HARVARD UNIVERSITY Library of the Museum of Comparative Zoology BREVIORA Museum 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.1 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). 1 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. 2 BRE VI ORA No. 488 124° 126° '28° Figure 1. The northwestern Molucca Sea showing the islands and banks (200 meter depth) and the locations of places mentioned in the text. 1987 A NEW FLYING LIZARD 3 Alter 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 Anolis 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 1 5 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 thornlike 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 4 B RE VI ORA No. 488 Manure*. lu'rfc.UeJl 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 lichenate pattern. The chest was washed with dull yellow. The throat fan was pale lemon- 1987 A NEW FLYING LIZARD 5 yellow — almost white — 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 170919 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 oi grey than do laige adults. Inger and Musters concur that Draco of both the lineatus and 6 BREVIOIU No. 488 vo/ans 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 170917 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. 1. 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 170930), the larg- 1987 A NEW FLYING LIZARD 1 Table 1. Seven ways in which species of Draco from Sulawesi differ. volans lineatus biaro 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 biaro 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 desciibed 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 BRE VI ORA No. 488 I have examined six specimens of 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 D. 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 Ahaetulla prasina, the skinks Mabuya multifasciata and Lamprolepis smaragdinus, and three geckos. Hemidactylus frenatus and Gehyra mutilata are abundant Indo-Pacific human commensals. Gymnodactylus jellesmae is rare in collections and seems to have been previously known only from Sulawesi. 1987 A NEW FLYING LIZARD 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. 1972. The anoles (Sauria: Iguanidae) of the Lesser Antilles. Bulletin of the Museum of Comparative Zoology, 143(1): 1-1 15. Musters, C. J. M. 1 983. Taxonomy of the genus Draco L. (Agamidae, Lacertilia, Reptilia). Zoolische Verhandelingen 199: 1-120 + 4 plates. BREVIOR A Mu sen in ol Comparative Zoology US ISSN 0006-9698 Cambridge, Mass. 2 September 1988 Number 489 NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. V. AN O LIS DAN I ELI, A NEW SPECIES OF THE LATIFRONS SPECIES GROUP AND A REASSESSMENT OF ANOLIS APOLLINARIS BOULENGER, 1919 Ernest E. Williams1 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. propinquus Williams, 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 A. 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 A. biporcatus Wieg- mann. Material since made available to me from Cundinamarca and Antioquia now makes it quite clear that my 1970 material was 1 Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- setts 02 1 38. 2 BREVIORA No. 489 composite, and that only the Cundinamarca specimens were A. apollinaris. Fortunately, the illustrations in Williams (1970) are of Munich specimens from Cundinamarca, which are true A. apollinaris. 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. apollinaris and the new species belong to the alpha section of the genus (Etheridge 1960) and must be referred to the Anolis 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 A. apollinaris 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. apollinaris 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 of A. apollinaris 106 mm) and he compared the species with A. solifer and A. copei (both synonyms of A. biporcatus), which are approximately this size. Unfortunately, Dunn did not compare apollinaris 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 Gunther (1859), describing Anolis fraseri, included a specimen of 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. apollinaris with the two synonyms of A. 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 111 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 Serna 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 A. 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. Serna coll., 28 May 1972; CSJ 441, M. A. Serna and H. Echeverri coll., 23 March 1983; CSJ 720, M. A. Serna and H. Echeverri coll., 26 July 1985. Yarumal: ICN 5998 (formerly CSJ 168), collector and date of collection un- known. Referred Specimen. “Western 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 ANOLIS DANIELI 5 semicircles, which is abruptly larger (rather than all 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 BREVIOR.4 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, 0 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 of A. danieli there is no data on color in life. The best information (translated) has been provided by Marco Antonio Serna 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 patternless, 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 ol 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 15 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 danieli' s 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 B REV 1 ORA 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 (C SJ 435) is known from El Retiro, 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 1 1 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. apollinaris 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— A. 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 B REV 10 R A 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. Serna and H. Echeverri for providing the newer specimens of A. 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 of 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. 1 885. 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: 68-8 1 . 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. Williams, E. E. 1 966. South American anoles: Anolis biporcatus and Anolis fraseri compared. Breviora Museum of Comparative Zoology, No. 239: 1-14. . 1970. South American anoles: Anolis apollinaris Boulenger, 1919, a relative of A. biporcatus Wiegmann (Sauria, Iguanidae). Breviora Museum of Comparative Zoology, No. 358: 1-1 1. . 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, 481^183. In R. Huey et 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 center anteroposteriorly diating from 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 3-4 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 {lh supracili- shorter and these by smaller ary ) margin these by subgranular se- ries conical scales followed by granules Ear 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 behind it largest next su- praorbitals, slightly smaller behind interpari- etal Scales between 2—4 2-5 no interparietal interparietal and semicir- - cles Scales between 0-1 0-1 1 suboculars and supralabi- als 1988 ANOLIS DANIELI 15 Table 1. Continued. apollinaris danieli propinquus Supralabials to 7-8 7-9 7 below center of eye Trunk scales swollen, keeled with inter- spersed granules; middorsals slightly enlarged or 2 rows dis- tinctly so swollen, keeled or pyramidal, sur- rounded by gran- ules; 2 -A mid- dorsal rows enlarged granular, con- vex, subequal Ventrals Femoral scales smooth or keeled, juxtaposed to imbricate unicarinate, multi- carinate near knee smooth, subimbri- cate to imbricate unicarinate, multi- carinate at knee smooth, juxta- posed or sub- imbricate 4th toe lamellae 23-29 23-27 25 Dewlap in male only large in male and female; smaller in female hatchling; no fe- male known Dewlap scales densely scaled, scales bluntly keeled densely scaled, keeled scales crowded but a series of 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 6 1 12 3 117 unique type, a SVL 2 94 2 104 hatchling Table 2. Latifrons group anoles with long legs and green bodies. 16 BREVIORA No. 489 03 & u a c a. on a o & to a .a a 5 a a a> on on K 45 45 Si a £■ a a. on a ■♦**» a a Si T3 45 45 1) -54 I (N T3 45 45 2 E a ’S ■g 3 * « _r x 1 2 1 S C/3 HD jD 13 CD X -4—* o o 6 03 X cd . c cd Cd C/3 cx 0 0/ U 1/5 3 0 13 0 C/3 H 3 <£: >> G C ^ X G 1> 3 S Uh ^ a u •♦-» '0 u jd 3 C/3 3 •D (D *G G cd Wh 1 cx G O Uh •♦-* X G *c CD cd u 00 O > 0 O u -♦-* cd c/3 > X C/3 13 o C/3 'O cd ^ 3 2 CO o 3 45 W OfiCS C/3 c/3 c G cd 00 G G X O 45 CO X - ° E n £ x .£ ts -c X 45 c o x C/3 G O cd O 5— CJ 00 ^ * -§ s o 5 -r 1 3 15 3 — C 2x3 C S o O _ 35 ^ X> SR >> C 3 45 2 So x' 3 « 'C G T3 G cd -*-• G CD 3 C/3 C/3 O O £ +-> X V- 3 Wh >. . S c -g « -C •£ Z § « 5 g o «3 2 P> £ a 2 -c <3 •o - £ 3 3 ~ o t; “ i- 3 E £ u -O C S 3 3 2 cd d> 0/5 13 d> « g cd S E = £• n «3 2 2 Ji a5’§*jS.S E & Vh i O g ■o •’"> 22 00 T3 •C £ d> ox) 3 > S3 o c 2j u o o 00 c 00 I 3- T3 £ d> OS O SI O c 03 a ■^4 a § <4 00 H cd o 3 =3 2 o cd ».r\ a' 0 s E 8 3 3 3 *-» u r 3 a VO I rn I 00 •C X) E x> 3 C/3 •o C3 1 c jd 13 a pari- 1 0/5 u D, X) G 0 O G 0/5 0/5 c 0^ T3 0/5 0/5 X) XJ 03 0 G d) G G 13 .2 8 X3 > 3 > 3 C/5 0/5 JD 13 2 H posed, no en- middorsal rows frequent inter- taposed, mid- taposed, larged slightly enlarged spersed granules dorsals not or middorsals middorsal rows slightly enlarged not enlarged Table 2. Continued. 18 BREVIORA No. 489 03 •S Co J3 <3 3 cs- 03 ■> i C <4> 4> 5! 3 & 3 Cl. 1 i 3 3 K 4l 2 H X •'" 3 O ^ C/3 O d> -*-» 03 O C a X) 03 D H, S •fi ’S d> 03 C/3 O o C a x> ^ T3 > 41 41 41 .Mm' O ^ X 2 ef T O S ° 5 E * 03 -*-» X r£ T3 ■“ (U cd -*-* X 3 C/3 O a 'C x E X3 41 d> ■4-* o3 „ G £ ‘C 2 o ^ 'il-S -1 ES 3 •-H 41 X 4i 1- -4-» 41 « e a c 2 'C C § -8 41 cd O X m (N 41 O $ a ° c 3 •rt 41 x 4i •*-* 41 «< E £ c 2 ■d c § -0 u 3 a u 3 ■-> 41 x 4i M >-* 41 ^ E £ 3 2 'C « cd C 41 3 c ° 3 C/3 13 o C/3 03 t- o E d> Uh 00 u cd C/3 d> X d> C/3 VO (N a +-* d> G c r 13 O C/3 C/3 (N c/3 i 2 U c/3 03 in >. o u *— I ’“x 2 3 41 * _T C T3 41 -tJ 2 o 41 c ^ 03 >3 « 3 CJ 03 03 £ O "i U 41 3 Cd 41 is 41 | O 3 O. J3 03 41 41 41 03 3 Z * 2 m 4i C/3 T3 o ^ Vx C/3 c 3 ■ 3 O C/3 C/3 o '{i. 41 41 _. 33 C 73 g E « 41 o C/3 C/3 13 S G oo ,rH (N . --h 03 t-j 4j ■w -o >. ^ “ §8.2 8? d) £J> Cd O o3 si s * d) o o o E C/3 d> o a jd xi o $ Q C/3 13 o C/3 O- 03 41 Q C/3 13 o C/3 Id G cd +-» s cu X E E c o Wx (N Ov • pH c o «— i •H* C c 03 E *o o X ^ O 2 o c 41 > 3 X > 40 cC H E H j X > 40 s 1988 ANOLIS DAN1ELI 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 tenor nasal sepa- anterior nasal rated from ros- rated from rostral separated from tral by one by one or two scales rostral by one or two scales scale 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 or subgranular followed by a double series of smooth series of smooth squarish scales scales, irregu- lar in size scales Ear moderate moderate moderate Loreal rows 5-9 5-8 3-5 Interparietal moderate moderate not differentiated 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 0 0 U suboculars and suprala- bials Supralabials to 6-9 7-12 /-y below center of eye 20 BREVIORA No. 489 Table 3. Continued. fraseri insignis microtus Trunk scales smooth with inter- spersed granules, none or two mid- dorsal rows en- larged smooth, juxta- posed one to 3 middorsal rows enlarged smooth or slight- ly rugose; flank scales rhomboi- dal; flat, mid- dorsal scales elongate, rather irregular in shape Ventrals smooth or keeled, juxtaposed to im- bricate smooth, juxta- posed to sub- imbricate smooth, juxta- posed or imbri- cate Femoral scales unicarinate, multi- carinate at knee smooth wrinkled, not keeled 4th toe lamel- 18-24 23-27 20-22 lae Dewlap large in both sexes large in both sexes large in both sexes Dewlap scales small, smooth densely scaled, scales small, very weakly keeled very weakly and densely scaled Postanal scales present in males, sometimes obscure, absent in females Scales poste- keeled smooth smooth rior to vent Tail crest never present in any species Tail SVL ca. 2x ca. 2x ca. 2x Maximum 6 1 16 3 153 3 111 SVL 9 102 9 135 9 104 1988 ANOLIS DANIEL1 21 Figure 1 head. Anolis danieli, new species, ICN 5997 (holotype). Dorsal view of 22 BREVIORA No. 489 Figure 2. head. Anolis danieli, new species, ICN 5997 (holotype). Lateral view of Figure 3. Anolis danieli, new species, ICN 5997 (holotype). Ventral view of head. 1988 ANOLIS DAN1ELI 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 DANIELI 25 T A. apollinaris V A . da n ie I i O A . f r aser i O A. frenatus • A. latifrons # A. princeps © A. purpurescens Figure 7. Map of distribution of the Anolis latifrons species group in Colombia. BREVIORA Museum 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. MA C ULI VENTRIS BOULENGER, 1898 AND A. MEDEMI, A NEW SPECIES FROM GORGONA ISLAND Stephen C. Ayala1 and Ernest E. Williams2 Abstract. Anolis maculiventris Boulenger, 1898, the widespread Pacific low- land fuscoauratus group anole of northwestern South America, is redescribed on the basis of a series of specimens from the region of the type locality 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- like 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, 1 908, A. fuscoauratus Dorbigny, in Dumeril and Bibron, 1837, A. maculiventris Boulenger, 1898, A. ortoni 1 929 Pepperwood Lane, Petaluma, California 94952. 2 Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138. OCT 3 1 1988 2 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 FUSCO AURATOID ANOLES 3 Cope, 1868, and A trachyderma Cope, 1875. Two additional taxa, A. tolimensis Werner, 1916 and A mariarum Barbour, 1932, were treated as synonyms of A. antonii but are now known to be valid species. A. ortoni has subsequently been considered a member of the pentaprion 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-Pena 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 fuscoauratus 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 1 6 Colombian localities for A. maculiventris, including six localities (93 specimens) visited by Ayala’s group, and 1 0 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 ANOLES 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 Narino (Figs. la-f). 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 44-49 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 ANOLES 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 patternless 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). 1988 TWO FUSCOAURATOID ANOLES 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 1979. 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 Castano coll., 22 May 1979. Diagnosis. A beta Anolis of the fuscoauratus 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 ANOLES 11 Figure 3. James Lazell’s drawing of MCZ 78944 shows the boldly accentuated pattern of the series preserved by Federico Medem. 12 BREVIORA No. 490 about 2 x snout-vent length. No enlarged postanal scales (except in MCZ 78947). Measurements. Holotype. Head length 15 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 ANOLES 13 A 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, listed originally as A. “ fasciatus is a juvenile A. chocorum Williams and Duellman, a rain forest species found sporadically between Costa Rica and western Colombia; BMNH 1926.1.20.107, listed 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 “ binotatus ” 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 3-4, 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 differences between Anolis medemi and Anolis maculiventris. Character Anolis medemi Anolis maculiventris Basic color rufus, red- or orange brown brown, olive-brown, brown-gray Pattern usually evident, often prominent; dark bars on sides and back of head, body, tail and legs sometimes prominent, usually none at all; dor- sal surfaces seldom barred, if so not promi- nent Vertebral region series of 9-1 1 more or less prominent spots or bars between head and tail often dark; usually little or no trace of dark spots or bars between head and tail; some- times dark spots at top of lateral bars on flanks Rank region usually 5-6 darker brown bars, with no tendency to black bars if present may be nearly black; usually al- most completely absent Back of head Dark bar over head be- tween eyes present, usu- ally prominent; occipital spot present little or no pattern; bar between eyes often not prominent; occipital 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 ANOLES 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 A. 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 Ayala, 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 at. (eds.), 16 B REV I ORA 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. Soc. 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. II. 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. BREVIORA Museum of Comparative Zoology Cambridge, Mass. 25 March 1991 Number 491 MAY 0 1 1991 HARVARD LARVAL DEVELOPMENT/ RELATIONSHIPS, AND DISTRIBUTION OF MANDUCUS MADERENSIS, WITH COMMENTS ON THE TRANSFORMATION OF M. GREYAE (PISCES, STOMIIFORMES) David G. Smith,1 Karsten E. Hartel,1 and James E. Craddock12 Abstract. Larval development of Manducus maderensis 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. 1 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. 2 Woods Hole Oceanographic Institution, Woods Hole, Massachusetts 02543. 2 BREVIOH4 No. 491 MATERIALS AND METHODS The primary material examined by us is from the Woods Hole Oceanographic Institution (WHOI) collections housed at the Mu- seum of Comparative Zoology (MCZ), Harvard University. Ad- ditional material from CAS, IOS, ISH, UMML, URI, and USNM was also examined. All institutional acronyms follow Leviton et al. (1985). Counts and measurements follow Hubbs and Lagler (1964), and photophore terminology follows Weitzman (1986:227). Counts of bilateral structures were made on the left side. Specimen size is given as standard length (SL) in mm. Drawings were made with the aid of a Zeiss SV-8 stereo microscope with a camera lucida attachment. DESCRIPTION OF MANDUCUS LARVAE Manducus maderensis The larvae of M. maderensis can be distinguished from the larvae of other elongate “gonostomatids” by the combination of: (1) pronounced annular mucosal folds along the intestine, (2) a dorsolateral row of chromatophores, and (3) fewer than 40 anal- fin rays. The smallest specimen, 10.0 mm (MCZ 82191, Fig. 1A), is completely untransformed although notochord flexation is com- plete. It is moderately elongate with a body depth approximately 9% of SL. The dorsal and anal fins are fully formed. The dorsal fin is located posterior to the midpoint of the body and slightly anterior to the anal-fin origin. The pectoral fin has a well-devel- oped base and blade but lacks rays. The pelvic fins are just be- ginning to develop and are located slightly less than one dorsal- fin length anterior to the dorsal-fin origin. The preanal length is 70% of SL. The gut consists of a short esophagus, a short sac-like stomach, and a long intestine. The intestine is externally visible and has pronounced annular mucosal folds. A series of 22 dark chromatophores runs longitudinally along the dorsal part of the body, just ventral to the dorsal midline, from the level of the pectoral fin to the end of the caudal peduncle; this series of chro- matophores will be referred to as the dorsal series. There is a 1991 MANDUCUS, DEVELOPMENT, RELATIONSHIPS Figure 1. Manducus maderensis. A) Larva, 10 mm SL (MCZ 82191). B) Transforming larva, 18 mm SL (MCZ 82194). Drawn by L. Meszoly. 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, 18.0 and 18.5 mm (MCZ 82194 [Fig. 1 B], and 82193), 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 1 8 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 82178). At 24 mm (MCZ 82177) 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- 6 BREVIOR.4 No. 491 .jjO ^3 05 03 G cd H N £ u 2 ">! (N ^ nn N £ O 2h 5 T3 — (S3 03 X) c 03 03 c 03 . > < U < CQ K Oh > < O ca. 44 Table 2. Selected counts and measurements for transforming larvae of Manducus greyae. 1991 MANDUCUS, DEVELOPMENT, RELATIONSHIPS N vo fN oo N in O or 5 fN ' OO N m vJ Tt 5 fN ^ 00 o OT ^ fN ^ OO Pi vo U Tf S fN ^ OO N ^ pi vo or o? fN ^ OO N 01 Pi vo O OT "S fN ^ 00 o o a oo o ofr of fN — i fO of I I m m m o — i in in fN fN — < m in + tn m fN ov o (N ro O — 1 — ' or in I I ro — 1 ro or d o o Pi > < > < O Pi _l cu d cd < + c •«-» o cd Vh cd X3 O tT E 19 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 BREVIOR.4 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 a/., 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, IOS). 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 91350, 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 1 8 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 MANDUCUS, 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 IOS 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#1 3* (2:18-21) 17°48'N, 25°29'W, 515-600 m, 0950 hrs.; 7089#2 1 * (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) olf 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 : 1 73) WH 1 9 1/66, 2 1°00'S, 30°00' W, 200 m, 2000 hrs.; 3 1 3/68* ( 1 : 1 77) WH 8-III/68, 26°1 0'N, 1 9°26'W, 580 m, 2233 hrs.; 1 125/68* (1:56) WH 20-III/68, 13°56'S, 27°38'W, 580 m, 2255 hrs.; 1665/71* (1:1 57) WH 443/71, 21°35'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.; 28 19/7 1* (1: 125) WH 498- III/71, 1 7°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, 21°17'N, 85°22'W, 124 m, 0020 14 BREVIOR.4 No. 491 hrs.; 54303 (1:110) Oregon 2007, 7°34'N, 54°49'W, 445 m; 56952 (1:145) RHB3052, 1 1°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°1 3'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°31'W, 475 m, 0055 hrs.; 82183 (1:61) RHB2946, 9°03'N, 5 1°05'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, 0 1 20 hrs.; 82 1 86 (2:22 & 52) SUN2083, 1 7°08'N, 27°55'W, surface, 000 1 hrs.; 82187 (2:25 & 27) SUN2 101, 26°37'N, 4 1°1 8'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, 11°00'N, 40°10'W, 500 m, 2045 hrs.; 82191 (1:10) RHB2966, 12°21'N, 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 82 1 93; 82 1 97 (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°21'N, 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°1 5'N, 46°50'W, 100 m, 0300 hrs.; 88257 (1:24) JEC7712, 0°01'N, 37°40'W, 80 m, 2235 hrs.; 88258 (1:18) 1991 MANDUCUS, DEVELOPMENT, RELATIONSHIPS 15 JEC7705, 3°08'N, 42°52'W, 25 m, 0 1 30 hrs.; 91350(1 :220) Ocean Acre 12-55N, 32°11'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°01'N, 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°51'E, 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; 24871 1 (1:27) Ocean Acre 1-18C, 32°10'N, 63°48'W, 100 m, 0145 hrs. Manducus greyae MCZ 75518 (1:43) GRH1046, 12°38'S, 148°55'E, 3,240 mwo, 1740 hrs.; 82462 (1:22) GRH101 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, 15 1°05'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 BREVIOJU No. 491 pha” (4), Maurolicus muelleri (269), Photichthys argenteus (1), Pollichthys mauli (79), Valenciennellus tripunctulatus (553), Vin- ciguerria attenuata (1,059), V. nimbaria (1,807), V. poweriae (&25), 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 IOS), 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 (USF), 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 17845). 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, Sternoptychidae, and associated Stomiiform groups: Develop- ment and relationships, pp. 184-198. In 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. Craddock, 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 MANDUCUS, 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. In 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. 1982. 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. Gibbs, 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. Mukjhacheva, V. A. 1978. A review of the species of the genus Diplophos Gunther (Gonostomatidae, Osteichthyes) and their vertical and geographical distribution. Trudy Institute of Oceanology, 111: 10-27 (English translation). Ozawa, T., and K. Oda. 1 986. 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. ORA MCZ B R E'X Museum of G^^:p,a-yatiYe Zoology UNIVERSITY US ISSN 0006-9698 Cambridge, Mass. 25 March 1991 Number 492 A PERUVIAN PHENACOSAUR (SQUAMATA: IGUANIA) Ernest E. Williams1 and Russell A. Mittermeier2 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 Hernandez-Camacho, 1988, from Gutierrez, Department of Cundinamarca). Specimens or species known or suspected to be from adjacent 1 Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- setts 02138. 2 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 Hernandez-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 EEW 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 3 Figure 1 . Phenacosaurus orcesijuv., MCZ 165211. 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 of Anolis over many years (see also Figs. 1-4). 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. 4 BREVIOK.A No. 492 Figure 2. P. orcesi juv., MCZ 165211. Dorsal view of head. Figure 3. P. orcesi juv., 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 5 < Figure 4. P. orcesi juv., 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 corner 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 7 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 is absent in all juvenile Phenacosaurus { MCZ 14165: P. heteroder- mus, 30 mm SVL; EPN 2218: P. sp. 38 mm SVL). It also lacks the differentiated large round flat flank scales separated by gran- ules, characteristic of P. heterodermus, 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. heterodermus [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 9 10 BREVIORA No. 492 Figure 6. P. orcesi Paratype, USNM 166533. Dorsal view of head. 1991 A PERUVIAN PHENACOSAUR Figure 7. P. orcesi Paratype, USNM 166533. Ventral view of head 12 BREVIOK.4 No. 492 Table 1 . Comparative scale variations in the Peruvian phenacosaur and orcesi and heterodermus. Peruvian phenaco- saur orcesi 57 hetero- dermus Scales between second canthals 4 4 3-6 Postrostrals 4 2-3 3-6 Scales between supraorbital semicircles 0 0 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 0 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 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 of P. 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 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 21 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 : (1) 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. 78°00' 77° 30' 77e00' 14 BREVIOH4 No. 492 Figure 9. Left: Map to show the Venceremos area in relation to Peru as a whole. Right: Detail map to show Venceremos in relation to other localities in northern San Martin and adjacent Amazonas, Peru. 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. 10) 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 BREVIOIU 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 Phenacosaurus. Caldasia, 3(1 1): 57-62. Hellmich, W. 1 949. Auf der Jagd nach der Paramo-Echse. Deutsche Aquarien- und Terrarien-Zeitschrift, 2(5): 89-91. Lazell, J. D., Jr. 1 969. 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 BRE Yu 1 1,, 0 R A useunm of Cditi^araltiYe Zo UNivdfRsrj y US ISSN 0006-9698 olo Cambridge, Mass. 25 March 1991 Number 493 NEW COLOSTETHUS (AMPHIBIA, DENDROBATIDAE) FROM SOUTH AMERICA Juan A. Rivero1 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 Serna, 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 Serna, 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 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. 493 Figure 1. MCZ-A 100217, holotype of Colostethus 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, 1988) with •A-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 3A 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 BREVIOIL4 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 12 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. 8 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 IOS 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 LF 8.70 7.90 7.20 7.93 LT 8.90 8.30 7.35 8.18 LFT 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; IOS = breadth of interorbital space; UE = breadth of upper eyelid; ED = eye diameter; DT = tympanic diameter; LF = length of femur; LT = length of tibia; LFT = 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- 991 NEW COLOSTETHUS FROM SOUTH AMERICA i u. OX) co 00 NO o r-~ ■u- 00 ON ro (N as Nf >70 o CN 00 NO Os >o 70 ro — i CO CO oo Tf 00 q q rt N- Os > < cd >TN CN 00 00 (N 70 O in NO OO in On it 00 CN (N o NO ON >7"N — • ON CO CO 00 CO Os q N" >n q On O o CN r-* r- co 70 o >70 NO ro OO o co NO r- , CN (N q co q o q q o rn o it) o O O o IZN *TN o o >70 CO >70 SD tT o r- . — . CN CN 00 CN iq q in q VO q o *— < CO CO CO q q a lS~) q in q ON a o u o o r-* CN ON ON d (N d ni d n «ZN in o o o >70 N* 00 in >o ■it •— i N" NO VO C CN o OO tq q >n q rs m — 1 00 NO CO CO 00 N" 00 q Tf in 00 ca O co r-* oo d ni 70 >70 CN r- r-' NO Os CN NO 00 *— < VO l— i 70 NO NO o ON it CN ON (N N- ■n CN co o On co oo ITN q (N in q >70 CO CO oo CO as. q N" in N" ON c A JU O o ON •—1 r-’ NO d — 1 70 O CN N- as 70 o >70 >70 N- OO cn N" NO 00 NO r-~- cn o no q in q q o C<0 q CO q oo N" 00 q q q ON o OO ON d d ni d ni d d CO d CN d d o o o — ' d d d d CO .O 0) o CN u NO l/^ O >tn o o in 70 o O o >70 N- TN O >/N O o o >70 o o >70 CN o o 00 00 CO "nT VO CO VO C3 CN CN ‘T'N tq q r~~ 70 in >70 o ti* N- o as a NO VO tq >o co q m m »— < (N q CO CO CO N- r-~ co N" N- ON O Tf oo oo d > oo o Q C/5 H z > > > oo H 15 ■*— * C^J No. AS QQ hJ C/3 H z SOI W Q H q H t GO \ CQ C/3 w UJ N H w Q q \ Q C/3 ui H t q q u I W q P w Q q d q X DC D Q w w q q q For key to characters see Table la. 10 BREVIORA 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 xh 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 COLOSTETHUS FROM SOUTH AMERICA 1 1 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 BREVIOR.4 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 idiome/us 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 idiornelus 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 Figure 3. MCZ-A 89108, 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 lA size of tympanum; a short, oblique 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 IOS 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 LFT 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 LFT/SV 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 89106 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. 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 V2 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 3A 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 B RE VI ORA 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. fu/iginosus and C. nexipus, are quite different from C. macu/osus, and so is a species from western Ecuador, which will be described elsewhere. Colostethus paradoxus, sp. nov. 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 V2 free; choanae small, rounded; canthus rostralis rounded but an- 1991 NEW COLOSTETHUS FROM SOUTH AMERICA 21 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 BREVIOH4 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 Vi 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 Serna (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 Narino (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 Latin facies, 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- Figure 6. MCZ-A 94751, 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 % 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 Y-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 BREVIOH4 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). Table 3. Colostethus faciopunctulatus measurements and proportions. 1991 NEW COLOSTETHUS FROM SOUTH AMERICA 27 OD cd u Cn r- cn cn in in m 00 r- r- in oo cn CN 00 oo cn CN CN CN cn »— < d oi o o d o o •— H d o d < CN in m in o o O in O O m o o o oo r- cn o N" cn o r- o 00 r- m in in cn CN — 1 cn so q SO o m cn q m Os q >n q o N- CN 00 00 cn n so q q SO q m cn 00 •o- 00 in St N- cn q in Os Os CN CN cn cn — J X d Os o d d d d X d o d X OS CN O in o O in O o o m o o >n Os 00 so in CN o os in cn SO in CN o q s C > m q ITS q r- SO m cn q q 00 cn >n N- q n- cn Os OS CN CN CN cn X X q o o d d d d X d o o X Os CN CN r- oo cn CN CN cn cn O d o^ d d d d d d d d 1 CN _ < SO O o o O o O o o o o o so cn o (N u- in t" o St N- r- CN so q Os cn CN CN 00 CN o q m cn q cn Os SO n- in q q — * r-’ C cn (N CN cn cn X o o 00 o d *— H d d X d d d d Os CN C/5 CN O o o o O O o o o o >n o m so cn o Os o o in so cn oo OO in CN 00 q 00 so q i-H m — H m cn cn q q r- in m q q ~ cd r- cn CN r- oo cn cn CN cn cn CN q o d d d d d d d d d q 2 Os CN Os O o o o in m m o o o >n SO Os oo Os in 00 so r-- CN Os r- — ^ r- so CN CN CN q — 1 > C/3 o Q C/3 H Z > > > oo H ft o C/3 H SOI C/3 C/3 q W UJ C/3 q •*—> cd Z > 03 q z w Q H q H m w H Q Q q H t q u C/3 X X W q X w Q q q q X X D Q U4 w q X X q For key to characters see Table la. 28 BREVIOR.4 No. 493 The Rio Azuela species also has white specks on the upper lip, 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 ofKansas, 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 identification de los Colostethus (Amphibia, Dendrobatidae) de Colombia. Caribbean Journal of Science, 24(3- 4): 137-154. B R Muse mm l'8R%y 2 4 /99p E VM R Ni^£Rsity of Comparative Zoo 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. Laurent1 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 of Liolaemus is discussed. INTRODUCTION The bewildering diversity of the genus Liolaemus Wiegmann has been well documented by L. Muller, 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- 1 Investigador Principal del CONICET, PRHERP-Fundacion Lillo, Miguel Lillo 251, 4000 Tucuman, ARGENTINA. 2 BREVIOJE4 No. 494 Table 1. Characters of the subgenera Liolaemus and Eulaemus. Liolaemus (395 <5<5, 356 $2) Eulaemus (313 <53, 313 22) Preanal pores 0-7 3-12 Jt= 2.19 x = 6.40 <5 in 91% of sped- >4 in 92% of speci- mens, the series gen- mens, the series gen- erally shorter than the erally longer than the 1st toe (12.4% of ex- 1st toe (2.29% of ex- ceptions) ceptions) Nostrils lateral latero-dorsal Distance between upper inferior (3.37% of ex- superior (7.03% of ex- border of subocular ceptions) ceptions) and lip compared with distance between nasal plates Upper labials generally flat and long, generally high and the 4th below eye, short, 5th, 6th, or 7th with posterior border below eye, with pos- oblique terior border vertical Range Chile (>50 taxa) northern Chile (6 taxa) southern and western southern and western Argentina ( 1 8 taxa) Argentina (25 taxa) Bolivia (3 taxa) Bolivia (6 taxa) Peru (3 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 of 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 of 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. walked Shreve are names that have been applied to a large set of Andean populations from Peru southward to Catamarca Province, Argentina. The form walked 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 walked. 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 syn types 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 BREVIOR.4 No. 494 However, a comparison of the syntypes with Bolivian 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 may 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 ofL. 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 (Dumeril 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 LIOLAEMUS 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, 11,300 to 13,600 feet,” and L. annectens orientalis Muller, 1923 (synonymized with multiformis simonsi Boulenger, 1902, by Burt and Burt, 1931), type locality “Oberer Pilcomayo, zwi- schen Tarija and S. Lrancisco, 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 sig- 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. 6 BREVIORA No. 494 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 7 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. 8 BREVIOIL4 No. 494 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, O — L. polystictus, sp. nov. • = L. robustus, sp. nov. S = Type of L. signifer. ® = Lectotype of L. multiformis. Liolaemus robustus, new species (PL I, Figs. 1-4; PL II, Fig. 3) Holotype. One male (FMNH 34242/H) from Junin, Depto. Junin, collected by K. P. Schmidt. Paratypes. PERU: Depto. Junin: Same data as holotype: FMNH 34242/1-23, 34247, 11 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 LIOLAEMVS 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- 19 dorsal scales (instead of 18-30) 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 internasals 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, 41 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 BREVIOR.4 No. 494 very faintly keeled or smooth, 14 in head length. Flank 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, 19 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 1 1 (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 (3 1 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 (41 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 Table 2. Meristic characters of Liolaemus robustus. (7 SS, 7 2$) <33 22 Scales around midbody Dorsal scales between occiput and levels of anterior border 48-61 (Jc = 53.05) 50-60 (Jc = 54.43) of thighs Ventral scales between post- 44-59 (Jc = 52.28) 51-56 (Jc = 53.14) mentals and vent Lateral scales between anterior 63-74 (Jc = 68.59) 72-78 (Jc = 74.14) and posterior limtjs 49-68 (Jc = 58.40) 56-68 (Jc = 62.14) Scales in the 1 5th verticil of tail 18-22 (Jc = 20.28) 19-23 (Jc = 20.86) Gular scales between ears 29-35 (Jc = 32.04) 28-32 (Jc = 30) Hellmich’s index 14-17 (Jc = 15.40) 13-15 (Jc = 14) Lamellae under 4th finger 14-17 (Jc = 15.63) 14-16 (Jc = 15) Lamellae under 4th toe 18-21 (Jc = 19.78) 19-21 (Jc = 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 = 111.47), from 109 to 126% (mean = 117.73) 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 BREVIOR.4 No. 494 T able 3 . Comparative variation of meristic and morphometric characters IN LlOLAEMUS ROBUSTUS, SP. NOV., L. POLYSTICTUS, 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 + 2 or 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 25-49 frontal region as % of length of 5 dorsal scales ( x = 16.9) (x = 31) (x = 23.8) (x = 35.72) Width of head in 19.6-24.6 19.70-23.7 18.3-22.3 % of snout-vent length (x = 21.68) (x = 21.25) (x= 19.98) Distance between 85.2-1 10.1 70.9-107.6 posterior borders of eyes in % of head height (x= 95.49) (x = 84.9) Length of 4th toe 20.5-38.6 31.6-50 nail in % of width of 5 ven- tral scales (<3<5 only) (x = 30.02) (x = 40.81) Distance between 80-124 62-99 the pubic sym- physis and the vent as % of ear- eye distance (22 only) (x = 98.84) (x = 78.65) 1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 13 Table 3. Continued. signifer robustus polvstictus williamsi (= multiformis) (N = 15) (N = 17) (N = 15) (N = 32) Minimum distance 90-142 56-87 65-139 between nasals (x = (Jc = 70.8) (Jc = 95.7) in % of mini- mum distance between supra- ocular scales 1 13.31) Rostral height in % 43-65 ■ 50-74 48-62 of eye-lip dis- tance (Jc = 50.57) (Jc = 60.62) (Jc = 55.85) Length of 5 dorsal 67-104 29-71 scales in % of ear-eye distance (jc = 79.4) (Jc = 49.58) Eye-lip distance in 47-70 37-58 % of subocular length (jc = 54.83) (Jc = 49.05) Length of 1 st finger 51-83 40-65 (without claw) in % of length of 5 dorsal scales (jc = 65.95) (Jc = 54.14) the norm for the entire genus Liolaemus, but the other features might characterize a valid 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 syn types 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 1 3%) 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). oo z 14 BREVIOR.4 No. 494 y'V © s- 0® JO - H ® ® ®e o* '■'"•0 « d <3 o * ® \ -sb>, • * ® Jr-. * ® ’ \ X X \ . 'v \ V. vJ' ® ®« ® % ® ® • ® ® o . o o _ o . <3 JSL o o *r Figure 2. Scatter-diagram of width of head (ordinates) and snout-vent length (abscissa). Measurements in tenths of a millimeter. • = Liolaemus robustus <5, and O = 2. x = L. signifer <3, and = 2. S = Type of L. signifer. 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- ern 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 ofHuancavelica, 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 161 157-58, 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- 70-1- 60 • .V X XX XX _ X X ••X X X X e • • X X ~Q0 100 120 X X X X 140 Figure 3 . Scatter-diagram of distance between posterior eye borders (ordinates) and head height (abscissa). Measurements in tenths of a millimeter. • = Liolaemus polyst ictus, 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. x 30- 25- x 20- X X X X X X 15-F 45 50 55 60 65 70 X X 75 80 ^5 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 BREVIORA No. 494 80- X • •• x# • •* 30 4 -3 T- 40 50 60 70 80 x x 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 1 3 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 Liolaemus polystictus, sp. nov. (9 66, 8 22, JUVENILES NOT INCLUDED). 66 22 Scales around midbody 57-70 (Jc = 61.34) 63-68 (Jc = 64.87) Dorsal scales between occiput and level of anterior border of thigh 55-63 (Jc = 59.22) 57-70 (Jc= 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 (x = 62.78) 53-76 (Jc = 63.75) Scales in 1 5th verticil of tail 1 7-20 (JC = 19.40) 19-22 (Jc = 20.87) Gular scales between ears 27-32 (Jc = 29.56) 28-32 (Jc = 29.88) Hellmich’s index 16-21 (jc = 17.78) 14-21 (Jc = 18.38) Lamellae under 4th finger 17-18 (Jc = 17.67) 15-19 (Jc = 16.25) Lamellae under 4th toe 20-24 (Jc = 21.56) 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, 4 1 between ear opening and front leg, 32 scales between ear openings, 63 scales around the body, 61 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 15th 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 BREVIOR.4 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 1 1 (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 X X X X X X X X X X • • • 20 X X X • # 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- 10 4 * 1 1 1 — 16 21 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 BREVIOR.4 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 para type 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 931 9— 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 BREVIOR.4 No. 494 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 1 8-29% 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 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 internasals. 26 BREVIOR.4 No. 494 Table 5. Meristic characters in Liolaemus williamsi, sp. nov. (1 1 68, 1 1 22, JUVENILES NOT INCLUDED). 66 22 Scales around midbody 54-64 (Jc = 59) 57-67 (Jc = 60.09) Dorsal scales 48-62 (Jc = 54.64) 49-65 (Jc = 57.82) Ventral scales 67-77 (Jc = 70.82) 71-78 (Jc = 74.55) Lateral scales 66-75 (Jc = 70.36) 66-77 (Jc = 72.36) Scales of tail’s 1 5th verticil 17-21 (Jc = 19.36) 18-22 (Jc = 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 internasals and the anterior frontal, 12 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 corner 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, 17 in the 15th 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 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 (21 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 B REV I ORA No. 494 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- U-f- 1 -| T 1 1 30 40 50 60 70 80 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 B RE VI ORA 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 l’holotype de Proctotretus signifer Dumeril et Bibron, 1837 (Reptilia, Iguanidae). Bulletin du Museum national d’histoire naturelle, 4C 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. Contribution 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 L. alticolor Barbour 1909 L. atacamensis Muller & Hellmich 1933 L. austromendocinus Cei 1974 signifer group L. andinus a. andinus Koslowsky 1895 a. poecilochromus Laurent 1986 L. aymararum Veloso, Sallaberry, 1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 31 APPENDIX. Continued. Subgenus Liolaemus L. bellii b. bellii Gray 1845 b. araucaniensis Muller & Hellmich 1932 b. moradoensis Hellmich 1950 b. neuquensis Muller & Hellmich 1938 L. bibronii Bell 1 843 L. bisignatus Philippi 1860 L. bitaeniatus Laurent 1984 L. buergeri Werner 1907 L. capil/itas Hulse 1979 L. ceii Donoso-Barros 1971 L. chiliensis Lesson 1828 L. coeruleus Cei & Ortiz 1983 L. constanzae Donoso-Barros 1961 L. copiapensis Muller & Hellmich 1933 L. curicensis Muller & Hellmich 1938 L. cur is Nunez & Labra 1985 L. cyanogaster c. cyanogaster Dumeril & Bibron 1837 c. brattstroemi Donoso-Barros 1961 L. donosoi Ortiz 1975 L. duellmani Cei 1978 L. elongatus e. elongatus Koslowsky 1896 e. petrophilus Donoso-Barros & Cei 1971 L. exploratorum Cei & Williams 1984 L. ftzgeraldi Boulenger 1899 L. fuscus Boulenger 1885 L. gracilis Bell 1843 L. gravenhorstii Gray 1845 L. hellmichi Donoso-Barros 1974 L. hernani Sallaberry, Nunez & Yanez 1982 L. kriegi Muller & Hellmich 1939 L. kuhlmanni Muller & Hellmich 1932 Subgenus Eulaemus signifer group Navarro, Iturra, Valencia, Penna & Diaz 1982 L. disjunctus Laurent 1990 L. dorbignyi Koslowsky 1898 L. eleodori Cei, Etheridge & Videla 1985 L. fabiani Yanez & Nunez 1983 L. famatinae Cei 1980 L. fittkaui Laurent 1986 L. forsteri Laurent 1982 L. griseus Laurent 1984 L. huacahuasicus Laurent 1985 L. islugensis Ortiz & Marquet 1987 L. jamesi Boulenger 1891 L. montanus Koslowsky 1898 L. multicolor Koslowsky 1898 L. nigriceps Philippi 1860 L. orientalis Muller 1923 L. ortizi Laurent 1982 L. polystictus Laurent 1991 L. puritamensis Nunez & Fox 1989 L. robustus Laurent 1991 L. ruibali Donoso-Barros 1961 L. schmidti Marx 1960 L. signifer s. signifer Dumeril & Bibron 1837 5. annectens Boulenger 1901 L. williamsi Laurent 1991 fit zinger ii group L. boulengeri Koslowsky 1898 L. chacoensis Shreve 1 948 L. darwinii Bell 1843 E. donosobarrosi Cei 1974 L. fitzingerii f. fitzingerii Dumeril & Bibron 1837 f. canqueli Cei 1973 L. irregularis Laurent 1986 L. melanops m. melanops Burmeister, 1888 32 BREVIORA No. 494 APPENDIX. Continued. Subgenus Liolaemus Subgenus Eulaemus L. lemniscatus Gravenhorst 1838 L. leopardinus l. leopardinus Muller & Hellmich 1932 /. ramonensis Muller & Hellmich 1932 /. valdesianus Hellmich 1950 L. lorenzmuelleri Hellmich 1950 L. monticola m. monticola Muller & Hellmich 1932 m. chiilanensis Muller & Hellmich 1932 m. villaricensis Muller & Hellmich 1932 L. nigromaculatus Wiegmann 1834 L. nigroviridis n. nigroviridis Muller & Hellmich 1932 n. campanae Hellmich 1950 n. minor Muller & Hellmich 1932 n. nigroroseus Donoso-Barros 1966 L. nitidus Wiegmann 1834 L. paulinae Donoso-Barros 1961 L. pictus p. pictus Dumeril & Bibron 1837 p. argentinus Muller & Hellmich 1939 p. chiloecnsis Muller & Hellmich 1939 p. major Boulenger 1885 p. talcanensis Urbana & Zungia 1977 L. platci Werner 1898 L. robertmertensi Hellmich 1964 L. sanjuanensis Cei 1982 L. schroederi Muller & Hellmich 1938 L. tacnae Shreve 1 94 1 L. tenuis t. tenuis Dumeril & Bibron 1837 t. punctatissimus Muller & Hell- mich 1933 m. xanthoviridis Cei & Scolaro 1980 L. ornatus Koslowsky 1898 L. rothi Koslowsky 1898 L. uspaliatensis Macola & Castro 1982 1992 OVERLOOKED LIOLAEMUS SPECIES FROM PERU 33 APPENDIX. Continued. Subgenus Liolaemus Subgenus Eulaemus L. variegatus Laurent 1984 L. velosoi Ortiz 1987 L. walked Shreve 1938 E. zapallarensis z. zapallarensis Muller & Hellmich 1933 z. ater Muller & Hellmich 1933 z. sieversi Donoso-Barros 1954 BREVIORA _ I JtjD P -A 1QQO Museum of Comparative Zoology US ISSN 0006-9698 U IN LV f- F? Q i ~r Au Cambridge, Mass. 31 January 1992 Numbee?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. Williams1 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- 1 Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138. 2 B REV I ORA No. 495 Figure 1. Anolis lamari, holotype. Dorsal view of head. 1992 ANOLIS LAMARI 3 Figure 2. Anolis lamari, holotype. Lateral view of head. 4 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, tigrinus, 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 5 I i Figure 4. Anolis lamari, holotype. Lateral view to show pattern. 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 LAMARI 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 from 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 Senor 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 of 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 ruizi from the eastern slopes of the Cordillera Oriental in the Departments of Boyaca and Casanare, and lamari, 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 tigrinus ) 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 16 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 6 153). 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. 80° 70° 60° Figure 6. Colombian distribution of the tigrinus superspecies. MACARENA marks the presumed locality of an undescribed member of the tigrinus superspe- cies. 1992 ANOLIS LAMAR1 1 1 The locality has previously been regarded as doubtful, because it was thought to be too distant from the other Colombian species of the tigrinus 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. tigrinus, 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 A. richardi. In continental South America I count 1 8 described 1992 ANOLIS LAMAR1 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 173111. D. A. 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, macro/epis, 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 BREVIOR.4 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 the 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, shep/ani, 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 LAMAR1 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 A. 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 of 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 species— except one, Anolis santamar- tae— 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 ANOLIS LAMAR I 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, 1 982 (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 so/itarius, 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 of santamartae, even if santamartae did have 1992 ANOLIS LAMAR1 19 Figure 1 2. 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 B RE VI ORA 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 of 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 LAMAR I 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. 13) 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 of 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 BREVIOHi 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 A. 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 if 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 LAMARl 23 it. It would then consist of all South American alpha 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 a/. (1988). I 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, and H. I. Rosenberg. 1988. Formal taxa, species groups, and perception of the genus Diplodacty/us (Reptilia: Gekkonidae). Zeitschrift fur zoologischen Systematik und Evolutionsforschung, 27: 44^18. Etheridge, 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., and E. E. Williams. 1986. Una nueva especie de saurio para la Cordillera Oriental de Colombia (Sauria; Iguania). Caldasia, 15(71-75): 51 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, 481^483. 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 Y I 0 R A Museum of Comparative Zoology US ISSN 0006-9698 Cambridge, Mass. 2 February 1994 Number 496 A NEW SPHAERODACTYLUS (SAURIA: GEKKONIDAE) FROM BEQUIA, GRENADA BANK, LESSER ANTILLES James Lazell1 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 of 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 “Sphaerodactylus 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. 11.3 ± 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, fiat, 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 vincenti, MCZ 79723, Cane Garden, St. Vincent. 4 BREVIORA No. 496 Figure 2. Type locality of Sphaerodactylus 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 0 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 Sphaerodactylus v. vincenti (and S. 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 BREVIORA No. 496 Paratypes. MCZ 175142, 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 BANK SPHAERODACTYLUS 7 reducing the light V on the tail base. The two females set extremes ol 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 differences 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 21 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 79716-19, 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 Mest) 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. Sphaerodactylus 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.” S. 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. mo/ei of Trinidad, Tobago, and the adjacent mainland of South America. I have examined 1 9 specimens of S. 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 S. 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. kirbyi does not fit into the elaborate pattern of geographic variation in S. vincenti 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. S. 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. V. 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 to josephinae 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— moni lifer. However, ocellate S. v. festus effectively 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. kirbyi agrees with S. 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- Table 1. Some diagnostic characters and variation in the Sphaerodactyl 10 BREVIOR4 No. 496 uj tu x u w &n u DC Oh H s o es CL. D O oC o P s 03 3 >H 05 1 03 2 O z o p < z 2 < u L/3 UJ H < Z + c/3 + UJ . r> U <* CC *-h UJ Q Z 1 5 Q UJ H d < O UJ H <■ 5 z O tt < Oh UJ UJ DC < C/3 < H UJ C/3 Q i UJ ffl < X y DC < X 52 X £ 0 C/3 UJ X X UJ h H U Z 2 < X 0 w (/3 o X H W H < U 5 z oi < u w -I (N d »— H CN Tf oo r- II > Q CL. P +1 cn +1 X- +1 in "hi 00 +1 ± 2 ± 1 ± 2 +1 +1 r\ + H in v * SO Q < d d Os 00 ^ ^ ^ d r- z U d d o _■ _ ; < on +1 +1 -hi -hi -hi H cn J < /■s m 00 +l +1 +1 cn -hi UJ 1/3 cC o Q Q d oo 6 o >n _ \ o X — i •— H — - Os d r-^ i—l y—t — H H 2c n cn 0$ UJ H s ' — 1 ^ — * c/f UJ £ hJ UJ < > O 0$ on o hJ * * H < c SO m (N * G s •S adamas festus pheristus i§ UJ H X UJ UJ U 0 H C3 H kirby •S 5 5 K 53 X Cl. CX cr i n — -- — in m m cn — rf o ri — • — | m m 00 1 ON 1 I tJ- m 1 — — — i d d — < — < — i T3 n — «■' -> — ■" s — •/ s — ' s — ' " — ^ s — x in NO — r- o o rr O & °o r .V ^ • ■ ^ o o ° o ’cPo ° o o 0 o ° ^ =°°° ° A v'. -1:- • • . • / • v. !•% • o o %.*• * • • - % • • / Narrower Head Wider Plastron t Wider Head Narrower Plastron 50 100 150 200 250 300 Carapace Length (mm) 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. ob/onga, 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 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 Figure 7. Lectotype of Chelodina novaeguineae (BMNH 1946.1.22.36, sub-adult female measuring 137 mm carapace length), from Katow, S.E. New Guinea [= Mawatta, Binaturi River, Western Province, Papua New Guinea]. 14 BREVIORA No. 497 ,82t .8 .78 .76 A C. novaeguincac ■ C. pritchardi a C. rcimanni j ■ ■ ■ ■ A A A A A A U' ^.72 U A A A A A A A A. ■ A A* A Broader Carapace A AA & A A A 1 . A A Aa .68 80 100 120 140 160 180 200 220 240 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 > Fern > 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 100 MM. C. NO VAE G UINE.AE INCLUDES only New Guinean specimens, no Australian ones. Feature C. longicollis (n = 37) C. pritchardi (n = 43) C. novae- guineae (n = 51) C. reimanni (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. pritchardi. 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 1 5 1 mm. The largest specimen of New Guinean C. novaeguineae I ex- 1994 NEW CHELODINA FROM PAPUA NEW GUINEA 17 Figure 1 0. Dorsal, ventral, and lateral views of skull of Chelodina novaeguineae (AGJR-T 504, 178 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 BREVIORA 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, 15 C. novaeguineae (12 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 .24r 22 S£ S -4 o C. longicollis a C. novaeguincac ■ C. prilchardi * C. reimanni & 3 C/2 18 is .16 wc ss 1 /l-l U .14 - H .12 A Broader Triturating Surface O O O O O O 80 100 120 140 160 180 200 Carapace Length (mm) 220 240 Figure 12. 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. reimanni 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 horny 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 B RE VI ORA No. 497 I80r 160 140 X 120 o> C — Z 80 c/5 3 X! 5 60 100 40 20 80 o C. longicollis a C. novaeguineae ■ C. prilchardi A C. reimanni More Robust Skull O 100 A A A O oo o o 120 140 160 180 200 220 240 Carapace Length (mm) 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 CHELOD1NA FROM PAPUA NEW GUINEA 21 ,22t -^1 OX) c J 18 = .16 £ .14 j£ .12 OX C/3 • * c/) CO S .08 o C. longicollis a C. novaeguineae ■ C. pritchardi a C. reimanni Grcalcr Relative Mass Skull plus Mandible j* .06 o .04 cs “ .02 o4— 80 A A A O O o O O o 100 120 140 160 180 200 Carapace Length (mm) 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. pritchardi 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. longicollis and C. pritchardi slightly more angular. The premaxillae are usually fused into a single small premaxilla in New Guinean C. novaeguineae ( 1 1 of 12 skulls) and C. reimanni (2 of 2); unfused but very small in Australian C. novaeguineae (3 of 3), and unfused and large in C. longicollis 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. longicollis 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. longicollis 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; IOW = 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 12 AND 13 FOR ANALYSIS OF SKULL MEASUREMENT RATIOS. Species Mus. Sp. No. SL SWT SWM SDM SD IOW 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 novaeguineae 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 1 17939 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 1 1.4 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 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. Feature C. longicollis (n = 7) C. pritchardi (n = 6) C. novae- guineae (n = 15) C. reimanni (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 IOW/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. longicollis, 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, 1983; 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 and 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. dt _ TW x SWT x SD RI~ 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 CHELOD1NA 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 (1950). Atlanto-axial (Cl 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 BREVIOR.4 No. 497 30 28 G E 26 2 24- 22- -o £ 60 60 w u 00 CO L. u > < 20 18 16 . I . 1 1 siebcnrocki ^ "expansa" A A A parked A rugosa expansa A oblonga "longicollis" pritchardi Q W) novaeguineae longicollis o sleindachneri 1 1 26 28 30 32 34 36 Average Egg Length (mm) 38 40 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. 16). 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 .95 .9 eg u, ■£ .8 bo c o -4 .75 s 4= T3 .7 £ .6 .55' 26 28 30 32 34 36 38 40 Average Egg Length (mm) 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 al., 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- A parked A siebenrocki pritchardi o y\rugosa steindachneri longicollis ^ O O^"longicollis” oblonga ’’expans a” A A ex pans a novaeguineae 28 B REV I ORA No. 497 120H J= u 3 u x u -a c — u N M 60 w 1 10- O 100- 60 > < * 60 > < 90- 80- 70- 60- 50- ■ novaegumeae • pritchardi A rugosa ▼ siebcnrocki □ parked O steindachneri 3 oblonga ▼ T 3 □ O O o 40-1 140 160 180 200 220 240 260 Carapace Length (mm) of Female 280 300 Figure 1 7. 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- Table 5. Average length and width of eggs for various species of Chelodina. CLa/qx = maximum carapace length (mm). Values: mean ± standard deviation (range) (mm). Chelodina “ longicollis" and “ expansa ” (in quotations) are THE SUBGENERIC GROUPINGS OF LEGLER, WHERE HE COMBINES DATA FROM SEVERAL DIFFERENT SPECIES. 1994 NEW CHELODINA FROM PAPUA NEW GUINEA NO o- On O C > TD 3 C/5 C CU hJ -C TD £ NO o N o ,, N r- o o r- SO o* q OO ON q 00 O n q q q q 00 in r- rsj d q 00 OO OX q O' q r-’ q (N o OO ON — H OO OX SO OX OX H SO OX^ o> ■F— < OO — 4 ON *— * rs) d 1 *— » ox s— ^ W' — 1 T— < (N ^ ' .g OD C w/ m w' m s— ^ (^X m ^ ^ o ^ ^ O 3" ^ S m ^ ^ 00 S V m q q Os Os q 00 00 OO d cn so d o rn d (N * ^3 5* q So to s: C/D O p S o 1 ? c to C5. o * to u u u U u U U u U 0 u u 30 BREVIOR.4 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 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 Figure 1 9. Alternative phylogenetic relationships of Chelodina subgeneric group “A”. See Figure 1 1 for preferred hypothesis. 32 BREVIOR.4 No. 497 derived features between C. pritchardi and C. longicollis, and is encumbered by the less parsimonious double loss of chelid fo- ramina in the novaeguineae-reimanni lineage and the pritchardi- longicollis 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 of 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 CHELODINA FROM PAPUA NEW GUINEA 33 Crumly, John Iverson, John Carr, Russ Mittermeier, and Ernest Williams. 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: New South Wales: Armidale (nr.), UNLV s/n ( 1 5); Sydney, AMNH 14151; Sydney, 60 mi S, UMMZ 130161, 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, 1 10483, 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; Ali Vil- lage, Aramia River (8°05'S, 142°55'E), USNM 213490; Boze, Binaturi River (9°05'S, 143°01'E), AGJR-T 504; Daru Roads (9°03'S, 143°12'E), MCZ 142500; Emeti, Bamu River (7°48'S, 143°15'E), MCZ 138102; Fly River at Strickland River junction (7°35'S, 141°25'E), MCZ 53758-61; Giringarede, Binaturi River (9°03'S, 1 42°57'E), MCZ 1 53930; Katatai (9°0 1 'S, 1 43° 1 8'E), MCZ 138101,1 42495, 1 54340; 34 BREVIORA No. 497 Katow (= Mawatta, Binaturi River) (9°08'S, 142°55'E), BMNH 1946.1.22.36 (lec- totype C. novae guineae): 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, 141°30'E), MCZ 134392; Mabaduane, Pahoturi River (9°1 7'S, 142°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; Onomo, Oriomo River (8°52'S, 143°10'E), PNGM 23510; Peawa, On- omo River (8°55'S, 143°12'E), AMNH 104010; Tarara, Wassikussa River (8°50'S, 141°52'E), AMNH 58410; Togo, Pahotun 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 pritchardi: Papua New Guinea: Central Province: Bore, Kemp Welch River (9°53'S, 147°46'E), MCZ 1 73543; 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 Jaya: Merauke (8°25'S, 140°28'E), AGJR-T 746, 1299-1300, 1325, 1614-1619, 1642; No data: MTKD 14603. Chelodina steindachneri: Australia: Western Australia: Marloo Station, MCZ 33501; Mundabullangana, 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 CHELODINA 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 scripta). 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 Emydura (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. Eine 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(1 1): 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 BREVIOK4 No. 497 Shaw, G. 1794. Zoology of New Holland. Vol. I. London, J. Davis. 33 pp. Siebenrock, F. 1914. Eine 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. 1901. Ueber Reptilien und Batrachier aus Ecuador und Neu-Guinea. Verhandlungen der Zoologisch-Botanischen Gesellschaft Wien, 51: 593-603. Williams, E. E. 1950. Variation and selection in the cervical central articulations of living turtles. Bulletin of the American Museum of Natural History, 94: 510-561. BREVIOR A Museum of Comparative Zoology US ISSN 0006-9698 Cambridge, Mass. 2 February 1994 Number 498 CHELID TURTLES OF THE AUSTRALASIAN ARCHIPELAGO: II. A NEW SPECIES OF CHELODINA FROM ROTI ISLAND, INDONESIA Anders G. J. Rhodin1 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 pritchardi 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, Chelodina reimanni 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, 1 Museum of Comparative Zoology, Harvard University, Cambridge, Massachu- setts, and Chelonian Research Foundation, Lunenburg, Massachusetts. 2 BREVIOR.4 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. pritchardi 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 held 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 BREVIOFL4 No. 498 the specimens and relinquishing claims to the formal description. These 18 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 from northern Australia, 54 C. longicollis 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 (partim) Ten Kate, 1894:689 Chelodina novae-guineae (partim) Lidth de Jeude, 1895:1 19; De Rooij, 1915:315 Holotype. MCZ 176730 (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 AG JR 450 from the personal preserved collection of Rhodin. Figure 2. Holotype of Chelodina mccordi (MCZ 176730), adult female mea- suring 197.5 mm carapace length, from Roti Island, Indonesia. 1994 NEW CHELODINA FROM ROT1 5 6 BREVIORA 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 194 mm carapace length, pur- chased from native villagers by Frank Yowono in Kupang, west- ern 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 Boulenger, 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 Boulenger, 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 7 8 BREVIORA 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 km2, 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 9 10 BREVIOR.A 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 southwestern shore, 4 km south of Tudameda] to “Baa” [= modern Ba’a along the northern shore]. The locality “Danau Naloek” does not appear on modern 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, southwestern 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 al., 1974) with relatively wide head, wide 1994 NEW CHELOD1NA FROM ROT1 1 1 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 novaeguineae 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 from 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 from 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 BREVIORA No. 498 1794), Chelodina novaeguineae Boulenger, 1888, Chelodina pritchardi Rhodin, 1993, Chelodina reimanni Philippen and Grossman, 1990, and Chelodina steindachneri Siebenrock, 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 (1895) interpreted this as a normal pattern. One of Yowono’s specimens 1994 NEW CHELOD1NA 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 = unkjvown 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 1 17.5 124.5 67.0 26.3 AGJR 454 m 151.0 1 16.5 47.0 1 15.5 123.5 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 -Cl scute border. Ml typically half as wide as M2. In C. pritchardi 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, Head Width / Carapace Length Carapace Width / Carapace Length .8751 .85 .825- .8- .775- .75 .725 .7 .675 .65 .625 O O O O O O O O O O O O 0° O o o O O ■ o Broader Carapace ° o o o° ■ . A A ^3 ■ ■ ■ ■ O O ■ A A A ° 3 A <1 ■4 < O A cA A A AA o \ o o o O O o AO ▲ aA aAAA $ A ^ AaoA ^ A ^ A A A O O A o C. longicollis ® C. mccordi A C. novaeguineae ■ C. pritchardi A C. reimanni 40 60 .221 — - — m 80 100 120 140 160 180 200 220 240 .21- .2- .19- .18 .17- .16 .15 .14- .13 .12 B A O O A a ^ o o o o A a A © o ° °@ o o Wider Head o o C. longjcollis o C. mccordi A C. novaeguineae ■ C. pritchardi A C. reimanni 3 a 3 o o £>■ ■ o o o 40 60 80 100 120 140 160 180 200 220 Carapace Length (mm) 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). Composite Ratio (CW x HW x 1/PW) / CL Plastron Width / Carapace Length ,6r .575 .^5 .525 .5 .475 .45 .425 O O O O O O Broader Plastron o o o o °° ® ° o%g „§ o OO O o °C o o o o o o o o o 2 A A A m<* ■ a‘ £ A » 9 A^3^»AAa ^ A ® » AA r, A # A .375 .35+ O C. longicollis o C. mccordi A C. novaeguineae ■ C. pritchardi A C. reimanni 40 60 80 100 120 140 160 180 200 220 24C I) .38 .36 .34 .32 .3 .28 .26 .24 29 O O A 3 O O o O ° A ° i o o Broader Carapace Wider Head Narrower Plastron O O o o, § * 3 ® A A A A A Ad a jj v_ °o fY* o C. longicollis o C. mccordi A C. novaeguineae ■ C. pritchardi A C. reimanni cPo ° ° o o ° o 3 2 3 ® ® 3 A O O O O C P . 18J — 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 BREVIOR.4 No. 498 l.4r -a r? "3 c *5b U cs ■c — -a £ 1.2 .8 .6 CS C ’5b £ -41 o C. mccordi A C. novaeguineae ■ C. pritchardi Wider Marginal Number 1 Narrower 3 A A A A A A & 3 3 O 3 3 3 3 3 3 3 An > Abd > Fern > Pec ^ Gul. C. pritchardi usually with Pec > Fern, C. novaeguineae with either Pec > Fern or Fern > 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 of 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 B RE VI OK4 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. PRITCHARD1 N = 17 AND C. NOVAEGUINEAE N = 19. Ratio C. pritchardi (n = 43) C. mccordi (n = 17) C. novaeguineae (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 162.5 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 218 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, 7 C. longicollis, 1 5 C. novaeguineae ( 1 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; IOW = 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 IOW OW PtW TW AG JR 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. Ratio C. pritchardi (n = 6) C. mccordi (n = 4) C. novaeguineae (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 IOW/OW .470 ± .037 .503 ± .056 .570 ± .055 Skull Depth / Skull Length Triturating Width / Skull Length 20 BREVIOR.4 No. 498 .22 7 A 18 .16 .14 Broader i Triturating Surface O C. longicollis ® C. mccordi A C. novaeguineac ■ C. pritchardi a C. reimanni A A O A A A A A A A A A A 3 3 3 3 .12 O O O O '80 O 100 ,42r B 120 140 160 180 200 220 240 .38 .36 o C. longicollis o C. mccordi A C. novaeguineac ■ C. pritchardi a C. reimanni .34 .32 .3 Deeper Skull A A 3 .28 .26 O O O O 3 O O 3 3 .24* '80 100 120 140 160 180 200 Carapace Length (mm) 220 240 Figure 8. (Part 1). Skull morphometries. Graphs plotting morphometric vari- ation for four species of Chelodina subgeneric group “A”, showing the relationships of: A. maxillary triturating width ratio (TW/Skull Length); B. skull depth ratio (SD/SL). Robusticity Index (TW x SW x SD) / SL Tympanic Skull Width / Skull Length ,85f .825 .8 .775 .75 .725 Wider Skull O C. longicollis o C. mccordi A C. novaeguineae ■ C. pritchardi * C. reiinanni .675 .65 O A O O A a ch o 3 3 .625* — 80 1801 — 100 120 140 160 180 200 220 240 D 160 140 More Robust Skull 120 100 o C. longicollis ® C. mccordi A C. novaeguineae ■ C. pritchardi a C. reimanni 80 60 40 20 O O O O A A aa A 3 m O O 3 3 3 80 100 120 140 160 180 200 220 Carapace Length (mm) 240 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. 1994 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- neae and 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 of Chelodina 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 porosus 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 turtle Amy da 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 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 0 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 of 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 1 89 1 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 CHELODINA 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 Illinoisan-Riss glaciation, about 170,000 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. longicollis, C. pritch- ardi, and C. novaeguineae to C reimanni, the most derived mem- 28 BREVIOR.4 No. 498 reimanni novaeguineae mccordi pritchardl longicollis steindachneri Figure 10. Hypothesized phylogenetic relationships of the six currently rec- ognized species of Chelodina subgeneric group “A”. The monophyly of Chelodina “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 phytogeny, 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 of Chelodina 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 modern 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 longicollis, C. novaeguineae, C. pritchardi, C. reimanni, and C. steindachneri all 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 Aoki, 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: 449-452. . 1 889. 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. Kjrsch, 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. Thorne (eds.). 30 BREVIORA No. 498 The Origin of the Australians. Canberra, Australian Institute of Aboriginal Studies. Cox, C. B. 1970. Migrating marsupials and drifting continents. Nature, 226: 767-770. De Roou, N. 1915. The Reptiles of the Indo-Australian Archipelago. I. Lac- ertilia, Chelonia, Emydosauria. Leiden, E. J. Brill. 334 pp. Doutch, H. F. 1972. 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: 1 1 1-1 16. 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. In 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 ( Trachemvs scripta). 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. 154 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. Eine 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. 1994. 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 ROT1 31 siebenrocki Werner, 1901. Bulletin of the Museum of Comparative Zoology, 147(11): 465-488. 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 IUCN convened at Caracas, Venezuela, 21 to 28 October 1984. IUCN Publ. NS. Shaw, G. 1794. Zoology of New Holland. Vol. I. London, J. Davis. 33 pp. Siebenrock, F. 1914. Eine 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. 1950. 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 Museum of Comparative Zoology US ISSN 0006-9698 Cambridge, Mass. 2 February 1994 Number 499 AN ECOLOGICAL STUDY OF THE ENDEMIC HISPANIOLAN ANOLINE LIZARD, CHAMAELINOROPS BARBOURI (LACERTILIA: IGUANIDAE) Glenn Flores12, 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 1919 by K. P. Schmidt, the anoline lizard 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 1 Present Address: Robert Wood Johnson Clinical Scholars Program, Yale Uni- versity School of Medicine, IE-6 1 SHM, P.O. Box 3333, New Haven, Connecticut 06510-8025. 2 To whom reprint requests should be addressed. 2 BREVIORA No. 499 are the relationships of Chamaelinorops-, it has been argued that the genus is either very derived (Etheridge, 1960; Wyles and Gor- man, 1980), 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 Chamaelinorops 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 15 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. 1994 ECOLOGY OF CHAMAELINOROPS BARBOURI 3 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 Cana 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 (1,140 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 Cana 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 0 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 (t- 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 1994 ECOLOGY OF CHAMAELINOROPS BARBOVRI 5 T able 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 0 1 2 3 4 5 Males 35 1 1 Females 1 25 1 1 Juveniles 2 1 Unsexed adults 2 Totals 1 64 3 1 1 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 6 B REV 1 ORA No. 499 Table 2. Types of terrestrial perches chosen by Chamaelinorops BARBOURI. 1 On rocks Under rocks On dead leaves Under dead leaves Bare 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 ( < V2" in diameter). (See Materials and Methods Section in text for addi- tional EXPLANATION OF PERCH DIAMETER CATEGORIES.) Ground covered Under- ground Rock by leaf 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 7 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 of Anolis 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. singu/aris (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, 1994 ECOLOGY OF CHAMAELINOROPS BARBOURI 9 Table 4. Thermal biology data for Chamaelinorops barbouri. 1 TB = body temperature; Ta = air temperature; N = number of individuals observed; X ± SD = MEAN PLUS OR MINUS ONE STANDARD DEVIATION; Rs = SPEARMAN’S COEFFICIENT OF RANK CORRELATION OF BODY AND AIR TEMPERATURES. N tb ta x ± SD Range x ± 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. singulars 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, vTB - 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 of 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 o t Anolis. Hatched lines represent shaded habitat. 10 B REV I ORA No. 499 Figure 3. A plot of the body temperature and air temperature data for Cha- maelinorops barbouri. The solid line represents the isothermal line (body tem- perature = air temperature, TB = TA). The dashed line is the linear regression ol body temperature and air temperature, for which the equation is provided. The correlation coefficient (rs) = 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 of Chamaelinorops. We often attempted to observe individuals in the held, 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 familiar with Chamaelinorops escape behavior, which usually consists of a very brief scampering dash or series of hops (of no more than several 1994 ECOLOGY OF CHAMAELINOROPS BARBOURI 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 B REV 1 ORA No. 499 Table 5. Chamaelinorops barbouri and Anolis humilis compared. Data FOR A. HUMILIS IS FROM Fitch (1973, 1975). Feature Chamaelinorops barbouri Anolis humilis Snout-vent length Usually <50 mm <45 mm Color Light brown/tan with variable pattern Dark brown/olive with variable pattern Dewlap Small, black with yellow edge Large, red with yellow edge Modal perch On ground, mostly leaf litter, marked shade preference Often leaf litter, but most- ly above ground on but- tressed roots of large trees; marked shade preference Body proportions Head relatively short, body compressed, long limbs, very long tail Stout body, short limbs and tail Scales Middorsal scales greatly enlarged, keeled, 4-10 rows Middorsal scales greatly enlarged, keeled, 8-12 rows Foraging behavior Sit-and-wait predator (?) on ground Active forager on ground and buttressed roots of large trees Defensive behavior Primarily crypsis, via im- mobility, start and stop flight; always flees on ground, never climbs 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 of basking behavior (Fitch, 1973, 1975). However, the two anoles differ markedly in a number of other features which directly reflect the greater specialization of 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 of Chamaelinorops appear to be rigidly 1994 ECOLOGY OF CHAMAELINOROPS BARBOURI 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 BREVIOH4 No. 499 column of Chamaelinorops, 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- norops 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 Chamaelinorops in response to bird-like whistles and clicks, suggesting Chamaelinorops 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 parvifrons and very large centipedes common under rocks. It is clear from our discussion of Chamaelinorops 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. Chamaelinorops 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 Chamaelinorops resulted in indi- viduals moving out of sun patches too quickly for us to note basking. However, although we actively searched for Chamaeli- norops 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 Chamaelinorops, 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 Chamaelinorops, constitutes an insignificant proportion of this lizard’s daily activities. 1994 ECOLOGY OF CHA MA ELINOR OPS BARBOUR1 17 Given that Chamaelinorops 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 Chamaelinorops 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. Chamaelinorops occurs exclusively in closed forests where patch- es of sun are few and far between; furthermore, Chamaelinorops shows a predominant preference for shade within such forest habitat (Fig. 1). Hence, the distance necessary for shuttling be- tween sun and shade in Chamaelinorops 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, Chamaelinorops, 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 Chamaelinorops 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), Chamaelinorops clearly maintains a body temperature well above air temperature, with the value of xTB - xTA 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 Chamaelinorops 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. 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C/3 o cj o 0 CJ 1 o O 0 5 * <5> 5o - — o q — — n m n — — x ^ in ^ _ tN - M U r* rs ,.m i'i O' UJ UJ u u ri — . — o rNt in n A in n 'T 00 po ff| O' ^ ^ ^ E ^ - IN (N -3 -J CQ CQ CQ wtu JXII U U W od od od 2 n p> ri o rs — — ; r*s vs ^ — — - r*s m m m vn vs — — — — *o n oo 'i — oo 0* r m m »o o J - - n O' >o UJ UJ U CJ — O' rs — ! S? r~~ oo -J X ui w u 3 £ z z ^ W-, £ VS VS -t rs in /, CQ CQ vs 5 5 £ oi o; t (n f- . “ n^K M h- >T) - ^ 3 . cn rs cs tt ”5 c-« P P P g? S S S: - _J J J j CD ffl ^ So u u u ni I I11 i-~ CJ O U CJ * * Z J — T r-T oT rn e* »©* * ^ O' — fN - — © — n m (A >2 <0 in - - [2 QjJ r«S - ws r- — » ® S S 2S oo E ^ S r- ro f- r- t=: 22 S ou — i — : cq -J-JjXcamcQ UlUJ-T- UIUJmUJIIt UUJ CJ (J o * * § oc 'O q 5 3 O'. - n n ^ n 7 ^ M 00 00 00 ^ ^ i/, J J J J 03 Cfl £ UJ UJ uj UJ X X x UUUUcda:^ m oo q rs rn r+t £ £ £ -J -J CO UJ UJ X U U o' r-' q' vs O n ^ r-4 rr r*s — , r-4 VS — E — r-4 rs 0 J JO UJ UJ rS U U 'O r-* o? Q M M rS m -T vs wS rR — — 04 OO 00 - s s s -3 J Is UJ UJ _J U U uj . . CJ 'O o n 'O n m r>i m — — r*-| ^ m - — w 2 O 5d w w 5 uu“ rrr-oo u u - o ^ Soo ^ oo 0 — or o m — — • fo n i *0 m On NO O' — ON — r^ — — — m — m r4 m t*-, - - - - n n r. in in ri - ■ ri oo - r-< c-4 r*', c^\ c*} m /i in it on U < < < < cn 0) 8. on 5 3 ■6 <3 C 1 C 3> T5 _ 3 S' I I a s. I *0 o 2 <0 3 § ! J5 3 3 I 5 u ^ 1 ^ -3 S 3 3 S. I I 3 51324, KEH8485, 112356, RHB2537, 112369, RHB2540, 112370, RHB2541 12357, RHB2542 Appendix 2. Continued. 38 BREVIORA No. 507 0) J3 c 3 c 2 0) Cm -o 0) -O £ 3 3 00 o & a o N U 2 oo m 00 *^I S fN (N -3 CO CD 0 ad ad 1 ¥ s' n M M r- cs c* J CQ CQ UJ X X U ad ad */■> Q O n ^ ^ 00 *5 ^ D rs n £2 CQ LQ X X CJ * c* "'T oT (N rO rO On /-S r* 2 -T g 00 h- - on x! -3 CQ CQ S UJ X X x U ad ad £ a s rs r-> *^n — • — d n 'O m -^r -o ^ 1 f h 5 ?S T J J J JQ-- — UJUJLQUJ — — — — — — — CJ CJ CJ CJ ^ ^ qq* fk” V oT V . . . . 2 — rs ro ■? ^ vn o> n n - q O' O' •/“>«*> •/‘i */“> O-. '-O ’C ^ — _ — — r^r>irsr^ -M(N^-fflCQCDa3CQCnfl2 SSnS^pI^JMsS KOOOOOCC^^^'V’T^^ r^r-D-r^^^ — -— — — — — -J-3_3_)^XXXXXXX LUtUUJUjy — CJ CJ CJ CJ O , ^,* V o* rs oo oio^’t5555555 o — — — — — oodadadadadadad — ad 3 00 u CO o o 8. CO s o B o B. <0 0 1 55 I *0 3 *0 to 2 y C\ 5 S. c !3 -J C B -0 Appendix 2. Continued. 39 1999 AZORES DEEP-SEA FISHES 0) _o 3 c 2 0) TJ « v/~> m 'O O o 7 - rs ri *t O O' O vr, — — — (N (N fN CD CQ CQ CD 23 CQ X X X X X X cd. ccL ceL ad cZ « cf — V o'" V oo fN o O U OO OO O' O ■ _J UJ o r- r-~ r~ r- r~- o — r-~ 2 2 2 2 2 2 o ^ o m — 00 o 5 x u 1 CD g g 4 i 5 5 o; oo oo ^ $ Q 3 $ 3 w. S 2 § 2 § m _ ov ^S5 —3 r- uj j U uj .U © • t m O ZL tt r-< 22? — — rr x — — — I o >Zi 00 00 Z oC v“T c/J — - rs vn m v") rr _ O — — «/“l (O O 2 00 oo ^ r- oo -i jq ri] m J u 3 “ r-i c-i . S? 3 - - 2 » $ 2 2^-2 2 ^ O a 2 2 § - CO CO “ sf * V> O V-) oor'nhv-l-^-'ft’TO'(Z^'iZrf: « co M 9, io o ,* rs^rr Z] — r-i ?S *«t -n- r>* 5 XOCMMgOOO'^^^^ x O ^ £ 00 OO S O o O' S r-r-r-r-oo — — — rHrsrsil^ r- — ^ 2° r- u- ^ £ jjjjTODcQaicnacDS j oa S -j -J ^ S ? Z 7 WUJWWjillllllT U1 X t m UJ UJ ^ 5 5 S UCJCJO^aCa'a£aC:rCacDa' U aC ^ 'O t r-~ — „ ,75 «o vr> vr> 'T'vrvvrv'rvv^ooooooo vr, o O vr, m v-) o O O O C A u < < ? T r- C9 c/) 22 *0 8. CO 1 o s» «3 Jk J5 .2 •*» .2 -5 o -J a I » s s §■ 1 Appendix 2. Continued. 40 BREVIORA No. 507 T 3 lA ^ fN >0 U = = = 3 8 S vi v ^ © o o v> o Q oo — X CD CD WIX U cC aL 8 8 - — . m — O on m — o o 00 OO M O ^ ^ r^- i — m oh J J j CD CQ CD W WWIXX U CJ CJ od od ad o cn* vo" — * On* m* 3 /, fN O U s5 = 3 3 3 £ 8 3 V v vr, © o © vn J CD CD UJ X X CJ at ad SIN 'O C4 CN — i o o s » s $ jDaS UJ X X 03 CJ at at X - . at O' — ^ « 1^1 N M ^ — © © $£ -33^ v-> © O m — — — o M fN U 5 — n n ij 00 00 00 o >o r- 1^ - in rs fo J J j CQ CQ CQ 3 WWujixXt UUUqCoCoC^ D'* Q* 3T* O'* t"* — * vf rr«A)w-i — 'or^r-' — — — oo 0 o o — — — TT Vi V V v-1 VA) o O O O rs O. v, r* So m tt 00 J ^ /i r-~ 00 1 _I X CD S UJ UJ X x CJ U at ^ 00* fN* TT* CN* V 'O m O O — — NT '•T Vi v, © o N N rs CD CD CD XXX ad ad ad Q K O' On V) C-4 03 CD CD XXX ad 0 d ad '■o oT oT s s ; . V — 00 a — in , J O' Os Vi , 00 — — CD U CD M -3 X X X CD “ o' ai Qd X <->... Cud - « 5° C* g m — O O © v, _ _ _ o ' * 3 O 3 N O Ov V 3 ^ i/-' V m bo w ^ ^ ~ j “ “ 5 a a W X X I T U ad ad ad £ X ad r- n n n 00 O' O' -3 CQ 2 w x S U ad 00 00 $ g 2f ~ n 3 ^ o o 00 ““ — — r* m ■ “ V* NO 2 ^ 3 IN £ 2 2 « j C3 Z J UJ x 5 £ 3 * 1/5 § Sftfgrf x « S S3 - 2 2 2 § I S ON -3 CD UJ X U ad _ r*-> m m v, © 2 g V o © o o o K S § » ” ” s ^ _j _j co x UJ UJ x u CJ CJ ot 5j no* O'* Vi* m* VO VO — Q — — 3 m — —mm v m o o < hJ < < c n U O' ^r — 00 _ r- C/3 .S£ o 8. C/3 C 1 S I O O) §• U 0 1 <«i § js - a *^3 •2 ^ a so J g i 5 Ci, .0 2 5 ^ a a ■c g „a IS O' 1 £ ■'T r+i TT VG T tt" 00 00 CH rn 'O «/"T w-i 'O V~l TT r*1 -J -3 < < hJ < < < < < r- o o VO . Tj- oo oo "T a\ r» On -1 LU u CO X at 3 4J 1 o. < a « 2 4> -O £ 3 e 00 _o B a o N O LU o 2 I 33 — u vT «"> On -T- 00 TT <-»■> -J LU CJ $ - ^ rs £0 - -J — r-* LU CO CO U X X . at at '© s Ov 5! ^ X -v 5 z — h "g © X , a 9 — r- O V .<-> p 22 2f £ CO x at CO i-% «■£ < X Q O u h-* 8 £ J LU u CO X at —vi r* — << J < * *3 ‘5k c cn s- cn cj ■S * 1 -5 72 0* CT3 V o 2 *u -3 U « a -o -n a. o 5 o «« 3 c a »■* •3 :a •— w 2 .2 c« u E o 3 <« <- &> •2 c E & L* •w w ■a o p u C3 e* U o> CJ o o> r-^> H CJ 7 v i o M ui s 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 1 8 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 \2 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 lost 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: F2M = 10.8, P = 0.002). Elevation explained most of this variance (sim- ple r = 0.76, partial r2 = 0.35), whereas island area contributed very little to the model (simple r — 0.51, partial r2 = 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 (r2 = 0.51, F311 = 3.79, P = 0.04). In this model, island area (simple r = 0.22, partial r 2 = 0.25) and dis- tance to a potential source population (simple r = -0.007, partial r2 = 0.21) explained most of the variance, whereas the contri- bution of elevation was small (simple r = 0.50, partial r2 = 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- 8 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 shortest). Gorda (WHICHEVER DISTANCE WAS Island No. of Species Area Elevation (km2) (m) Distance to Potential Source 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 0 429 177 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 0 138 155 6.8 (Tortola) Great Thatch 2 123 187 0.7 (Tortola) Scrub 0 97 141 3.7 (Tortola) Great Tobago 0 87 147 4.0 (Jost Van Dyke) Mosquito 0 50 95 17.7 (Virgin Gorda) Great Dog 1 33 89 11.1 (Virgin Gorda) Necker 0 30 32 22.0 (Virgin Gorda) Frenchmans Cay 2 24 131 0. 1 (Tortola) Little Thatch 0 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. albilabris in a riverbed by Old Hill west of White Bay and in a marshy site in the town of Great Harbour (MCZ 110990-1). On Anegada, we found several concentrations of L. albilabris 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. albilabris 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 schwartzi We located E. schwartzi on six of the 1 7 islands visited: Virgin Gorda, Great Dog, Beef, Frenchmans Cay, Tortola, and Jost Van LEPTODACTYLID FROGS IN THE BRITISH VIRGIN ISLANDS. 10 BREVIORA No. 508 x o u. on Q X O U uj X D Z < J on £ PJ Z Q UJ 2 Di o u Q UJ 2 x E z o u UJ (N UJ _J m < H C/3 -o u o o CJ ^ £ 0) O n- o' O C T3 ■— o U 3 cd cd PJ cd PJ cd PJ cd" _l PJ TO c cd t-c cd 2 _E U 00 CL a C/3 cd C/3 PJ . T3 (N oo 00 Q\ ON — - C c O cd £ 2 S 2 = C (U 2 I cd J o PJ \r. PJ cd" PJ m ON ON ON ON o nJ C/3 On CJ i i m r- > u 3 O o 00 CJ o o 00 -*-> CJ 4— > CJ CJ o CJ o CJ o CJ o CJ o u o 4—* CJ o 20 i m i r7_ (N 02 02 GO (N 1 (N 1 — 1 1 7 o o o m a o a H 1999 BRITISH VIRGIN ISLANDS’ LEPTODACTYLID FROGS 1 1 Q UJ D Z H Z o U (N W CQ < H C/3 _ c/3 TO J-H o o o ^ 1) W „ ^ 00 -*! C 2° 3 c/5 Os T? <+-. ^ 3 O __T ^ _c "aj O N 03 03 C c3 n r- ON C/3 3 a £ J O x3 3 O C/3 U- (D "3 3 in -*— > o O o O Os Os ON O o 3 c n to C/3 o C/1 V o «. s< a Cj >3 c/T d d cj d) £• 3 Cj "§ o II 3 "S 03’ CjJ PJ ^ 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 anti liana, located near the peak of the ridge that extends along the length of the island. In addition, on the night of 10-1 1 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 of E. 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). Eleutherodactylus 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: F348 = 1.67, P = 0.19; Tortola: F2A9 = 3.07, P = 0.06; Virgin Gorda: F, 57 = 0.18, P = 0.67). Similarly, there were no significant differences in SVL of E. schwartzi among years (Tortola: FU6 = 0.01, P = 0.90; Virgin Gorda: F236 = 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 (F2 14) = 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 (F271 = 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, n = 51; 1993-96 combined) on Guana, 1.3 g on Virgin Gorda (SD = 0.2 g, n = 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 BREV/ORA No. 508 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. albilabris, ■ E. antillensis, ▲ E. schwartzi, ;i- E. cochranae. 1999 BRITISH VIRGIN ISLANDS’ LEPTODACTYLID FROGS 15 £ E 35 T 30 -- 25 - .c O) c _0> C 3 O C CO 20 -• 15 -- 10 -- 0 Guana □ E. antillensis ME. schwartzi 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, n — 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 of 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 17 Q Z < < J o H C6 O H < Q aS O O z 3 OS > z 0 s os 1 a: C*3 Q < i ^2 On 55 On w -J z -< § Si K o § o as oj £ s Oj -J id u. 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