaller 
laterally. The largest, also the most medial on both sides, in nar- 
row contact with the semicircles in the male holotype on the right. 
In the male paratype all, including the largest, of the medial su- 
praoculars in contact with all the semicircles. The female paratype 
obscure. 

Parietal area: Low ridges separating the rounded convex su- 
pratemporal area and the relatively depressed and flat parietal 
table, converging straight back to end in low bosses that leave a 
narrow notch between them. The scales of the ridges very bluntly 
keeled and intermediate in size between the mostly large smooth 
scales of the supratemporal area and the uniformly small and 
strongly pustulate scales of the parietal table. 

The interparietal scale, inferred to be such because of its shape 
and anterior medial placement, small, rhomboid in the male ho- 
lotype, more irregular in shape in the female and male paratypes, 
all specimens without a parietal eye. In the male holotype and 
the male paratype interparietal separated from the supraorbital 
semicircles by 1 scale on each side, almost as large or larger than 
itself. In the female the interparietal in contact with both adjacent 
scales of the semicircles. Four or 5 scales between the interparietal 
and the nape scales. 

Lateral head scales (Fig. 2j— The lateral head scales all smooth 
except for the lateral faces of the posterior canthals, which are 
pustular like their dorsal surfaces. 

One to 3 rows of loreal scales, counted just in front of the 
preoculars. The anterior of these always in single rows. The total 
number of loreals only 6 on 1 side, 7 on the other in the male 
holotype, the male paratype 6 on both sides, 8 on both sides in 
the female paratype. Two preoculars on each side in all specimens. 



BREVIORA 



No. 504 




1996 



PHENACOSAURUS IN WESTERN VENEZUELA 




Figure 3. Phenacosaurus tetahi, holotype, MBLUZ R-215: Lateral view of head. 



Eight supralabials to below the center of the eye in the male 
holotype, 9 in the male paratype, 10 in the female paratype. Six 
to 9 postoculars on each side arch round the back of the orbit, 
the uppermost in contact dorsally with the posteriormost scale of 
semicircle of its side and laterally with the posteriormost super- 
ciliary. 

The temporal region divided into supra- and infratemporal by 
an intertemporal ridge, covered by 2-4 scales. The ridge strongly 
convex but not shelf-like. 

The supratemporals larger, but variable in size close to the 
parietal ridges, smaller near the intertemporal ridge. The infra- 
temporals are larger close to the intertemporal ridge, smallest in 
a narrow zone in the center of the region, then again larger, almost 
as large as the uppermost infratemporal scales in a band from the 
ear to the comer of the mouth. 

The ear small, inconspicuous, smaller than many of the scales 
surrounding it, smaller also than the inferred interparietal. 

Ventral head scales (Fig. iy*— The mental incompletely divided, 
in contact with only 2 diiferentiated sublabials between the in- 



Figure 2. Phenacosaurus tetarii, holotype, MBLUZ R-215: Dorsal view of head. 



BREVIORA 



No. 504 




Figure 4. Phenacosaurus tetarii, holotype, MBLUZ R-215: Ventral view of head. 



1996 



PHENACOSAURUS IN WESTERN VENEZUELA 




Figure 5. Phenacosaurus tetarii, holotype, MBLUZ R-215: Lateral view of 
entire animal. 



fralabials (male holotype and male paratype). Five sublabials on 
each side in contact with the infralabials in the male holotype, 
the first sublabials only by their comers. In the male paratype 2 
sublabials are in contact with the infralabials, 3 on the left. The 
female paratype obscure. The swollen medial gulars grading pos- 
teriorly into swollen gulars less than half their size. 

Trunk (Figs. 4 and 5): The middorsal scales, a dorsal crest of 
Type 3 sensu Lazell (1969, fig. 1), a single series of vertebral scales, 
swollen keeled cones, at irregular intervals each such scale sep- 
arated from other vertebrals by a pair of flat or slightly swollen 
paravertebrals that meet middorsally. In the male the sequence 
of the 2 types of middorsals is nearly regular until the region of 
the sacrum, where the cones are in contact. About 4-5 rows of 
paravertebrals, rounded and flat, varying somewhat in size, in 
contact or slightly overlapping and showing little or no intervening 
skin. Below this on each flank a zone of distinct granules, mostly 
fully separating round flat scales that differ little from the par- 
avertebrals in size but tend to be somewhat smaller. Below this, 
beginning at about the middle of the flank patches of slightly 
larger round flat scales in partial contact appear; they appear to 
be concentrated in irregular areas that are more lightly pigmented 
than the rest of the flank. Ventrals smooth and strongly imbricate, 



10 



BREVIORA 



No. 504 




Figure 6. Phenacosaurus tetarii, holotype, MBLUZ R-2 1 5: Flank scales behind 
shoulder. 



bluntly pointed and about equivalent in size to the round flat 
scales of the flanks. 

Limbs (Fig. 4): All limb scales smooth, differing primarily in 
size. The scales of the anterior face of upper arm and of the thigh 



Figure 7. Phenacosaurus tetarii, male paratype, MHNLS 664: Dorsal view of 
head. 



1996 



PHENACOSAURUS IN WESTERN VENEZUELA 



11 




12 



BREVIORA 



No. 504 




Figure 8. Phenacosaurus tetarii, male paratype, MHNLS 664: Lateral view of 
head. 



distinctly larger than those of the posterior face, especially on the 
thigh, where the posterior scales are almost granular. The scales 
of the lower arm and tibia not differing much in size on the 
anterior and posterior surfaces. The digital scales of both hands 
and feet with the dorsal and ventral surfaces lamellar— wider than 
long, and overlapping distally— throughout the length of the digits. 
The adhesive lamellae under phalanges ii and iii of the fourth toe 
23 in the male, not determinable in the female. 

Tail (Fig. 4): Tail curved and apparently prehensile. All caudal 
scales, except those at the very tail base, keeled, most sharply 
ventrally. At the base the crest scales keeled cones as large as and 
very similar to those of the middorsal sacral area. Farther back 
on the tail the crest scales become more elongate, lower, and 
smaller and toward the end of the tail indistinguishable from the 
other scales of the tail except that they are on the middorsal line 
of the tail. As on the dorsum, the series of caudal crest scales is 
interrupted by the medial juncture of paravertebral caudals, but 
in this case the paravertebrals are keeled and the interruptions 
are few and at highly irregular intervals. 

Dewlap (Fig. 4): Dewlap in both sexes relatively small, extend- 
ing posteriorly a little farther than the level of the axilla, densely 
scaled, the swollen scales about the size of the ventrals. 

Color in Life. (The patterns shown in Figure 4 are long after 



1996 PHENACOSAURUS IN WESTERN VENEZUELA 



13 




Figure 9. Phenacosaurus tetarii, male paratype, MHNLS 664: Ventral view of 
head. 



14 



BREVIORA 



No. 504 




Figure 10. Phenacosaurus tetarii, male paratype, MHNLS 664: Lateral view 
of entire animal. 



preservation.) The color of the holotype was described shortly 
after preservation, when the darker tones had been accentuated 
by the preserving liquid. From the parietal region to the tip of 
the tail, the dorsum was dark brown (sepia No. 119) (Smithe, 
1975). There were five rounded blotches on the dorsum from the 
nape to the insertion of the hindlimbs, the distances between these 
spots being about 5 mm. These spots were light cream (light drab 
No. 1 1 9C), but reference to photographs (slides) of the live animal 
shows that in life the color was much more yellow (buff-yellow 
No. 53), as was the color of the belly and the throat. The head 
from the tip of the snout to the beginning of the parietal region 
was a dark olive green (grayish yellow No. 430). The limbs have 
the color of the dorsum but with some transverse bars of an orange 
color (ferruginous No. 41), especially on the anterior limbs. There 
was a white band on each side of the head running from the 
supralabials and loreals to the flanks at midbody. Belly light, the 



Figure 1 1. Phenacosaurus tetarii, male paratype, MHNLS 664: Flank scales 
behind shoulder. 



1996 PHENACOSAURUS IN WESTERN VENEZUELA 15 




16 BREVIOHA No. 504 

scales lightly spotted with black and brown. The tail dark brown 
(sepia No. 1 19) with lighter transverse bands. 

The dewlap is yellow. 

Sex Dimorphism. There are problems in inferring sex dimor- 
phism with only three specimens available. The males (two spec- 
imens) (>80 mm) appear to be larger than the female (one spec- 
imen) (69 mm), which is certainly mature because it laid an egg 
(15 X 10 mm). The male holotype was much more brown than 
green, whereas the single female was a general light green in color. 
This apparent color difference is problematic because there is just 
one other specimen (the MHNLS paratype) for which the color 
in life has not been reported. 

Habitat. The female paratype was collected on a shrub; the 
male holotype was encountered on spongy lichens and dead leaves. 
Both specimens were collected in stunted forest in the ecotone 
between cloud forest and the shrubby paramo. The low trees have 
very tangled and leathery branches, and the region is cloudy dur- 
ing the greater part of day. Phenacosaurus tetarii probably inhabits 
exclusively the higher cloud forest and the lower limits of the 
paramo at elevations between 2,200 and 3,000 m. 

Distribution. This is a species endemic to the Sierra de Perija 
and restricted to the higher altitudes that constitute the border 
between Colombia and Venezuela and known from two localities 
22 km apart, the Paramo del Tetari and the massif of Cerro 
Pintado. 

Phenacosaurus euskalerriari, new species 

Holotype. MBLUZ R-308, adult male, collected by Jon Ugarte, 
March 22, 1991, in the canyons of Mesa Turik, Sierra de Perija, 
Estado Zulia, Venezuela (72°44'27"W, 10°22'23"N), 1,600 m el- 
evation. 

Paratype. MCZ 1 7475, adult female, collected by Javier Zabala, 
March 17, 1991, in the Campamento de la Gran Depresion of 
Mesa Turik, Sierra de Perija, Estado Zulia, Venezuela (72°42'48"W, 
10°24'10"N), 1,700 m elevation. 

Etymology. Euskalerriari signifies, in the Basque language, "of 
the Basques." The name is proposed for this new species in honor 
of the expedition "Vasco-venezolana Turik 1991," during which 
these specimens were collected. 



1 996 PHENACOSAURVS IN WESTERN VENEZUELA 1 7 



Diagnosis. A phenacosaur closest to P. orcesi but differing in 
the shorter length of the interparietal, the aspect of the scales 
surrounding the interparietal, the size of the uniform flank scales 
and the higher lamellar count, and possibly in the blue dewlap 
(blue is a very unusual dewlap color, and the color of the orcesi 
dewlap is unknown). 

Description. As in the previous description, this description is 
primarily based on the male holotype; the female is mentioned 
only when differences are clearly visible. 

Head: Dorsal head scales (Fig. 72^)— Antorbital area: Scales 
smooth or weakly rugose, smaller toward the tip of the snout. Six 
squarish or rectangular postrostrals. Circumnasals in broad con- 
tact with the first supralabials of each side and separated from 
the rostral by the outermost postrostrals. Dorsally 4 scales be- 
tween the circumnasals. 

Canthal scales, 6 on each side, the anteriormost in contact with 
the circumnasal of its side. Two scales between the second can- 
thals in the male holotype, in the female paratype 3 scales. The 
frontal depression very shallow, formed by parts of 4 large scales 
in the male holotype, parts of 6, only slightly smaller scales, in 
the female. 

Orbital area: All scales of the semicircles heavily tuberculate, 
except the 2 most posterolateral in the male holotype. The an- 
teriormost and posteriormost of each side form prominent bulges 
in front and behind the orbit, and the lateral edges of these and 
other scales of the semicircles slightly raised in rounded ridges, 
circumscribing the supraocular area. 

The scales of the supraocular area smooth or slightly rugose. 
The largest supraocular on each side in contact with the supra- 
orbital semicircles. The supraoculars grading in size mediolater- 
ally. Superciliaries small and smooth, grading in size from larger 
anteriorly to granular posteriorly. 

Parietal area: The converging boundary ridges of this area begin 
as strong tuberculations on flat or only slightly convex scales 
adjoining the posteriormost scales of the semicircles. The tuber- 
culate or pustulate boundary ridges converge to meet posteriorly 
in 2 small raised blunt bosses at midline. Notching in this case 
minimal, merely the groove between the 2 bosses. 

Lateral head scales (Fig. 7 i^ — Most lateral scales smooth, but 



18 



BREVIORA 



No. 504 




1996 



PHENACOSAURUS IN WESTERN VENEZUELA 



19 




Figure 13. 
of head. 



Phenacosaums euskalerriari, holotype, MBLUZ R-308: Lateral view 



the lateral faces of 2 canthals, the scales of the intertemporal ridge 
and the postoculars more or less heavily tuberculate, some suboc- 
ulars weakly tuberculate. In the male, 2 rows of loreals— total of 
8 — on each side, arranged as follows, counting from the front a 
single row of 5 scales, increasing in size posteriorly, a double row 
of 2 scales one precisely above the other, then again a single row, 
1 scale underneath the single preocular. In the female on the right 
side, precisely the pattern seen in the male; on the left side, how- 
ever, only 6 loreals, with the single preocular in contact not only 
dorsally with the second canthal, but also in ventral contact with 
fifth and sixth supralabials. 

One preocular on both sides in both specimens. Three sub- 
oculars on both sides in the female, and also on the right side in 
the male, but 4 on the left. Both posteriormost suboculars in the 
female with 2 or 3 tubercles. Seven heavily tuberculate post- 
oculars. The suboculars broadly in contact with the supralabials. 
Seven supralabials to below the center of the eye on both sides 
of both specimens. 

The intertemporal ridge moderately prominent, covered by 4 
scales, each with 1 or more tubercles. Supratemporals and infra- 
temporals smooth and flat, moderate in size. 



Figure 12. 
of head. 



Phenacosaums euskalerriari, holotype, MBLUZ R-308: Dorsal view 



20 



BREVIORA 



No. 504 




1996 



PHENACOSAURUS IN WESTERN VENEZUELA 



21 




Figure 1 5. Phenacosaurus enskalerriari, holotype, MBLUZ R-308: Lateral view 
of entire animal. 



Ear conspicuous, oval or ovoid, the greater dimension vertical, 
larger than adjoining scales, smaller but not greatly smaller than 
interparietal. 

Ventral head scales (Fig. 7 ^y) — Mental semidivided in male with 
a nearly transverse posterior margin. Five postmentals (2 of them 
differentiated sublabials) between the infralabials. Three or 4 scales 
in the sublabial series on each side are in contact with infralabials. 
The postmental gulars larger than the more posterior central gu- 
lars but smaller than the rows of labials just medial to the sub- 
labials. 

Trunk (Figs. 15 and 16): No dorsal crest in either specimen, 
but on the nape patches of very low subconical scales. At mid- 
dorsum and meeting in the midline hexagonal smooth scales only 
little larger than and very little different from the uniform flank 
scales. Ventrals smooth, larger than dorsals, subimbricate, and in 
transverse rows. 

Limbs (Fig. 15): All limb scales smooth, but differing in size. 
Those of the anterior faces of upper and lower arms and of thigh 



Figure 14. Phenacosaurus euskalerriari, holotype, MBLUZ R-308: Ventral 
view of head. 



22 



BREVIORA 



No. 504 




Figure 16. Phenacosaurus euskalerriari, holotype, MBLUZ R-308: Rank scales 
behind shoulder. 



and tibia about as large as flank scales, their ventral and posterior 
faces more nearly granular. The digital scales of both dorsal and 
ventral surfaces are lamellar— wider than long, and overlapping 
distally throughout the length of all digits. The adhesive lamellae 
under phalanges ii and iii of fourth toe 21 in the male and 25 in 
the female. 

Tail (Fig. 15): The tail strongly compressed, curved, apparently 



1996 PHENACOSAURUS W^ESTERN VENEZUELA 23 

prehensile. Large postanals in the male only. A distinct crest of 
keeled swollen scales begins at the base of the tail, where the scales 
are small then distinctly larger beyond the tail base, but decreasing 
gradually in size to tail tip. Lateral scales smooth and small. Four 
midventral rows of keeled scales begin about 30 scales behind 
postanals. 

Dewlap (Fig. 15): Dewlap posterior to the insertion of the arms 
in both sexes. Edge scales are smaller than ventrals in the male. 
The lateral scales are in rows separated by relatively wide wrinkled 
areas of naked skin. 

Color and Pattern. The male is shown by a slide to have a 
general emerald green coloration (emerald green No. 163) marked 
irregularly with brownish (dark drab No. 1 1 9BB), very likely 
highly cryptic in its habitat. Head brownish in the parietal area 
and at the tip of the snout. On the flanks more prominent brown 
blotches that project on the belly as triangles that resemble the 
peaks of a mountain range. The tail is banded green and brown. 
The limbs present the same pattern as the body. 

The dewlap is light lead-colored blue. 

Habitat. This species inhabits the forests of Perija and has been 
encountered at approximately 1,700 m elevation, both specimens 
on bushes in scrub/dwarf forest. It appears to be a species of the 
cloud forests below 2,000 m in elevation, and its coloration of 
green and brown in life suggests a cryptic aspect appropriate to 
its silvicolous habit. In view of the altitudinal amphtude of the 
cloud forests in which it lives, the species could be distributed 
between 1,600 and 2,500 m elevation, uniquely in very humid 
forests. It is possible then that it is parapatric with the larger 
species of higher and more open formations. 

Distribution. P. euskalerriari is known only from the Mesa 
Turik (1,600-2,300 m), a limestone meseta, located between the 
Rio Apon and the headwaters/origins of the Rio Palmar on the 
Venezuelan slopes of the Sierra de Perija, Estado Zulia, Venezuela. 

Phenacosaurus nicefori Dunn 

Phenacosaurus nicefori Dunn, 1944, Caldasia, 3: 59. Type: ILS 64. 

Type Locality. Pamplona, Norte de Santander, Colombia. 

Diagnosis. Smaller than Phenacosaurus heterodermus and P. tetarii (maximum 
SVL: male 63 mm, female 58 mm) and differing from the giant species and P. 



24 



BREVIORA 



No. 504 




1996 P//£'A^/lC05'/lC//?L/5' IN WESTERN VENEZUELA 25 



heterodermus by fewer large (as does P. tetarii) round flat scales on the flanks and 
with the posterior notch between parietal crests especially wide, wider than any 
other species in the genus. 

Description. Head: Dorsal head scales (Fig. 77,)— Antorbital area: Six to 8 can- 
thals; if 7 or 8, 1-2 small canthals have been intercalated in the series. The third 
canthal largest. One small scale or none between anteriormost canthal and the 
circumnasal. Four to 5 squarish scales border the rostral posteriorly. The circum- 
nasal on each side in broad contact with the first supralabial and separated from 
the rostral by 1 postrostral or in contact. Four scales between the circumnasals 
dorsally. Small scales behind the circumnasals occur in 1 or 2 rows medial to the 
anterior canthals. 

Orbital area: The scales of supraorbital semicircles always at least weakly tu- 
berculate. There are 1 or 2 pairs in contact, or 2 scales on one side may contact 
with one on the other. The scales of the supraocular area, which are always smooth, 
decrease in size laterally. Two or 3, rarely 4, of the larger supraoculars are in 
contact with the semicircles medially. These scales are separated from the super- 
ciliary margin by 2-3 granular rows. Superciliaries are mostly subgranular, but 
the 1 or 2 anteriormost of the series are slightly larger. 

Parietal area: Lyre-shaped and tubercular lateral parietal ridges arise from the 
scales that are in contact with the posteriormost scales of the supraorbital semi- 
circles and terminate in bluntly swollen boss-like scales separated by low, wide 
notch, relatively wider and lower than in other species of the genus. All scales of 
parietal table distinctly lower than the bounding parietal ridges and more or less 
strongly tuberculate. Interparietal with or without an eye, larger than the very 
small ear, round, subrhomboid or subhexagonal, are separated from the semicircles 
by 1 or 2 scales or in contact. Scales lateral to the interparietal tend to be about 
as large as the interparietal. Four to 6 scales intervene between interparietal and 
the notch, which is filled by a transverse row of 2-3 smooth scales. 

Lateral head scales (Fig. 18) — There are 1 or 2 rows of loreals; if 2, the upper 
row is posterior only or is intercalated at intervals above the lower row. Total 
loreals vary from 4 to 9. Only 1 preocular is present, usually small and in contact 
only with the anterior subocular and the second canthal; if larger and additionally 
in contact with the sublabial series, it perhaps implies a fusion of a lower loreal 
with the preocular. There are 4-5 suboculars and 6 to 8 usually tuberculate post- 
oculars. Seven to 9 supralabials extend to below the center of the eye. 

A moderately prominent intertemporal ridge is covered by 3 or 4 scales. Su- 
pratemporals smooth, mostly small, largest toward the parietal ridges. Infratem- 
poral smooth, mostly largest near the intertemporal ridge and toward the comer 
of the mouth. 

Ventral head scales (Fig. 7 9^ — Mental semidivided, in contact with 4 postmen- 
tals between the infralabials: 2 sublabials, 1 on each side and 2 medial gulars. 



Figure 17. Phenacosaurus nicefori, KU 181131, Betania, Tachira, Venezuela: 
Dorsal view of head. 



26 



BREVIORA 



No. 504 




Figure 18. Phenacosaurus nicefori, KU 181131, Betania, Tachira, Venezuela: 
Lateral view of head. 



Three or 4 of the sublabial series on each side in the contact with the infralabials. 
Central gulars smooth, small, longer than wide or as wide as long, juxtaposed, 
becoming larger polygonal adjacent to the sublabial series. 

Trunk (Figs. 20 and 21): Dorsal crest variable, absent or interrupted (Type 3 
of Lazell) or consisting of adjoining keeled scales on the middorsum. On the nape, 
the crest usually comprises small cones irregularly arranged. Flank scales are 
smooth, juxtaposed or even subimbricate, sometimes weakly separated, variable 
in size, but the larger round scales are relatively few. Ventrals smooth, imbricate 
to subimbricate, in transverse rows. 

Limbs (Fig. 20): Limb scales smooth, including supradigitals. All subdigitals 
lamellar. Lamellae under phalanges ii and iii of fourth toe 1 5-22. 

Tail (Fig. 20): Tail weakly compressed. Enlarged postanals present in the male. 
A Type 3 or Type 4 crest on the tail, with the crest scales much larger than laterals, 
about as large as 2 ventralmost rows. Lateral scales are weakly rugose, becoming 
distinctly keeled before midlength. The 4 ventralmost caudal scale rows are keeled, 
the 2 medial ventral rows largest, a bit larger than the scales of the tail crest. 

Dewlap (Fig. 20): In the male extending a short distance posterior to the level 
of the axilla, densely scaled, with crowded rows of scales. In the female represented 
by a densely scaled fold extending only to the level of the axilla. 

Color in Life. There is just one description of color in life for a specimen that 
we have examined. It is by William Duellman for a specimen (KU 1811 30) from 
Betania on the eastern slopes of Cerro Tama, Estado Tachira, Venezuela: "Dorsum 
grayish tan to dark brown. Labial region and venter creamy white. Dewlap creamy 
white with pale orange stripes." A slide shows that the dorsum is banded. 



Figure 19. Phenacosaurus nicefori, KU 181131, Betania, Tachira, Venezuela: 
Ventral view of head. 



1996 



PHENACOSAURUS IN WESTERN VENEZUELA 



27 




28 



BREVIORA 



No. 504 




Figure 20. Phenacosaurus nicefori, MCZ 67979, Pamplona, Norte de Santan- 
der, Colombia: Lateral view of entire animal. 

Habitat. This species inhabits the high-Andean humid forest in an altitudinal 
band between 2,000 and 2,600 m above the level of the sea in the Colombian 
Cordillera Oriental and in Venezuela in the Paramo de Tama, which occupies 
parts of two states, Estado Tachira and Estado Apure. 

Individuals perch on small shrubs and bushes or on moss. 

Material Examined. VENEZUELA; Estado Tachira: Betania, altitude 2,150 m: 
CVULA IV 0898-2 18V-219-VJEP; CVULA IV 0899-220-VJEP; KU 181130- 
132; Paramo de Tama, altitude 2,400 m: MCN 4529, FMNH 5684; COLOMBIA: 
Departamento Norte de Santander: Pamplona. 

MUSEUM ABBREVIATIONS 



CVULA = Coleccion de Vertebrados de la Universidad de los 
Andes, Merida, Venezuela 

KU = Kansas University, Museum of Natural History, Lawrence, 
Kansas, USA 

MBLUZ = Museo de Biologia de la Universidad de Zulia, Ma- 
racaibo, Venezuela 

MCN = Museo de Ciencias Naturales, Caracas, Venezuela 

MCZ = Museum of Comparative Zoology, Harvard University, 
Cambridge, Massachusetts, USA 

MHNLS =^ Museo de Historia Natural La Salle, Caracas, Vene- 
zuela 



1996 



PHENACOSAURUS IN WESTERN VENEZUELA 



29 




Figure 2 1 . Phenacosaurus nicefori, Betania, Tachira, Venezuela: Flank scales 
behind shoulder. 



ACKNOWLEDGMENTS 

The Venezuelan authors express their gratitude to Javier Rome- 
ro and Lionel Lanier as guides during the Cerro Pintado and 
Paramo del Tetari expeditions, respectively. Angel Viloria thanks 
the Union de Espeleologos Vasios (UEV) and the Sociedad Ve- 
nezolana de Espeleologia (SVE) for allowing his participation in 
the "Expedicion Vasio-Venezolana Turik 1991." Fieldwork was 
partly supported by funds from the Faculty of Sciences of the 
University of Zulia via the MBLUZ, the Division de Estudios 
Basicos Sectoriales, and the Division de Investigacion. Joris La- 
garde (SVE) made available the Kodachrome of P. euskalerriari. 



30 BREVIORA No. 504 

All of the authors are grateful to Maria Jose Praderio and Ve- 
ronica Ponte at the Museo de Historia Natural La Salle, Caracas, 
to Jaime Pefaur at the Universidad de los Andes, to Fernando 
Navarrete at Museo de Ciencias Naturales, to William Duellman 
at the University of Kansas and to the artist Laszlo Meszoly. 
Brigitte Poulin assisted in proofreading the text. Publication costs 
were covered in part by a grant from the Wetmore-Colles Fund. 

LITERATURE CITED 

Aleman G., C. 1953. Contribucion al estudio de los reptiles y batracios de la 

Sierra de Perija. Memorias de la Sociedad de Ciencias Naturales La Salle 

[Caracas], 13: 205-225. 
DuMERiL, A. M. C, A>fD A. A. DuMERiL. 1851. Catalogue Methodique de la 

Collection des Reptiles. Paris, Museum National d'Histoire Naturelle. 224 

pp. 
DimN, E. R. 1944. The lizard genus P/jewacosawrw^. Caldasia, 3: 57-62. 
Hellmich, W. 1949. Auf der Jagd nach der Paramo-Echse. Deutsche Aquarien- 

und-Terrarien Zeitschrift, 2(5): 89-91. 
Lazell, J. D., Jr. 1 969. The genus Phenacosaurns (Sauna: Iguanidae). Breviora, 

Museum of Comparative Zoology, 325: 1-24. 
Myers, C. M., E. E. Williams, and C. W. McDiarmid. 1993. A new anoline 

lizard {Phenacosaurus) from the highlands of Cerro de La Neblina, Southern 

Venezuela. American Museum Novitates, 3070: 1-15. 
RuEDA A., J. v., AND J. I. Hernandez-Camacho. 1988. Phenacosaurus inder- 

enae (Sauria: Iguanidae), nueva especie gigante, proveniente de la Cordillera 

Oriental de Colombia. Trianea (Acta cientifica y tecnologica INDERENA), 

2: 339-350. 
Smithe, F. B. 1975. Naturalist's Color Guide. American Museum of Natural 

History, New York. 
Williams, E. E., Ar>iD R. Mittermeier. 1991. A Peruvian phenacosaur (Squa- 

mata: Iguania). Breviora, Museum of Comparative Zoology, 492: 1-16. 
Williams, E. E., G. Orces-V., J. C. Mattheus, and R. Bleiweiss. 1996a. A 

new giant phenacosaur from Ecuador. Breviora, Museum of Comparative 

Zoology, 505: 1-32. 
Williams, E. E., M. J. Praderio, and S. Gorzula. 1996b. A phenacosaur from 

Chimanta Tepui, Venezuela. Breviora, Museum of Comparative Zoology, 

506: 1-15. 



B R E V I O R A 

LIBRARY 

Museum of Comparative Zoology 

APR 5 1996 

us ISSN 0006-9698 

Cambridge, Mass. 18 April 1996 Number 505 



A NEW GIANT PHENACOSAUR 
FROM ECUADOR 

Ernest E. Williams,' Gustavo Orces-V,^ 
Juan Carlos Matheus,^ and Robert Bleiweiss'' 

Abstract. A new giant Phenacosaurus from the eastern Andes (La Bonita- 
Santa Barbara Region) of Sucumbios Province of Ecuador is described. It, like P. 
inderenae Rueda and Hernandez, 1991, differs from all other species in reaching 
a maximum size of more than 100 mm and differs from inderenae in the smaller 
size of the largest class of heterogeneous scales (flat flank scales interspersed with 
smaller scales and granules). In the density of the largest class of scales, it resembles 
heterodermus and differs from nicefori Dunn, 1 944, and tetarii Barros, Williams, 
and Vilora, 1 996. From all the remaining species, it differs in having heterogeneous 
scales. 

INTRODUCTION 

The first giant phenacosaur, Phenacosaurus inderenae (>100 
mm in snout-vent length [SVL]), was described by Rueda and 
Hemandez-Camacho (1988) from Gutierrez, in the southeast of 
the Department of Cundinamarca, Colombia, on the eastern slopes 
of the Cordillera Oriental of the Andes, syntopic or sympatric 
with P. heterodermus. 

Since that description, there has been an explosion of infor- 
mation concerning these lizards. New species belonging to several 
subgroups have been described: tetarii Barros, Williams, and Vi- 
loria, 1996, and euskalerriari Barros, Williams, and Viloria, 1996, 
both from the Venezuelan side of the Cerro de la Perija; neblininus 



' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 

^ Departamento de Ciencias Biologicas, Escuela Politecnica Nacional, Apartado 
17.01.2759, Quito, Ecuador. 
' Apartado 17-17-742, Quito, Ecuador. 
■* Department of Zoology, University of Wisconsin, Madison, Wisconsin 53706. 



2 BREVIORA No. 505 

Myers, Williams, and McDiarmid, 1993, from the Cerro de la 
Neblina, at the border of Brazil and Venezuela; and an unnamed 
juvenile (Williams and Mittermeier, 1991) from Venceremos, 
Department of San Martin, Peru. Two more require description, 
a second giant from Ecuador, formally described herein, and an- 
other small species from Chimanta Tepui, Venezuela (Williams, 
Praderio and Gorzula, 1996). 

Sorting out at least the similarities and differences of the several 
new described and undescribed species within the genus seems 
necessary at this time. The justification of the genus will be post- 
poned until a separate paper. 

It has only been possible to separate P. inderenae and the species 
described later by comparing the whole type series of inderenae 
with the whole type series described later. Both type series are 
small, four in the case of inderenae, eight (but one was found 
dead and is now a skeleton) in the new species later, but each 
comes from a well-defined local area. We find that the large, flat, 
round scales are consistently smaller in the Ecuadorean species 
as compared with P. inderenae and, on that basis, describe the 
former as a new species. To facilitate future research, we have 
described the holotype separately and in detail. We discuss vari- 
ation within the remainder of the type series of the new species 
in the same format as the description of the type, except that we 
describe a probable hatchling separately. We then deal with the 
variation in the type series of inderenae in the same style and 
format. 

The material of the new species is deposited in four museums: 
the type and the first found of the paratypes (now a skeleton) in 
the Museo Ecuatoriano de Ciencias Naturales (MECN), four of 
the paratypes in the Escuela Politecnica Nacional (EPN), two of 
the paratypes in the Museum of Comparative Zoology, and one 
paratype in the National Museum of Natural History (USNM). 

The new species, which is very close to P. inderenae in many 
characters in addition to size, is here named for the distinguished 
Brazilian scientist, Paulo Emilio Vanzolini. 

Phenacosaurus vanzolinii, new species 

Holotype. MECN 0309, adult male. 

Type Locality. ECUADOR: S La Alegria, at an elevation of 
2,360 m, ca. 14 km by road from La Bonita (77°37'42"W, 



1996 



NEW GIANT PHENACOSAUR FROM ECUADOR 




La 
Alegria ^'-, 



ECUADOR 



J »-»' 



Figure 1 . The regions of the type localities of Phenacosaurus inderenae (circle) 
and Phenacosaurus vanzolinii (diamond). 



0°27'30"N), Provincia de Sucumbios (formerly the northwest part 
of Provincia de Napo), Robert Bleiweiss and Juan Carlos Matheus 
coll. 15 March 1985. 
Paratypes. All ECUADOR: Provincia de Sucumbios: MECN 



4 BREVIORA No. 505 

0327: found dead in the road (now a broken skeleton), at the same 
locaUty as the type, Juan Carlos Matheus coll. 1986, sex unde- 
termined; EPN 2218: Sitio Las Ollas, 4 km S Sebundoy, on a 
road that was opened by heavy machinery when the road was 
constructed. Ana Almendariz and Alicia Arias coll. 21 May 1988, 
a juvenile male, possibly a hatchling; EPN 2219: Sitio Sebundoy, 
100 m from the concrete bridge over the Rio Chingual on the 
new road to La Bonita, Ana Almendariz and Alicia Arias coll. 
21 May 1988, aduh male; EPN 2221: Sitio Sebundoy, elevation 
1 ,950 m, 6 km N Escuela de Sebundoy, adult male; MCZ 175159 
(formerly EPN 2220): adult male with extruded hemipenes, same 
data as EPN 2221; MCZ 175160: Sta. Barbara (77°3r41"W, 
0°37'58"N), SE Parroquia El Carmelo. Relatives for Janira Re- 
gelado coll. 1988. Janira Regelado don. 1989, adult male with 
extruded hemipenes; USNM 293683: same data as MCZ 1 75 1 60. 

Diagnosis. A giant species, exceeding 100 mm in maximum 
SVL, resembling P. heterodermus, P. inderenae, P. nicefori and 
P. tetarii in the presence of large, round, flat flank scales that at 
least on the lower flanks are more or less widely separated by 
granules; differing from P. orcesi and P. neblininus, in this regard; 
differing from P. heterodermus, P. nicefori, P. orcesi, and P. neb- 
lininus in maximum adult size, in the posterior height of the 
casque, and in color pattern and from P. inderenae by smaller 
scales of the largest ffank class, as well as of larger posterior lateral 
gular scales. 

Description of Holotype. Head: Head with massive casque. 
Swollen rugose parietal crests on each side angle obliquely pos- 
teromedially to end in 2 knobs connected by an intervening notch, 
although lower than the knobs, high above the nape. 

Dorsal head scales (Fig. 2;— Antorbital area: Scales, pustulate 
posterolaterally, smooth anteriorly, moderately large except for a 
single small scale in the shallow frontal depression and one zone 
of small scales posterior to the circumnasals, another such zone 
on each side between the anterior canthals and the medial series 



Figure 2. Phenacosaurus vanzolinii, holotype, MECN 0309, Dorsal view of 
head. 



1996 



NEW GIANT PHENACOSAUR FROM ECUADOR 




6 BREVIORA No. 505 

of large scales. Circumnasals oval, nostrils slightly posterior with- 
in the scales, the acute end of each scale in contact with the first 
supralabial. A subtriangular postrostral separating each circum- 
nasal from the rostral. Five postrostrals. 

Frontal depression shallow. A rosette of larger scales, very weakly 
rugose or pustulate, around a small smooth scale about equivalent 
in size to the small scales of the snout. Three or 4 scales across 
the snout between the second canthals. Canthals swollen, their 
edges raised and very rugose, apparently covering bony encrust- 
ations, four on each side, the third largest and expanded medially, 
the anteriormost smallest, separated from the circumnasal of its 
side by 2 superimposed small scales. 

Orbital area: Scales of the semicircles very large, rugose and 
pustulate, 1 pair in broad contact. A single large pustulate scale 
anterior to this contact. Supraocular scales very much smaller, 
very finely shagreened, 4 scales on the left side, 6 on the right 
side in rather well-defined supraocular disks, the 2 largest scales 
in contact with the semicircles. The anteriormost superciliaries 
of both sides largest, subtrapezoidal, the whole surface exposed 
dorsally, the remaining 6 or 7 superciliaries smaller, quadrangular, 
in a single row mostly on lateral face of head, barely exposed 
dorsally. The remainder of the supraocular surfaces filled by scales 
smaller than those of the supraocular disks, larger than the su- 
perciliaries. 

Parietal area: Scales very variable in size and shape, moderate 
to small, most lightly rugose but distinctly pustulate. Lateral crest 
scales very rugose, indicating bony ornamentation underneath, 
arising abruptly from the low central parietal area, sloping upward 
toward the posterolateral bosses that are the borders of a moderate 
but narrow notch. Three smooth scales across the median notch. 
The several scales covering the two eroded posterolateral bosses 
also smooth. The posterior transverse ridge formed by the bosses 
and notch rises half again as high above the triangular nape scales 
as they are tall and projects slightly backward above the small 
interval before the crest scales begin. 

No parietal eye. The scale believed to be the interparietal iden- 
tified on position and shape. It is in the midline anteriorly, a 
rather small, narrow triangle anteriorly in contact with the sem- 
icircles. Lateral to this scale are apparent parietal scales, the largest 



1996 



NEW GIANT PHENACOSAUR FROM ECUADOR 




Figure 3. 
head. 



Phenacosaurus vanzolinii holotype, MECN 0309, Lateral view of 



scales in the parietal area, subtrapezoidal, in contact with the 
semicircles, posteriorly meeting behind the median "interpari- 
etal." The scales further posterior irregular in size, shape, and 
arrangement. A count of about 6 to 7 scales from the "interpa- 
rietal" to the notch at the posterior end of the casque. 

Lateral head scales (Fig. ij — All scales on lateral surfaces of 
head smooth, except the canthals, which are rugose and pustulate. 
Loreals in two rows, either the upper or the lower row interrupted. 
On right side, 5 large, 1 small scale in lower row, 1 large and 1 
small in upper row, total 8; on the left side, one large, 3 small in 
lower row, 3 large in upper row, total 7. 

Preoculars 2 right side, uppermost in contact with second can- 
thai, 1 left side, in contact with second canthal. Suboculars 4 on 
each side, broadly in contact with supralabials. Postoculars in 
double rows, 5 in each anterior row, larger than adjacent tem- 
porals, 3 in each posterior row, which meets the intertemporal 
bar. 

The upper temporals variable in size, small to subgranular. An 
intertemporal bar abruptly projecting, shelf-like, covered by only 
3 large scales, but 1 row of smaller scales above and 1 below on 
the base of the ridge. Lower temporals exhibiting 2 regions, the 
upper with smaller scales than the lower except that there are 
irregular small scales around the comer of the mouth. Supralabials 



BREVIORA 



No. 505 




1996 



NEW GIANT PHENACOSAUR FROM ECUADOR 




Figure 5. Phenacosaurus vanzolinii, holotype, MECN 0309, Lateral view of 
entire animal. 



elongate rectangles, 8-9 on both sides to below the center of the 
eye. 

Ventral head scales (Fig. ^j — Mental partly divided, deep, sub- 
pentagonal, not indented, in contact with 3 scales between the 
infralabials. Nine infralabials on right side, 1 on left. Very large 
first sublabials, each more than 5 x the size of the single median 
gular. Three somewhat smaller sublabials on each side, in series 
with the first, in contact with the infralabials. Total sublabials on 
each side five. Central gulars smooth, swollen, variable in size, 
largest adjacent to the sublabials. Posterolaterally, posterior to 
the sublabials, the lateral gulars mostly much enlarged, but very 
variable in size, in several distinct rows. 

Dewlap (Fig. 5): Edge scales convex, smooth, smaller than ven- 
trals. Lateral scales about the same size as the edge scales, in single 
rows separated by naked wrinkled skin. 



Figure 4. 
head. 



Phenacosaurus vanzolinii, holotype, MECN 0309, Ventral view of 



10 BREVIORA No. 505 

Trunk (Fig. 5): Dorsal crest of triangular scales, beginning on 
the nape 2 small scales behind the abruptly vertical casque, high 
anteriorly, only low above the sacrum. Nearly continuous in a 
single row, interrupted at irregular intervals by median contact 
of the paravertebrals. 

Flank scales very heterogeneous, with at least 3 classes of scales: 
(1) prominent very large round flat scales, (2) smaller round flat 
scales, no more than 'A the size of the largest, (3) small convex 
scales and granules as well as naked skin. The largest scales on 
the lower flanks. The paravertebrals, which belong to the first 
class of very large scales, in 1 or 2 rows, posteriorly in contact or 
slightly imbricating, anteriorly always separated by granules or 
naked skin. Other very large scales always separated by the other 
2 classes of scales, separated most widely on lower flanks. 

Axilla and groin granular. Ventrals smaller than the largest flank 
scales, averaging a little larger than the second class of flank scales, 
rather irregular in size and shape, imbricate or subimbricate. 

Limbs (Fig. 5): Anterior face of forelimbs mostly with separated 
flat smooth scales. Anterior face of hindlimb mostly with scales 
in contact. Posterior face of upper arm and thigh with granules 
separated or in contact, of lower arm and leg with small scales 
mostly in contact. Supradigitals smooth. Subdigitals all lamellar. 
Lamellae under phalanges ii and iii of fourth toe 25-26. 

Tail (Fig. 5): Strongly compressed, with a crest of larger sharply 
keeled, dentate scales. All scales keeled except a few rows above 
at the base of the tail and about 8 rows immediately behind vent. 
Postanals much enlarged (male). 

Color in life: No descriptions of the color in life of the type 
series exist. Fortunately, we do have slides of the holotype. It is 
revealed to be dorsally greenish in background with the underparts 
totally white. The large round scales are often yellow or lighter 
green. There is some vague tendency to banding. The head scales 
have an orangish cast when not overlaid by green smudging. A 
broad white band unites the supralabials. There is yellow banding 
on the limbs and digits. 

Variation: The Adult Paratypes. Head: The casque is high to 
very high posteriorly and its lateral crests are always significantly 
raised above the central parietal roof 

Dorsal head scales— IKniovb'\i2i\ area: Pustulations may extend 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 11 

far forward but never to the tip of the snout or the area between 
the circumnasals. Small scales may extend only alongside the 
fourth (anteriormost) canthal or reach the anterior end of the third 
as in the holotype. There are 3 postrostrals in EPN 222 1 and 4 
in all others. The rostral area is injured in 3 specimens (EPN 
22 19, MCZ 1 75 1 60, and USNM 293683), in all the circumnasals 
appear to be, as in the holotype, narrowly in contact with the first 
supralabial, and separated on each side from the rostral by one 
of the frontal depression there is always a rosette of scales sur- 
rounding a central scale or scales, but in EPN 2221 the central 
scale is relatively large and there is a small scale off center to the 
left. In EPN 2219, MCZ 175160, and USNM 293683 there are 
2 small or moderate scales at the center of the rosette. In MCZ 
175159 there are 2 rosettes, the posterior with a small scale, the 
anterior with a large scale in center but a small scale just off center. 
The large scales of the rosette may be pustulate or with none or 
few pustules. 

The canthals are always 4 and the anteriormost always sepa- 
rated from the circumnasals by at least 2 superimposed scales. 
The third is always largest and expanded medially. The first can- 
thai may or may not be expanded medially. These scales are nearly 
smooth in MCZ 175159, except for a few pustulations, but more 
or less wrinkled in the other paratypes, which are more or less 
heavily pustulate as well. 

Orbital area: The scales of the semicircles are always pustulate 
to heavily pustulate except the extreme posterolateral scales, which 
may entirely lack pustules. The supraoculars, on the other hand, 
and the superciliaries, are always smooth or very finely sha- 
greened. A single pair of the scales of the semicircles may meet 
medially or these may be wholly separated by 1 row of rather 
large pustulate scales. The supraoculars are relatively few in num- 
ber, the enlarged scales in 2 or 3 rows, with the largest scales 
medial and in contact with the semicircles, and one or more of 
the lateral enlarged scales in contact with the superciliaries. A few 
smaller scales, variable in size and number, fill the remainder of 
the supraocular area. The anteriormost 1 or 2 of the superciliaries 
in all of the type series fully exposed dorsally, larger, and sub- 
triangular or subtrapezoidal. The remainder of the single row of 
superciliaries is quadrate, subequal or the posteriormost again 



12 BREVIORA No. 505 

larger, and in 5 of 6 specimens barely exposed dorsally, mostly 
on the lateral surface of the head. In MCZ 1 75 160, the entire row 
is fully exposed dorsally, and the anteriormost element best de- 
scribed as quadrate like the other superciliaries. 

Parietal area: These scales in the paratypes are extraordinarily 
variable, not only in size and shape but also in rugosity and 
pustulation. EPN 2219 shows an extreme in pustulation with very 
little rugosity; MCZ 175160, on the contrary, is extreme in ru- 
gosity, which nearly conceals all pustulation. 

The lateral crests, as in the holotype, slope upward toward 
posterior bosses that, however, are either the lateral borders of a 
more or less deep notch or of a transverse occipital ridge. A notch 
hke that of the holotype occurs only in EPN 2219. In USNM 
293683 and EPN 2221, a series of 4 bosses unite to form a trans- 
verse ridge. In MCZ 1 75 160 and MCZ 1 75 1 59, there are 4 trans- 
versely oriented bosses but a narrow and deep notch. 

No parietal eye in any adult specimen. In EPN 2221 and MCZ 
175160, a smaller and shorter scale occurs in the position of the 
"interparietal" of the holotype, but the scales called "parietals" 
in the holotype are broken up in both specimens. In the 3 re- 
maining adult paratypes, the scales are so irregular or asymmetric 
that no interparietal is plausibly demonstrable. In consequence, 
counts from interparietal to the posterior crest of the casque can 
only be made in EPN 2221 (4 or 5) and MCZ 175160 (6 or 7). 

Lateral head scales— M\ lateral scales are smooth in the para- 
types except USNM 293683, which has the canthals pustulate as 
in the holotype. Only MCZ 175160 has a single loreal row on 
both sides. The total number of loreals in this specimen is 4 or 

5, the series grading in size anteriorly. On both sides the very 
large and long preocular has a dorsal vertical groove at the middle 
of its length, indicating that 1 loreal has fused with the preocular 
on each side. If so, the true total count of loreals would be 5 and 

6, respectively. The other paratypes have 2 rows, 1 row, upper 
or lower, always interrupted. The total number of loreals in these 
paratypes varies from 7 to 9. Preoculars are 2 in 3 of the paratypes 
as in the holotype. In EPN 22 1 9 and 222 1 , however, there is only 
1 preocular on each side. In all cases, the preocular is in contact 
with the second canthal. The suboculars vary from 3 to 4 in 4 
paratypes and are 5 only on one side of EPN 2219. Postoculars 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 13 



are in 2 rows behind the eye (on the bony ridge that is the junction 
of jugal and postorbital). The number of postoculars in each row 
varies from 3 to 5; only in MCZ 175159 does the number of 
postoculars rise to 6 in each of the 2 rows on the right side and 
to 6 on the posterior row, 7 in the anterior row on the left. The 
supralabials are elongate rectangles in all specimens, and the num- 
ber to the center of the eye varies from 7 to 10. 

The temporal area is in all specimens divided into supra- and 
infratemporals by a very prominent projecting, shelf-like inter- 
temporal ridge (marking externally the squamosal-postorbital bar 
that is the lower border of the upper temporal fenestra). The edge 
of the intertemporal ridge is covered by 2 large horizontally ex- 
tended scales in all paratypes. The number of scale rows entering 
the ridge base from the supratemporals or infratemporals varies 
from to 1 on the upper or lower side independently. 

The supra- and infratemporals are quite smooth. Where in the 
supratemporal region a scale overlaps the edge of the casque, the 
scale is always smooth on the supratemporal side, always wrin- 
kled, rugose or pustulate on the parietal side. Most of the supra- 
temporals are subequal and polygonal, but occasional smaller 
polygonal or narrow scales intervene posteriorly. The infratem- 
porals divide abruptly into 2 regions differing in the size of their 
scales. The abrupt size difference is coincident with the margin 
between dark and light pigment in this lower temporal region. 
This condition is consistent in all the paratypes as well as in the 
holotype, but in the paratypes as in the holotype, just below the 
intertemporal ridge larger scales again occur, and at the lower 
margin around the corner of the mouth quite small scales are 
found. 

Ventral head scales— The mental is always semidivided. It var- 
ies somewhat in width but is always as deep as it is wide, and not 
or only slightly indented by the sublabials or medial gulars. The 
first sublabials vary much in size and may be significantly different 
in size on the 2 sides of the same animal. However, they are 
always at least 4-5 x as large as the medial gulars that lie between 
them, although these also vary very much in size. The first sub- 
labials are in series with a row of sublabials, each of which may 
be as large or larger than the first. A total of 4-7, always counting 
the first, may be in contact with the infralabials. The central gulars, 



14 BREVIORA No. 505 

those that cover the throat posterior to the medial gulars that lie 
between the sublabials, are always smooth, mostly juxtaposed, 
swollen, elongate, and larger anteriorly, becoming smaller and 
more rounded, subimbricate in the center of the throat. Granules 
are visible between some of the central gulars. Those gulars that 
lie next to the sublabials are consistently larger than those oc- 
curring centrally. 

Lateral to and behind the sublabials and lateral to the insertion 
of the dewlap are lateral gulars. The anterior ones are tiny, barely 
separating the sublabials from the infralabials. These become larg- 
er posteriorly and, indeed, become as large as the posterior sub- 
labials from which they are distinguished by their orientation. 

Dewlap (all paratypes are males): Moderate, not extending pos- 
teriorly much beyond the insertion of the arms. Edge scales, 
smooth, imbricate, smaller than ventrals. Lateral scales, about 
the same size as edge scales, in very regular single rows, the scales 
well separated by wrinkled skin or by wrinkled skin with very 
occasional minute scales. 

Trunk: A dorsal crest of a single row of triangular interrupted 
at irregular intervals by 2 paravertebral scales joining across the 
midline, highest on the nape, lowest on sacrum. 

There are 3 classes of flank scales: (1) round, flat scales, varying 
in size but larger than crest scales; (2) round, flat scales much 
smaller than class 1 ('A their size or less; (3) convex granules, large 
or small, some almost as large as some class 2. One or more rows 
of paravertebrals of class 1 size are always in contact or separated 
by only 1 row of class 3 granules or scales. Class 1 scales lower 
on the flank, on the other hand, are always separated by 2-5 rows 
that include class 2 scales and/or smaller class 3 scales or granules. 

The ventrals are always smooth, flat or slightly convex, im- 
bricate and about the size of class 2 scales or slightly larger. 

Limbs: Anterior upper arm scales smooth, variable in size, 
sometimes imbricate. Posterior upper arm scales granular, jux- 
taposed. 

Anterior lower arm scales smaller, narrower, smooth, subim- 
bricate. Posterior lower arm scales granular, juxtaposed. 

Manus with weakly multicarinate scales, imbricate dorsally on 
carpus, palm scales narrower, subimbricate. Supradigitals mul- 
ticarinate. All subdigitals lamellar. 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 15 

Thigh scales anteriorly large, smooth, subimbricale proximally, 
imbricate at knee, posteriorly with narrow scales, subimbricate 
or juxtaposed. Tibial scales smooth, small, narrow and imbricate 
anteriorly, subgranular and subimbricate posteriorly. In EPN 22 1 9, 
larger scales are found among the granules. 

Pes with scales dorsally and on the sole indistinctly carinate 
and subimbricate. Supradigitals weakly multicarinate. All sub- 
digitals lamellar. Lamellae under phalanges ii and iii of fourth 
toe 24-28. 

Tail: Always strongly compressed and with a distinct crest of 
keeled scales, which, however, varies in height but, in most para- 
types, has a dentate appearance. The lateral caudal scales are 
always keeled, except dorsally at the base and ventrally for 5-10 
rows behind the vent. 

Variation: The Juvenile Paratype. The juvenile EPN 2218 re- 
quires a separate description. Many of the differences are surely 
ontogenetic, but it is not obvious that all of them are. Differences 
between this specimen and the remainder of the type series are 
italicized. 

Head: There is no casque but its margins are partially indicated 
by low ridges (lateral parietal crests) bounding the parietal area. 

Dorsal head scales — Aniorbixal area. There are no pustulations. 
Scales at the tip of the snout and 2-3 rows posterior to the cir- 
cumnasals and a few scales between the canthals and a larger 
median row of scales are small. A total of 7 postrostrals, 5 almost 
granular postrostrals in addition to the larger circumnasals. Each 
circumnasal is broadly in contact with the first supralabial of its 
side. There are 3 small scales between the circumnasals dorsally. 

The frontal depression is very shallow with large scales forming 
a rosette around small scales in the center. 

The canthals are 6 on the left side, 5 on the left, gently arched, 
not keeled, the first (posteriormost) widened medially on both 
sides, third largest, also expanded medially, the anteriormost on 
the left side in contact with the circumnasal, that on the right 
separated from the circumnasal by 1 small scale. 

Orbital area: The supraorbital semicircles are separated me- 
dially by a single row of scales only slightly smaller than those of 
the semicircles. One especially large supraocular in contact with 
the semicircles on each side. Two to 3 enlarged supraoculars in 



16 BREVIORA No. 505 

a second row, not in contact with the superciliaries. Other scales 
of the supraocular region smaller. Two or 3 short polygonal an- 
terior superciliaries followed by quadrate subgranular scales. 

Parietal area: A parietal eye indicated by a light spot in the 
hexagonal interparietal, the largest scale in the parietal area and 
separated from the semicircles by a large scale on the left side 
and by 2 small scales in a row on the right side. 

There are moderate-sized scales that abut laterally on the in- 
terparietal but they do not meet behind it and intervening scales 
of small or moderate size separate these from the semicircles as 
well as others that separate them from scales that cover the lateral 
parietal crests and are larger than any scales of the parietal area 
except the interparietal. 

The lateral parietal crests converge but do not meet. There is, 
instead, a wide gently convex medial area, presumably marking 
the position of the future posterolateral bosses and the median 
notch. Over this gentle convexity, 4 rows of small scales, here 
called "notch scales," precisely comparable in size to the nape 
scales, enter the parietal area to abut against the 4-5 rows of 
abruptly larger (moderate-sized) scales behind the interparietal. 

Lateral head scales— T\No loreal rows, 8-10 total loreals, 4 large 
and 4 small on the right side. 4 large and 6 small on the left side. 

Preoculars 2, counting on each side the upper scale that overlaps 
the loreal rows and excludes the lower preocular from contact 
with the second canthal. Five suboculars each side, broadly in 
contact with the supralabials. Eight to 10 supralabials to below 
the center of the eye. 

Temporals smooth, flat. Lower temporals variable in size, but 
a vaguely indicated division by scale size with the scales averaging 
smaller in the upper pigmented region and larger in the lower 
unpigmented region. A well-defined intertemporal row beginning 
with a single large elongate scale and continuing as a double row 
of slightly enlarged scales. Upper scales subequal with an abrupt 
transition at the lateral parietal ridges where the enlarged scales 
of the parietal area begin. 

Ventral head scales — M^nlSiX partly divided, in contact with 4 
postmentals between the infralabials, 2 medial gulars between the 
very large first sublabials. Three additional sublabials in series 
with the first on each side are in contact with the infralabials. 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 17 



Central gulars smooth, juxtaposed or subimbricate. Some of 
the posterior gulars next to the sublabials markedly enlarged, 
nearly as large as the last sublabial. Lateral gulars intervene be- 
tween the posterior sublabials and the infralabials. 

Trunk: A dorsal crest begins on the nape, seven scales behind 
the enlarged scales of the parietal area, as raised, still relatively 
small protuberant scales in 1 or 2 rows, rising to 3 broad-based 
triangular typically blade-like crest scales in sequence, behind 
which the crest is only indicated by a series of enlarged smooth 
oval scales, interrupted at intervals by paravertebrals joined over 
the midline. Flank scales heterogeneous with large round flat class 
1 scales, largest dorsally and there often in contact, on lower flanks 
smaller and most often separated by class 2 and 3 scales and 
granules. Axilla and groin granular. Ventrals smooth, flat, irreg- 
ular in size and shape, imbricate or subimbricate. 

Limbs: Anterior face of fore- and hindlimbs with imbricate or 
subimbricate flat smooth scales. Posterior face of upper arm and 
thigh with granules, of lower arm and lower leg with small scales. 
Supradigitals smooth. All subdigitals lamellar. Lamellae under 
phalanges ii and iii of fourth toe 25. 

Tail: Compressed, without a crest or a continuous middorsal 
row, smooth or with an occasional hint of keeling dorsally, none 
below, but the latter scales more convex. Postanals much enlarged 
(male). 

Dewlap: There is no trace of a dewlap. 

Food. Stomach contents from MECN 0327 were examined by 
James Carpenter (then at the Museum of Comparative Zoology). 
He reported: 

At least four taxa are present in the sample as follows: 

Order Lepidoptera, family Geometridae— an intact larva 
("looper" or "inchworm"). 

Order Coleoptera, family Buprestidae— elytra (forewings). 

Order Hymenoptera, Halictidae, tribe Augoclorini— head 
capsule, thorax, wing, parts of legs. 

Order Homoptera, Cicadidae— most of the large frag- 
ments, including a head capsule. 

In addition there are a number of eggs. These could be 
from the cicada. 



18 BREVIORA No. 505 

Ecology. The holotype was collected by Robert Bleiweiss. He 
here provides an expanded version of his field notes: 

Collected during the early afternoon along a road cut. The 
animal was moving slowly and deliberately down the stem 
of a large vine belonging to the family Gesneraceae that was 
overhanging the road bank along a forested section of the 
road. 

The weather was sunny and warm. It was probably between 
1 and 2:30 in the afternoon. The collection site was a few 
minutes drive down from La Alegria near a stream gorge and 
on the shady side of the road. 

Juan Carlos Matheus collected the second (damaged) specimen 
dead in the road near the same site. 

Ana Almendariz has reported on the times and sites of collec- 
tion of the remaining specimens that have exact data. All were 
collected between 10:00 and 12:30 a.m., and all were collected 
along roads. The days of collection were humid, but at the mo- 
ments of collection there was never rain but, instead, a bit of 
sun— "un poco de sol." 

Ana Almendariz has also provided a description of the general 
region where she collected (translated): 

While the region is within the 'cloud forest zone', it has been 
heavily cultivated, removing the trees that were of commer- 
cial value, then clearing the forest for field crops. Original 
forest is therefore confined to slopes and ravines difficult of 
access. The margins of the roads are covered with secondary 
vegetation, mostly Asteraceae. 

She reported also that the local people say that the "camaleon o 
camelion" is common in the fields of maize. Bleiweiss and Ma- 
theus collected at La Alegria at higher elevations with steep ter- 
rain. This area, the type locality, was still heavily forested in 1985. 

The ecology reflected by these experiences accord wells with 
the observations of Williams collecting heterodermus in the Sa- 
bana de Bogota and with the more extended observations made 
by Kenneth Miyata on 2 full days of study in August 1973 (Miyata, 
1983). 

Comparisons. Phenacosaurus vanzolinii requires comparison 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 19 

primarily with P. inderenae. The species are both giant, with P. 
inderenae significantly the larger, given the small series of both 
that are available. Both have rather similar coloration and both 
have highly diflferentiated flank scales— the three classes of scales 
described above— seen also in P. hetewdermus. Both differ from 
P. hetewdermus not only in size and the associated development 
of the casque but also in color pattern and details of squamation. 

For the comparison of the two giant species, we have available 
the whole of the type series of P. inderenae, three females and 
one male, generously loaned by Hemandez-Camacho and Rueda. 
For better comparison, we describe these specimens in the same 
detail as we have previously done for the type series of P. van- 
zolinii: one female, Inderena (IND)-R 2999, has already been 
quite accurately illustrated (fig. 1 in Rueda and Hemandez-Ca- 
macho, 1988). The single male, IND-R 3381, is illustrated in our 
Figures 6-9. 

Head: The head is casqued as massively and the posterolateral 
bosses of the casque are about as high as in any P. vanzolinii. 

Dorsal head scales (Fig. 7j— Antorbital area: There appear to 
be larger scales behind the circumnasals and between the canthals 
and the median larger scales and no small scales in the frontal 
depression. The scales between the circumnasals appear to be 
larger than in P. vanzolinii. Postrostrals vary from 3 to 5, 3 in 
the holotype female IND-R 3213, 4 in IND-R 3744 and 3381, 
and 5 in IND-R 2999, in the latter case including the circum- 
nasals, which are in broad contact with the first supralabials but 
only narrowly in contact with the rostral. In the 3 other specimens, 
the circumnasals are again broadly in contact with the first su- 
pralabials but separated on each side from the rostral by one of 
the postrostrals. 

In the frontal depression in all specimens the scales are smooth 
and large. There is no rosette surrounding a central small scale. 
Instead, there is a symmetrical arrangement of 4 large scales, 1:2: 
1 , and there are always 2 scales (3 or 4 scales in vanzolinii) between 
the second canthals across the frontal depression. 

The canthals are smooth or very bluntly keeled. There are 4 
asymmetrically in IND-R 2999, 4 left side, 5 right side, or 5 in 
all the others. On both sides of all specimens the anteriormost 
canthal is in contact with the circumnasals (separated by 2 su- 
perimposed small scales in vanzolinii). 



20 



BREVIORA 



No. 505 



78" 



77° 



10 Km 



Maldonodo'^ ^n 




Figure 6. The known localities for P. vanzolinii. 



Orbital area: In all 4 inderenae, the scales of the semicircles 
are wrinkled and vary from not to moderately pustulate, with 2 
or 3 pairs in contact medially ( 1 or none in vanzolinii). The supra- 
oculars (as in vanzolinii) are smooth or very finely shagreened. 
On both sides of all specimens the 2-3 largest scales are in contact 
with the semicircles, the 4 scales in the second row are in contact 
with the superciliaries, and a central scale is interposed between 
the first and second rows. As in 2 of the 6 paratypes of P. van- 



Figure 7. Phenacosaurus inderenae. paratype male, IND-R 338 1 , Dorsal view 
of head. 



1996 



NEW GIANT PHENACOSAUR FROM ECUADOR 



21 




22 BREVIORA No. 505 




Figure 8. P. inderenae, paratype male, IND-R 3381, Lateral view of head. 



zolinii, the superciliaries in IND-R 3381 and IND-R 2999 are 
fully exposed dorsally. In the holotype, the largest specimen, and 
IND-R 3744, the smallest, only the 2 or 3 largest superciliaries 
are visible dorsally, but in both of these the eyes are somewhat 
sunken in, and this feature may be an artifact. As in vanzolinii, 
the first superciliary on each side is larger and trapezoidal, and 
there are about 7-8 subequal posterior superciliaries. 

Parietal area: There is no parietal eye in any specimen. In 
contrast to vanzolinii, there is a recognizable interparietal in all 
cases, diamond-shaped, variable in size, in narrow contact with 
at least 1 of the 2 scales of the last conjoined pair of the semicircles. 
Two pairs of scales lateral to the inferred interparietal are always 
relatively large and regular but not always symmetrical in shape 
or size. However, the posterolateral pair does not meet behind 
the interparietal. Instead, in all specimens 1 or 2 smaller scales, 
narrow and triangular, intervene posteriorly. The lateral crest 
scales are wrinkled and slope rather gently upward (more steeply 
in the male IND-R 3381) posteriorly toward the posterolateral 
bosses, which are separated by a notch, deep in the male, shallower 
in the females. Five to 7 scales can be counted from the inter- 
parietal to and including a notch scale. 

Lateral head scales (Fig. 8J — As in vanzolinii, all the lateral 
head scales, other than the canthals, are smooth. The canthals 
may be weakly rugose but are never pustulate (as in some van- 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 23 



zolinii). The loreals are in all specimens in only 1 row (usually 2 
rows in vanzolinii, but 1 row on both sides in USNM 293683). 
The total number of loreals varies from 3 to 5 (7 to 9 in vanzolinii) 
(5 on both sides in the holotype of inderenae; 5 on the left side, 
4 on the right in IND-R 3381; 4 on the left side, 3 on the right 
in IND-R 2999; 3 on both sides in IND-R 3744). There is one 
very large preocular in series with the loreals and in contact with 
the second canthal. There are 3 or 4 suboculars, 4 on both sides 
in the holotype of inderenae, 4 on the left side in IND-R 33821 
(3-5 in vanzolinii), all broadly in contact with the supralabials. 
The postoculars are in 2 rows, 5 in the anterior row, 2 in the 
posterior row, the latter large scales that dorsally are in contact 
with the intertemporal ridge. The supralabials are 6 or 7 to below 
the center of the eye (7-10 in vanzolinii). 

The intemporal ridge is distinctly shelf-like in inderenae but 
less regular than in vanzolinii, varying in shape and squamation. 

The supratemporals are moderate, subequal, except on the pos- 
terolateral edge of the casque where they are smaller, narrower 
and vertically elongate. 

The infratemporal tend to be smaller in the pigmented area 
of the infratemporal region and are distinctly smaller ventrally at 
the comer of the mouth. However, the distinction between the 2 
zones of infratemporals— a smaller upper pigmented zone and a 
larger lower unpigmented zone— is decidedly blurred in inderenae 
as compared with the sharp distinction seen in vanzolinii. 

Ventral head scales (Fig. Pj— The mental is wider than deep, 
almost completely divided (semidivided in vanzolinii), and in- 
dented posteromedially (not or only slightly indented in vanzo- 
linii). The first sublabials vary as much or more in size as in 
vanzolinii and, again as in vanzolinii, may vary impressively on 
the 2 sides of 1 animal. In a unique instance (IND-R 2999), the 
left median gular (a gular in contact with the mental) is more than 
% the size of the left first sublabial. In all other cases in both 
species, despite the striking variation in sublabial size the first 
sublabials (and most of the succeeding sublabials) are much more 
to very much more than 4 x the size of any median gulars. 

The number of sublabials in contact with the infralabials com- 
pared for inderenae and vanzolinii, shown in Table 1, barely 
overlaps. 



24 



BREVIORA 



No. 505 




Figure 9. P. inderenae, paratype male, IND-R 3381, Ventral view of the head. 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 25 



Table 1 . Sublabials in contact with infralabials. 





Inderenae 


Vanzolinii 




2999 


8/6 


2221 


6/7 


3381 


8/7 


175160 


4/4 


3744 


7/8 


2219 


5/4 


3213 


7/9 


309 


5/5 






293683 


5/6 






2220 


5/6 



The lateral gulars of inderenae are much like those of vanzolinii, 
tiny anteriorly between the sublabials and infralabials, larger to 
much larger behind the sublabials, from which, as in vanzolinii, 
they are distinguished by orientation. However, while in vanzo- 
linii the lateral gulars may be large to quite large as they approach 
the insertion of the dewlap, the similar scales of inderenae are of 

2 sizes only, moderate and small, well intermixed. 

Dewlap (Fig. 10): As in vanzolinii, the dewlap of inderenae is 
moderate, not reaching much beyond the level of the insertion of 
the arms. There appears to be no appreciable sexual difference in 
this character, the dewlap of the single male of inderenae known, 
IND-R 3381, extending little or no farther back than that of the 

3 females. The scales of the edge are smooth, imbricate, smaller 
than ventrals in both sexes. The lateral scales, while in rows, are, 
unlike vanzolinii, not separated by wrinkled naked skin but by 
numerous small scales, variable in size with larger scales next to 
the row scales, often making it difficult to determine whether the 
rows are single, double, or indeed multiple. Some rows are in- 
complete. 

Trunk (Fig. 10): There is a single dorsal crest as in vanzolinii 
and, as in vanzolinii, it is interrupted at irregular intervals by 2 
paravertebral scales that abut across the midline. As in vanzolinii, 
it is highest on the nape. There are 3 classes of flank scales as in 
vanzolinii and heterodennus. However, unlike the other two spe- 
cies, the class 1 scales are often more than twice as high and wide 
as the crest scales are high. 

Ventrals in inderenae are smooth and imbricate, as in all phen- 
acosaurs, but in this species substantially larger than class 2 flank 
scales. 



26 



BREVIORA 



No. 505 




Figure 10. P. inderenae, paratype male, IND-R 3381, Lateral view of entire 
animal. 



Limbs (Fig. 10): Anterior upper arm scales smooth, strongly 
imbricate to subimbricate, some larger than ventrals. Posterior 
upper arm scales granular, separated. 

Anterior lower arm scales smooth, variable in size, smooth, 
subimbricate. Posterior lower arm scales granular subimbricate. 

Manus with smooth scales, imbricate dorsally and on palm. 
Supradigitals smooth, all subdigitals lamellar. 

Thigh scales anteriorly smooth, not as large as some upper arm 
scales, about as large as ventrals, imbricate proximally, subim- 
bricate at knee, posteriorly smooth, swollen, smaller but variable 
in size, partly imbricate, partly separated. Tibial scales smooth, 
imbricate to subimbricate. 

Pes with supradigitals smooth or very weakly carinate. All sub- 
digitals lamellar. Lamellae under phalanges ii and iii of fourth 
toe 23-31. 

Tail (Fig. 10): Strongly compressed with a dentate crest. Distal 
caudals keeled. 

Color in life: There is an elaborate description of the color in 
life of the holotype. Two slides of color in life also exhibit color 
change. There is also the slide of the Houston Zoo animal taken 
by Harry Greene. The impression is that of a dorsally red brown 
animal that can change the vague dorsal banding to black. 



1996 



NEW GIANT PHENACOSAUR FROM ECUADOR 



27 




Figure 1 1 . Photo in life of the holotype of Phenacosaurus vanzolinii. Robert 
Bleiweiss, photographer. 



DISCUSSION 
Comparative Ecology and Behavior 

Only Hellmich (1949) of those who have seen Phenacosaurus 
in the field has even the appearance of considering the genus as 
limited to paramo. In his case, his characterization of Phenaco- 
saurus as a "Paramo-echse" may refer only to the species, named 
from a single specimen, found on the ground among the Espeletia 
in the Paramo de Sumapaz (a paramo that overlaps the borders 
of Cundinamarca, Meta, and Huila). He was, as appears from the 
text of his discussion, quite aware of the considerable range of 
elevations and habitats in which the genus has been found. Al- 
though, on the evidence of Hellmich's single specimen, the genus 
is found in typical paramo, it is, in fact, known from a surprisingly 
broad range of elevations below those that support paramo under 
natural conditions. 

All specimens off. vanzolinii, for example, were collected be- 
tween 1,950 and 2,630 m. These elevations, all on the eastern 



28 



BREVIORA 



No. 505 




Figure 12. Photo in life of the no locality Phenacosaurus inderenae. Harry 
Greene, photographer. 



slope of the Andes, are in the forested zones corresponding to the 
humid premontane and montane formations of Holdridge ( 1 967). 
The forests are typically tall and often bathed in clouds for much 
of the day. The trees, including many tree ferns, are heavily laden 
with epiphytes and mosses. Although human activities have de- 
stroyed much of the natural forest vegetation along the road be- 
tween Santa Barbara and La Bonita, large patches still exist along 
the steep slopes above the road and in the steeper stream gorges. 
The scanty data from Bleiweiss's and Almendariz's field notes 
are surprisingly consistent on a number of points. Bleiweiss col- 
lected the type on a sunny afternoon around 1:00 p.m., after the 
clouds had lifted: "The animal was moving slowly and deliber- 
ately down the stem of an unidentified (herbaceous) Gesneraceae 
overhanging the [upper] bank along a forested section of the road." 
Similarly, Ana Almendariz reported that her specimens were col- 
lected along roads between 10:00 and 12:30 a.m. in humid but 
sunny weather. 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 29 

These observations are congruent with those of a group led by 
Williams that collected P. hetewdermus north of Bogota and with 
Miyata's (1983) more detailed account of the same species in the 
same area. It may well be characteristic of P. vanzolinii and its 
close relatives of the hetewdermus species group within Phena- 
cosaurus to be most active at midday and/or in relatively sunny 
weather. 

Zoogeography 
Robert Bleiweiss 

The discovery of a new species of Phenacosaurus is not sur- 
prising given that the remarkably diverse Andean herpetofauna 
remains poorly known (Lynch, 1986). We are nevertheless im- 
pressed by its discovery in the Santa Barbara region. There is 
nothing about the physical reliefer general climate of this section 
of the Andes to suggest that it might harbor endemic lizard species. 
Moreover, a large collection of forest frogs made by Bleiweiss and 
Matheus at the same time that they obtained the new Phenaco- 
saurus, although containing new Eleutherodactylus and Coloste- 
thus species, appears no more distinctive than similar collections 
from other high-elevation sites in the northern Andes (W. R. 
Heyer, personal communication). 

The discovery is, in fact, interesting primarily in showing how 
little we know about the Andean herpetofauna. The original im- 
petus for Bleiweiss's exploration of the Santa Barbara area was 
provided by his previous studies of geographic variation in the 
Andean hummingbird Helianthus exortis, a common resident of 
humid montane forests throughout Colombia and eastern Ec- 
uador. The highly variable female plumage of//, exortis resolves 
into a striking dimorphism of male-like and female-like individ- 
uals in southern Colombia and northernmost Ecuador around 
Santa Barbara (Bleiweiss, 1985a,b, 1991). Thus, avian patterns 
already suggest that the Santa Barbara fauna was distinctive and 
had some zoogeographic connection with southern Colombia. In 
the absence of data from other vertebrate groups, birds provide 
the only available context for evaluating the new giant phena- 
cosaur. 



30 BREVIORA No. 505 

Indeed, several characteristics of Santa Barbara's avifauna are 
noteworthy and parallel what is obtained in Phenacosaurus. A 
number of sources indicate that the avifauna is a mix of northern, 
southern, and even more typically western (Pacific slope) ele- 
ments. Van Sneidem's recent collection of birds from the Andes 
of extreme southern Colombia, near the border between Putu- 
mayo and Nariiio (0°3rN, 0°49'N), and thus close to Santa Bar- 
bara (0°23'N), documented northern range extensions for seven 
species (Fitzpatrick and Willard, 1982). All were previously known 
no farther north than the Napo drainage in Ecuador (about 0°20'S) 
(Meyer de Schauensee, 1971). Moreover, Bleiweiss and Matheus 
(in preparation) collected two White-faced Nunbirds {Hapiloptila 
castanea), the first east slope records for a species known previ- 
ously only from scattered localities on the Pacific slope. Bleiweiss 
and Matheus's own observations also indicate that at least two 
rare birds endemic to the east slope were actually common around 
Santa Barbara and La Alegria, the Collared Jay (Cyanolyca viri- 
dicyana) and Red-hooded Tanager (Piranga rubriceps), which ap- 
pears true also for their populations just across the Colombian 
frontier in Nariiio (Hilty and Brown, 1986). The phenacosaurs 
show the same ecological phenomena: The sympatric occurrence 
in the Santa Barbara area (at La Bonita) of a northern (heterod- 
ermus — see earlier) and one from a southern (orcesi) species group 
of Phenacosaurus suggests a faunal mixing zone. 

Admittedly, these patterns are inferred from few data and can 
only be regarded as provisional. They do, however, suggest that 
future collecting along this poorly known section of the Andes 
will prove fruitful. One distinctive feature of the Eastern Cordil- 
lera of the Andes south from Bogota to around Santa Barbara 
may bear on the patterns described earlier. Nowhere along this 
stretch are there peaks higher than 3,000 m (Vuilleumier, 1970). 
If high-elevation habitats are too limited in extent to support 
viable populations, then animals found just below in the forested 
zones may enjoy ecological release (Terborgh and Weske, 1975). 
This could explain this unusual mix and greater abundances of 
species along this segment. 

The significance of the new Phenacosaurus for our understand- 
ing of Andean speciation patterns must await further phylogenetic 
studies. Present evidence cannot distinguish whether the taxon is 



1996 NEW GIANT PHENACOSAUR FROM ECUADOR 31 

a recently derived species or a reliclual population of a once more 
widespread taxon. It is worth noting the Puffbird population of 
the Santa Barbara region is morphologically distinguishable from 
populations on the Pacific slope (Bleiweiss and Matheus, unpub- 
lished observations). The possibility therefore remains that the 
lizard and the bird have differentiated in situ. This unprepos- 
sessing section of the Andes may turn out to be an ecological and 
evolutionary "hot spot." 

ACKNOWLEDGMENTS 

We are specially indebted to Ana Almendariz and Alicia Arias 
for collecting five of the type series of P. vanzolinii and to Janira 
Regelado for providing two additional specimens. Without these 
specimens, the description of Phenacosaurus vanzolinii would 
have been extremely difficult or impossible. Without the drawings 
provided by Laszlo Meszoly, comprehension of the text would 
have been equally difficult or impossible. Brigitte Poulin assisted 
in proofreading the text. Publication costs were covered in part 
by a grant from the Wetmore-Colles Fund. 

LITERATURE CITED 

Aleman G., C. 1953. Contribucion al estudio de los reptiles y batracios de la 
Sierra de Perija. Memorias de la Sociedad de Ciencias Naturales La Salle 
[Venezuela], 13: 205-225. 

Barros, T. R., E. E. Williams, and A. Vilora. 1996. The genus Phenacosaurus 
(Sauria: Iguania) in western Venezuela: Phenacosaurus tetarii, new species, 
Phenacosaurus euskalerriari, new species, and Phenacosaurus nicefori Dunn, 
1944. Breviora, Museum of Comparative Zoology, 504: 1-30. 

Bleiweiss, R. 1985a. Iridescent polychromatism in a female hummingbird: Is 
it related to feeding strategies? The Auk, 102: 701-713. 

. 1985b. Variation and population structure of the Tourmaline Sunangel 

Heliangelus exortis exortis. American Museum Novitates, 2811: 1-14. 

1992. Widespread polychromatism in female sunangel {Heliangelus) 



hummingbirds. Biological Journal of the Linnaean Society, 45(4): 291-314. 
Dunn, E. R. 1944. The lizard genus P/zfrtaco^aMrMS. Caldasia, 3: 57-62. 
FiTZPATRiCK, J. W., AND D. E. WiLLARD. 1982. Twcnty-onc birds new or little 

known from the Republic of Columbia. Bulletin of the British Ornithological 

Club, 102: 153-158. 
Hellmich, W. 1949. Auf der Jagd nach der Paramo-Echse (Ein Beitrag zur 

Kenntnis der Gattung Phenacosaurus). Deutsches Aquaren und Terrarien 

Zeitschrift, 11(5): 89-91, 105-107. 



32 BREVIORA No. 505 



HiLTY, S. L., AND W. L. Brown. 1986. The Birds of Colombia. Princeton, New 
Jersey, Princeton University Press. 

HoLDRiDGE, L. R. 1967. Life Zone Ecology. San Jose, Costa Rica, Tropical 
Science Center. 

Lazell, J. D., Jr. 1969. The genus Phenacosaurus (Sauria: Iguanidae). Breviora, 
Museum of Comparative Zoology, 325: 1-24. 

Lynch, J. 1986. Origins of the high Andean herpetological fauna, pp. 478-499. 
In F. Vuilleumier and M. Monasterio (eds.), High Altitude Tropical Bioge- 
ography. New York, Oxford University Press. 

Meyer de Schauensee, R. 1971. The Birds of South America. Philadelphia, 
Livingston Publishing Company. 

MiYATA, K. 1983. Notes on Phenacosaurus heterodermus in the Sabana de 
Bogota, Colombia. Journal of Herpetology, 17: 102-105. 

Myers, C. M., E. E. Williams, and R. W. McDl\rmid. 1993. A new anoline 
lizard {Phenacosaurus) from the highland of Cerro de la Neblina, Southern 
Venezuela. American Museum Novitates, 3070: 1-15. 

Rueda, J. v., AND J. L HernAndez-Camacho. 1988. Phenacosaurus inderenae 
(Sauria: Iguanidae), nueva especie gigante, proveniente de la Cordillera Ori- 
ental de Colombia. Trianea (Acta Cientifica y Tecnologica INDERENA), 2: 
339-350. 

Terborgh, J., and J. W. Weske. 1975. The role of competition in the distri- 
bution of Andean birds. Ecology, 56: 562-576. 

Vuilleumier. F. 1 970. Insular biogeography in continental regions. I. The north- 
em Andes of South America. American Naturalist, 104: 373-388. 

WiLLL\MS, E. E., AND R. A. MiTTERMEiER. 1991. A Peruvian phenacosaur (Squa- 
mata: Iguania). Breviora, Museum of Comparative Zoology, 492: 1-16. 

Williams, E. E., M. J. Praderio, AND S. GoRZULA. 1996. A phenacosaur from 
Chimanta Tepui, Venezuela. Breviora, Museum of Comparative Zoology, 
506: 1-15. 



>^ 



B R E V I OR, A 

Museiim of Comparative Zoology 

us ISSN 0006-9698 

Cambridge, Mass. 1 8 April 1996 U N 1 V Number 506 



A PHENACOSAUR FROM CHIMANTA TEPUI, 

VENEZUELA 

Ernest E. Williams,' Maria Jose Praderio,^ 
AND Stefan Gorzula^ 

Abstract. A new species of the genus Phenacosaurus is described from Chi- 
manta Tepui, close to P. neblininus. It differs from P. neblininus (and other known 
phenacosaurs) in having the interparietal smaller than the ear and in having the 
circumnasal in broad contact with the rostral and only barely touching or not all 
in contact with the first supralabial. It also differs from neblininus in a generally 
darker coloration and having the belly with bold dark reticulation. 

INTRODUCTION 

Until Lazell (1969) described the new species Phenacosaurus 
orcesi from two localities in Ecuador, the anoline lizards separated 
as the genus Phenacosaurus had been known only from Colombia 
and from just over the border in Venezuela. A summary of new 
information has been reported in Williams et al. (1996). 

Now still another small but distinctive new species, represented 
by a unique specimen, deposited in the collections of the Sociedad 
de Ciencias Naturales "La Salle," Caracas, most similar to the 
other tepui species, P. neblininus, from Cerro de La Neblina, 
provides the easternmost representative of the genus from Chi- 
manta Tepui in Venezuela. 

The new species is named after the late Carlos Todd, long active 
in conservation work (Gorzula, 1987), who participated in the 
exploration of Chimanta Tepui that resulted in the discovery of 
the new species Phenacosaurus carlostoddi. 



' Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- 
setts 02138. 

^ Herpetologia, Direccion Ciencias Naturales, Fundacion Museo de Ciencias, 
Apartado 5883, Caracas 10 10- A, Venezuela. 
' Biosphere Consultants, 614 West Main Street, Newbem, Tennessee 38059. 



2 BREVIORA No. 506 

DESCRIPTION 

Phenacosaurus carlostoddi, new species 

Holotype. SCN 10351, adult female, coll. S. Gorzula and A. 
Farrera, February 1, 1984. 

Type Locality. The southern high plateau of Abacapa-tepui 
(05°12'N, 62°19'W) (CHIMANTA V.), Estado Bolivar, Venezue- 
la, 2,200 m. 

Diagnosis. A small phenacosaur closest to P. neblininus, but 
differing from it and all other phenacosaurs in having the ear 
opening larger than the interparietal, rather than smaller or much 
smaller, in having the circumnasal in broad contact with the 
rostral not at all in contact with the first supralabial, instead of 
having a scale intervene between the circumnasal and the sulcus 
between the first supralabial and rostral, and an apparently gen- 
erally darker coloration, and in having the belly with bold dark 
reticulation. 

It (and neblininus) differs from the orcesi group, to which neb- 
lininus was first referred, in the condition of the fourth toe. La- 
mellae (scales wider than long, distally imbricate) in the fourth 
toe are restricted to phalanges ii and iii. 

Description. Head: Casque indicated by distinct lateral and oc- 
cipital ridges. 

Dorsal head scales (Fig. 1): Antorbital area — ScdAts smooth or 
weakly rugose, small at the tip of the snout and posterior to the 
circumnasals and between the canthals and a median row of larger 
scales. Post rostrals 8, including the circumnasals and the ante- 
riormost loreals of both sides. The latter on both sides just exclude 
the circumnasals from the sulcus between the rostral and the first 
supralabial. Dorsally 4 scales between the circumnasals. 

Canthals 6 on the left side, 7 on the right, rounded or very 
bluntly keeled. On both sides the anteriormost canthal separated 
from the circumnasal by 2 scales, one behind the other. Six scales 
between the second canthals across the frontal depression. 

Frontal depression shallow, the scales within it all larger than 
those at the tip of the snout. 

Orbital area— Scales of the supraorbital semicircles large, smooth 
or lightly rugose, 2 pairs in contact. Scales of the supraocular area 
smooth or very weakly shagreened. On each side the 3 largest of 



1996 



PHENACOSAUR FROM VENEZUELA 




Figure 1. Phenacosaurus carlostoddi, holotype, SCN 10351: Dorsal view of 
head. The black areas on the parietal scales and on some scales of the semicircles 
represent the characteristic pustulations referred to in the text. 



the supraocular scales forming a medial arc, the 2 largest of these 
in contact with the supraorbitals, the third separated by granules. 
Lateral scales of the supraocular region somewhat enlarged me- 
dially, but always sharply smaller than the medial supraoculars 
and becoming granular at the superciliary border. One (right side) 
or 2 (left side) anteriormost superciliaries short, bluntly keeled, 
quadrate or wedge-shaped, and followed only by granules not 
distinguishable from the adjacent granules of the supraocular re- 
gion. 

Parietal <2r£'<2 — Scales on the boundary ridges of this area not 
significantly larger than adjacent nape or supratemporal scales. 



BREVIORA 



No. 506 




Figure 2. 
head. 



Phenacosaurus carlostoddi, holotype, SCN 10351: Lateral view of 



No pustulations or rugosities on the scales of the lateral and 
occipital ridges. About 3 rows of scales lateral and posterior to 
the interparietal and 1-2 rows anterior to it distinctly enlarged. 
These enlarged parietal scales and the interparietal itself distinctly 
smaller than the larger scales of the frontal depression. An indis- 
tinct parietal eye. The interparietal scale diamond-shaped, small, 
smaller than the rather large ear, and separated by 1-2 scales from 
the semicircles. All scales surrounding the interparietal swollen, 
rugose with irregular raised areas and abundant pustulation, char- 
acteristic of other phenacosaur species also, overlying rugosities 
on the underlying bone. Some scale borders difficult to see. About 
7 or 8 scales, decidedly irregular in shape and size between the 
interparietal and the subgranular nape scales. 

Lateral head scales (Fig. 2): Loreal rows 3 on each side. Total 
number of loreals 1 7 on right side, 1 4 on left. 

Preoculars 2 on each side, the uppermost in contact with the 
second canthal. Suboculars 4 on right side, 5 on left. Postoculars 
not well defined grading into the lower temporals. Seven to 8 
supralabials to below the center of the eye. 

Lower temporal scales slightly convex, smooth, juxtaposed, 
larger near the postoculars into which they grade. A weakly dif- 
ferentiated intertemporal zone of 1 row grading from the largest 
next to the postoculars to scales not distinguishable from nape 
scales. 



1996 



PHENACOSAUR FROM VENEZUELA 




Figure 3. 
throat. 



Phenacosaurus carlostoddi, holotype, SCN 10351: Ventral view of 



Upper temporals immediately above the intertemporal row 
small, flat, and smooth, subequal, abutting above on a zone de- 
marcated by a slight ridge, surrounding the parietal area. 

Ear opening on both sides, vertically ovoid, the narrower end 
above, its vertical dimension 6-8 times the height of the highest 
crest scales, relatively larger than the ears of any other species of 
the genus, larger than the interparietal. 

Ventral head scales (Fig. 3): Mental almost completely divided, 
very bluntly indented, in contact with 4 postmentals between the 
infralabials, 1 first sublabial on each side, each about 4-5 times 
the size of the 2 medial gulars, which are themselves somewhat 
larger than the gulars posterior to them. Two additional sublabials 
on the right side, 3 on the left, in contact with the infralabials. 

Central gulars small, smooth, swollen, juxtaposed, subrectan- 
gular, becoming somewhat larger and distinctly polygonal toward 
the sublabials. 

Trunk (Figs. 5 and 6): An indistinct and at intervals interrupted 
crest of triangular scales, sometimes in 2 rows, sometimes in 1 
row, always low, but of varying heights, and never much larger 



BREVIORA 



No. 506 





Figure 4. A subdigital view of the toes and hands of the holotype of P. carlo- 
stoddi, SCN 1035 1, to show that the fourth toe has the lamellar condition restricted 
to phalanges ii and iii. This feature has been confirmed in two paratypes of P. 

neblininus. USNM 322911 and 322912. 



than the paravertebrals. Paravertebral and flank scales subequal, 
flat or slightly swollen, round, weakly rugose, tending to be in 
transverse rows, in contact paravertebrally, separated on lower 
flanks by naked skin or, in part, by granules, grading into ventrals. 
Ventrals smooth, oval, in transverse rows, subimbricate or sep- 
arated by naked skin, larger than any dorsals. 



1996 



PHENACOSAUR FROM VENEZUELA 




Figure 5. Phenacosaurus carlostoddi, holotype, SCN 10351: Lateral view of 
entire animal. 



Limbs (Fig. 4): Upper arm scales smooth, larger and imbricate 
anteriorly and ventrally, smaller and juxtaposed or separated dor- 
sally, posteriorly, and ventrally. Lower arm scales keeled, imbri- 
cate anteriorly and ventrally, smooth, juxtaposed or subimbricate 
dorsally, posteriorly and ventrally. Thigh scales anteriorly keeled, 
imbricate, posteriorly smooth subgranular. Supradigitals rugose 
rather than keeled. The toe pad of the fourth toe restricted to the 
intermediate phalanges (ii and Hi). (Two or 3 scales are again 
wider than long at the insertion of the proximal phalanx of the 
fourth toe into the palm, but they are not believed to be lamellae. 
They are described as lamella-like.) All other toes are subdigi tally 
totally lamellar. Eighteen lamellae under phalanges ii and Hi of 
fourth toe. All fingers subdigitally lamellar. 

Tail (Fig. 5): Distinctly compressed. A single median crest (ex- 
cept on the tail base where the relevant scales are low and small), 
becoming larger, sharply keeled and conspicuously dentate about 
10 mm behind the hindlimbs. Lateral scales not keeled near base 
of tail, small, quadrate, rugose, becoming larger and distinctly 
keeled posteriorly. No enlarged postanals (female). Scales behind 
vent smooth, becoming keeled only 20-22 mm behind vent. 

Dewlap (Figs. 5 and 6): Posteriorly just reaching beyond the 
level of the insertion of the forelimbs. Edge scales much smaller 



BREVIOR.4 



No. 506 




1996 



PHENACOSAUR FROM VENEZUELA 





68* 


66» 


64' 




62° 


60° 




1 1 



1 


I 1 


) ^^' ATLANTIC 
_^!^S=!^;^^^i,/~^^ OCEAN 


100 KM 




^ 


^ 


-^J--^ 


T ^"^^ 




-V 


\ 


•V / 


( '^~'t^Zy^ 


- 


< 


^ 


ill t*''- 


% -:/:-(,-^V^ 




J 


• 


y// \ 




- 


^ 

^ 


p^ 




Chimanta ^-. ^\ / 




! 


J ^ ! 


^ • . 


\ r-'^'"' \ 


«v 


tVt 


— ^ > f 






/ 


)^ 


•4 


1 
/ 


74° 72° 70* 68° 66° 64° 62° 6OT 


12° 


' ^-^f^ Q ATLANTIC OCEAN 




/ 


\_ .,/^ s. 


\ 




'''\r^ \ ^ 








\ \ 


ID* 


. /' A ^— ^^.^-^i] 








\ i 




,■ \ ) VENEZUELA Htk 




\ 




^-_^^^^ ^ - - 




-\ ^■-' 1 i2<iii 




_ \ 




— -^_y^ 


9° 


^ 
<: 


r^-^^i^^^ 


>. 


/ . 














■-■• I 




r 


6* 

4° 


COLOMBIA 


i^M 






Cerro de 
la Neblina 




2- 


- 


>^ ^^' BRAZIL 



Figure 7. "Pantepui" ^^-^^w Mayr and Phelps (1967), showing the locations of 
Cerro de la Neblina and Chimanta Tepui. 



than ventrals, oval. Lateral scales abruptly larger, but still smaller 
than ventrals, in rows, triangular or trapezoidal. 

Size: Snout-vent length (SVL) 55 mm, tail 73 mm. 

Color in Life (from Gorzula's Field Notes). "Distinct black 
markings on an off-white background on the head. Dorsum with 
light brown markings. Bluish gray color on the gular fan when 
extended." 

Color as Preserved. Head and body black dorsally, with very 
vaguely indicated coarse lighter mottling. Suboculars and su- 



Figure 6. Phenacosaurus carlostoddi, holotype, SCN 10351: Ventral view of 
entire animal. 



10 BREVIORA No. 506 

pralabials light with spotting below middle of eye and below 
second canthals. Loreals mostly light on right side, mostly dark 
on left side. Light spotting in front, above, and behind arms and 
in axilla. Limbs above black, vaguely spotted or mottled. Throat 
with small black spots and streaks. Belly and anterior tail boldly 
reticulate with black, the reticulations broken at midline. Limbs 
below light centrally with black spotting on anterior and posterior 
margins. Posterior two-thirds of tail black. 

Habitat. Found at about 11:00 a.m. in a small crack in the 
sandstone, near the top of a deep crevasse on a very exposed rock 
escarpment. There were only some stunted Bonnetia roraimae 
scrub and patches of vegetation within a radius of about 100 m. 

Comment. Phenacosaurus carlostoddi would appear to be gen- 
uinely rare. From 1983 to 1987, 22 localities were explored on 
the Chimanta Massif Gorzula visited and collected the herpe- 
tofauna of 16 of these localities, and others made similar collec- 
tions at five of the remaining six localities. Collecting parties 
usually stayed 3-5 days at each locality. Gorzula has also collected 
amphibians and reptiles on the adjacent Angasima and Adanta 
tepuys, on Aprada Tepui, on Ptari Tepui, at a dozen or so localities 
on the Auyan Tepui Massif, at localities on the "Los Testigos" 
chain of tepuys, at three localities on Ilu Tepui, on Yuruani Tepui, 
and at two localities on Cuquenan Tepui. Gorzula has also col- 
lected at dozens of localities at intermediate elevations in the 
Gran Sabana. He reported the following: 

There was usually no problem in collecting "tepui species" 
once their habitat was known. Phenacosaurus carlostoddi and 
Atractus steyermarki were exceptions to this general rule. The 
only other high elevation anole was Anolis chrysolepis eewi, 
a short-legged rock-dweller, that turned up on widely sepa- 
rated tepuys at elevations above 1,700 m. 

Associated Species. Also collected from "Chimanta V" were 
Ololygon sp. (an undescribed species common around swamps 
on most tepuys in the Gran Sabana region) [now Scinax sp. 
fide Duellman and Wiens, 1992], Arthrosaura sp. (an unde- 
scribed species collected at various localities but only on the 
Chimanta MassiO and Stefania ginesi (very common in 
swamps and in adjacent Brocchinea hectioides, apparently 



1996 PHENACOSAUR FROM VENEZUELA 11 

endemic to the Chimanta Massif but with closely related 
species or subspecies on most other tepuys in the Gran Sa- 
bana region [Duellman and Hoogmoed, 1984]. 

DISCUSSION 
Ernest E. Williams 

It is especially necessary to begin to sort out the similarities 
and differences within Phenacosaunis with seven new taxa de- 
scribed since the last revision (Lazell, 1969), which recognized 
just three species. 

Three groups are currently recognizable in the genus: the het- 
erodermus group, the orcesi group, and the neblininus group: 

I. The heterodermus group (two subgroups) is defined by scale 
heterogeneity: the round flat enlarged scales intermingled with 
smaller scales and granules. All the subdigital scales of the 
hands and feet are always totally lamellar (wider than long 
and with a distal free edge), as in the species heterodermus. 

(1) The heterodermus subgroup sensu stricto (strongly het- 
erogeneous flank scalation, well-developed casquing, 
moderate to giant size) includes heterodermus Dumeril 
and Dumeril, 1851 (maximum SVL 76 mm), the Colom- 
bian giant inderenae Rueda and Hemandez-Camacho, 
1988 (maximum SVL 1 1 8 mm), and the Ecuadorian giant 
vanzolinii Williams, Orces, Matheus, and Bleiweiss, 1996 
(maximum SVL 104 mm). 

(2) The nicefori subgroup (weakly heterogeneous flank sca- 
lation, casquing dependent on size, small or near giant) 
includes nicefori Dunn, 1944, a species now known to 
be smaller than heterodermus (maximum SVL 63 mm) 
and tetarii Barros, Williams, and Viloria, 1996, (maxi- 
mum SVL 85 mm), a near-giant species tentatively re- 
ferred to nicefori by Aleman (1953) and Lazell (1969) 
and now shown by an additional two specimens to be a 
valid species. 

II. The orcesi group differs from the heterodermus group by the 
absence of heterogeneity in flank scales (the enlarged flat 
round scales). All subdigitals of the hands and feet are totally 
lamellar, as in the heterodermus group. 



12 BREVIORA No. 506 

The orcesi group (homogeneous flank scalation, relatively 
poorly developed casquing, small size) includes Ecuadorian 
orcesi Lazell, 1969 (maximum SVL 59 mm), and Venezuelan 
(and probably Colombian: the Sierra de Perija occurs on both 
sides of the border) euskalerriari Barros, Williams, and Vi- 
loria, 1996 (maximum SVL 53 mm) and a single juvenile 
from Peru (Williams and Mittermeier, 1991) (SVL 32 mm), 
which was left unnamed because of its juvenile status. 
in. The neblininus group again differs from the heterodermus 
group in the absence of heterogeneity in its flank scalation 
but is defined by the fourth toe of the hindfoot having the 
most distal and the most proximal phalanx nonlamellar (the 
distal scales narrow = nonlamellar and two or three of the 
most proximal lamella-like). All the subdigitals of the hands 
and four of the five digits of the feet are lamellar. 

The neblininus group (homogeneous flank scales, poorly 
developed casque, small in size) are confined thus far to two 
tepuys in south-central and eastern Venezuela: the neblininus 
Myers, Williams, and McDiarmid, 1993, from Cerro de la 
Neblina (maximum SVL 63 mm) and car lost oddi Williams, 
Praderio, and Gorzula, 1996 (this paper) (maximum SVL 55 
mm). 

All the species of Phenacosaurus are poorly known, some be- 
cause of the difficulty of collection, as may well be true of the 
neblininus group [Myers et al., 1993:12-14; S. Gorzula, see earlier 
under Habitat). For the heterodermus group, the difficulty may 
be quite different. Special difficulty in species discrimination re- 
sults from the extraordinary variability of P. heterodermus, as 
currently recognized. 

It is almost certain, however, that the present concept of P. 
heterodermus is an unresolved complex of sibling species. Old 
material, discolored to a muddy brown by formaldehyde that was 
too strong and, as well, from inexact localities, is nearly useless 
for discrimination of species. New material collected from precise 
localities and preserved in a fashion that does not obscure color 
and pattern will be necessary to solve this problem. Above all, it 
will also be necessary to find new characters. 

The fourth toe of the neblininus group is anomalous among the 
phenacosaurs. All the other phenacosaurs have all subdigitals 



1996 PHENACOSAUR FROM VENEZUELA 13 

lamellar. The ueblininus group overlaps the variation ascribed to 
Anolis totally. In the fourth toe of the neblininus group phena- 
cosaurs only the most restricted anoline toe pad— the subdigitals 
under phalanges ii and iii — is lamellar (wider than long, imbricate 
distally), as in some Anolis. In many other Anolis, some fraction 
of the subdigitals of the proximal phalanx is lamellar. Peterson 
(1983:270) cited Anolis aequatorialis as having half to two-thirds 
of the anterior portion of the proximal phalanx lamellar. Phena- 
cosaurus was cited as possibly "unique" in having lamellae on all 
the subdigitals of the proximal phalanx of the fourth toe. 

The discovery that the fourth toe of the neblininus group was 
anomalous for Phenacosaurus was very late, much too late for 
the fact to be recorded in Myers et al. (1993), indeed, well after 
the manuscript for this description of carlostoddi was "com- 
pleted." In fact, in only two of the paratypes of neblininus has 
this "anomaly" been verified. Peterson (1983) was writing 10 
years before the description of neblininus, before any but three 
species were considered valid. Peterson saw only hetewdermus, 
but of the nine species now current only the two show the fourth 
toe of the hindfoot as anything but totally lamellar. 

The paraphyly o{ Anolis relative to Phenacosaurus was sug- 
gested by Etheridge and de Queiroz (1988:312). Presumably the 
assumption of paraphyly would make carlostoddi and neblininus 
the most primitive known phenacosaurs, but this is not unequiv- 
ocal. The neblininus group still shows the presumed synapomor- 
phy of all the subdigitals of the hand and four of five of the feet 
(Fig. 4) being lamellar. Possibly the "anomaly" could be a ho- 
moplasious loss of the lamellar condition for the proximal pha- 
lanx of the fourth toe in only the neblininus group of phenacosaurs. 
Perhaps a totally lamellar condition of all the fingers and toes was 
the original condition of all the anolines. 

Perhaps the reverse of the Etheridge and de Queiroz supposition 
is true. Phenacosaurus is not derived from within Anolis. Instead 
of the Venezuelan tepui species being most primitive, and the 
Colombian species being most derived, with P. orcesi and related 
forms being intermediate, the evolutionary scene might be very 
different. Instead, the totally lamellar condition of the subdigitals 
of the hands and the feet may be primitive for anoline lizards, 
the heterogeneous flank scalation primitive for the genus Phe- 



14 BREVIORA No. 506 

nacosaurus, and a well-developed casque primitively restricted 
to large phenacosaurs. 

Intriguing as this discussion might be, it would obviously be 
inappropriate to append this extensive and still incomplete work 
to a species description. 

ACKNOWLEDGMENTS 

The authors would like to thank artist Laszlo Meszoly, whose 
illustrations are, as always, excellent. Brigitte Poulin assisted in 
proofreading the text. Publication costs were covered in part by 
a grant from the Wetmore-Colles Fund. 

LITERATURE CITED 

Aleman G., C. 1953. Contribucion al estudio de los reptiles y batracios de la 
Sierra de Perija. Memorias de la Sociedad de Ciencias Naturales La Salle 
(Caracas), 3: 205-225. 

Barros, T., E. E. Williams, and a. L. Viloria. 1996. TYiegenus-Phenacosaurus 
(Squamata: Iguania) in western Venezuela: Phenacosaurus tetarii, new species, 
Phenacosaurus euskalerriah. new species, and Phenacosaurus nicefori Dunn, 
1994. Breviora, Museum of Comparative Zoology, 504: 1-30. 

DuELLMAN, W. E., AND M. S. HooGMOED. 1 984. The taxonomy and phylogenetic 
relationships of the hylid frog genus Stefania. Miscellaneous Publication, 
University of Kansas Museum of Natural History, 75: 1-39. 

DuELLMAN, W. E., AND J. J. WiENS. 1992. The status of the hylid frog Ololygon 
and recognition o{ Scinax Wagler, 1830. Occasional Papers of the Museum 
of Natural History, University of Kansas, Lawrence, 151: 1-23. 

Dunn, E. R. 1944. The lizard genus Phenacosaurus. Caldasia, 3: 57-62. 

Etheridge, R. E., AND K. DE QuEiROz. 1988. A phylogeny of the Iguanidae, pp. 
283-367. In R. Estes and G. Pregill (eds.), Phylogenetic Relationships of the 
Lizard Families. Stanford, California, Stanford University Press. 

GoRZULA, S. 1987. Homenaje a Carlos Todd. EDELCA, Revista de CVG/Elec- 
trificacion del Caroni, C.A. (Caracas). Aiio Xll/Segunda Epoca/No. 6: 21. 

Lazell, J. D., Jr. 1969. The genus Phenacosaurus (Sauria, Iguanidae). Breviora, 
Museum of Comparative Zoology, 325: 1-24. 

Mayr, E., AND W. Phelps. 1967. The origin of the bird fauna of the South 
Venezuelan highlands. Bulletin of the American Museum of Natural History, 
136: 273-327. 

Myers, C. W., E. E. Williams, and R. W. McDiarmid. 1993. A new anoline 
lizard {Phenacosaurus) from the highland of Cerro de La Neblina, Southern 
Venezuela. American Museum Novitates, 3070: 1-15. 

Peterson, J. E. 1983. The evolution of the digital pad in Anolis. Comparisons 
among the anoline genera, pp. 245-283. In A. G. J. Rhodin and K. Miyata. 
(eds.). Advances in herpetology and evolutionary biology. Museum of Com- 
parative Zoology. 



1996 PHENACOSAUR FROM VENEZUELA 15 



RuEDA, J. v., AND J. I. Hernandez-Camacho. 1988. Phenacosaurus inderenae 
(Sauna: Iguanidae), nueva especie gigante, proveniente de la Cordillera Ori- 
ental de Colombia. Trianea (Acta Cientifica y Tecnologica INDERENA), 2: 
339-350. 

Williams. E. E., and R. Mittermeier. 1991. A Peruvian phenacosaur (Squa- 
mata: Iguania). Breviora, Museum of Comparative Zoology, 492: 1-16. 

Williams, E. E., G. Orces-V., J. A. Matheus, and R. Bleiweiss. 1996. A new 
giant phenacosaur from Ecuador. Breviora, Museum of Comparative Zool- 
ogy, 505: 1-32. 



JAN 2 6 2000 



B R E V I O Ii.A 

Nl niseMnn oi V^onnparative /^oology 



.RD 
SITY 



us ISSN 0006-9698 



Cambridge, Mass. 21 September 1999 



Number 507 



DEEP-SEA PELAGIC FISHES FROM THE AZORES 

(EASTERN NORTH ATLANTIC) DEPOSITED IN THE 

MUSEUM OF COMPARATIVE ZOOLOGY 

FiLIPE M. PORTEIRO,' KaRSTEN E. HaRTEL," 

James E. Craddock,--^ and Ricardo S. Santos' 

Abstract. The meso- and bathypelagic fishes collected by the Woods Hole 
Oceanographic Institution from the Azores Economic Exclusive Zone and stored 
at the Museum of Comparative Zoology are reviewed and show a total of 9,260 
specimens from 120 collections composing 153 species in 56 families. Twenty- 
eight species in 12 families are new to the area and are documented here in detail. 
A full list of all species and localities is included. 



INTRODUCTION 

The mesopelagic and bathypelagic fishes account for more than 
35% of the total number of species reported from the Azores 
(Santos et ai, 1997). These two groups are diverse in the area, 
and various species can be found in the open ocean as well as in 
the narrow regions around the islands. 

Interest in understanding the oceanography of the Azores re- 
sulted in a number of research programs in the region over the 
past 150 years. However, it was only with the expeditions of 
Travailleur and Talisman (Vaillant, 1888) that detailed reports on 



' Departmento de Oceanografia and Pescas, Universidade dos Acores, PT-9900 

Horta (Acores) Portugal, filipe@dop.uac.pt. 

- Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, 

USA. 

3 Woods Hole Oceanographic Institution, Woods Hole, MA 02543, USA. 



2 BREVIORA No. 507 

deep-sea fishes began. Late in the last century and during the first 
decades of this century, the scientific cruises of Prince Albert I 
of Monaco produced the beginnings of an inventory of the oce- 
anic ichthyofauna of the Azores (e.g., Collett, 1896, 1905; Ri- 
chard, 1905; Zugmayer, 1911, 1933; Roule, 1919; Roule and An- 
gel, 1924). Other important meso- and bathypelagic material from 
the Azores was collected by the Michael Sars North Atlantic 
Deep Sea Expedition (e.g., Murray and Hjort, 1912; Koefoed, 
1956, 1958; Bolin, 1959) and by several Danish expeditions with 
Dana (e.g., Jespersen, 1915; Regan and Trewavas, 1929; Ege, 
1930, 1953; Nafpaktitis, 1968; Bertelsen et al, 1976). Since then, 
several other countries have been involved in oceanic research 
programs in the Azorean region (e.g., France, Germany, Portugal, 
Russia, Sweden, and the United States). 

In particular, several oceanographic expeditions to the Azores 
region were conducted by the Woods Hole Oceanographic Insti- 
tution (WHOI) between 1928 and 1984. These cruises were 
aimed at investigations of the pelagic domain and the material 
collected is now archived at the Museum of Comparative Zoology 
(MCZ), Harvard University. Most of the specimens were fully 
identified at WHOI during ecological and biogeographical stud- 
ies, but only selected parts of the collection have been used in 
publications (e.g., Borodin, 1931; Haedrich, 1964; Nafpaktitis et 
al., 1977). Arruda (1997) produced a checklist of Azorean fishes 
primarily based on literature references. Many of his records are 
confirmed by the specimens cited in this paper. 

We had the opportunity to study the eastern Atlantic collection 
housed in the MCZ, and based on those studies, we record the 
first documented occurrence of some meso- and bathypelagic 
fishes in the Azores region. In addition, we present a complete 
list of the fishes collected in the Azores by the WHOI expeditions. 

MATERIAL AND METHODS 

Following Santos et al. (1997) we consider Azorean waters as 
the Azores Economic Exclusive Zone (EEZ, Fig. 1). A complete 
listing of all the collecting stations and associated data of the 10 
WHOI cruises done in the EEZ are shown in Appendix 1. These 
stations are also plotted in Figure 1 . The station data are arranged 



1999 



AZORES DEEP-SEA FISHES 




R/V Atlantis; C. O'D. Iselin 09-17.07.1928 

R/V Atlantis, W.C. Schroeder, 1931 

RA' Atlantis. 1947 

RA Atlantis, 1948 

R/V Delaware II, Cr 63-04, 14-17.05.1963 

RA' Atlantis II, Cr 13, 20-27.09.1964 



RA' Atlantis II, Cr49, 24-27.06.1969 
RA' Chain, Cr 105, 05-10.07.1972 
RA' Atlantis II, Cr 78, 05.09.1973 
RA^ Atlantis II, Cr 101. 05-10.07.1978 
RA' Oceanus, Cr 158, 25.08-02.09.1984 



Figure 1. 



chronologically by cruise and referenced by field numbers. The 
field numbers refer to the collectors as follows: CODI refers to 
collections made by C. O'D. Iselin, RHB to collections made by 
Richard H. Backus and James E. Craddock, SUN to surface col- 
lections that accompanied the RHB stations, CEL to collections 
made by Charles E. Lea, and KEH to collections made by Karsten 
E. Hartel. The identification and distribution of the species found 
in the WHOI collections were reviewed and compared with San- 



4 BREVIORA No. 507 

tos et al. (1997), Whitehead et al. (1984-1986), Hureau and Mo- 
nod (1973), Quero et al. (1990a), and numerous primary works. 
In addition, we compared the specimens with related material in 
the MCZ collections. 

All records of species new to the area are outlined in the fol- 
lowing section. Appendix 2 lists all of the remaining species 
found in the WHOI collections from the Azorean EEZ. Some 
rare species caught just outside the EEZ are also reported. Full 
data on MCZ collections are available at http:// 
www.mcz.harvard.edu/fish. We follow the family nomenclature 
and arrangement of Nelson (1994). Fishes that could only be 
identified to genus or family are usually not included in Appendix 
2 except in cases of rarity or of special interest. 

RESULTS 

The WHOI/MCZ collections from the Azores were taken at 
120 stations fished from the surface to about 1,000 meters, usu- 
ally with an open net (Appendix 1, Fig. 1). A computer search 
of the database at MCZ (MUSE) from the EEZ produced 1,198 
lots and more than 9,260 specimens with 153 species in 56 fam- 
ilies. The majority were mesopelagic (ca. 88%), followed by epi- 
pelagic (ca. 9%), and a small number of bathypelagic species. 

Myctophidae is the largest family represented with 41 species 
from 646 collections and 70% of the specimens. Other families, 
such as the Stomiidae, Stemopthychidae, Gonostomatidae, Pho- 
tichthyidae and Paralepididae, are reasonably well represented, 
both in terms of their specific diversity and total numbers (see 
Appendix 2). 

Benthosema glaciate is the most numerous fish in any individ- 
ual collection because it was abundant at two stations (RHB 2535, 
RHB 2536) made in the northern area of the EEZ. However, 
Gonichthys coccoi, Lobianchia dofteini, and Lampanyctus pusil- 
lus are the most commonly caught fish in the collections (Ap- 
pendix 2). Of the 20 most abundant species, 65% belong to the 
Myctophidae. However, fishes in other families (e.g., Argyrope- 
lecus hemigymnus, Vinciguerria attenuata, or Stomias boa) are 
also common. Larvae represent fewer than 20% of the total num- 
ber of fishes. 



1999 AZORES DEEP-SEA FISHES 5 

The collections add 28 species in 12 families to the Azores 
ichthyofauna (see below). Bathylagidae and Anotopteridae are 
new families to the area. 

NEW RECORDS OF FISH SPECIES FOR THE AZORES 

Bathylagidae 
Bathylagus bericoides (Borodin, 1929) 

Comments. Occurs in tropical, subtropical, and temperate wa- 
ters of all oceans. Although reported from the northeast Atlantic 
(Cohen, 1984a), the MCZ material represents the first document- 
ed records from the Azores. 

Azores Material. 61912, R/V Atlantis II 49, RHB1917, 
35°34'N 2r54'W, 135-140 m, 0045-0219 h, 25.VI.1969, 1:48 
mm SL; 61909, RA^ Atlantis II 13, RHB1039, 40°47'N 28°4rw, 
450-500 m, 2030-2350 h, 20.IX.1964, 2:29-1 16 mm SL; 61913, 
RA' Atlantis II 49, RHB1928, 38°05'N 26°N29'W, 192-202 m, 
0231-0408 h, 27.VI.1969, 1:105 mmSL; 149499, RA^ Atlantis II 
101, CEL7833, 36°37'N 25°04'W, 50-170-80 m, 2146-2345 h, 
09.VII.1978, 1:91 mm SL. 

Bathylagus greyae Cohen, 1958 

Comments. This species is generally tropical (Cohen, 1984a), 
but we have found numerous records from the Azores and adja- 
cent waters. 

Azores Material. 66131, R/V Atlantis II 49, RHB1922, 
36°32'N 23°49'W, 135-140 m, 0046-0221 h, 26.VI.1969, 1:90 
mm SL; 149498, RA^ Atlantis II 101, CEL7837, 37°48'N 
23°39'W, 350 m, 2211-0045 h, 10-1 1.VII.78, 1:67 mm SL; 
150907, RA^ Chain 105, RHB2537, 42°17'N 29°59'W, 225-255 
m, 0017-0135 h, 05.VII.1972, 1:33 mm SL; 150908, RA^ Chain 
105, RHB2540, 39°55'N 29°03'W, 60-70 m, 2135-2250 h, 
05.VII.1972, 2:62 mm SL; 150909, RA' Chain 105, RHB2550, 
35°03'N 24°30'W, 290-310 m, 0112-0235 h, 08.VII.1972, 1:45 
rmn SL. 

Bathylagus longirostris Maul, 1948 

Comments. Known from tropical and subtropical waters of the 
Atlantic, Pacific, and Indian oceans (Cohen, 1984a). Originally 



6 BREVIORA No. 507 

described from Madeira. Although these are the first two records 
from the Azores, the species is common just south of the area 
based on MCZ material. 

Azores Material. 66184, RA^ Chain 105, RHB2552, 34°17'N 
24°05'W, 60-70 m, 2158-2305 h, 08.VII.1972, ?; 91356, RA^ 
Chain 105, RHB2553, 34°12'N 24°01'W, 125-135 m, 2355-0105 
h, 08-09.VII.1972, 2:90-125 mm SL. 

Opi sthoproctidae 
Dolichopteryx spp. 

Comments. Two forms of this genus, based on distinctive post- 
larvae, are included here. One of these is probably D. longipes 
which has been reported from Madeira (Cohen, 1984b). 

Azores Material. 66370, RA^ Chain 105, RHB2543, 39°10'N 
28°33'W, 250-260 m, 0540-0815 h, 06.VII.1972, 1:27 mm SL; 
66348, RA^ Oceamis 158, KEH8477, 37°35'N 32°55'W, 600 
mwo, 0204-0325 h, 30. VIII. 1984, 1:29 mm SL; 66345, RA^ At- 
lantis II 13, RHB1041, 39°24'N 27°11'W, 220-300 m, 1405- 
1830 h, 21.IX.1964, 1:22 mm SL. 

Rhynchohyalus natalensis (Gilchrist and von Bonde, 1924) 

Comments. This species was caught on the western edge of the 
EEZ and is included here because of the broad range suggested 
by other MCZ specimens from the Canaries (MCZ96900, 
MCZ66428) and from the western North Atlantic as far north as 
39°28'N (MCZ60718). The only other northeast Atlantic record 
is from the stomach of a tuna fish at Madeira (Maul, 1946; Cohen, 
1984c). 

Material. 66429, RA^ Atlantis II 13, RHB1046, 39°30'N 
35°58'W, 425-500 m, 0915-1237 h, 27.IX.1964, 1:37 mm SL. 

Alepocephalidae 

Bellocia koefoedi (Parr 1951) 

Comments. This endemic North Atlantic bathypelagic species 
lives between 2,500 and 5,850 m (Markle and Quero, 1984; Mar- 
kle and Sazonov, 1990). An MCZ specimen was caught at the 



1999 AZORES DEEP-SEA FISHES 7 

southwest corner of the EEZ close to the type locality. There are 
few records of this species for the northeastern Atlantic. 

Azores Material. 36636, R/V Atlantis II, ATLAN4715, 
35°37'N 30°57'W, 3,200 mwo, 1312-? h, 16.VII.1947, 1:135 mm 
SL. 

Platytroctidae 
Sagamichthys schnakenbecki (Krefft 1953) 

Comments. A 24-mm SL postlarva found in the MCZ material 
from the middle of the EEZ confirms Quero et al. (1984) map- 
ping. This species was not indicated for the Azores in Matsui and 
Rosenblatt (1984). The eastern Atlantic MCZ material includes 
more than 20 records of postlarvae and juveniles from 16°N to 
56°N, with some just to the east of the Azores. MCZ material 
also includes several records from off the east coast of North 
America that have not been reported previously (e.g., 
MCZ75508). 

Azores Material. 128329, RA^ Chain 105, RHB2538, 40°42'N 
29°15'W, 560-600 m, 0950-1225 h, 05.VII.1972, 1:27 mm SL. 

Gonostomatidae 
Cyclothone livida Brauer 1902 

Comments. Recorded from southern Iceland to Angola (Bad- 
cock, 1984a; Quero et al, 1990b). One MCZ collection docu- 
ments its occurrence in the Azores. Additional MCZ material 
shows that C livida, formerly thought to be an eastern Atlantic 
endemic (Miya and Nishida, 1996), actually ranges across the 
Atlantic to 55°W between 3°N and 12°N. 

Azores Material. 143321, RA^ Atlantis II 49, RHB1924, 
37°04'N 24°42'W, 750-830 m, 0808-1005 h, 26.V1.1969, 1:34 
mm SL. 

Cyclothone pseudopallida Mukhacheva 1964 

Comments. A wide-ranging species in all tropical, subtropical, 
and temperate oceans. C pseudopallida was expected to be in 
the Azores, but these are the first confirmed records. In the Ca- 



8 BREVIORA No. 507 

naries, this species is a nonmigrant and is restricted to depths 
between 550 and 800 m (Badcock, 1970). 

Azores Material 142489, Kf\ Atlantis II 13, RHB1036, 
42°20'N 29°09'W, 470-520 m, 1600-1830 h, 19.IX.1964, 1:34 
mm SL; 142501, RA^ Atlantis II 49, RHB1919, 35°56'N 22°40'W, 
650-750 m, 0708-1030 h, 25.VI.1969, 3:34-37 mm SL; 142502, 
RA^ Atlantis II 49, RHB1924, 37°04'N 24°42'W, 750-830 m, 
0808-1005 h, 26.VI.1969, 3:34-37 mm SL. 

Sternoptychidae 
Argyropelecus gigas Norman 1930 

Comments. A widespread species in all oceans. Badcock 
(1984b) infers that the species should occur in the Azores. How- 
ever, based on the MCZ material, it is rare in the area, with one 
specimen, but it is progressively more common to the south and 
east. 

Azores Material. 137182, RA^ Atlantis II 49, RHB1919, 
35°56'N 22°40'W, 650-750 m, 0708-1030 h, 25.VL1969, 1:16 
mm SL. 

Polyipnus polli Schultz 1961 

Comments. The species is included based on a single specimen 
caught at the northwest edge of the Azores, which is the north- 
ernmost record. This eastern Atlantic endemic is also known from 
the Grand Meteor Bank (Badcock, 1984b), the Canaries 
(MCZl 35280), and further south (Quero et al, 1990c; Harold, 
1994). 

Material. 149500, RA^ Atlantis II 101, CEL7816, 42°05'N 
35°35'W, 1,000 m, 2355-0240 h, 03-04. VII.78, 1:25 mm SL. 

Stomiidae 

Aristostomias grimaldii Zugmayer 1913 

Comments. This species is reported for the Atlantic, Pacific, 
and Indian oceans but is rarely caught in the eastern North At- 
lantic. According to Goodyear (1990) this is the second record 
of A. grimaldii for that area. The MCZ material shows a concen- 



1999 AZORES DEEP-SEA FISHES 9 

tration of this species along the east coast of North America, 
between 40°N and 25°N. 

Azores Material. 150372, RA' Atlantis II 101, CEL7833, 
36°37'N 25°04'W, 350-170-80 m, 2146-2345 h, 09.VII.1978, 1: 
85 mm SL. 

Astronesthes gemmifer Goode and Bean 1 896 

Comments. An Atlantic, Pacific, and Indian Ocean species that 
occurs south of 45°N. Gibbs (1984a) maps the possible occur- 
rence in the Azores area; however, the MCZ material probably 
represents the first documented records. 

Azores Material. 133383, RA^ Atlantis II 49, RHB1919, 
35°56'N 22°40'W, 650-750 m, 0708-1030 h, 25.VI.1969, 1:25 
mm SL; 133387, RA^ Chain 105, RHB2555, 34°00'N 22°55'W, 
470-520 m, 1002-1215 h, 09.VII.1972, 1:20 mm SL. 

Astronesthes micropogon Goodyear and Gibbs 1970 

Comments. A member of the "cyaneus" group as defined by 
Goodyear and Gibbs (1970), who noted that 35°N is the northern 
limit of its range. We know only one record from the Azores. 
The specimen is the northernmost record from the eastern Atlan- 
tic. 

Azores Material. 91742, R/V Atlantis II 49, RHB1918, 
35°36'N 22°05'W, 169-175 m, 0231-0409 h, 25.VI.1969, 1:73 
mm SL. 

Astronesthes neopogon Regan and Trewavas 1929 

Comments. A. neopogon is restricted to the North Atlantic and 
was described from just outside the Azorean EEZ. The MCZ 
specimen seems to be only the third northeastern Atlantic record 
since Maul (1956) described specimens from Madeira. 

Azores Material. 149502, RA^ Atlantis 7/101, CEL7832, 36°N 
25°W, 350-190 m, 0050-0345 h, 09.VII.1978, 1:81 mm SL. 

Eustomias macrurus Regan and Trewavas 1930 

Comments. The species is known from the Atlantic, Pacific, 
and Indian oceans. This is the first record for the eastern Atlantic 
north of 6°N. It probably indicates that E. macrurus is more wide- 



10 BREVIORA No. 507 

ly distributed in the eastern Atlantic than previously thought 
(Gibbs and Barnett, 1990). It is widespread in the western North 
Atlantic from 30°S to about 42°N (Gibbs and Barnett, 1990). 

Azores Material. 149504, RA/ Atlantis II 101, CEL7838, 
37°47'N 23°52'W, 350 m, 0045-0347 h, 11. VII. 1978, 1:70 mm 
SL. 

Melanostomias valdiviae Brauer 1902 

Comments. The species is known from all oceans. In the east- 
em Atlantic it occurs to about 35°N (Gibbs, 1984b), but this is 
the first confirmed record from the Azores. Additional specimens 
of the species might be expected in the southern part of the area 
based on its known distribution. 

Azores Material. 132155, RA^ Atlantis II 49, RHB1918, 
35°36'N 22°05'W, 169-175 m, 0231-0409 h, 25.VI.1969, 1:143 
mm SL. 

Stomias brevibarbatus Ege 1918 

Comments. A widespread North Atlantic species that Gibbs 
(1984c) did not report from the Azores. MCZ material and Goad 
(1986) record it as far north as Canadian waters in the western 
Atlantic. MCZ records show the species occurring in the Azores, 
with other records to the west and southeast of the area. This 
species was recently reported in the Gulf of Mexico (Sutton and 
Hopkins, 1996). 

Azores Material. 128677, R/Y Atlantis II 49, RHB1928, 
38°05'N 26°29'W, 192-202 m, 0231-0408 h, 27.VI.1969, 1:129 
mm SL. 

Trigonolampa miriceps Regan and Trewavas 1930 

Comments. This rarely caught species is usually thought to be 
antitropical in the Atlantic Ocean, but we have seen an MCZ 
specimen from just off the Gulf of Guinea at 0°30'S. Gibbs and 
Barnett (1990) and Gibbs (1984b) note the species far north of 
and far south of the Azores. The MCZ material adds the first 
records for the Azores based on three specimens, all from the 
northern portion of the area. 

EEZ Material. 114690, RA^ Atlantis II 13, RHB1039, 40°47'N 



1999 AZORES DEEP-SEA FISHES 11 

28°4rw, 450-500 m, 2030-2350 h, 20.IX.1964, 1:50 mm SL; 
62186, RA^ Chain 105, RHB2542, 39°47'N 28°57'W, 165-175 
m, 0102-0222 h, 06. VII. 1972, 1:30 mm SL; 149503, RA/ Atlantis 
II 101, CEL7818, 40°50'N 34°11'W, 1,000 m, 2120-0040 h, 04- 
05.VII.1978, 1:45 mm SL. 

Paralepididae 
Uncisudis quadrimaculata (Post 1969) 

Comments. The species is endemic to the eastern central At- 
lantic (Post, 1990). The MCZ material contains four lots of larvae 
from 39°N. Two of these are from within the EEZ and two from 
just to the west. The samples were caught near the surface at 
dusk or at night. 

Azores Material. 68562, R/V Atlantis II 13, RHB1043, 
39°28'N 31°00'W, 20-35 m, 1920-2332 h, 23. IX. 1964, 5:18-24 
mm SL; 68563, RA^ Atlantis II 13, RHB1045, 39°34'N 33°37'W, 
15-32 m, 1930-2325 h, 26.IX.1964, 2:33-38 mm SL. 

Anotopteridae 
Anotopterus pharao Zugmayer 1911 

Comments. This species is antitropical in the Atlantic and Pa- 
cific oceans (Maul, 1973a; Post, 1984). Adults are rare in collec- 
tions, probably due to gear selectivity (Heemstra, 1990). We in- 
clude the species based on two postlarvae from the Azores. We 
have seen an additional postlarva (MCZ73016) from just outside 
the EEZ and one 265-mm SL juvenile (MCZ43141) found in the 
stomach of an Alepisaurus brevirostris at 40°17'N 36°07'W 
(Haedrich, 1964). 

Azores Material. 73018, R/V Atlantis II 49, RHB1924, 
37°04'N 24°42'W, 750-830 m, 0808-1005 h, 26. VI. 1969, 1:15 
mm SL; 73020, RA^ Chain 105, RHB2549, 35°08'N 24°25'W, 
40-50 m, 2340-0102 h, 07.V11.1972, 1:22 mm SL. 

Myctophidae 
Hygophum reinhardtii (Liitken 1892) 

Comments. A widely distributed, bipolar species found across 
the subtropical and tropical Atlantic Ocean to about 40°N (Naf- 



12 BREVIORA No. 507 

paktitis et al, 1977). The MCZ material contains the first con- 
firmed records and shows that the southern Azores region is the 
northern edge of the range of this species in the eastern Atlantic. 
H. reinhardtii appear to be quite common south of 38°N within 
the Azores, and based on Nafpaktitis et al. (1977) the species 
would be expected to occur there. All the specimens were caught 
near the surface at night by neuston net. 

Azores Material. 114801, RA^ Atlantis II 49, SUN 191 8, 
35°36'N 22°05'W, surface, 0000-0330 h, 25.V1.1969, 2:35-45 
mm SL; 114802, RA^ Atlantis II 49, SUN1920, 36°23'N 23°35'W, 
surface, 0000-0400 h, 25. VI. 1969, 8:15-39 mm SL; 114803, R/ 
V Atlantis II 49, SUN 1920, 36°23'N 23°35'W, surface, 0000- 
0400 h, 25.VI.1969, 2:27-30 mm SL; 114804, R/\ Atlantis II 
49, SUN1921, 36°27'N 23°42'W, surface, 0000-0415 h, 
25. VI. 1969, 3:24-31 mm SL; 114805, RA^ Atlantis II 49, 
SUN1921, 36°27'N 23°42'W, surface, 0000-0415 h, 25.VI.1969, 
4:20-29 mm SL; 114810, RA^ Chain 105, SUN2545, 37°23'N 
26°33'W, surface, 0000-0215 h, 07.VII.1972, 1:30 mm SL; 
114811, RA^ Chain 105, SUN2546, 37°20'N 26°30'W, surface, 
0000-0345 h, 07.VII.1972, 4:30-38 mm SL; 114908, R/V At- 
lantis II 49, RHB1920, 36°23'N 23°35'W, 63-65 m, 2045-2218 
h, 25.VI.1969, 2:37-40 mm SL; 114993, RA^ Chain 105, 
RHB2552, 34°17'N 24°05'W, 60-70 m, 2158-2305 h, 
08.VII.1972, 1:13 mm SL. 

Lampadena anomala Parr 1928 

Comments. L. anomala occurs in subtropical and tropical wa- 
ters of the Atlantic, Pacific, and Indian oceans (Nafpaktitis et al., 
1977; Bekker, 1983). It is uncommon throughout all of its range. 
Swinney (1991) reports the occurrence of the species south of 
Madeira. We here confirm one specimen from the Azores and 
note others from southeast of the area (MCZ 96916). 

Azores Material. 149506, R/V Oceanus 158, KEH8489, 
38°06'N 29°44'W, 400 mwo, 0010-0103 h, 02.IX.1984, 1:62 mm 
SL. 

Lampanyctus cuprarius Taning 1928 

Comments. A bipolar subtropical species (Nafpaktitis et al., 
1977) that is found principally south of 42°N on both sides of 



1999 AZORES DEEP-SEA FISHES 13 

the Azores EEZ. MCZ material confirms this species for the 
Azores. 

Azores Material. 112931, R/V Chain 105, RHB2553, 34°12'N 
24°0rW, 125-135 m, 2355-0105 h, 08.VII.1972, 2:46-65 mm 
SL; 151346, RA^ Atlantis II 101, CEL7838, 37°47'N 23°52'W, 
350 m, 0045-0347 h, 11. VII. 1978, 4:45-57 mm SL. 

Lepidophanes gaussi (Brauer 1906) 

Comments. A bipolar subtropical species that is found princi- 
pally south of 37°N that was expected to occur in the Azores 
since the area is on the northernmost part of its range (Nafpaktitis 
et al, 1977). MCZ material establishes this species for the 
Azores. 

Azores Material. 109522, RA^ Atlantis II 49, RHB1916, 
35°30'N 2r46'W, 39-41 m, 2229-0003 h, 24-25.VI.I969, 8:42- 
48 mm SL; 109523, RA' Atlantis II 49, RHB1919, 35°56'N 
22°40'W, 650-750 m, 0708-1030 h, 25.VI.I969, 3:40-43 mm 
SL; 109524, RA^ Atlantis II 49, RHB1920, 36°23'N 23°35'W, 63- 
65 m, 2045-2218 h, 25.VI.1969, 3:42-44 mm SL; 109591, RA' 
Chain 105, RHB2551, 34°49'N 24°32'W, 700-740 m, 1620-1845 
h, 08. VII. 1972, 1:45 mm SL; 109592, R/V Chain 105, RHB2552, 
34°17'N 24°05'W, 60-70 m, 2158-2305 h, 08.VII.1972, 2:40-43 
mm SL; 109671, RA' Delaware 7/63-04, DL63-04:012, 36°57'N 
24°50'W, 180 m, 1730-1815 h, 12.V.1963, 1:42 mm SL. 

Loweina rara (Lutken 1892) 

Comments. L. rara is known from tropical areas of the Atlantic, 
Pacific, and Indian oceans (Nafpaktitis et al, 1977; Bekker, 
1983). In the Atlantic, the Azores is on the northern edges of its 
range. HuUey (1984) shows a locality near the Azores, which is 
confirmed by four stations in the MCZ material. 

Azores Material. 102770, RA^ Delaware II, DL63-04:012, 
36°57'N 24°50'W, 180 m, 1730-1815 h, 12.V.1963, 1:41 mm SL; 
151184, RA^ Atlantis II 101, CEL7822, 39°41'N 32°19'W, 350- 
170-75 m, 2347-0140 h, 05-06.VII.1978, 2:41 mm SL; 151185, 
RA^ Atlantis II 101, CEL7823, 39°46'N 32°09'W, 350-175-75 
m, 0150-0340 h, 06.VII.1978, 1:41 mm SL; 151186, RA^ Atlan- 



14 BREVIORA No. 507 

tis II 101, CEL7829, 3/°54'N 27°22'W, 350-145 m, 0140-0340 
h, 08.VII.1978, 1:39 mm SL. 

Myctophum nitidulum Garman 1899 

Comments. Occurs in the Indian, Pacific, and Atlantic oceans 
as a common tropical/subtropical species (Nafpaktitis et al, 
1977). Its presence in the area was expected, considering its dis- 
tribution. The Azores is the northeastern edge of the species' 
range, but it does not appear to be common in the EEZ. 

Azores Material. 106197, R/V Atlantis II 49, SUN 1921, 
36°27'N 23°42'W, surface, 0000-0415 h, 25.VI.1969, 3:29-57 
mm SL. 

Notoscopelus caudispinosus (Johnson 1863) 

Comments. Occurs in the Atlantic, Pacific, and Indian oceans. 
Questionably referred to a tropical/subtropical pattern (Nafpaktitis 
et al, 1977). This species is more common in the western North 
Atlantic. The Azores represents the northern boundary in the east- 
em Atlantic, but it is apparently not common in the area. 

Azores Material. 104073, RA^ Chain 105, RHB2552, 34°17'N 
24°05'W, 60-70 m, 2158-2305 h, 08.VII.1972, 1:106 mm SL; 
104093, RA^ Chain 105, RHB2554, 34°08'N 23°59'W, 480-520 
m, 0115-0235 h, 09.VII.1972, 1:60 mm SL. 

Melamphaidae 
Scopelogadus mizolepis mizolepis (Giinther 1878) 

Comments. S. mizolepis is a bathypelagic species that inhabits 
the Atlantic, Pacific, and Indian oceans, but it was not previously 
known for the northeast Atlantic (Ebeling and Weed, 1963, 1973; 
Maul, 1973b, 1986). This subspecies is widespread in the western 
North Atlantic and in the eastern tropical Atlantic to South Africa 
(EbeUng and Weed, 1963, 1973; Ebeling, 1986; Maul, 1990). The 
juvenile specimens reported here from the western edge of the 
EEZ might well be waifs from the western Atlantic. Ebeling iden- 
tified these specimens in 1963. 

Azores Material. 42891, RA^ Atlantis Iselin, CODI/ATL119, 
40°05'N 35°10'W, 1260-1440 m, ? h, 07.IX.1928, 2:19-24 mm 
SL. 



1999 AZORES DEEP-SEA FISHES 15 

Epigonidae 
Microichthys coccoi Riippell, 1852 

Comments. Seven specimens of this rarely collected fish were 
caught in Azorean waters by the WHOI expeditions. It is inter- 
esting to note that these specimens were the only members of the 
genus Microichthys taken in thousands of WHOI trawls in the 
North Atlantic. The comparison of these juveniles and postlarvae 
with other epigonid specimens and illustrations shows that they 
do not belong to Epigonus telescopus since they lack the typical 
diagonal pigment slash on the caudal penducle. The MCZ spec- 
imens agree very closely with those figured as Epigonus teles- 
copus in Kofoed (1952:1, pi. IIA; see Tortonese, 1986). Our re- 
cords confirm this taxon in the Azores, but they are not the first 
for the area since all four specimens observed by Koefoed (1952) 
were caught at the EEZ. 

Azores Material. 149630, RA^ Atlantis II 101, CEL7822, 
39°41'N 32°19'W, 350-170 m, 2347-0140 h, 05-06.VII.1978, 1: 
13 mm SL; 149629, RA^ Atlantis II 101, CEL7829, 37°54'N 
27°22'W, 350-145 m, 0140-0340 h, 08.VII.1978, 1:21 mm SL; 
149631, RA^ Atlantis II 101, CEL7834, 36°36'N 25°07'W, 350- 
170 m, 2355-0142 h, 09-10. VII. 1978, 1:19 mm SL; 149632, RJ 
V Atlantis II 101, CEL7837, 37°48'N 23°39'W, 350 m, 2211- 
0045 h, I0-11.VI.1978, 2:17-20 mm SL; 149633, RA^ Atlantis 
II 101, CEL7838, 37°47'N 23°52'W, 350 m, 0045-0347 h, 
11. VII. 1978, 1:23 mm SL; 149634, R/V Atlantis II 101, 
CEL7840, 38°49'N 22°20'W, 350-180 m, 2155-0047 h, 11- 
12.VII.1978, 1:18 mm SL. 

CONCLUSION 

The species reported here for the first time add about 5% to 
the total Azorean fish fauna and more than 15% to the known 
mesopelagic fishes (Santos et al., 1997). The WHOI/MCZ Azo- 
rean collections represent more than 75% of all species known 
from the region and are certainly the most extensive documen- 
tation of mesopelagic fishes of the area. 

The oceanic ichthyofauna of the Azores is primarily subtropi- 
cal/temperate with a few species from subpolar cold water and 



16 BREVIORA No. 507 

warmer tropical waters. The archipelago is located at the bound- 
ary between temperate and subtropical water masses, since the 
Azores Front meanders through the area around the 35°N sepa- 
rating the two regions. The influence of the Azores Front on the 
biogeographic patterns of the Azores area has not been assessed. 
Backus et al. (1977), using collections of myctophids, created 
zoogeographic regions and provinces for the Atlantic. They in- 
clude the Azores in the Azores-Britain Province of their North 
Atlantic Temperate Region. The definition of this province, how- 
ever, raises some questions because faunistic and oceanographic 
gradients were observed throughout the area considered. Further 
evaluation of the role of mesoscale oceanographic structures as 
physical boundaries for mesopelagic fish will be of interest for 
the future development of an actual biogeographic model. 

ACKNOWLEDGMENTS 

We thank John Kelly, Deborah Triant, and Peter Mclntyre for 
their assistance in the MCZ collection and Ana Martins for her 
critical review of the manuscript. The collecting, sorting, and 
transfer of materials to the MCZ and computerization of the 
WHOI collection was largely supported by the U.S. National Sci- 
ence Foundation between 1961 and 1995. This paper was par- 
tially supported by the Funda^ao Luso-Americana para o Desen- 
volvimento (FLAD) (Proc. 3.L/A. V/P 561/96). We thank R. H. 
Backus, L.P. Madin, and G. R. Harbison, who were chief scien- 
tists on the cruises that collected the bulk of the material. 

LITERATURE CITED 

Arruda, L. M. 1997. Checklist of the marine fishes of the Azores. Arquivos 
Museu Bocage 3(2): 13-164. 

Backus, R. H., J. E. Craddock, R. L. Haedrich, and B. H. Robinson. 1977. 
Atlantic mesopelagic zoogeography. Memoir of Sears Foundation for Marine 
Research 1(7): 266-287. 

Badcock, J. 1970. The vertical distribution of mesopelagic fishes collected on the 
Sond Cruise. Journal of Marine Biological Association of the United King- 
dom 50: 1001-1044. 

. 1984a. Gonostomatidae, pp. 284-301. /// P. J. Whitehead, M.-L. Bauchot, 

J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east 
Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 510 p. 

. 1984b. Sternoptychidae, pp. 302-317. In R J. Whitehead, M.-L. Bauchot, 



1999 AZORES DEEP-SEA FISHES 17 

J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east 

Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 510 p. 
Bekker, V. E. 1983. Myctophids of the World. Moskva, Akademyia nauk SSSR. 

248 pp. (In Russian) 
Bertelsen, E., G. Krefft, and N. B. Marshall. 1976. The fishes of the family 

Notosudidae. Dana-Report 86: 1-1 14 -I- 1 pi. 
BoLiN, R. 1959. Iniomi, Myctophidae. Report on the Scientific Results of the 

"Michael Sars" North Atlantic Deep-Sea Expedition 1910 4(2, 7): 1-45. 
Borodin. N. A. 1931. North-Atlantic deep-sea fishes. Bulletin of the Museum of 

Comparative Zoology 72(3): 55-89. 
COAD, B. W. 1986. The shortbarbel dragonfish, Stomias brevibarbatus new to the 

fish fauna of the Atlantic coast of Canada. Canadian Field Naturalist 100(3): 

394-395. 
Cohen, D. M. 1984a. Bathylagidae, pp. 392-394. //; P. J. Whitehead, M.-L. Bauch- 

ot, J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east 

Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 
. 1984b. Opisthoproctidae, pp. 395-398. In P J. Whitehead, M.-L. Bauch- 

ot, J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east 

Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 510 p. 
. 1990. Argentinidae, pp. 235-238. In J. C. Quero, J.-C. Hureau, C. Karrer, 



A. Post, and L. Saldanha (eds.). Check-list of the Fishes of the Eastern Trop- 
ical Atlantic. Vol. I. Lisboa, JnictAJnesco. 510 p. 

CoLLETT, R. 1896. Poissons provenant des campagnes du yacht L'Hirondelle 
(1885-1888). Resultats des Campagnes Scientifiques Accomplies sur son 
Yacht par Albert ler Prince Souverain du Monaco, Fascicule X, Imprimerie 
du Monaco. 198 pp. + 6 pi. 

. 1905. On some fishes from the sea off the Azores. Zoologischen Anzeiger 

28(21/22): 723-730. 

Ebeling. a. W. 1986. Family Melamphaidae, pp. 427-431. In Smith, M. M., and 
P. C. Heemstra (eds.). Smith's Sea Fishes. Macmillan South Africa (Publ.) 
Ltd. Johannesburg xx -I- 1047 pp. 

Ebeling. A. W., and W H. Weed III. 1963. Melamphaidae III: Systematics and 
distribution of the species in the bathypelagic fish genus Scopelogadus Vail- 
lant. Dana-Report 60: 1-58. 

. 1973. Order Xenoberyces (Stephanoberyciformes), pp. 397-478. In D. 

M. Cohen, J. W. Atz. F H. Berry, J. E. Bolke, R. H. Gibbs, E. A. Lachner, 
G. W Mead, and D. Mehinan (eds.). Fishes of the Western North Atlantic. 
Memoir — Sears Foundation for Marine Research. Vol. 1, No. 6. 

Ege, V. 1930. Contributions to the knowledge of the North Atlantic and Mediter- 
ranean species of the genus Paralepis Cuv. A systematical and biological 
investigation (Sudidae, Paralepis). Report on the Danish Oceanographical Ex- 
peditions 1908-1910 to the Mediterranean and Adjacent Seas 2(Biology, A. 
13): 1-193. 

. 1953. Paralepididae I (Paralepis and Lestidium). Taxonomy, ontogeny, 

phylogeny, and distribution. Dana-Report 40: 1-184. 



18 BREVIORA No. 507 

GiBBS, R. H. Jr. 1984a. Astronesthidae, pp. 325-335. In P. J. Whitehead, M.-L. 
Bauchot, J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.), Fishes of the 
North-east Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 510 p. 

. 1984b. Melanostomiidae, pp. 341-365. In P. J. Whitehead, M.-L. Bauch- 
ot, J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east 
Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 510 p. 

. 1984c. Stomiidae, pp. 338-340. In P J. Whitehead, M.-L. Bauchot, J.-C. 



Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east Atlantic 
and the Mediterranean. Vol. I. Paris, Unesco. 510 p. 

Gibes, R. H., Jr., and M. A. Barnett. 1990. Melanostomiidae, pp. 308-337. In 
J. C. Quero, J.-C. Hureau, C. Karrer, A. Post, and L. Saldanha (eds.). Check- 
list of the Fishes of the Eastern Tropical Atlantic. Vol. I. Lisboa, Jnict/Unesco. 

Goodyear, R. H. 1990. Malacosteidae, pp. 338-340. In J. C. Quero, J.-C. Hureau, 
C. Karrer, A. Post, and L. Saldanha (eds.). Check-list of the Fishes of the 
Eastern Tropical Atlantic. Vol. I. Lisboa, Jnict/Unesco. 

Goodyear, R. H., and R. H. Gibbs, Jr. 1970. Ergebnisse der Forschungsreisen 
des FFS "Walther Herwig" nach Siidamerika. X. Systematics and zoogeog- 
raphy of Stomiatoid fishes of the Astronesthes cyaneus species group (Family 
Astronesthidae), with description of three new species. Archiv fur Fischer- 
eiwissenschaft 20(2/3): 107-131. 

Haedrich, R. L. 1964. Food habits and young stages of North Atlantic Alepisau- 
rus (Pisces, Iniomi). Breviora, 201: 1-15. 

Harold, A. S. 1994. A taxonomic revision of the stemoptychid genus Polyipnus 
(Teleostei: Stomiiformes) with an analysis of phylogenetic relationships. Bul- 
letin of Marine Science 54(2): 428-534. 

Heemstra, P C. 1990. Anotopteridae, pp. 142-143. In O. Gon. and P C. Heemstra 
(eds.). Fishes of the Southern Ocean. Grahamstown, J. L. B. Smith Institute 
of Ichthyology. 462 pp. -I- 12 pis. 

HULLEY, P A. 1984. Myctophidae, pp. 429-483. //; P J. Whitehead, M.-L. Bauch- 
ot, J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east 
Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 

Hureau, J. C, and Th. Monod (eds.). 1973. Check-list of the Fishes of the North- 
eastern Atlantic and of Mediterranean. Vol. I-II. Paris, Unesco. xxii -t- 
1014 pp. 

Jespersen, p. 1915. Sternoptychidae (Argyropeleciis and Sternoptyx). Report on 
the Dana Oceanographical Expeditions 1908-1910 to the Mediterranean and 
Adjacent Seas 2(Biology, A2): 1-41. 

KoEFOED, E. 1952. Zeomorphi, Percomorphi, Plectognathi. Report on the Scien- 
tific Results of the "Michael Sars" North Atlantic Deep-Sea Expedition 1910 
4(2, 2): 1-26. 

. 1956. Isospondyli. 1. Gymnophotodermi and Lepiphotodermi. Report on 

the Scientific Results of the "Michael Sars" North Atlantic Deep Sea Ex- 
pedition 1910 4(2, 5): 1-21. 

. 1958. Isospondyli. 2. Heterophotodermi. 1. Report on the Scientific Re- 



1999 AZORES DEEP-SEA FISHES 19 

suits of the "Michael Sars" North Atlantic Deep-Sea Expedition 1910 4(2, 
6): 1-17. 

Markle, D. E, and J. C. QuERO. 1984. Alepocephalidae, pp. 228-253. //; R J. 
Whitehead. M.-L. Bauchot, J.-C. Hureaii. J. Nielsen, and E. Tortonese (eds.). 
Fishes of the North-east Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 
510 p. 

Markle, D. F, and Y. I. Sazonov. 1990. Alepocephalidae, pp. 246-264. In J. C. 
Quero, J.-C. Hureau, C. Karrer, A. Post, and L. Saldanha (eds.). Check-list 
of the Fishes of the Eastern Tropical Atlantic. Vol. I. Lisboa, Jnict/Unesco. 
1492 p. 

Matsui, T. and R. H. Rosenblatt. 1984. Review of the deep-sea family Platy- 
troctidae (Pisces: Salmonifirmes). Bulletin of the Scripps Institution of Ocean- 
ography. Vol. 26. vii -I- 159 pp. 

Maul, G. E. 1946. Monografia dos peixes do Museu Municipal do Funchal. Or- 
dem Iniomi. Boletim do Museu Municipal do Funchal (2, 2): 1-22. 

. 1956. Additions to previously revised Orders or Families of fishes of the 

Museu Municipal do Funchal (Stomiatidae, Astronesthidae, Paralepididae). 
Boletim do Museu Municipal do Funchal 9(24): 75-96. 

. 1973a. Anotopteridae, p. 102. In Hureau. J. C, and Th. Monod (eds.). 

Check-list of the Fishes of the North-eastern Atlantic and of Mediterranean. 
Vol. I. Paris, Unesco. 510 p. 

. 1973b. Melamphaidae, pp. 343-346. //; Hureau. J. C. and Th. Monod 

(eds.). Check-list of the Fishes of the North-eastern Atlantic and of Mediter- 
ranean. Vol. I. Paris, Unesco. 510 p. 

. 1986. Melamphidae, pp. 756-765. In P J. Whitehead, M.-L. Bauchot. J.- 

C. Hureau. J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east At- 
lantic and the Mediterranean. Vol. II. Paris. Unesco. 517-1007. 

. 1990. Melamphaidae. pp. 612-618. In J. C. Quero. J.-C. Hureau, C. 

Karrer, A. Post, and L. Saldanha (eds.), Check-list of the Fishes of the Eastern 
Tropical Atlantic. Vol. I. Lisboa. Jnict/Unesco. 

MiYA. M., AND M. Nashida. 1996. Molecular phylogenetic perspective on the 
evolution of the deep-sea fish genus Cyclothone (Stomiiformes: Gonosto- 
matidae). Ichthyological Research 43(4): 375-398. 

Murray, J., and J. Hjort. 1912. The Depths of the Ocean. London. Macmillan 
and Co. (1965 reprint), xx + 821 pp. -t- 4 maps -t- 9 pi. 

Nafpaktitis, B. G. 1968. Taxonomy and distribution of the lanternfishes, genera 
Lobianchia and Diaphus, in the north Atlantic. Dana-Report 73: 1-131. 

Nafpaktitis, B. G., R. H. Backus, J. E. Craddock, R. L. Haedrich, B. H. Rob- 
ISON, AND C. Karnella. 1977. Family Myctophidae. pp. 13-265. In Fishes 
of the Western North Atlantic. Memoir — Sears Foundation for Marine Re- 
search. Vol. 1. No. 7. 

Nelson. J. S. 1994. Fishes of the World (3rd. edition). John Wiley, and Sons, 
Inc., New York, xvii -I- 600pp. 

Post, A. 1984. Anotopteridae. pp. 509-510. In P J. Whitehead, M.-L. Bauchot, 



20 BREVIORA No. 507 

J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east 
Atlantic and the Mediterranean. Vol. I. Paris, Unesco. 

1990. Paralepididade, pp. 373-384. In J. C. Quero, J. C. Hureau, C. 



Karrer, A. Post, and L. Saldanha (eds.). Check-list of the Fishes of the Eastern 
Tropical Atlantic. Vol. I. Lisboa, Jnict/Unesco. 

Quero, J. C, T. Matsui, R. H. Rosenblatt, and I. Sazonov. 1984. Searsiidae, 
pp. 256-267. //; P J. Whitehead, M.-L. Bauchot. J.-C. Hureau, J. Nielsen, 
and E. Tortonese (eds.). Fishes of the North-east Atlantic and the Mediter- 
ranean. Vol. I. Paris, Unesco. 

Quero, J. C, J.-C. Hureau, C. Karrer, A. Post, and L. Saldanha. 1990a. 
Check-list of the Fishes of the Eastern Tropical Atlantic. Lisboa, Jnict/Unes- 
co. Vol. I-III: 1492pp. 

Quero, J. C, J. C. Njock, and M. M. de la Hoz. 1990b. Gonostomatidae, pp. 
283-292. //; J. C. Quero, J.-C. Hureau, C. Karrer, A. Post, and L. Saldanha 
(eds.). Check-list of the Fishes of the Eastern Tropical Atlantic. Vol. I. Lisboa, 
Jnict/Unesco. 

Quero, J. C, J. C. Njock, and M. M. de la Hoz. 1990c. Stemoptychidae, pp. 
275-282. //; J. C. Quero, J.-C. Hureau, C. Karrer, A. Post, and L. Saldanha 
(eds.). Check-list of the Fishes of the Eastern Tropical Atlantic. Vol. I. Lisboa, 
Jnict/Unesco. 

Regan, C. T, and E. Trewavas. 1929. The fishes of the families Astronesthidae 
and Chauliodontidae. The Danish "Dana" — Expedition 1920-22 in the North 
Atlantic and the Gulf of Panama 5: 1-39. 

Richard, J. 1905. Campagne scientifique du yacht "Princesse-Alice" en 1904. 
Observations sur la faune bathypelagique. Bulletin de ITnstitut du Musee 
Oceanographique du Monaco 41: 1-30. 

ROULE, L. 1919. Poissons provenant des campagnes du yacht "Princesse-Alice" 
(1891-1913) et du yacht "Hirondellell" (1914). Resultats des Campagnes 
Scientifiques Accomplies sur son Yacht par Albert ler Prince Souverain du 
Monaco, LII. 190 pp. + errat. -I- 7 pi. 

RouLE, L., and F Angel. 1924. Notice preliminaire (IVe et demiere) sur les larves 
et les alevins de Poissons recueillis par S. A. S. le Prince Albert I de Monaco 
dans ses croisieres (annees 1905 a 1915). Bulletin de ITnstitut Oceanograp- 
hique (Fondation Albert ler de Monaco) 451: 1-7. 

Santos, R. S., F M. Porteiro, and J. P. Barreiros. 1997. Marine fishes of the 
Azores: Annotated checklist and bibliography. Arquipelago. Life and Marine 
Sciences, Supplement 1, Ponta Delgado. xxviii -I- 244 pp. 

Sutton, T T, and T L. Hopkins. 1996. Species composition, abundance and 
vertical distribution of the stomiid (Pisces: Stomiiformes) fish assemblage of 
the Gulf of Mexico. Bulletin of Marine Science 59(3): 530-542. 
SwiNNEY, G. N. 1991. Lampadena anomalu Parr (Myctophiformes, Myctophidae) 

in the NE Atlantic. Journal of Fish Biology 38(6): 959-960. 
Tortonese, E. 1986. Apogonidae, pp. 803-809. In P J. Whitehead, M.-L. Bauch- 
ot, J.-C. Hureau, J. Nielsen, and E. Tortonese (eds.). Fishes of the North-east 
Atlantic and the Mediterranean. Vol. II. Paris, Unesco. 



1999 AZORES DEEP-SEA FISHES 21 

Vaillant, L. 1888. Poissons. Expeditions Scientifiques du Travailleur ct du Tal- 
isman pendant les annees 1880, 1881, 1882, 1883. Paris. 403 pp + xxviii pi. 

Whitehead, M.-L., Bauchot, J.-C. Hurhau, J. Nielsen, and E. Tortone.se (eds.). 
1984-1986. Fishes of the North-east Atlantic and the Mediterranean. Vol. I- 
III. Paris, Unesco. 1473 pp. 

ZuGMAYER, E. 191 1. Poissons provenant des campagnes du yacht Princesse- Alice 
(1901-1910). Resultats des Campagnes Scientifiques Accomplies sur son 
Yacht par Albert ler Prince Souverain du Monaco, Fascicule 35. 174 pp. + 
6 pis. 

. 1933. Liste complementaire des determinations faites par M. Zugmayer. 

Appendice, pp. 73-85. In Roule, L., and F. Angel (eds.). Poissons provenant 
des campagnes du Prince Albert ler de Monaco. Resultats des Campagnes 
Scientifiques Accomplies sur son Yacht par Albert ler Prince Souverain du 
Monaco, Fascicule 86: 1-115. 



22 



BREVIORA 



No. 507 



N 
U 
W 

z 
< 

o 

N 
< 

tu 
I 

H 



1/2 

tu 

C/5 



O 
O 

a: 



X 

f- 
o 

u. 
< 

< 
Q 



X 

5 
z 
w 

Q. 

a. 



00 






a 
Q 



00 

c 

o 



03 



O 

Z 



Q 



c^csccrtc^c^rtc^ 



?3:^c3Mc3car5:3:3Sc3c3 

-aT3-a-OTr-a-OT3-aT3T3-T3 
oooooooooooo 
ZZZZZZZZZZZZQQQZQQZZZZZZZ 











o 














































"^• 














































<u 


« 


-- 


a 


rt 




rt 




u 


a 


a 


o 


^ 


O 


§ 


1^ 


O 






in 




1) 


U 


D 


'■5 

u 
3 


■a 
o 
Z 


o 

00 

n 


o 
Z 


O 
O 
ri 


:3 

■a 

O 

Z 


o 

rl 

in 


o 

Z 


o 

00 


'J 

u 
3 


r3 

■o 

O 

Z 


C3 
O 

Z 


1 
O 
1^ 


1 
in 

IT) 


1 

O 

ITi 


ir, 

O 
in 


1 
O 
r| 


1 
o 


O 


O 


1 




3 


O 

u 
3 


3 


C/5 


— 


1^. 


ri 




OO 


-t 


*^ 


r*", 


t 


(^ 


rj 


O 


O 


r) 


o 


GO 


on 


C/0 



^^^^^^^^^^^^^^^^^^^^^^^^^ 



so 

o 
ON 



o — 



in 



o o o in o 
O D in r) -^ 



r) 00 On — 

m O — -^ 



-*---- 



o 
o 



O I^ o 

— mo 



in — OONOON'tinONCNONONONOOt^t^vO — — 'r<-) — — 
r<"< r<-i r<-i r^ n r) n tN (N m (N CN <N r) O) ri r) r<-, f<-i rr-, r<-, r<", 



1^ 

o 



zzzzzzzzzzzzzzzzzzzzzzzzz 



oooNint^i^t^oor~-inininosor<^r~-r--^'^ 
inTrO'*ror<-iinr)in(Nriinr)00^(Nno 



00 (^ '^ 00 t~~ '^ 

rj ro r^ r) (^ m 



0\ 


o 


o 


1^ 


in 


>n 


t^ 


.^ 


NO 


r^ 


1^ 


NO 


r<i 


^ 


t 


r<l 


1^ 
in 


NO 


00 
00 


-Nf 


m 


r<-i 


<^, 


m 


— 


'^ 


ON 




o 


00 


o 












O 

o 

5 


o 
o 

5 


5 


ON 


ON 


ON 


00 

ON 


nO 

o 
o 


o 


O 


o 


in 

o 


z 
< 


Z 

< 


Z 

< 


z 

< 


o 

m 


o 


3 


1 

r<1 


1 


-1 


-1 


-1 


-1 


nD 


\D 


NO 


NO 


NO 


u 


<j 


u 


H 


h- 


f- 


H 


-J 


-J 


-J 


-J 


-1 


u 


u 


u 


< 


< 


< 


< 


Q 


Q 


Q 


Q 


Q 



nO(N-^— 'OnOOnoOOnOnOnOnOnOn 



NO 


r- 


00 


On 


o 


— 


rl 


f<-i 


'I- 


in 


m 


(^ 


(^ 


m 


t 


^ 


■^ 


•^ 


^ 


^ 


o 


o 


o 


O 


o 


o 


O 


o 


o 


o 



o 



in 

o 



oQoacopapamBQpQpamzzz 
ft;Qia;cEiciioioiQiciiCESc/5c/5c>n 



00 

ri 

ON 

_>. 

"3 



I 

ON 



C<1 

NO 
ON 



^ 



— r~ 

ON ON 



00 

On 



I 



NO 

ON 



D 
CO 

[^ 

I 
o 



U 



dJ 




(/I 




CA 




1) 




> 


c 






j: 








11 




y; 


> 


CC 


^ 



s: s: 
-S -S 



1999 



AZORES DEEP-SEA FISHES 



23 



Q 
u 
—I 

z 

F 
z 
o 
U 



5 
z 

ou 

Cu 
< 



OJj 






D. 



OJj 

c 
o 
_1 



C3 



O 

Z 
"3 



C3 

Q 



ZZZZQZZZZQZZZZZZZZZZZZZZZ 



o 
I 



— o "/^ o 



I 
o 



o o 



I I 



o 
I 

m m iri 



I I I 



r^ On UO O _ - - 

oo — — >oo — — 



o o 

rl 00 
I 

o o 



n "/^ (N 



in ri 

o 



l^ O 



n 00 



I I 



O 

(N 

I 

O r) 






u 
o 



o 

C3 



O — — 



33333333333 



^^^^^^^^^^^^^^^^^^^^^^^^ 



On nD 



O 



riON'nnr^'/^OONONvo 
'^ '^ in ■^ ri rJ r<^ rJ r^ ■* 



inmoiONroinOON 
O r<~i '^ T)- o) r) rn r) 



<N (N CS (N fN r) M (N r) M ri ri r) (N r) (N n fN fN (N (N (N n (N (N 

ZZZZZZZZZZZZZZZZZZZZZZZZZ 



o ^ vo (^1 1^ D r- 
r<-, r<^ m ri rj r^i m 



o o o 



r) ro m r<^ in (^1 r-i r<^ (^ O in O O O o) 

nOOOOOOOOOOOOOOUuuutJuvJvv-'w 

ininininin>ONDNONor-r--ooooooininininNONOvor--oooooo 



r^imr'%(^mfOr<^r<^rn 



inNDt^ooovO — dr^'*in\Di^ooinNDt^ooo^nin\Ot^oo 
— — — — — ri ri ri r) ri ri ri rJ oi — . — — — ci ri ri ri ri r) o) 

mmmmmmmmSSmmmcQzzzzzzzzzzz 

XXXXXXXXXXXXXXOOODDDOOOOO 



ON 
NO 
ON 



c 

3 

ri 

I 
Tf 
ri 



;^ 

u 

u 

u 
u 

Qi 



ON 

u 

u 



s: 
3 



24 



BREVIORA 



No. 507 



cn) 



Q 






C 

o 



Q 
22ZQZZZZZZQZZZQZZZQZZZZZZZ 



in O in O O in >ri 

sO m in O t^ ON (^ 

o — n ^ O " 

I I I I I 

in o o 



o 

I 

in in O 

_ , . , , _ _ (^ vo in 

O — ri in O O — r) 



o 
I 



vo ri ri >.D ^ 



o O m in 
c) vo t^ -^ 



o 

I 

o 

o 



o o 
— ' ^ 

I I 



in o o 
m rj (N 
—■mm 

I I I 






in O 



vo ri 00 
O — •* 






3 3 3 3 3 3 3 

c/5 c/5 on 00 oo CO on 



^obNinf^Ot^rooooaN—'inorJin-^ONin— 'OONcnooor<^ 
OOin-^00'nr<^r<imm'^Mror<^OOininOOinOOr<-i(-<-i 

oooooooooooooooooooooooooo 
OOONONONONOOOO\OvOin^^"*^'*'*(^(NOOONONON^VO 

r<-i rn r) r) r-J ri ri ri ri ri fN (N cN r) tN ri rj rj r) m (^ n ri n rj ri 



zzzzzzzzzzzzzzzzzzzzzzzzzz 



Q 

u 

D 

z 

H 

z 

o 
u 



Q 

z 

Ou 

< 



m 

J 



o 

z 



vO — I^ in — ' 00 r<^ 

ri n — ' m in D rt 



vc^t^rjin— 'r-Oooo — "^oomCTNt^nooo 
r)(N— '■^inin'^— '(NfNr)— 'OO^-^— 'OO 

OOOOOOOOOOOOOOOOOOOuuuuuuu 

r-jo)MOONONOsONt^r~vo>ninin-^'*'^'^'^(NCNrJONON(^t^ 



m vO I^ 00 
m r<-) r^ r^ 



O— ■nm^'sOi^oooNO— 'r)r<iTi-inin>ot^O — "+ 
'^■^'*'*"*"*'^"*'*i^i/^''^''^''^''^<^<^<^'*'^'* 
ininininininininininininmininininininininininininin 
ri r) r) ri ri o) n ri ri ri ri ri ri ri ri r) ri ri ri n rj rj <^i o) rj rt 

D^ X nC X £ 03 






m m CQ ,z z Z Z 
n d: d: D D D D 



z 



z 



ON 



Q 



^ 

3 



I 
in 

o 



in 

o 



S3 
a; 



1999 



AZORES DEEP-SEA FISHES 



25 



D 
W 
D 
Z 

P 

z 

o 
U 



X 

S 
z 



Oil 



Q 



a. 
Q 



OJJ 

c 

o 



r3 



o 
Z 

2 






ZZZ2ZZZQZZZQZZZZZQZZZZZQZ 



I/; 

u 

u 





















ir-l 


ir, 




















O 


O 


O 
00 
























t^ 


1^ 








Q 


o 










00 


00 


— ^ 














Q 








o 


6 


1 




O 


O 


vO 


vC 


w-) 




O 


O 


1 

o 


1 

o 


1 

O 






(L> 


U 


aj 


<U 


Q 








OS 




(N 


(N 


o 


o 


t 




Ov 


OS 


r- 


t^ 


t^ 


o 




y 




3 




^ 


o 




o 


? 




rj 


1 


1 


^ 


't 


'"' 




"j" 


7 


1 








u 


u 


t4- 




c 


o 
q 


o 


1 




6 


6 


7 


O 


O 


6 


6 


O 




o 


O 


o 


6 


6 


1 


o 


3 


3 


3 


3 


n 


o 


o 


Q 


ir-i 


in 


■T) 


iri 


O 


ri 


ri 


in 


O 


im 


m 


ir, 


Vi 


m 


o 


lO 


00 


00 


00 


C/^ 


U-l 


— ' 


00 


't 


ri 


r^ 


r<~i 


r<-i 


r<"i 


^ 


— 


— ' 


r~, 


^ 


rn 


r<~i 


r>-i 


r<~j 


t^t 


'~^ 


r^, 



^^^^^^^^^^^^^^^^^^^^^^^^^ 



ro t~- ^ 

o o o 



r^ 00 in ON 
c — rj r<-, 



o — inin^O — ■^<NONOsON00ONr-jr<-)ri-i(Nin 
r<~)Tt(NOO — OO — — 'Or^r^r^r^r^tNin 

^^V^voV^r<^(N<N<N — oor-r-r-mmmminin^ 



__. , , _ r»l 

r) tN (N r^ ro (^ (^ rn rn rn r<^ (^. ro r^J ri rt rj N rj (N 04 (N rJ fN 



zzzzzzzzzzzzzzzzzzzzzzzzz 

^j^I—^^onovon---— ^inr--voK vcoooooo 
— — Ttr+T]-inminininin>nr<^mr^ir<-;mo;^ 

o o o o o o o o o o ° o_ ° ° °_ ° ° 

ONONONoooooor-r-r-in^vovo^vor~-r- 



/— ^ 


vr 


00 


1^ 


ON 


o 


r) 


^ 


ri 




n 


— 


o 


m, 


^— 


^- 







o 





o 








1^ 


in 


ITi 


^ 


r~ 


O 


O 


o 


c^ 


(^ 


C^ 


f^ 


m 


't 


1- 


^ 



\D 00 a\ <^\ Q 00 a^ C> '^ n rn ^ 'T, -^ t> CO o^ (D ^ r\ (T-i ^ tr, ^ ^ 

't ^ '^ mi O — — ri rj rj r) r) rj r) rj ri ri (^ r^. r^i r<-i (^. <^ i^ <;^ 

ininmmONoooooooooooooooooooooooooooooooooooooooo 



rj (N ri n n [^ t^ 

Z Z Z Z £ J " 



D 'Z:i D 



U 



(^ 1^ i^ t^ t^ 1^ 

J J J J J J J 
uj w ua pj PJ w u 



r^ t^ 1^ 

J J -J 









1^ 



PJWWUUJUPJWUUUJW 



EoSoJT^EoOiUUUUUUUUUUUUUUUUUUUU 





00 




r^ 


m 


ON 


r- 


"■^ 


ON 


>. 








3 


,4_j 


^-^ 


(1) 


O 


00 


1 


in 


in 


o 


o 



00 o 



-2 -2 



26 



BREVIORA 



No. 507 






Q 



ZQZZZZZZZZZZZZZZZZZZ 



Q 



o 

On 



O 

00 



O 
On 



O 
00 



o 

uo O 



O 

E 



o o 

c, o 



o o 



E S 



o 

E 

o 
c 

o 



o 
5: 

E 

o 
o 



o 



o 

o 



VO ITj — — Tt — . 



o 

E 

o 
o 



o 
o 
o 



oc 

c 
o 
-J 



OOONiTjinirjV — — 'OfNCN^^ONOOON^K 

O r) O •/"; iTi IT) iTi ri rj (N (N (N (N O O m r^i "* lO 



(N (N (N (N r<-j Ci r^ (^ m (^ m f<^ (^ (^ m (^ n ri r-i (^1 



O O On ON ON VO 



o 

u 

a; 



zzzzzzzzzzzzzzzzzzzz 



Q 

D 









ONTft--r-~>o(^>nir) — oor^iTino— 'OvO'^ 

■^T)-r<^r<^rnr<^OOOininirimr<^— '-hOO 



r-~oooooor~-i^r^t~-0\ONONOooooooooooooooooo 



H 
Z 
O 

U 



5 
z 
u 
a. 
a. 

< 



o 
Z 



00 ON o — IT' :£' i^ 

r*-i r'-i ^ ^ "^ f^ r~~ 

00 00 00 GO 

I^ t^ 1^ t^ 

J J J 



ooonO — rir"/Ttir, vor^ooONO 
r^i^oooooooooooooooooooooN 



^ '+ ^ ^ rf ^ Tt 



^ ■* Tf 'I- Tt Tl- Tt 



^ -^ 

oooooooooooooooooooooooooooooooo 



uuwwwuwwuujo-ipauLiwoLiujpqwtLiu 



00 
ON 






ex, 
u 

CO 
fS 

I 

00 
3 
< 



1999 



AZORES DEEP-SEA FISHES 



27 



u 


UJ 




X 


N 




i- 


"^ 




z 


, 






Qi 




Q 






UJ 


i/; 




H 


Z 




QC 


u 




2 


S 




u 


u 




c^ 


s 




C/3 
UJ 


00 






u. 
O 




0. 


Qi 


< 


C/5 


UJ 




>^ 


m 


1^ 


S 


D 


_i 





Z 


D 


_1 

o 

o 

N 


GO 


Q 
< 

Q 




Z 


z 


UJ 

> 


o 

5 


< 

tin 

UJ 


< 


UJ 


J 


DS 


_1 




< 


_1 


Z 


Cl 


O 


UJ 


S 


u 


> 


o 


u. 


_j 


u 


O 


D 


u. 


a: 


Z 


o 


UJ 

m 


< 


5 


s 


5 


3 


z 


(/5 




UJ 




U 


< 
> 


UJ 


>< 


< 


X 


5 
z 


J 


z 


UJ 


Uj' 




0. 


O 


N 


Q. 
< 


< 


W 


— 
X 
H 


S 




< 


a: 


Z 


Z 


O 




UJ 


N 


Q 


S 


< 


UJ 
Q 





UJ 




_1 


I 


w 


f- 


u 


> 

UJ 

Q 


S 


z 


o 

u. 


o 
z 


Q 


to 




,1-^ 


uu 


< 


hJ 


y 


UJ 


C/5 


0. 
[/5 


< 
O 


5 
S 


I 


H 




C/D 






< 


UJ 

z 


UJ 

O 

z 

< 


on 


t/3 


Qi 


Ou 


< 




UJ 

Q 


1/) 

z 

o 

H 




ri 


UJ 
c/5 




>< 








C/5 




a 


D 




z 


O 




1 


> 




<« 


a. 





c 

3 
C 

12 



e 

3 
C 

GO 

_o 

a 

o 

N 
U 






0) 

Q. 



I 



o 
z 






X 

a: 






9 

00 

X 
LU 



X 



ffi 

X 

a: 






' .^ 00 






;^ u 



r- — 

. "P <^ 

oo 2 Cl 

r^ ^ oc 

oo — f»^ 



" 2: 

CD r-* „* 



— .00 



s5 S 



to Ov UJ 



U X 



O CQ 

oo X 
1- oL 



CO 

X 

a: 






CD m 

X I 
a: a: 



CQ 

I 

or: 



ffl 

X 
a: 



—  r^ 



CO 

X 



I 2^ 



CO 
I 
a: 



00 CO 

^5 



I 

of 


1^ 


00 




W-) 




«/^ 


(Vt 


00 




CO 


o 


i 


X 


^ 


-o* 


rs* 


— 

z 


f*^ 


r~4 


o 


TT 


< 


cn 


« 


^ 


X 


V 


< 



m 

X 
a: 



CQ 

I 
a!. 



«s « 



-O CQ «A 

S X § 



< 

-J 



< < J 



< < 



< < -J < 



8 5 



- X: o o 

T a T <^ 

oo ^ v<!) — 






oo -^ 



n m (N ^- "/^ 



fs f«^ r^ — 



o i: 



r^ ^ 



ON o 



(N (N ^ 



\o — — rs — 









8 

Ik 




^ -2 a s ? 

SP 3 S 'C 'C 




a- 



II 1 1 

aq ^ a; i>^ 



= I 



V 

z 



■3 

•a 

-3 i 

C S 

O U 

U Z 



4> 

n 

-3 



C^ 



■a 



1 I 

o a 



n 

•a 
:a 
n 



41 

•c 






.a 

a 
O 



^ s 






28 



BREVIORA 



No. 507 



D 
U 

D 
Z 

H 
Z 

o 
u 



ri 



C 



<U 



3 
C 

bO 

§ 

o 

N 

U 



11 

if 

oc _ ^ 

z: =3 "^ 

i^ = ;c 

iz :£ cc 

. X 

#*: ■:^ Dd 

■=■ '■^ - 

'^ 5 — 

o- » o 

r~ «/•, r- 

5 = 5 
rr — <~^ 

. o a: 



CO r^ 

X » 

O! CD 

I 

r a: 

s ^- 

- S 

S - 



5 " 

a; I 



^.2 



-5 ^ 

- _ • - o 

OO -, TV — 

I 2 ;:; ™ 

Q i " I 

. -> , r< 

<^, (-^ -o ^ 

f  C r-~ — 

f, T 00 

r— -T -Jn 



- 2: 



CO 

^ o^ o 
9° r" ^ 






-J CO _j 
^ X UJ 



oo" r* 



00 



00 -T — m 



< 



< < 







00 r- 




s?; 


'^ 




„ 






as: 




V-. 












v^, f-% 


<»i 




r^ 






r- — 




0^ — 






a 






J CO 




■n -T 


■V 








LU X 




— _ 


— 










o a: 




s s ^' 




1 






5t '~^' 




ss 


r4 




m 






o r~- 




J -J 


CO 




X 








Ci u; 


X 




a. 










o u 


o; 










, 










§ 






r- w-i" 




- oo' 


r4 




o^ 
















« 






_1 CO 

LU X 




1 - 






00 










00 (-. 


S 




I 










" _1 


1^ 
CD 




UJ 








-J UJ 

go 


X 

u 




.-1 03 


o< 










00" 




£i 


CTv 




ss 




0\ '^ 


r>t 




I's 


CQ 




U uj 


r^ 


TT "• 






« j:? 


X 




ou 


CQ 

X 


r< 


t— ' 


»*i 


X 00 
UJ . 


r-* 




S:' "' 


ot 


00 3 


0^ 




■■^9. 






00 T 


00 


^ o 


CQ 
X 


m 

X 

a: 


ii 


i 

CO 
X 
a:: 


00 

I 

LU 


OO 0^' 

-J _i 

UJ UJ 


CO 

X 


55 

< _. 

- m 


1*1 




ii 

^ 




T* 






8 S 





v^ 


oT "^ 


CT- 


T 




r^ 




r^ 


^ rs 


-O 






</-i 


f^ _i 


m 


CQ 


w-v CQ 




1 


^ R s 


< 


^ 


X 


IS 
< 


< 


J 


< 








_j 










-1 



— rj 



00 ;o p. p_ 00 :§ 

« f? ?? f? 2: ^ -^ 

[j _j -a _j CD CQ 9; 

[2 W UJ LJ X X — 

cj O tj o a; o; z 

. m* ^ o 00* "^ t<0 

— »/-i t/l -o ^ f~* 

■^ 2^ 5^ '?''■* 00 

5" <v Q O "^ "^ "^ 

Q J^. iF, J^, r^ r-^ 00 

— ""'.'*' 

-J" <N m rn — S^* j^' 

_ 00 00 00 '^ '^ >^ 

Q r- r- r- — — >1 

5 _j -J J m CQ ?^ 

2 UJ LU UJ X X CQ 

5 U U O a: oi X 

o - . - . - a; 

_J «*J Q V-, ^ — 

o Q P f; ■* 

ojj — m 

^ f^' — ■^* -v' r; r^ 

f^ f^ f^ fi -^ *^ lA 

_ 00 00 00 O ^ ^ 

. -I _i J m m g 

■^ UJ UJ UJ X X =: 

g U U O a; a: J 

L_ r-^ w^^ <T o^ 0> «' 

< « j; ;o K — o 

2 S S S ' " S 

Q -A ^ ■?. ^ r^ ^ 

o ~ ~ ~ ~ . . - 

CJ f^H* ^' 1^' 00' ^ c; 

fN r^ m m — ^ 

- 00 00 00 00 <^ ^ 

— r- t^ r- r- — r^ 

rj _] _j _1 _j m CQ 

O UJ UJ UJ UJ X X 

r-. u o (J (J o: a: 



6 



Q 
Z 

u 

Q. 

a. 
< 






__ o 



CO 
U 

U 

C/3 













5 
















s 


fc 




1 

1 






i 


1 


i 


1 
5 


3 

^5 


•i. 


Q 


•0 


s 


c^ 


•0 


Q 


^ 


13 


<;> 


%> 


^ 


%> 


1 


Q 


1 


t 

5 


§ 


•^ 


1 


1 


1 


^ 

2 


5 





^ 
u 





s 




i 





5s 


Ss 


>. 


;v 











09 


U 








U 












P 


S 






to 


<3 





la 


C 






3 


lO 





^ 


Q 


-<5 


•^ 





!^ 


« 


u 






-c 


<b 


J? 


I 


2? 






^ 


'^ 



■3 
ft 



IK 

e 



V 

a 

■3 

u 

i^ 

a. 
o 
c 



c/5 



1999 



AZORES DEEP-SEA FISHES 



29 



Q 

Z 

H 
Z 

o 
u 



r) 



D 
Z 

u 

0. 

< 



s 

C 
•T3 



(U 

E 

3 
C 

GO 

O 

B 
a 
o 

N 
U 



c/3 
U 



C/) 

'3 

00 



>o W-. op 

r-* oo r^ 

oo ^ O 

r^ oo — — ("-* rvi rs 

-1 X CO CD oa oa CO 

^ LU X X X X X 

U :^ a: Qc: a; oi ai 




00 r- ^ ^ ^  

o <> w^ «-i >; 

-o^xaamoQcoS 
ujluluxxxXt: 



I tju •• 1 -^ (-1 r-i r- 1 

. 2\ Ov oo m ■^ -T 

' O O "^ 'C 'O *o 

I »A *n m f*i 1^ r*! 



WW ffi <^ ^ w-T op* 

-1 X CD CO 3 m 

tU Ul I I T I 

O ii a; oi S^ g 



o 



_ o 



«0 

3 

s 

I 

s 

1 
I- 



2 
5 

-Si 
I- 




in 

3 



8 
I 






u 

•r 
o 



30 



BREVIORA 



No. 507 



Q 
U 

D 



E 

3 
C 

t+3 



0) 

£ 

3 
C 

60 

t 

o 

N 

U 



■O — <> 1^ 00 <*> 

f»i ■v — rn ^ v% 

o o 1^ v% t/i »n 

— — — fvi r^ r< 

CQ CQ CQ CQ CO ffl 

X X X X X X 

»: O^ Cll o^ of c^ 



3 — 00 t 
00 r< r* 



I ^ 1^ 



O ^ "^ "^ "-^ ' 

00 '*^ **> ''^ '*^ I 

S ^' 5 ?^" s 

2 o o *^ "^ 

r ?S 



00 — — r-« r< 

X CS CQ CQ CQ 

UJ X X X X 

U ct. Qi u t^ 






^ X 



r-i r^ >n "C 

X X ;3 r^ 
ai a: "^ CD 
. . I 
■o ><-> K ai 

>^ ^ 5^ - 



?: 2? 



g s * 

00 *" _ 

c s ^■ 



^2S 



1^ m 



CO i: - CQ 



CD CQ 






r.* 



I X CO 



.00 - . 00 — 

p~ r^ o r4 ro 00 

^ T 00 '^ "O ^ 

Ov 00 00 ^ (> 00 






- s S 

CQ CO rQ 

X X X 



• a <? 



" S -^ *^ 
ni (^ ^f 



2; '^ 

Es 

00 — 

00 — 
X CQ 
U X 

QO '^ 



00 



s § g , 

m CQ £ (M 

X X 5 CO 

a: Dd a; X 

. . - fti 

05 00 '^ *,* 

rn </% >0 </^ 

m -^ 3 S 

SCO op ^ 

rn <^ 00 



vt w*i tr, 



 r.* v^ 0\ 



1*1 



2; w^ r^ **i 

^ "T On »/^ »^ 

2 => - a r? 

ffi CO 

X X i X 

or: a: a: ^ 



X ca 



^s S 









f^ 


m rn 




r% 


r^ t^ 





"" 




f*l 


a 


§^ 


f^ 


w^ 




V, 


r^ 





ri m f^ * 



— *^ ""f .fi 
o* »^i "^ 5 

" ^ G ^ 



CT< m — 

s s ^ 

s 2 °° 

a: 2: c-r 

S X - 

3 - X 

00 M (^ 



:#■?;: 



§ i^' § #■ 9 

P C r^ S SS 



U tj M M 3^ u 



- r- r— — -^ ■n' 

^ W-l (*1 «-i 1.0 ^0 * 

'o f*i m ^ -^ n' ■^ 

rn 5- 00 00 00 00 » 

r^ o rn rn m rn ^ 

00 00 —. — — — — 



«/~i — r^ r* 
0\ m m r-n 



? g X 

- ^ n; 

-- 00 

pi 

—J — 00 

UJ ca 

U X 000 

_ Oi r^ o* 

O . '^ 'T' 

T rn — — 

«ri — CD to 

— 00 a: oi 



< < 



< < 



o 
u 



r4 '— 






X 

5 
z 

w 

CL 

Q. 

< 



C/5 



m ^ 



» "" 



— 'O 



_ o^ 



U 
CO 



5 

3 
& 






a 
s 



<3 

s 

<3 



c 

a. 

E 'E 

s a 
60 .60 

c s: 



I 



.5<i 



3 
■5 



I 






s 



■s: -5 
o o 

oq oq 



■3 

B 
O 



1999 



AZORES DEEP-SEA FISHES 



31 



D 

UJ 
D 
Z 

Z 

o 
u 



X 

5 
z 

0. 

< 



c 

3 
C 



XI 

E 

3 
C 

00 

_o 

B 
a 
o 

N 

U 



en 



CQ CQ CO 

III 
at ad. a 



S s 2; « ;f '; 

00 00 CT- "^ "^^ "^ 

fZ fZ _- r-t r~i <^ 

_j _i 03 :n CO 3: 

UJ UJ I X X X 

o o o; a: a; a; 



- _ ^^ 



OO O -Jv OC r^ 

«**! tr, r^ r^ f^i 

»> — r^ r-« »-* 

s s a s s 



s ri 2? , 

-) CD CO S 

'^ = X £ 

u o: ot: I 

•^ j:' S = 



.'1,^- 



c^ '^ 3 !C; 2 :::^ 

r^ r*-, ^ f^ ^ "^ 

OO 00 O ""i ^'^ '''i 

fSr r- _ r^ r^ /^ 

_1 J 21 CD CQ X 

UJ UJ X X X X 

U O a: oi oi a: 

V ■n' 00* r-" ^ — 

— CH ~- r^ r-t f, 
O O f^ r-J r^i r-* 

— — ON Q 0^ Q 
l/^ */% rM f*i r^ f*! 



(^ ''^ ^ "^ 9£ rt 

r^ r^ -^ ^ C- ^ 

(ir (S: — — r^ rn 

J J CD CQ CD 21 

UJ UJ X X X X 

U U aC 0^ al o£. 



^3 

• o 

IT °a 
Zi ^ 

UJ ^ 

US 

— o\ . 



CQ 

X 

a: 



r-< */"■ r^ oo rn 

r^ — 00 ^'^ o^ 

O O r^ — r^ <"-* 

— — 0^ Q o-. o 

i^ lAi r^ <*i 



f*! r-T 



00 2; r^ ;C' 

m -T- X m 

oS»=x 

S j; — r-. 



S «■ ° t; 

ujSXx uiujxxx = i r1 






^ -o _ r- t 
f^ <^ r-i ^. 



— _ — <7^ — — OnO 
V, I/", f*^ r-i w^ vr. r^ r^i 



O- O- ^ 



(J CQ 

S o; S 

S <-j c-< 



CQ 
X 
a! 



s ss: 



r5 



O 



S 



oo r^ O 5 

_J — 0\ 1 
UJ CO 00 ^ 

U I r^- 

- ^ 'C 



S i 



< < 



? 



u 



— i5\ rj I* 

v-i — CD ^ 

•^ ^C -F 0\ 

■^ 2 i "^ 

— 00 oc — 



CO 


„J 


X 


UJ 


a: 


O 






s 


!;< 




Ov 



Ul ^ 



^ ;7 



r-« — 

OO 9s 

J 5 

UJ X 

0\ — 



< < < < 



<N 


O 


CM 


o 


•■^ 


0\ 




o 
in 



— ri — ^ 



— fS — lO 



C/3 



s 

o 
"S 






■I 

O 

"a 

-5 



^ Vj 

•S "^ 
5 J 









o 

5 
<^ 



2 
c 

'5 s 






^ ^ a. 



a 

60 



o 



^ 5 



32 



BREVIORA 



No. 507 



Q 
Id 
D 
Z 

P 

z 
o 
U 



X 

Q 
Z 

u 

CL 

< 



X) 

E 

3 
C 



£ 

3 
C 

60 

B 

c3 
O 

N 
U 



c/3 



c/> 


§ 


fl> 


-Q 


O 


?? 


^ 


5 


C/3 


o 



<*i Qo «^ vi^ r^ (N 

*n rn — m »n ■«■ 

go 00 ON *n >*^ tri 

t^ r- — rM fN r4 

-3 -J CQ CD m GO 

w uj X I X r 

<-> O Qc: c.; a: or: 

;$ :£ w* * <5 f^* 

O O 00 00 00 00 

^ ^ o 2; ov 0\ 

^ ^ rN r-* f^ r^i 

C! Cl 5^ SI '^ 'T 

2P op (5 o tri w-i 

, '^ — ::: <^ <-^ 

r^ -J en £ cc cQ 



X 
a: 



is?: 

rn r-i (^ 

= CQ CB 
S ^ X 



X 

a: 



CQ 

I 



r- "^ — 

00 1^ QC 
S «ri w. 



U U 



I 5 3: I 



s s » =■ a s 

2 2 s? » K J 

•^ •^ ?:' S s; s 

— — 0\ ^ Oi &■ 



r-4 w-1 12 

r^ m 3 

00 00 o 

f^ f^ z: 

_1 _J to 

UJ LU X 

(J o o; 

•«T r~' oo' 



■^ S S 

S^ "^ ■^ 5 
X r-^ r^ ■> 

03 CO CQ S 

X X X CO 

a; o; a: X 

. . .ai 

o o *^ 

.— r^ r^ w-i 

CO oo oo r^ 

^ O 0\ 00 

CM r^ r^ ^ 



i * ; - 

O- R r.- S 



-J .A 

UJ fN 

tj CD 

I 

oo" aL 

9. « 



CD O W-, 
X — f^ 

Od CD CQ ' 

. X X ; 



. 




X - 


■A 


»N| 


^ r- 












*5 


CO 


CO 


X 


I 


00 I 


u 


a£ 


2 a; 



oo p; 



sS 



;£ S § S S S 5 

oo 00 00 c^ "^ "^ ^ 

r^ 1^ r^ — r-j f^ ij 

-1 _1 _) CO 03 CO S 

UJ LU UJ X X I ^ 

u u u a: o; a; ^ 



0> r>* ^ r- ^ ^ 

r^ ■^ o <n 1^ m 

o ^ 00 o oo 00 

^ O On <> ^ U\ 

"^ •'^p r^ f^ rvj r-t 



LU ffl 

U X 

. a: 



-J cc 



— SO 0^ — — — 



X — "^ 

;3 On W-. m 

QC — r^ w-i 

. CQ CD "^ 

v^ =: tii r-* 

r- -C X rn 

00 -^ 

'O . . Qi 

- r^ o- 

■O 0^ 1-^ f^ 

1^ r- ^/^ 00 

O 00 r-~ v-^ 

— -O o r- 

CQ 'O 

•^ w-T r-t" — 

OS -^ "^ «/-i 

. O </-■ V-, 

^y — r-i r^ 

t^ CQ CQ 00 

£ X X X 

^ U Qi. ai. 



CO 

X 



Si 

O CO 
^ ex 



o CD 
■o* V" 



_) 


-J 


< 


-J 


J 


J 


_1 


< 


< 

-1 






oo 








I 


00 


o 


IW 


w-i 


_ 


o 




»o 


vTi 


_ 


£ 



— r- -^ 



^ 



1 

5 


1 


.5 




1 

1 

3 


C<5 


1 


1 

\ 




1 
1 

8 


i> 


%> 


u 


Q 


a 


<0 


ifl 






1 


:0 


^ 


a 

<3 


1 


s 


§■ 


§■ 


to 


1 
s. 


1 


1 


1 


1 


1 


a' 


1 


1 


^ 

H 

^ 


-Si 

1 



j; ft 
£ 2 

I 1 



44 

n 

•3 

3 
O 
O 

Z 



n 

■3 

'q. 

I. 
n 

0. 



1999 



AZORES DEEP-SEA FISHES 



33 



S3f 



Q 
U 
D 
Z 
H 
Z 

o 
U 



ri 



JO 

r- 
C 
3 
C 

T3 



0) 

£ 

3 
C 

00 

o 

N 

U 



X 


P 


UJ 
1^ 


2? 




00 




X 






m 


*^ 




;si ;si 5 





^ oc r^ 


^ 


m 


LU X "^ 


m 


I 


U LU ^- 


X 


oi 


_^^S 


ai 




(*i" cT o ^ ^' 

f^ ^ s; S3 5 

tuluXXi tUujX — 
f 1 ''"■"< 2 U U a: a: 



UJ LU Ul I 

O U U I 



f*l X> t^ (?N 0\ r*l 

Q Q Q 0^ ^ ■^ 

— -B 5 ^ o o "^ 

I ^n r^ r^ (^ to to r^ 



T 000 



^ to 

<j, — r-* r^ 

<o 03 CS 03 

S X I X 

r^ o^ a: Qd 



< 

-J 



< < 






a 
z 

UJ 
Q. 

< 



on 



o 

a. 
c/2 




4j 

-5 
>^ 
o 

3 
<^ 

s 

O 

05 



I 

5 



it 



S 



«< « s 



3 
o 
S 
O 



.a -a 

= 1 

.a 2 

a. u 



34 



BREV/ORA 



No. 507 



Q 
u 
D 
Z 

P 

z 

o 
U 



X 

5 
z 

u 
a. 

0. 



C 

•T3 



s 

3 
C 

(30 

o 

N 

U 



C/3 

u 



■Or-»o — «*i— O 0Of*l 

gpoopOOO*/". v-i »Aw^ 

-J— .—JCDCDmffl CQCO 

'^i T r^i r^' -^" -«' — ' ■-' "" 
"^ DC OO 



s. O 


f*, 


X 


oo 


^ 


On 




r^ m 






fN 


r^ 


«~) 


r^ 




rs CH 






r-4 


r^ 


r-* 


r^ 




r^ r-j 


■" "" 


~* 


~~ 












i^ 


o' 


o" 


? ¥ 


?" 






00 


oc 


O 


o 


W-, 


•Ti 




0\ •'^ 










r^ 


r^ 




— fM 


J _J 


-J 


CD 


CQ 


CD 


CQ 


O 


CD CQ 


u u 


LU 


X 


X 


I 


X 


^ 


X X 


J U 


u 


o; 


cd 


ai 


Of 


CD 

X 


oi aC 




■4 

o 


w 


00 




^ 


ii 






f-J 


rM 


r^ 


r-i 






^ >^. 


v-i 


r^ 


r^ 


r^ 


c^ 




c^ rM 






























"~ 


< 00 


r-' 


-> 


m" 


»X" 


r-" 




•* r^ 


m 


r^ 


■v 


m 


■^ 




00 


00 


o 


o 


w-i 




v^ 




r- 






r^ 


r^ 


w^ 


J _; 


-J 


CD 


m 


CO 


CD 


r^ 


CD CD 

i5 


J UJ 


UJ 


X 


X 


X 


X 


CC 


J U 


U 


o: 


al 


of 


cd 


5 


i r^ 


^' 


o" 


rs 


'O" 


o" 




 T 


T 




o 






V 


s o 


O 


00 


r- 




CTs 








r>t 


CH 


rst 


r^ 


Ov 




*/-. 


fN 


fv) 


r^ 


r^ 


rs 














r^ 



— o 



oo' r~-" rn 'O r^ 00 o' „' __' ■^C _ 

gOOOOOOOON-^^^ 00 "^'^ 



_J 03 CD CD CD ^ 
UJLUWXXXX-?^ 



UJ X X 

O cd i 



00 — (^ ^ — 
rn T o O o 
o o f^ -- - 



i 2j '*^ "^1 X) o\ 



00 ^ «^ — r*i f*i 



0\ CM — r- 
r-« — ■v r-^ 
OO o O *^ 



— r^ 



-^ o 


o 


m 




CD 


CD 


us 


I 


X 




X 


X 


a: 


ai. 




ex: 


or 


_ -1 














o 


i 




CT\" 


1 












. Cn* 
















■A 






f 


ss 


O 


»<-) 




o 




_1 00 


CD 


m 


1/^ 


CQ 


ffl 


"d 


X 


^ 


ca 

X 


X 
of 


X 

a: 


_-u 


o 


1 


i 


s 


c^ 



C; (^" jtl J:* a-" 



oo S o ^ ^ 



UJ _J 
CJ UJ 



'. s 



CQ CO rvi 

X I CQ 

o; o; I 

r^ o ■© 



»^ oo r^ 



m 

X 

a: 



o s? S 



''^•Q 



>ri — 



r-i -o o — 



Vi oo ly-t 



UJ UJ X X X U] 

^ (J oi a; S U 



00 »/-. 

-J CD 
UJ X 



Ov <*> 00 r^ »/% r- 

^ ^/^ r- 00 — w~i 

O O -^ -T O O 

— . — . O O O — C3 

"■^ *^ f^ r-4 c^ v-1 <vt 



oo — 



< < 



1- oo 



V) 

o 

ex 
oo 



I 

■i 



I 



be 



I 

I 



cS 



a 

1 



a 



I 

I- 
Si 



o 
S 

a 



Q Q 






1999 



AZORES DEEP-SEA FISHES 



35 



Q 
u 

D 



E 

3 
C 

T3 



E 

3 
C 

GO 

o 

N 
U 




I 3 •> m 

t/*i <A w^ 



r r- — r* r^ r>* 

-J -] CD CQ CQ CQ 

U) UJ X X X X 

o u a: a: o: a; 



— t^ S ! 






O -T r- r-* I 



_3_i_j-j-CDCDCaCD 
LUUJUJujXXXX 



— o 



i§ -O ^ ^ ON 



a S S K 5s 5 

LU UJ -^ -r X ^ 
O U o^, i d c*; 



r- — f^ 
o — r- 



w-T 



— — — — 00000000,- 

* »r.»Ai,^, — — — *— S 






r- — r-| -^- — - ' 
w, v-i — _ — "" 



-J -J 
UJ UJ 



r- t/i — '5 ~" 92  - 

rn 00 ^ "^ ''' ^ "T 

t~~ <x> — ^ ^ ^ ^ 

_J -T- CD CD 03 CC r4 

UJ i X X X X CQ 

U -^ a; ^ Qi Qi X 



?■; S o ■o" 5 C^ 

(i^ r^ CQ On r^ CM 

_! _1 I S ™ ™ 

UJ UJ a: 5 X X 

u u - 5 a: oi: 



SS 



0^ •«■ 



r~ tt r- — 



rj o Q — «"» 

■O <5\ CT> '^ -" 
" 00 00 00 00 



o o r- „- 
2 Z !1 ^ 



— — — — 00 !/■, >^, 



0« 00 <o 

oo o 

J -J m I 

U4 bJ X 

U U a: 



2: o ^ 

| = s 



•O f^ Ov — — 



-J -J I 

UJ UJ lo 

tj u 2 



— v. 1^ ^ 



«/^ r^ ^ 
r^ m -ty 



UJ UJ UJ 

U O U 



On O 



X ^ 

a: m 
. I 

§-■ 
. o 

?S 
X m 
a: I 

o S 



I" 
a: ad 



w^ *^ >/^ 



■O O 



§'^ 



OO — -^ — 2; 
— (*i m TT '^ 

00 00 00 00 ^ 

r-r-r-r; — — r-;r-» 

_J_J_J_3CDCDC0CC 
LULUUJuJXXXX 



■^ ^ ;^ « ;=• ^' g « 

_jj«t|a3cQcDaa _)_j_jcQq3 
X luiu^XXXx ujluujXx 





o' 


— m 


■O* — ' 


no 


o' •^' 


*/-r '3^" 


_- 


^ oC 


fW 


r-* 


T 


oo' r^' 


c^■ —  




i^ 


r- r- 










rt^ 


r^ w-i 




'■^ 


<> 




S-i! 






■O 00 












































1/-I 


«/-l 


»ri ^/^ 








«/-i >^ 






HZ 


— 









H 
Z 

o 
U 



ri 

X 

5 
z 

a. 



on 
U 






i 



S 



00 



3 

c 

t 



I 






.1 






36 



BREVIORA 



No. 507 



Q 
U 
D 
Z 

H 
Z 

o 
u 



X 

5 
z 

UJ 

a. 
< 



X) 

£ 

3 
C 



0) 
X) 

S 

3 
C 

o 

N 

U 







< 



I 






C/3 



o 

o 

o 

I 






i 
I 

I 
I 



I 

a 

I 



1999 



AZORES DEEP-SEA FISHES 



37 



D 

z 

p 
z 

o 
U 



rj 



0) 

£ 

3 
C 



E 

3 
C 

00 

_o 

a 
o 

N 
U 



-1 m 



P5 S 

UJ I 

U a; 



f-t -.J 

00 w. 

^ m 

UJ I 

o" <X" 



X 



O »^ r^ '^ Q — 

<N **^ ^ r^ n •'^ 

Poo O Ov -A W, 

r- ir — *^ f^ 

-1 -^ 3 CO ffi m 

UJ UJ i X I I 

O tj S oi at en: 

oo' o" Q ri Ov' r-T r^' ' 
00 ^ 1^5-0 •:>  
^- O r' ^ — 



an 00 ^ — 00 

rn n r^ 7 -T 

00 od 00 Ov «^ •^ 

r^ f^ r- — r^ r4 

-1 -1 -1 m m m 

UJ UJ UJ X z X 

U U U oi a: a! 



_ - g n n n 



rn oc — — 

m (-5 rn ^ 

■^ f^ r*^ r^ 

w^ •/^ •^ — — — 



v^ r^ — r^ r- 3 

r^ ri . . <^ '^ -Y 

X 00 C3 ^ "^ ''^ ' 

r- r- ^ — r^ r^ 

_J ^ oo CQ 00 03 

UJ UJ R X X X 

u cj [jj o: a; b: 



•^ 'O PS; 

O rn — — Z- 



_ 00 r^ 

^/^ r^ ^ 

— — — m rn r*! 
(T) «i ^ — — — 

— — ^^ — — — 

m* — " "^. — * '«' Ci* 

r^ r^ r- C:? '? ^ 

00 00 r^ o^ -^ w^ 
r- F- So — r^ r^ 
J _] " CQ 03 m 

UJ UJ -J- X X X 

u u UJ oe: oi a: 



o — . — ^ — 

— I— r^ r- (-< o 

r-^ — . — • rO -O ^ 

— — r^ rn 1*^ (*i 

ir^ w-i — — — — 



5 S? 



1^ 00 — (^ 0\  



ff 99 *r^ ff £ 



— r- p- fs« 



CO 



*ri «r, — 



S5- 

-J -3 li X CO 03 21 
UJ UJ [n tJJ X X X 

r-* r-* — " oC (*r o* tt" 

00 On — Q r^ g Q 

— — f*^ ?S m ^ ^ 

— — — — (*i r^ f*) 

«~, 1/^ <^ </^ — — — 



00 00 ^ ' 

r- F- r- 00 '^ f^ "^ 

_j _j _] -j: 03 oi «-_ 

UJ UJ UJ tjj X X S 

U U U ^ a; C3ii ^ 

■o" V o' V 00* r~' °^ 

— o r^ ~. — " • 

2 Z — ^ S S f^ 

^ C^ «o v^ — — P; 



CH r- v, _ r^ 3 • 

r^ rn r^ ^ fTi ? r>< 

?? ?? PS p? S ?i ?: p^ 

ujujujujxxai r- 

— r-* ov* n" r-* ^' „■ ^ 

if^r^';. o 

«r, ^^v% — — ^ ^ 

oo' <o ■^" q" S CJ 0-' _ 

oooooooer;:^^ 2 

_3_3-J_jCD£acO 2j 

UJUJUlUJXXx UJ 

UCJUOotfoiS Q 

*^ rn O^ r4 --o *^ O"' 3o' 

— ^^r^^ — — O 
m — f*^rnr*mi-', r-* 

— — — — rn<*ir-, — 



*^ S *X -J 

P^ F^ r-« r^ 






-1 _J 03 03 




UJ UJ X X 




U U a: oi 




r-' g' '^' o* 

O f»i •^. (- 




«-■« fl '*. <-, 




_. — rj ^ 








— — — — 




r-- r-- d 2 

r^ r^ e> i 

00 00 *^ '^ 






r~ r- -T r^ 




_j _: o CO 




UJ UJ r*. X 




U U ° ai. 




r-' OS* Q .^' 




r^ m rr (*i 




oo r-t 

^ tr, r'j — . 














m* "X f; 




r^ r^ • '^^ 




* ?5 S^ 






_] _j g? m 




UJ UJ _3 X 




U U UJ oi 




- -'-',' 




r^ C r-< JS 




(*i r^ i-n m 




_ __ f*-, CM 




«/-i tr, — — 












2 S o 2 




_i _j [^ I 


23 


UJ UJ 72 UJ 


X 


o u uu: 


OL 


w-r oo' — ' -T* 


r~" 


r^ fs* f. r^ 


»r, 


fn m m f*-. 


r". 




r^ 


^/^ V-, «-, vr, 





< < 



— oo 



< < 



Q 
Z 

u 

0. 

< 



C/2 



c/i 

(1> 

C/3 



2 

I ^ 

I- I 



1 



I. 






"5 

a 
3 a 



s. 

S 2: 
-J -J 






38 



BREVIORA 



No. 507 



H 

3 
C 

13 



E 

3 
C 

B 

c3 
O 

N 
U 



Q 
Ui 
D 
Z 

H 

Z 
O 

U 



X 

5 
z 

a. 
< 



C/3 

u 



sss ?; f? ps ^ ri !2 5 i g 2 2 :c 

-ICDCO -J-l-l-ll-J 

UJXX QJUIUJLU . — — 

^Uac U U U O 5 ^. „■ ^- ^- ,- ^- V 

- . . . - . . ^'^i — ; — r-* m n ■? W-, 

goooQ o«r-i<*ir^ — oO■o^»'^v*lV^^/^ 

^OOOO — f^rstm— Q-jCDoacQfflCQCQ 

^^qH ai at o^ U oL 

* 2 E? i*r 6\* ■^" o* H .^" .fi ^* 00 "* ■'' t^" 

r-r»<r>i (^r--r^r-o'^ — — — — . — _ 

-JCQCO _;-J-J_J^Z 

UJXX LUUlUJtuy — 

CJ od Qi O U U O p _ ;^- V ;5 r.- go r; 

^^OOO r^r^r^m o — mcDtDCDCDCD 

... ..... 0^ 

f^GCJ iSrSPr^SooS SS 

..->acD _:_j_i_)i§ — — 

UJXX UJUJUJUJUJ— — — _ — 

O oi a: CJ U U U :^ - ^: ^- ^- _■ r-.- .:.- 

O — r^ ONf'iO^O'*i — — rsr-irsfs 

w-i — — ^/^^/-,,/^,/-l,/-,rnXXXXXI 

^ ai qL Qi :^ OH u 

r^ fi 0\ r~. oc* v© r^" r-* '^' * *'"' 00 ^' oC rn v©" 

r^^"TX — rMi-nr^oo m-tfr-r-oooo 

oeo»^?c ocoooooo^ — 5" '*"' ^ ^ ^ ^ 

r-— r^-o p-r-r~r-0Cr~.O — O — — — 

-jccmca _;_j_:_3Xr^ 

UJXX-r UJUJUJtjJUJ — ""*"""""■" 

---. ..... •^ — r^-n-^^Q 

;;;rCiC:; ;7;:=;:?::;^7:?xxxxxxx 



3 



CA 

C/3 



I 



! 

I 



S 






ft 



Ik 

ft 

a 

§ 

I 

I- 



1999 



AZORES DEEP-SEA FISHES 



39 



Q 

D 
Z 

H 
Z 

o 

u 



s 

C 

12 



E 

3 
C 

GO 

_o 

o 

N 
U 

2 



r^r»ir*l"Trn-Tr<r-t*T'T«'^ rnr-i r^^n — n — ^ 

t--r~r-r^ — — — — r^r^r^o — — -o ■"r~^»*~*^'^-» 

-j-j-j-jccmaacQcacQcQ* cDcaJ -J-Jo^?^Z4 

uiUJUJljJxXXXXXX^ xxlC ^^iXXDD'^ 

.......... .X .X ,.-J-..£ 

■o^m^'~~r^ — -Or^oOi-w' ^ '^ r*' l^oQ^r-r-*-J 

^ — — ^ "^ r~- 1 — F- r^ r^ r^ ^ r-- r^ ^^ — — "^ '* ' ^i i^i v^ 

. , _ . . . ,£, - . . - - - O . - O . . -"T o - . o 

OOOOOOOO — 0^0^*'^i*/~'W~l^/^ QO'^f~~ Q00O»»— OiJv*-. 

UJUJLULU;^XXXXXXJ^UJXS LUUJ_J;:^55rH 

. . . , oo" 5 - - i . . u ^- . . z) 

_ — — — ^-^~-ooooa^'>o^J^ —rjog __Tr*'^'0«/-. w-i 

_ — — — or~-r-t--i~-r-~r-fr' r~-ao — __o •'••'". 'H' 

»ri»riv^<r^ — OOOOOO^ */-iOp w-ii/-» — — OOt'n 

~1 "1 '^* ""- " " ~ "". ~ 2 12 — '^' "". ""- S 

r-T 00* r^' oo* ^ C:* "^ f^ O Q O . — ' — . oT i-t" - - 2 "^* *^' . 

oooeoooo-^ooNO--. --,*n,/-, oo*^^ t^ oooo^,— oovis 

r-r~r-i— o — — — f^r^rniy-i |C:— -v-i C2 r-r-oo™ — —^ 

UJLULULU'oXXXXXXCD ujXCQ ^ UJUJHc^55z 

cjuuu-ia:a:a:a:3;a:x uoix i 00^.55005 

lAoooor- -o — -^OToo • — rn - , "^ON-r-n-ooo 

— — — — ONf^aooc'^^'^*^ — r^too o — — ^o'O'^^ 
— •■^ — .^CT^r-r-l~-^-^-^-CT^ —r-*^ y, — — ~■_«^^w^"T 

_ — __0 — — — — — — o _ — O TT — — „f*i — — O 

LLlLUUJUJ^iXXXXXXx ^^X lI^ t^tiJuj^55t 

■^' r— " — 'O f*^ Ov" rK' r«-r Ov* f*^ l~-' «o' o" t~-* -O On V" 00* — * O* o' 00* <*! 

— — r-*r^ — — r^o — '^f^oo mr^oo r- r^ r^ tt --q r^ r^ 0\ 
_ — — — — r^r-oooONO^'O — — .r^ — — — — >Zv-,'0'^ 

— — — — — r-r-r-r-r-r-~r~- — r-r— ^ .« — — ./Sw-itni/% 



< < 



f 






Q 
Z 

u 

< 



00 



o 

O. 
C/3 






u 
-5 



— 




5 


L. 




a 


^ 


a 


rr 








'^> 


C;. 


u 


5 

60 


^ 

%; 


1 








C 


s: 


c 


^ 




s 


"C 


Q 


-^ 




5; 


lA. 


53 


^ 





•« 


ii 


1 









40 



BREVIORA 



No. 507 



Q 
U 
D 
Z 

H 
Z 
O 

u 



c 

3 
C 

12 



0) 

E 

3 
C 

ao 

i3 

o 

N 
U 



ftj 



•O "^ 'O w^ r>* r^ 

r^ (n T*! — ^ w^ 

00 00 O o «r^ >/% 

r— r— « .^ r^ r^ 

-1 _] 03 03 m 03 

UJ UJ X X X X 

*^ O QC a: a; oi 

— J^J oo" ^* o *t" 



;^ ;2! o o o o 

•^ ^ — * "^ — * On" 

rN r*i ^ ^ ^ T 

00 00 00 o »n «/% 

r- r- r- — r-t r^ 

-J _j J ca 03 m 

tu uj uj X X X 

u u o oc: a; a: 



::i?' 



. «^ ^/^ o o o 



rn r^ OQ — O OO 

r-t m Ps "^ T 'T 

00 00 00 "^ '^ ""' 

r- r- r~- — f^^ <^ 

_J _J _j 03 03 CQ 

tu LU uj X X X 

o cj cj ck: oi q; 

oo — " «/-r o* oo" r-i* 

•^ w^ w^ r-i \o r- 

= = z? S R 

•/■. V-, U-, O O O 



1/-, O Q 

00 — 1^ 

X m 03 

U) I X 

U ct c^ 






J CO 03 
Uj X X 

CJ a: Qd 



>o <^ <^ 
O _ o O 

■i?l „ T -"T 



00 <> ■> ^ 

_j CD 03 r^ 

UJ X X CO 

tj Qi Ot: X 

^ — V-. . 

ir\ r^ rn f^ 

— O O 5?- 

— ■»r -T i^v 

W-, O O f^ 



00 r^ r- 
— (*t r^ 

00 oo 00 



_1 _J _J to CO CD 2 
ai Lu uj I X X T 
o u u a; Qi q; i 



'^ 2, 

"" 5 ;s "^ 

_) X en £0 

UJ u X X 

u ui q; i 



r^ 


r~ 


■o 


r^ 
















W-. 


O 








O 


m 


03 


r^ 


I 


X 


X 


2 


U 


cd 


a: 


Q 




fS 


m 


P 


S 


m 


^ 






o 


o 


r1 


^ 


















CM 


'-^ 






r^ 






o 


O 










r^ 


-1 
U 


03 


03 


03 


X 


X 


T 


o: 


Qd 


a; 


U 
















•o 


ss 








O 


o 



LU ^ 



CQ CD QQ r~4 

X X X CQ 

ot; o<; X 

... a: 

Soo r^ „- 

r- o op 

r- oo oo «A 

m rn rn 5? 

o o o r^ 



03 

X 



S 5 '^ o^ 

» 2 _ ?Q '^ o 

-1 E£ S 2 CQ m 

tjLj X X X X X 

o cd a: a; S 2 



?? 


o. 


Ov" 




-1 


03 

X 


2 


5 


u 


tt: VI 


r^ 


^ 




•jT 


r/1 












m 




■*r 














"■ 










o 










-1 

fll 


o 

OQ 

X 


^ 




u 


Of 


wj 


A 



- ^ S S! 
_ m C3 ^ 



r? n r 



p; 


o 




^ 


-1 

t) 


I 


w 


r* 










R 


g 


s 












«/■, 


o 




f-1 



_ — — o 

s s I s 

_1 _J m -, 

ui u X c; 

o u a: 5 



o "^ 0\ r- — l/^ 00 
»^ w-i — >p r- r- i/"! 



■« 



■« r4 00 
— oo F- 



^S r 



i/^w^wiOCDOO i/^"^. OO 



— OO f- Ov 

^ f^ f*l (^ L . 

•^ o o o o 



■O 0\ t/^ f*i 

•o »o — o 

— — ^ f*! 






< < 



ri 
X 

5 
z 

LU 
Q. 
Q. 

< 









55 



<3 

J 

O 





<o 


«) 


■? 


1 

60 


1 


5 


J 


1 


•« 


«> 


c 


a 


VJ 


<o 


3 


a 


a 








«) 


<u 


^ 


^ 


9w 


la. 












(J 


<> 


§ 


?? 


S 








^ 


:| 


:| 



S s 



I 

O 
II 

a, CO* 



5 

s 

§• 



1999 



AZORES DEEP-SEA FISHES 



41 



Q 
tu 

D 
Z 

z 

o 
U 



C 



0) 

£ 

3 
C 

o 

"B 

a 
o 

N 
U 



(^ 



a 

X 



OQ 

z 













00 


X 
















X 


u 
















a 


:>^ 


















oo' 
















r-" 


















00 


s 
















r>4 


'H' 
















-o 




wS 








r^ 


§ 




i 






O 


5; 




p 




-3 

U 


-J 

u 
U 


u 
U 


Q 


QQ 






3 




^' 




c^' 








<*f 




g;' 


1 


7. 


K 


??' 


s 


i 


K 


^ 




9 


m 


^ 


(N 


IS 





CQ 


ue 




^ 


a: 


c^ 




X 
a! 


1 




1 


i 

v^ 


i 




r- 










w-C 




•/^ 










r~ 




•V 


Cv 


<?c 


6\ 


On 


r-i 














OO 




s 


^ 


S 


5 


S 


V 


CO 

X 


o 

CD 

X 


5 

5 


< 

5 


< 

Q 


< 


CQ 


o 


o 


O 


o 


X 






u 


u 


U 


tj 


C£ 


o" 


ri 













— ^ — «^ 



00 — — — 



w-1 « W-) — ~ ~ 






< < 



<<_)<<<<<<< 



3 



? 



t^ O « ^ o -, •"" 

oooo'To-'~'~?T'72p5<^ 
rJ — — <s 2. '^ 



5 
z 

tu 

a. 
< 






_ _ r) >^ (s 



^- . — f*i «n — 






(^ — r4 fN 



o 

C/5 




I 

a ^ 

3 5 3 

ill- 
-^ u u 




i 

«> 





g. 











s. 


-5 


'5 


? 


"5 


a 


a 


1 


^ 


1 


^ 


1 


a 


<U 


<3 


c 


-C 


es« 


.^ 


^ 


-2 


5 






"S 


^ 


p 


(a. 


s. 


^ 


5^ 


o 





u 


^j 


a. 


|>2 


V2 



2 

3 

o 



§> 



^ Q 



•r: 

a 
•Si 

5 

I 

s 




it 



3 
'u 

■i-l 



42 



BREVIORA 



No. 507 



D 
u 

Z 

H 
Z 

o 
u 



X 

5 
z 

PU 

a. 

a. 

< 



E 

3 
C 



E 

3 
C 

a 
o 

N 

U 






CO 

o 

CO 



-1 



m 

X 



X 


2 


ai 


5 




00 


r^' 




0^ 


oo' 


r- 




l/~» 
r- 


3? 




00 


-T 




s 


i 



« 5 



i- F^ 



§ 

Fi 

- »■< 

- r- O 

5 2S 2f 



UJ CQ 
U X 






r-1 O- «^ 

OS — — 

— vi r- — 






CQ 
X 



o * 



5?! P 

5 ia -" 
°^ B o 

sis 



< J < < < 



< J 



T - 
2 -o 






— — — Tt 






W^ — ,- 



■» — _ 









3 



a 
a 



5, 
a 






















;2 








'G 






ev 


CQ 


n 




V 


C 
O 


2 




4> 

"3 


e 


13 

•a 

c 




■3 


■? 




44 


£ 


o 

a. 




■3 
•a 
s 


:a 

.a 


E 




"2 


a. 


1. 


u 


n 


u 


g 


c 


c 


O 


u 
< 


c3 




c3 


c!5 


o 
Z 



4^ 

"2 M 

= I 

o o 



B RE V I O R A 

jyi 11 s e ui nn ol L^ompairafi ve Zoology 



us ISSN 0006-9698 



Cambridge, Mass. 



20 April 2000 



Number 508 



NEW RECORDS AND DISTRIBUTIONAL AND 

ECOLOGICAL NOTES OF LEPTODACTYLID FROGS, 

LEPTODACTYLUS AND ELEUTHERODACTYLUS, FROM 

THE BRITISH VIRGIN ISLANDS 

Kristiina E. Ovaska', Jeannine Caldbeck-, and 
James Lazell, Jr.^ 

Abstract. Information on distribution and habitat use of frogs in the British 
Virgin Islands is needed for assessing population trends and status and for elu- 
cidating biogeographic patterns. We discovered 10 new populations of the four 
known species of Leptodactylidae on five of the 17 islands visited: Eleuthero- 
dactyhis antillensis on Great Camanoe, Great Thatch, Jost Van Dyke, and Beef 
Island; E. schwartzi on Beef Island, Frenchmans Cay, and Jost Van Dyke; E. 
cochranae on Great Thatch and Jost Van Dyke; and Leptodactylus albilabhs on 
Beef Island. We confirmed all but three previous island records: E. cochranae and 
L. albilabris on Virgin Gorda and an unidentified Eleiitherodactylus, known only 
from the stomach of a snake, on Peter Island. The earlier E. cochranae record is 
probably in error, but L. albilabris and Eleutherodactylus seem to have disap- 
peared from Virgin Gorda and Peter Island, respectively. The mean body size of 
adult males of E. antillensis and E. schwartzi was smaller on Virgin Gorda than 
on Tortola, and males of E. schwartzi were relatively large on the tiny (33 ha) 
island of Great Dog. On all islands except Tortola, E. schwartzi was almost ex- 
clusively associated with bromeliads. Island elevation and area explained 61% of 
the variation in the number of species when all 17 islands were included in the 



' Department of Forest Sciences, University of British Columbia, Vancouver, Brit- 
ish Columbia, Canada V6T 1Z4, and Biolinx Environmental Research Ltd., 1759 
Colburne Place, Sidney, British Columbia, Canada V8L 5A2; e-mail: kovaska@ 
jdmicro.com. 

- Thetis Island, British Columbia, Canada VOR 2Y0. 

^ Department of Herpetology, Museum of Comparative Zoology, Cambridge, Mas- 
sachusetts 02138, and The Conservation Agency, 6 Swinburne Street, Jamestown, 
Rhode Island 02835, U.S.A. MdZ 

LIBRARY 



FEB 2 2 2002 



LJ AD\/aRD 



2 BREV/ORA No. 508 

model. Elevation was the most important factor (partial r^ = 0.35), whereas area 
explained little of the observed variation (partial r- = 0.02). The availability of 
specific habitat features, such as aquatic breeding sites for L. albilabhs and retreat 
and nesting sites for Eleutherodactylus, are critical for populations on small is- 
lands. The distribution patterns in the British Virgin Islands do not indicate wide- 
spread extirpations or declines of frogs comparable to those observed in Puerto 
Rico and other parts of the world. 

. . . there is an urgent need to document the distribution and abundance 
of amphibians. 

Leonard (J 997) 



INTRODUCTION 

Precipitous declines in a number of anuran populations within 
the past few decades have led to local extirpations and even spe- 
cies extinctions (Mittermeier et ai, 1992; Pechmann and Wilbur, 
1994; Phillips, 1994; Blaustein and Wake, 1995). In Puerto Rico 
alone, three species of frogs (genus Eleutherodactylus, family 
Leptodactylidae) have disappeared within the past 20 years, and 
an additional seven show serious declines (Rivero, 1991; Joglar 
and Burrowes, 1996). Efforts to document and understand chang- 
es in anuran population and distribution characteristics are se- 
verely constrained by the paucity of baseline data, making it dif- 
ficult to distinguish between natural population fluctuations and 
those caused by human activities. Furthermore, in many cases, 
we simply do not know where populations occurred or still occur. 
Knowledge of habitat requirements and factors that limit the 
growth of populations is also incomplete for most species of frogs 
in neotropical areas. 

The British Virgin Islands (BVI), located on the easternmost 
portion of the Puerto Rico Bank in the Caribbean Sea, consist of 
about 50 islands, some of which are mere rocks or sand bars. 
During the last glacial maximum, the entire bank was united as 
a single land mass, which subsequently fragmented into numerous 
islands with the rising of sea levels (Heatwole et ai, 1981). Most 
of the islands have been isolated from each other and the rest of 
the bank for approximately 4,000-10,000 years (reviewed by La- 
zell, 1983). 



1999 BRITISH VIRGIN ISLANDS' LEPTODACTYLID FROGS 3 

Four species of leptodactylid frogs occur in the BVI: Lepto- 
dactylus albilahhs, Eleutherodactylus antillensis, E. schwartzi, 
and E. cochranae (MacLean, 1982). All but E. schwartzi, which 
is endemic to the BVI, are widespread on the islands of the Puerto 
Rico Bank and also occur in Puerto Rico itself (Rivero, 1978; 
MacLean, 1982; Schwartz and Henderson, 1991). 

Leptodactylus albilabris has a biphasic life cycle with aquatic 
larvae, whereas Eleutherodactylus species are completely terres- 
trial and have direct development. The distribution and habitat 
use patterns of all four species on the islands are poorly known, 
although other components of the herpetofauna of the BVI have 
received intensive attention over the past two decades (Mayer and 
Lazell, 1988; Lazell, 1983, 1991, 1995; Dmi'el et al, 1996). La- 
zell (1983) was aware of seven populations of leptodactylid frogs 
on four islands of the BVI. Mayer and Lazell (1988) added two 
new island records, including one for an islet of only 24 ha 
(Frenchmans Cay). Lazell (1991) reported a previously over- 
looked record for the 33-ha Great Dog Island (Heatwole et al., 
1981), bringing the total number of known populations to 1 1 on 
seven different islands. Ten additional islands that are larger than 
Frenchmans Cay had not been surveyed for frogs before our 
study. In many cases, the survey coverage of those islands known 
to support frogs was incomplete. 

Every October from 1993 to 1997, we investigated the distri- 
bution and ecology of leptodactylid frogs in the BVI. Based on 
surveys of 17 islands, we report on the distribution oi Leptodac- 
tylus and Eleutherodactylus species, including new island records 
for 10 populations. Our objectives were to (a) compile baseline 
data on the distribution, habitat use, and natural history of the 
frogs on different islands; (b) compare present distributions to 
historical records; and (c) examine the pattern of distribution in 
relation to predictions from island biogeography (MacArthur and 
Wilson, 1967; Lazell, 1983). 

METHODS 

Survey Methods 

Our operations were based on Guana Island, located ca. 0.5 
km north of the east end of Tortola, BVI. The survey periods 



4 BREV/ORA No. 508 

were 7-30 October 1993, 2-21 October 1994, 3-19 October 
1995, 8-28 October 1996, and 8-28 October 1997. During these 
periods, we also visited the following islands one or more times: 
Tortola (14-16 October 1993; 4-6, 13-15 October 1994; 6-8, 
14-15 October 1995; 11-12, 15-16, 19-20 October 1997), Beef 
Island (3-5 October 1995, 23 October 1996), Frenchmans Cay (7 
October 1995), Virgin Gorda (26-28 October 1993, 9-1 1 October 
1994, 17-18 October 1996), Jost Van Dyke (11-12 October 
1995), Great Dog (10-11, 16 October 1996; 21 October 1997), 
Great Camanoe (12 October 1996), Scrub (13 October 1996), 
Mosquito (16-17 October 1996), Anegada (20-21 October 1996), 
Cooper (22-23 October 1996), Peter (24-25 October 1996, 25- 
26 October 1997), Great Thatch (26-27 October 1996), and Great 
Tobago (17-18 October 1997). We also present data for Necker 
and Little Thatch, where residents have been listening for frogs 
for several years and one of us (JL) spent several rainy nights 
(three nights in October 1993 on Necker and one night in October 
1996 on Little Thatch). 

We used visual encounter surveys, auditory transect surveys, 
and night driving methods to locate frogs (Heyer et ai, 1994). 
We walked along trails in likely habitats after sunset listening for 
calls of males, and we scanned the ground and vegetation with 
headlamps for frogs. In 1996, we also played recorded advertise- 
ment calls of E. antillensis and E. schwartzi to induce frogs to 
call. In 1995 and 1996, the use of a car allowed us to cover longer 
distances on larger islands (Tortola, Beef Island, Anegada, Virgin 
Gorda); we stopped every few minutes to listen for frog calls. 
For each new island record, we collected at least one voucher 
specimen, which was deposited in the Museum of Comparative 
Zoology, Cambridge, Massachusetts (MCZ). 

In 1993, we systematically recorded information on each 
Eleutherodactylus heard or seen during those surveys carried out 
on foot and noted the following for each frog captured: species, 
sex, calling or not (for males), gravid or not (for females), snout- 
vent length (SVL), weight, microhabitat (ground, tree or bush, 
bromeliad, agave, herbaceous vegetation), and perch height. In 
1994 and 1995, we obtained comparable information only for 
frogs included in a separate study on vocal behavior. In 1996, we 



1999 BRITISH VIRGIN ISLANDS" LEPTODACTYLID FROGS 5 

measured the body size of E. schwartzi on Great Dog and Virgin 
Gorda to examine the hypothesis suggested by initial observations 
that the frogs on Great Dog were relatively large. We also mea- 
sured the body size of a sample of E. antillensis on Guana in 
1996. 

To obtain additional information on habitat use and dispersion 
of Eleutherodactylus, we set up auditory transects in October 
1994 on three islands (Guana, Tortola, and Virgin Gorda) and in 
October 1996 on Guana. In 1994, there were two transects on 
Guana, two on Sage Mountain, Tortola, and one on Gorda Peak, 
Virgin Gorda. On Guana, Transect 1 was in the north of the island 
along a ridge where E. antillensis appeared to be abundant, and 
Transect 2 was near the southwest tip of the island where an 
isolated patch of the species occurred. 

The transects followed the course of relatively straight sections 
of existing trails or paths, which marked the middle of the tran- 
sect. In 1994, each transect was 150 m long and 6 m wide. In 
1996 on Guana, Transect 1 was 815 m long and Transect 2 was 
300 m long. We increased their width from 6 m to 10 m, because 
previous observations indicated that we could accurately record 
all calling frogs within 5 m from the center of the transect. We 
placed a flag every 5 m in the center of the transect to divide it 
into sections of 3 X 5 m (in 1994) or 5 X 5 m (in 1996) on each 
side of the transect. 

In 1994, we recorded the presence/absence of arboreal and ter- 
restrial bromeliads with a crown diameter >10 cm in every 5-m 
X 3-m section of the transect. In 1996 on Guana, we measured 
habitat variables only for Transect 1 . The variables measured for 
each 5-m X 5-m section were: (a) sum of crown diameters of 
bromeliads (none, not present; low, <30 cm; moderate, 30-100 
cm; high, >100 cm), (b) percentage of ground covered by leaf 
litter, (c) depth of leaf litter/humus (measured for 152 or 47% of 
the 5-m X 5-m sections), and percent vegetation cover at heights 
of (d) <1 m, (e) 1-2 m, and (f) >2 m. The depth of the leaf 
litter and humus in each section was the average of three ran- 
domly located measurements obtained by poking a pencil in the 
leaf litter and measuring the depth of penetration. We estimated 



6 BREVIORA No. 508 

the percentage of ground covered by vegetation at different ver- 
tical layers and by leaf litter visually. 

To survey frogs, two observers walked along the midline of 
the transect after sunset and recorded the number and species of 
calling males in each section of the transect. We traced the exact 
location of frogs only when this was required to verify their pres- 
ence within the transect boundaries. In 1994, we surveyed the 
transects for frogs on Guana on four consecutive nights (17-18, 
18-19, 19-20, and 20-21 October). Transect 1 was surveyed 
twice each night on two nights and three times on one night to 
obtain information on the consistency of the number of calling 
frogs within nights. Transect 2 was surveyed once on 18 October. 
In 1996, we surveyed Transect 1 once each on 18 and 25 October 
and twice on 28 October. Transect 2 was surveyed once (on 18 
October). We surveyed both transects on Tortola twice on 13 
October 1994 and the transect on Virgin Gorda once on 10 Oc- 
tober 1994. We checked transects only on nights when rain had 
fallen during the 24-hour period prior to the search, and condi- 
tions were favorable for calling. 

Data Analysis 

We used a multiple regression analysis to examine the effects 
of island size and elevation on the number of species present. We 
also applied multiple regression to a reduced data set that ex- 
cluded both islands that contained the full complement of four 
species (Tortola and Jost Van Dyke) to include the distance from 
potential source populations in the analysis. The source for island 
size and elevation was Lazell (1983). The distance to the nearest 
potential source population was measured as the shortest distance 
between an island and either Tortola, Jost Van Dyke, or Virgin 
Gorda, whichever distance was shortest. 

We used one-way ANOVA to examine differences in body size 
of adult male E. antillensis and E. schwartzi among years. We 
also used ANOVA to compare SVL of E. antillensis and E. 
schwartzi among years and islands. 

We calculated the variance/mean ratio as an index of dispersion 
of calling males of E. antillensis in 5-m and 50-m sections of 
Transect 1 on Guana in 1996 and used the X" test to determine 



1999 BRITISH VIRGIN ISLANDS" LEPTODACTYLID FROGS 7 

whether the pattern was significantly different from random 
(Krebs, 1989). We performed a multiple regression analysis to 
examine the effects of habitat variables, measured in each 5-m X 
5-m section of the transect, on the number of calling frogs on 
Guana on 28 October 1996, when the number of frogs was the 
greatest. We also performed the same analysis using the number 
of 5-m X 5-m sections with frogs (1) and without frogs (0) as 
the dependent variable. The value for each section of the transect 
in the second case was determined based on whether calling frogs 
were found within a transect section during any of the four sur- 
veys in 1996. 

DISTRIBUTION 

Species Diversity 

The number of species per island varied from zero to four 
(Table 1 ). Only two islands, Tortola and Jost Van Dyke, contained 
the full complement of four species. One island had three species, 
three had two, and four had one. We found no frogs on the re- 
maining seven islands. 

Area and elevation explained 60.7% of the variance in the 
number of species among islands (multiple regression: F. ^ = 
10.8, P = 0.002). Elevation explained most of this variance (sim- 
ple r = 0.76, partial r- = 0.35), whereas island area contributed 
very little to the model (simple r = 0.51, partial r = 0.02). When 
the two islands with the full complement of species were deleted 
from the analysis and the distance to nearest potential source 
population was added as an independent variable, the model was 
marginally significant (r- = 0.51, F, ,, = 3.79, P = 0.04). In this 
model, island area (simple r = 0.22, partial r- = 0.25) and dis- 
tance to a potential source population (simple r = -0.007, partial 
r- = 0.21) explained most of the variance, whereas the contri- 
bution of elevation was small (simple r = 0.50, partial r- = 
0.001). 

Leptodactylus albilabris 

We found L. albilabris on four of the 17 islands: Beef, Tortola, 
Jost Van Dyke, and Anegada (Table 2). This species had not been 
previously documented from Beef (372 ha), separated from Tor- 



BREVIORA 



No. 508 



Table 1. Number of species of frogs in relation to island area, elevation, 

AND distance FROM A POTENTIAL SOURCE POPULATION. ISLAND AREA AND ELEVATION 
ARE FROM LaZELL (1983). DISTANCES FOR EACH ISLAND WERE MEASURED EITHER 
FROM TORTOLA, JOST VaN DYKE, OR VIRGIN GORDA (WHICHEVER DISTANCE WAS 

SHORTEST). 











Distance to 




No. of 


Area 


Elevation 


Potential Source 


Island 


Species 


(km-) 


(m) 


Population (km) 


Tortola 


4 


5,444 


521 


NA 


Anegada 


1 


3.872 


8.5 


32.5 (Tortola) 


Virgin Gorda 


2 


2,130 


414 


1 1 .7 (Tortola) 


Jost Van Dyke 


4 


840 


398 


NA 


Peter 





429 


177 


5.5 (Tortola) 


Beef 


3 


372 


244 


0.1 (Tortola) 


Great Camanoe 


1 


337 


187 


2.1 (Tortola) 


Guana 


1 


297 


266 


0.5 (Tortola) 


Cooper 





138 


155 


6.8 (Tortola) 


Great Thatch 


2 


123 


187 


0.7 (Tortola) 


Scrub 





97 


141 


3.7 (Tortola) 


Great Tobago 





87 


147 


4.0 (Jost Van Dyke) 


Mosquito 





50 


95 


17.7 (Virgin Gorda) 


Great Dog 


I 


33 


89 


11.1 (Virgin Gorda) 


Necker 





30 


32 


22.0 (Virgin Gorda) 


Frenchmans Cay 


2 


24 


131 


0.1 (Tortola) 


Little Thatch 





24 


100 


0.5 (Tortola) 



tola by a ca. 100-m wide channel. Our attention was first called 
to the presence of L. albilabris on this island by Dr. Gregory 
Mayer, who reported hearing calls and locating tadpoles, which 
were inspected by one of us (JL), in temporary pools among rocks 
in scrub vegetation several years ago. We did not locate this site 
but found L. albilabris in muddy ditches around the airport (MCZ 
124777-81, 125954). In 1995, we located several males calling 
from inside tufts of grass and from small cavities in the mud 
banks close to the water's edge, as well as many metamorphosed 
juveniles. We did not hear calls of L. albilabris east of the airport. 
On Tortola, we heard calls of L. albilabris from roadside ditch- 
es throughout the island and from small pools on Sage Mountain 
(MCZ 107339, 1 10992-5, 1 17677). On Jost Van Dyke, we heard 



1999 BRITISH VIRGIN ISLANDS" LEPTODACTYLID FROGS 9 

L. albilabhs in a riverbed by Old Hill west of White Bay and in 
a marshy site in the town of Great Harbour (MCZ 1 10990-1). 

On Anegada, we found several concentrations of L. albilabhs 
in an area called the Slob, ca. 1.5 km northwest of the airport 
(MCZ 125953). The frogs were in wet areas under dense shrubs 
on coral-limestone substrate covered by leaf litter and humus. 
Several males were calling from land crab holes, and we also saw 
many metamorphosed juveniles. 

We did not find L. albilabhs on any of the other islands, in- 
cluding Virgin Gorda, which we visited in three different years. 
In 1993 and 1994, our surveys were confined to Gorda Peak, but 
in 1996, we spent many hours driving around the island after 
sunset during and after rain. Extensive pools were present on 
Gorda Peak in 1993, but these were dry in 1994 and contained 
little water in 1996. Roadside ditches, where these frogs com- 
monly occurred on Tortola, contained water, but we detected no 
frogs. Small, temporary freshwater puddles were present on Great 
Camanoe. 

Eleutherodactylus antillensis 

We found E. antillensis on eight of the 17 islands visited: Vir- 
gin Gorda, Great Camanoe, Guana, Beef, Tortola, Frenchmans 
Cay, Great Thatch, and Jost van Dyke (Table 2). The species has 
not been previously reported from Great Camanoe, Great Thatch, 
or Beef (MCZ 132823). In addition, we have found no previous 
records of E. antillensis from Jost Van Dyke, although MacLean 
(1982) reported the distribution of the species to encompass "all 
major islands" of the Virgin Islands. On Jost Van Dyke, calling 
males of E. antillensis were patchily distributed in areas west of 
White Bay toward Old Hill and east to Great Harbour, including 
the town site (MCZ 124786). On Great Camanoe, we located 
frogs in the hills on the southwest portion of the island (MCZ 
125949). On Great Thatch, we found E. antillensis throughout 
the densely vegetated south slope of the island (MCZ 125950). 

Eleutherodactylus schwahzi 

We located E. schwartzi on six of the 17 islands visited: Virgin 
Gorda, Great Dog, Beef, Frenchmans Cay, Tortola, and Jost Van 



10 



BREVIORA 



No. 508 



CO 

Q 

Z 
< 

-J 

z 

5 

> 

X 

b 

a: 
03 

H 
Z 

izi 

O 

o 
a: 
[I. 



>■ 

H 

< 

O 

o 



UJ 

-J 

O 
u. 

Q 

o 

U 
UJ 

a 



< 



D 
Z 
< 

Q 
U 

E 
z 
o 
u 



Q 
a 

z 
o 
u 
\a 

a: 



UJ 
-I 
CQ 
< 







-o 










c 










rt 










N 










c 










M 






y. 




^ 






o 
o 




00 






(U 










a; 




'-~. 






W5 




d 






3 




a. 






^O 




yi 






> 




-* 






OJ 






y 








"3 


u 


00 


■-I- 




^-^ 




— 


o 






ir, 






n 


ri 


00 




OJ 


00 


00 


0^ 


"^ 


ci 


o 


c^ 




^ 


3 




"^ 


, 


^ 


O 




, 


3 




cn 


3 


3 


O 


nj 






C3 


ty. 

-a 






C3 


o 

a 


3 


ID 




s s 


X 


X 



o 
U 

o 
Z 



o 
o 

o 



X) 



o 

1) 
a 
on 



3 

o 
U 



o 

OJ 



-a 

3 



UJ 









00 
00 



N 

-a 

3 
ca 

(U 
>, 

C3 









W 



•o 

c 

c<3 



XI 

o 






=S u 





ir, 


ri 


00 


00 


r^ 


G^ 




^^ 






3 


3 


O 


<^ 


tfi 




•n 




3 

a: 



d 



PQ 



o 


1 


1 






[ 




CS 


1 










y. 




1 






Z 










ON 




PJ 












W 






ON 
ON 


ON 
ON 


'X. 










ON 






















' ' 


— 


u 


vO 


r^i 




NO 


"" 




















r^. 








ON 


ON 


-* 


ON 








r^ 


'^ 


>/", 


nO 


r^ 






ON 


rj 


O 


o 


ON 


ON 


On 


O 






On 


Ov 


On 


On 


On 


ITj 




ON 





— 




ON 




I^ 


NO 


ON 


ON 


Ov 


On 


ON 


On 


nO 


— 


>> 






— ^ 




ON 


ON 






1— ' 






ON, 


O^. 




u-i 


>r, 


o 


o 


^ 


O 


>/-, 


ON 


ON 






^ 








O^ 


o 


> 





O 

00 


o 




00 


— " 






o 


o 


o 


O 


'-) 


^ 






nC 


1 


1 


3 

on 




o 


^ 


o 


'«> 








o 


o 


o 


O 


o 


T\ 


-* 


CN 


(N 


^^ 




^- 





o 


o 


— « 


ON 


00 


00 





— H 


, 


, 




1 


vi 


^^ 


1 


1 






r<~, 


ri 


^^ 


— 


rj 


ir-, 




1 


nC 


00 




O 


1 


1^ 


o 


— 


ri 


1 


1 


1 


1 


1 


1 


r<-, 


'^ 


1 


1 




tN 


O) 


On 


— 




n 


— ' 


r- 


ri 


ON 


00 


00 


r<^ 


r) 




^ 


NO 



c 
n 



>n 


t~~ 


^— 


ON 


, 


ON 


n 


— 


1 


' 




o 


— 


O 



o 

1) Oij 

3 .t: 
< > 





u 






o 






c 






rt 




01) 


S 




o 


e« 




Q 


U 


nJ 










r3 


3 

03 


u 


Um 


3 


a 


o 


a 



T3 




3 
















y 


rt 


(^^ 


o 


U 


r 


IJ 


r, 


oa 


h- 



1999 



BRITISH VIRGIN ISLANDS' LEPTODACTYLID FROGS 



11 



Q 



H 
Z 

o 
U 



< 



o 
o 
<u 
Qi 
■j: 
3 

> 



<U 

u 

u 
3 
O 



D. 
C/5 



3 "^ S T3 

.2 sal 

> ii ^ 

(U ID ■" U. 

Q- Z 



o 
o 

3 



O 

Z 



■5 



tjj 



9^ — u 
o^ ^ w 



00 
00 



rt O _' 
C/5 ^ — 



o 



2 -a 



c 
o 

OJ 
T3 
C 

o 

3: 



a 



 S u 

OJ 



C3 

u 



in 

On 



s 

o 
x: 
H 

-a 
c 

N 



U 0^ 

00 — 



W I -J 



o 
U 

W PJ 

c/; rt rt 

u u u 



1/3 


vO 


vO 






i/-^ 


Q 


ON 






ON 


m 


nD 

ON 


ON 
ON 


> 

u 









ON 
ON 


On 






3 

CO 


ri 

1 


vO 
1 


s 


U 




1 




-t 


>r-, 




vO 


^^ 




rl 


Ol 


■n 


r) 










U 


j= 






■a 




y. 


u 










r3 


73 




-0 


■^ 




C 





c 

:5 


_2 


s 




C 

1» 




y; 


1— ( 


(X 




u 
U. 


a 






o 

OJ 

7; 



'c 
3 



ta 






a 

-4 
-J 



PJ 



d 



^ 


3 


s^ 


T3 


^ 


s 


^ 


^ 










'^ 


*^ 


5 






-^ 


« 


t« 


tu 


<+- 


= 






12 BREVIORA No. 508 

Dyke (Table 2). The presence of this species on Beef, Frenchmans 
Cay, and Jost Van Dyke was previously undocumented. On Beef, 
we found E. schwartzi along the road that transects the island and 
in a patch of terrestrial bromeliads {Bromelia pinguin) along a 
path that diverges from the main road near its northern end (MCZ 
124782). 

On Frenchmans Cay, we heard calls of E. schwartzi from gar- 
dens along the road east from the bridge to Tortola (MCZ 
124783). On Jost Van Dyke, we heard calls of scattered E. 
schwartzi from gardens, pastures, and gullies in and around Great 
Harbour (MCZ 124785). 

In 1996, we confirmed the presence of E. schwartzi on Great 
Dog Island (MCZ 125946-8), an islet of 33 ha, first reported by 
Heatwole et al. (1981). Numerous frogs were present in a ca. 13- 
m X 16-m patch of bromeliads, Hohenbergia antillana, located 
near the peak of the ridge that extends along the length of the 
island. In addition, on the night of 10-11 October 1996, we heard 
a single male calling near the beach in dense vegetation on the 
south side of the island ca. 500 m from this patch. We located 
five egg clutches ofE. schwartzi on 10 October within bromeliads 
(Ovaska et al, 1998). 

We observed numerous E. schwartzi on Sage Mountain, Tor- 
tola, and on Gorda Peak, Virgin Gorda, and we also heard calls 
and observed frogs in other areas of these two islands (MCZ 
107340-1, 115830-8, 117567-9, 117688-92, 119247-51, 
116273, 124784, and U.S. National Museum of Natural History 
329482-91). 

Eleutherodactyhis cochranae 

We located E. cochranae on three of the 17 islands visited: 
Tortola, Jost Van Dyke, and Great Thatch (Table 2). The species 
has not previously been reported from Jost Van Dyke or Great 
Thatch. Based on advertisement calls by males, E. cochranae was 
the most widely distributed and abundant frog species in the areas 
surveyed on Jost Van Dyke (MCZ 124787-8). These included 
areas west from White Bay toward Old Hill and east to Great 
Harbour. Calling males were perched on cacti, trees, and arboreal 
and terrestrial bromeliads. 



1999 BRITISH VIRGIN ISLANDS' LEPTODACTYLID FROGS 13 

On Great Thatch, we surveyed the southern slope of the densly 
vegetated island and captured E. cochranae (MCZ 125951). On 
Tortola, we captured E. cochranae on Sage Mountain (MCZ 
116269-71) and also heard advertisement calls from other for- 
ested locations, including sites near sea level. We did not hear 
calls of E. cochranae on Frenchmans Cay, a 24-ha islet separated 
from Tortola by a channel <10 m wide, although males were 
calling in adjacent areas on Tortola on the same night. 

We also did not find E. cochranae on Virgin Gorda, although 
we searched for it several times in 3 years (1993, 1994, and 
1996). MacLean (1982) lists this species from Virgin Gorda, but 
we have been unable to locate a voucher specimen or any other 
report of its occurrence there. 

BODY SIZE OF ELEUTHERODACTYLUS 

The SVL of calling males of E. antillensis did not show sig- 
nificant differences among years on any of the islands examined, 
although males tended to be smaller on Tortola in 1994 than in 
1993 and 1995 (Guana: F34S = 1.67, P = 0.19; Tortola: F,^^^ = 
3.07, P = 0.06; Virgin Gorda: F, „ = 0.18, F = 0.67). Similarly, 
there were no significant differences in SVL of E. schwartzi 
among years (Tortola: F, ,5 = 0.01, P = 0.90; Virgin Gorda: F^j^ 
= 0.38, P = 0.68). The data for all years were therefore combined 
for analyses of interisland differences. 

The average SVL of adult male E. antillensis varied among 
Guana, Tortola, and Virgin Gorda (F. ,4, = 24.9, P < 0.001; Fig. 
2). Males on Virgin Gorda were smaller (x = 27.2 mm) than 
those on Guana (x = 29.3 mm) and Tortola (x = 29.2 mm). The 
average SVL of calling males of E. schwartzi also differed among 
islands (F.j, = 29.4, P < 0.001; Fig. 1). Males were the smallest 
on Virgin Gorda (x = 22.3 mm), largest on Great Dog Island (x 
= 25.6 mm), and intermediate on Tortola (x = 23.8 mm). 

The average weight of calling males of E. antillensis was 1 .7 
g (SD = 0.1, « = 51; 1993-96 combined) on Guana, 1.3 g on 
Virgin Gorda (SD = 0.2 g, « = 58; 1993-94 combined), and 1.7 
g on Tortola (SD = 0.3, n = 52; 1993-95 combined). The average 
weight of calling males of E. schwartzi was 0.9 g (SD = 0.2 g, 
n = 16; 1993 and 1994 combined) on Tortola, 0.8 g (SD = 0.1 



14 



BREVIORA 



No. 508 




CARIBBEAN SEA 



GUANA 






SCRUB 



BEEF 



^18°30'N 

GREAT 
TOBAGO 


LITTLE 
TOBAGO 




/J 



GINGER 



H^ COOPER 



'SALT 



(? 



S^ PETER 



FRENCHMANS 
THATCH 



NORMAN 



GT. THATCH 



64°30'W 



\ 



2 4 



8 km 



Scale 



18°20'N 64''50'W 
_^ ^ 



Figure 1. Map of the British Virgin Islands indicating major islands and those 
mentioned in the text. Insert shows the position of the these islands in the Carib- 
bean. # L. alhilahris, I E. antillensis, A E. schwartzi, -:= E. cochramie. 



1999 



BRITISH VIRGIN ISLANDS' LEPTODACTYLID FROGS 



15 



30- 




Guana 



Tortola 



Virgin Gorda 



Great Dog 



Figure 2. Snout-vent length (SVL) of calling males of Eleutherodactylus an- 
tillensis and E. schwartzi from Guana, Tortola, Virgin Gorda, and Great Dog. 
Mean, top of bars; 1 SD, vertical lines. 



g, n = 39; 1993, 1994, and 1996 combined) on Virgin Gorda, 
and 1.2 g (SD - 0.2 g, « = 17; 1996) on Great Dog. 

Both species were sexually dimorphic with respect to body 
size, females being larger than males. The SVL of 14 female E. 
antillensis measured in 1993 was 33.8 mm (SD = 4.6 mm, range 
= 28.0-43.2) and their weight was 2.7 g (SD = 1.2 mm, range 
= 1.2-4.8 g; all islands combined). Ten female E. schwartzi were 
31.2 mm in SVL (SD = 3.0 mm, range = 25.5-35.5 mm) and 
weighed 1.9 g (SD = 0.4 g, range = 1.2-2.7 g). 

HABITAT USE BY ELEUTHERODACTYLUS 

Eighty-nine percent of all male E. schwartzi (n = 45) and 74% 
of male E. antillensis (n = 171) located in October 1993 were 
perched in vegetation <2.5 m high while calling (data for Tortola, 
Virgin Gorda, and Guana combined). The remaining 1 1% of call- 
ing E. schwartzi and 26% of E. antillensis were perched higher 
than 2.5 m and thus were out of our reach. We did not capture 
E. cochranae in 1993, although we audiotaped calls of this spe- 
cies on Tortola. In 1994, we captured nine E. cochranae (eight 



16 BREVIORA No. 508 

males and one female) at heights below 2.5 m on Sage Mountain, 
Tortola, but traced most calling males to perch sites well above 
our reach in trees. In contrast, we frequently observed E. coch- 
ranae (calling males, noncalling adults, and juveniles) in vege- 
tation <2.5 m high in October 1995 after high winds associated 
with Hurricanes Louis and Marilyn in September had visibly al- 
tered the habitat, knocking down many trees and stripping leaves 
off of those left standing; however, we did not systematically 
record perch heights. 

While calling, males of E. antillensis were most frequently 
perched on leaves or branches of trees and shrubs on Guana (60% 
of 94 observations), Tortola (68% of 53 observations), and Virgin 
Gorda (84% of 79 observations; data for 1993-95 combined for 
all islands). Males also called from herbaceous vegetation (Tor- 
tola: 28%; Virgin Gorda: 4%), terrestrial and arboreal bromeliads 
(Guana: 14%; Virgin Gorda: 4%), and agave plants (Guana: 
24%). On Tortola, calling E. schwartzi were perched on trees or 
shrubs (75% of 16 recordings) and herbaceous vegetation (25%). 
In contrast, the majority of observations of calling E. schwartzi 
on Virgin Gorda were from bromeliads (84% of 45 recordings), 
followed by trees and shrubs (13%) and herbaceous vegetation 
(2%). 

When examined in relation to the availability of bromeliads 
along auditory transects in 1994, the distribution of calling males 
of E. antillensis and E. schwartzi differed significantly from ran- 
dom on Virgin Gorda but not on Tortola (Table 3). On Virgin 
Gorda, males of E. schwartzi were restricted to sections of the 
transect that contained bromeliads. In contrast, male E. antillensis 
were not associated with bromeliads either on Virgin Gorda or 
Tortola (Table 3). On Guana, male E. antillensis were found ex- 
clusively in sections of Transect 2 containing bromeliads, but the 
relationship was not statistically significant based on habitat avail- 
ability, due to the small sample size (Table 3). The frogs were 
most abundant on Transect 1, where bromeliads were present in 
every section, thus precluding a similar analysis. 

On Guana in 1996, calling males oi E. antillensis were aggre- 
gated among 5-m X 10-m sections of the transect during all but 
one check (Table 4). On a larger scale, when the transect was 



1999 



BRITISH VIRGIN ISLANDS' LEPTODACTYLID FROGS 



Q 
Z 

< 

-J 

o 
o 

< 

D 
Oi 
O 

o 



o 

0!! 



z 
o 



ft; 
I 

s 

Q 
Z 

< 



lo On 
D; On 



-J 

2 

si 

u 
o 

O 

U 

< 

o 



< 

u 

o 

a: 
u 

D 



en 
u 

00 

< 



UJ 

oa 
O 

H 

u 
O 



< 

a 



X 



[A 

-a 



.2 C 

o a. 
oa 



o 
z 

C 



c 





in 

o 


o 




d 
A 


d 
A 


o 


« 


— ' 


d 
V 


d 
V 


d 
V 






ly-i 



in 



rt 


'TT 00 o 


ON o o 


o o 


1 


(^, — vO 


r) r<-i r-) 


-^ vO 




^^ 





ri o NO 



00 -H O 

-^ « r- 



o vo 



(N 00 Tf 

_ _ rj 



-H ON O 

-^ -^ in 



o '^ 



•2 ■■- 



T3 
i- 

O 

O 

c 
'5b 



o 

d 
Z 



■2 



c 

.2 ?r 



+ 



^ s 



(U - - _ - _ 

tj_^ S_^ ti^ tj_^ tj_j ^^ 



lA 

c 
o 

4—1 

o 



c .2 



o o o 



o o 



Z bZZZ §ZZ 



T3 

.2 

E 
o 



o 

c 

u 
a 






o 

c/3 

c 



O 

'■a 



c 
o 






c 

t 

>n 
X 



13 
1) 

*-» 
C 
3 
O 

U 



18 



BREVIORA 



No. 508 



< 

-1 



< 
z 

< 
-J 

a 



uu 
f- 



u. 
f- 



X 2 

< § 

o O 

< 5 









_) 
< 



o 
z 

-1 
< 

u. 

o 

z 

o 

a: 

UJ 
Q. 
(/) 



LU 

09 
< 



c 

o 



C/2 



o 



H 




U 


■o 


U 


u 


or) 


■a 


Z 

< 


'> 




i5 








o 


H 


1) 


U 


■Ji 


U 


c 


IZl 


C3 


z 

< 


f^ 



— I/; 



1/5 



ii ^ 



X 

o 
c 

■n 

'> 
5 

C 



X 



c 



5 c^ 



> 



X 



c 






C3 
> 



Q 



o o o 


o 


o o o 


o 


odd 


d 


V V V 


V 



On ri ri 

(^ ^ ON 



■-D 00 rj 
ri ri sd 



iri >n IT) 

o o o 

d d d 

V V V 






o 
A 



^ ^ C) — 



vO 00 — 
On O 00 



ON 



ri 



1) 



00 >o 00 
— rj (N 



O 

cc 



CJ 

u 









c 
o 



> 

3 
O" 

> 

o 

a 



C 






c 
o 

■Ji 



H 



1999 BRITISH VIRGIN ISLANDS" LEPTODACTYLID FROGS 19 

divided into 50-m X 10-m sections, calling males were highly 
aggregated during each check. The habitat attributes measured 
explained little of this dispersion. The number of calling males 
was significantly correlated with the habitat variables both when 
only data for the night with the most calling frogs (74 frogs on 
first transect check on October 28; F(, ,43 = 2.95, P < 0.01) were 
included and when each section of the transect was scored based 
on whether or not it was used by frogs during any of the checks 
(F(, ,43 = 5.67, P < 0.001). In both cases, the correlations were 
weak (r- = 0.11 and 0.19 for the two models, respectively). The 
partial r- for the habitat variables in the better, second model 
ranged from 0.007 to 0.054 and were the highest for the sum of 
crown diameters of bromeliads (0.045) and percent substrate cov- 
ered by leaf fitter (0.054). 

DISCUSSION 

We found 10 previously unreported populations of leptodac- 
tylid frogs on five islands (Great Camanoe, Beef, Frenchmans 
Cay, Great Thatch, and Jost Van Dyke) and confirmed all but 
three of previous records from the BVI. Demonstrating the ab- 
sence of a species is always problematic, and these small frogs 
are inconspicuous when not calling and could be missed easily. 
The month of October, however, is generally favorable for locat- 
ing frogs, because, together with November, it has the highest 
average rainfall per month (6.44 and 6.57 inches of rain in Oc- 
tober and November, respectively, based on weather records from 
1960 to 1984 obtained from Water and Sewage Department and 
Planning Division, Road Town, Tortola, and compiled by A. 
Swain). Rainfall is probably the most important factor affecting 
activity by Eleiitherodactylus species in the BVI, although activ- 
ity is also likely to take place on humid, rainless nights. It was 
not always possible, however, to time our visits to the different 
islands during or immediately after rain. Our confidence that we 
located all species is greatest for small islands that we visited 
repeatedly, such as Guana. We are also highly confident that there 
are no native frogs on either Necker or Little Thatch, because no 
frogs have ever been seen or heard there either by us or by res- 
idents. The only amphibian ever found on Necker was the intro- 



20 BREVIORA No. 508 

duced Hyla (Osteopilus) septenthonalis, which was collected 
there on 19 October 1993 from a crack in a recently imported 
wooden beam (MCZ 119258). 

Two of the three previous records that we failed to confirm 
were from Virgin Gorda {E. cochranae and L. albilabris; 
MacLean, 1982), and the remaining record was from Peter Island: 
an unidentified Eleutherodactylus found in the stomach of a 
snake, Liophis (Alsophis) portoricensis (Henderson and Sadjak, 
1996). This snake (MCZ 37303) was collected by Chapman Grant 
on 14 August 1932. The frog, uncataloged, was sent to the late 
Albert Schwartz for identification, but R. W. Henderson (personal 
communication) subsequently was unable to locate it in 
Schwartz's materials. 

On Peter Island in 1996, we walked throughout the inhabited, 
eastern part of the island at night, and in 1997 we spent a rainy 
night on the south side of the western part of the island investi- 
gating a verdant gully, which to us appeared the best site for 
locating frogs. Eleutherodactylus seems to have disappeared from 
Peter Island at some time since 1932. On Virgin Gorda, we cov- 
ered much of the island at night in the rain in three different 
years, including likely habitats on Gorda Peak (but excluding the 
roadless, easternmost portion of the island). We have found no 
records other than MacLean (1982) of either E. cochranae or L. 
albilabris, nor have we been able to locate voucher specimens. 
Furthermore, MacLean et al. (1977) do not report E. cochranae 
or L. albilabris from Virgin Gorda, raising suspicions about the 
1982 listings. We conclude that the record for E. cochranae on 
Virgin Gorda is in error and that there is no evidence that the 
range of this species extends east of Tortola. We cannot, however, 
conclusively dismiss the possible former presence of L. albilabris 
on the island based on accounts of residents, who remember 
"ditch frogs" in and around Spanish Town many years ago before 
the extensive ponds were drained for the construction of a marina 
and a hotel. 

When all 17 islands visited were considered, elevation and area 
explained much of the variation (61%) in the number of species 
among islands, with elevation being the most significant factor. 
The importance of elevation in biogeographical patterns of small 



1999 BRITISH VIRGIN ISLANDS" LEPTODACTYLID FROGS 21 

islands was emphasized by Lazell (1983), and our data support 
this hypothesis. Our data also show that even very small islands, 
such as Frenchmans Cay, a mere 24 ha, can support at least two 
species of frogs. Frenchmans Cay is relatively high, 131m, which 
might allow it to support more species than expected based on 
area alone. Both Frenchmans Cay and Beef, however, are sepa- 
rated from Tortola by narrow, bridged channels. Frogs dispersing 
in the rain can easily cross such bridges (JL, unpublished data 
from New England and China). Therefore, the number of species 
on these islands may reflect repeated colonizations from Tortola 
rather than permanent populations. The reconfirmation of E. 
schwartzi from Great Dog Island, an islet of only 33 ha located 
at least 3 km from the nearest potential colonization source (Vir- 
gin Gorda), shows that this species can persist on very small 
islands, provided suitable moist microhabitats, such as bromeli- 
ads, are present. 

Leptodactyliis albilabris, which has an aquatic larval stage, can 
be expected to be absent from islands that do not have suitable 
water bodies for breeding. Apart from temporary pools on Gorda 
Peak, drainage ditches (mostly paved) in Spanish Town on Virgin 
Gorda, and small freshwater puddles on Great Camanoe, we did 
not observe potential aquatic breeding habitats on the islands 
where we failed to locate this species. 

Stewart and Pough (1983) showed experimentally that the 
availability of retreat and nest sites can limit population growth 
of E. coqui in Puerto Rico. Terrestrial and arboreal bromeliads, 
plants that hold moisture in their leaf axils, may provide such 
sites for terrestrially breeding forest frogs. Of the three species 
of Eleutherodactylus that we studied, E. schwartzi was most 
closely associated with bromeliads, an association also pointed 
out by Schwartz and Henderson (1991). On all islands except 
Tortola, we found E. schwartzi almost exclusively in terrestrial 
and arboreal bromeliads. Broader habitat use on Sage Mountain, 
Tortola, can be explained by the relatively high rainfall and dew 
that this highest point in the Virgin Islands receives. On Great 
Dog, we found E. schwartzi nests with egg clutches only in a 
small patch of terrestrial bromeliads, which most likely facilitated 
the persistence of the population. 



22 BREV/ORA No. 508 

Eleutherodactylus antillensis and E. cochranae used a variety 
of microhabitats in addition to bromeliads. Most male E. antil- 
lensis called from perch heights <2.5 m in vegetation. Previously, 
Rivero (1978) and Henderson and Schwartz (1991) also noted 
that males often called from low vegetation. We observed E. 
cochranae using cavities in tree trunks and branches for calling, 
retreat, and nest sites (Ovaska and Caldbeck, 1997, and unpub- 
lished data). Most calling male E. cochranae were high in the 
trees, thus limiting our access to this species. Schwartz and Hen- 
derson (1991) stated that males call from 1 m (3 ft) above ground 
to high in the trees. According to Schwartz and Henderson, the 
species occurs primarily in xeric forests. On Tortola, however, we 
found E. cochranae together with E. schwartzi and E. antillensis 
in mesic forest on Sage Mountain. The habitat on Great Thatch 
was also mesic, and only Jost Van Dyke could be characterized 
as mainly xeric. 

Eleutherodactylus antillensis was the most widespread of the 
three species. Although not associated with bromeliads on the 
relatively wet islands of Tortola and Virgin Gorda, the presence 
of bromeliads appeared to become more important with increas- 
ing aridity. On the relatively dry island of Guana, the frogs were 
associated with sites that contained bromeliads and abundant leaf 
litter, although these factors explained only a little of the spatial 
dispersion of frogs along transects. Abundant leaf litter might be 
important for breeding, as all nests of this species that we have 
found have been under leaf litter (Ovaska and Caldbeck, 1997, 
and unpublished data). 

The mean body size of adult males of both E. antillensis and 
E. schwartzi differed among islands. Woolbright (1989) found 
that the growth of male E. coqui in the field ceased after repro- 
ductive maturity was attained. Furthermore, the period of growth 
could be extended in the laboratory under conditions that were 
unfavorable for breeding, thus resulting in greater maximum body 
size. Therefore, frogs that live under social or environmental con- 
ditions that favor the early attainment of reproductive maturity 
can be expected to be relatively small. The selective pressures 
responsible for the observed patterns in body size among islands 
cannot be resolved from our data, and studies that specifically 



1999 BRITISH VIRGIN ISLANDS" LEPTODACTYLID FROGS 23 

address this question are desirable. The differences were consis- 
tent among years, indicating that the operational factors are per- 
sistent over time. 

There are six islands larger than Frenchmans Cay in the BVI 
that have not been surveyed for frogs (Prickly Pear, Ginger, Salt, 
Norman, and Little Jost Van Dyke). All, however, are relatively 
dry and might not be suitable for frogs. Additional populations 
that were undetected by us may also continue to be discovered 
on the islands that we surveyed. Nevertheless, our study provides 
baseline data that may become increasingly important because of 
regional and global changes in climate patterns. 

ACKNOWLEDGMENTS 

Arnold Grobman generously shared his knowledge of frogs of 
the BVI and together with his wife opened his home to us, fa- 
cilitating fieldwork on Virgin Gorda. We are grateful for able 
assistance and company provided by Gad Perry and Kate Le- 
Vering, who spent many a rainy night helping us to locate frogs 
under less than ideal conditions. Wenhua Lu accompanied us on 
several occasions, and many volunteers helped with the surveys 
over the years. Gregory Mayer pointed us to possible sites with 
L. albilabris on Beef Island. Our work was financed personally 
and by The Conservation Agency, through grants from the Fal- 
conwood Foundation. 

LITERATURE CITED 

Blaustein, a. R., and D. B. Wake. 1995. The puzzle of declining amphibian 
populations. Scientific American, 272( April): 52-57. 

Dmi'el, R., G. Perry, and J. Lazell. 1996. Evaporative water loss in nine insular 
populations of the lizard Anolis cristatellus group in the British Virgin Is- 
lands. Biotropica, 29: 111-116. 

Heatwole, H., R. Levins, and M. Byer. 1981. Biogeography of the Puerto Rico 
Bank. Atoll Research Bulletin, 251: 1-63. 

Henderson, R. W., and R. A. Sadjak. 1996. Diets of West Indian racers (Colu- 
bridae: Alsophis): Composition and biogeographic implications, pp. 327-338. 
In R. Powell and R. W. Henderson (eds.). Contributions to West Indian Her- 
petology: A Tribute to Albert Schwartz. Ithaca, New York, Society for the 
Study of Amphibians and Reptiles, volume 12. 

Heyer, W. R., M. a. Donnelly, R. W. McDiarmid, L.-A. C. Hayek, and M. S. 



24 BREVIORA No. 508 

Foster. 1994. Measuring and Monitoring Biological Diversity. Standard 
Methods for Amphibians. Washington, Smithsonian Institute Press. 364 pp. 

JoGLAR, R. L., AND P. A. BuRROWES. 1996. Declining amphibian populations in 
Puerto Rico, pp. 371-380. In R. Powell and R. W. Henderson (eds.). Contri- 
butions to West Indian Herpetology: A Tribute to Albert Schwartz. Ithaca, 
New York, Society for the Study of Amphibians and Reptiles, volume 12. 

Krebs, C. J. 1989. Ecological Methodology. New York, Harper & Row Publish- 
ing. 654 pp. 

Lazell, J. 1983. Biogeography of the herpetofauna of the British Virgin Islands, 
with description of a new anole (Sauria: Iguanidae), pp. 99-1 17. In A. Rhodin 
and K. Miyata (eds.). Advances in Herpetology and Evolutionary Biology. 
Cambridge, Massachusetts, Museum of Comparative Zoology. 

. 1991. The herpetofauna of Guana Island: Diversity, abundance, rarity, 

and conservation. Departamento de Recursos Naturales de Puerto Rico Pub- 
licacion Cientifica Miscelanea, 1: 28-33. 

. 1995. Natural Necker. The Conservation Agency Occasional Paper, 2: 



1-28. 

Leonard, W. P. 1997. Review: Frog and toad calls of the Pacific coast: Vanishing 
voices. Herpetological Review, 28: 56. 

MacArthur, R. H., and E. O. Wilson. 1967. The Theory of Island Biogeography. 
Princeton, New Jersey, Princeton University Press. 

MacLean, W. p. 1982. Reptiles and Amphibians of the Virgin Islands. London, 
Macmillan Education Limited. 54 pp. 

MacLean, W. P., R. Kellner, and H. Dennis. 1977. Island lists of West Indian 
amphibians and reptiles. Smithsonian Herpetological Information Service, 40: 
1-47. 

Mayer, G., and J. Lazell. 1988. Distributional records for reptiles and amphib- 
ians from the Puerto Rico bank. Herpetological Review, 19: 23-24. 

Mittermeier, R. a., J. L. Carl, 1. R. Swingland, T B. Werner, and R. Mast. 
1992. Conservation of amphibians and reptiles, pp. 59-80. /// K Adler (ed.), 
Herpetology: Current Research on the Biology of Amphibians and Reptiles. 
Oxford, Ohio, Proceedings of the First World Congress of Herpetology, So- 
ciety for the Study of Amphibians and Reptiles. 

OvASKA, K., AND J. Caldbeck. 1997. Courtship behavior and vocalizations of the 
frogs Eleutherodactyhis antillensis and cochranae in the British Virgin Is- 
lands. Journal of Herpetology, 31: 149 155. 

Ovaska, K., J. Caldbeck, and J. Lazell. 1998. Eleiitherodactylus schwartzi 
(NCN). Reproduction. Herpetological Review 29: 97. 

Pechman, J. H. K., and H. M. Wilbur. 1994. Putting declining amphibian pop- 
ulations in perspective: Natural fluctuations and human impacts. Herpetolo- 
gica, 50: 65-84. 

Phillips, K. 1994. Tracking the Vanishing Frogs: An Ecological Mystery. New 
York, St. Martin's Press. 244 pp. 

RiVERO, J. A. 1978. The Amphibians and Reptiles of Puerto Rico. Mayagiiez, 
Puerto Rico, Universidad de Puerto Rico. 148 pp. 



1999 BRITISH VIRGIN ISLANDS' LEPTODACTYLID FROGS 25 



1991. Divagaciones sobre las especies de coquies en peligro de extincion 



y las causas posiblcs de esa situacion. Departamento de Recursos Naturales 
de Puerto Rico Publicacion Cientitica Miscelanea, 1: 54-55. 

Schwartz, A., and R. W. Henderson. 1985. A Guide to the Identification of 
Amphibians and Reptiles of the West Indies Exclusive of Hispanola. Mil- 
waukee, Michigan, Milwaukee Public Museum. 165 pp. 

. 1991. Amphibians and Reptiles of the West Indies: Descriptions, Distri- 
butions, and Natural History. Gainesville, Florida, University of Florida Press. 
720 pp. 

Schwartz, A., and R. Thomas. 1975. A check-li.st of West Indian amphibians 
and reptiles. Carnegie Museum of Natural History Special Publication, 1: 1- 
126. 

Stewart, M. M., and E H. Pouch. 1983. Population density of tropical forest 
frogs: Relation to retreat sites. Science, 221: 570-572. 

WOOLBRIGHT, L. L. 1989. Sexual dimorphism in Eleutherodactylus coqui: Selec- 
tion pressures and growth rates. Herpetologica, 45: 68-74. 



5376 



J 



Harvard MCZ Library 




3 2044 066 302 837