Deposition of Specimens

Contractual agreements with Profauna and Inparques provided that one half of the collection, including all holotypes and half of the species represented by single specimens, would be cataloged in the collection of the Museo de la Estación Biológica de Rancho Grande (EBRG) in Maracay, with the other half going to the amphibian and reptile collections of the American Museum of Natural History (AMNH) in New York. In order to assure an equitable split, specimens were divided after identifications, sex, and maturity were determined at the American Museum; EBRG numbers were then assigned and cross-referenced to CWM field tags. We thank Francisco J. Bisbal, Director of the Rancho Grande Museum, for facilitating the cataloging process and for helping in other ways.

Most specimens were collected by a herpetology team comprised of the two of us and John Daly, but obliging colleagues of other disciplines also provided specimens. All of this material is cataloged as collected by the 1994 AMNH–Terramar Expedition—an abbreviation for the official name “The Robert G. Goelet American Museum of Natural History–Terramar Expedition to Auyantepui”.

Specimens collected by J. Celsa Señaris, a guest in two of our camps, were retained by her for the collection of the Museo de Historia Natural La Salle (MHNLS) in Caracas, with half of her specimens also going to EBRG. Except for Oreophrynella (fig. 12), this material adds no species and did not contribute to the present report.

Measurements, Webbing Formulae, Tadpoles

Measurements of less than 12 mm were taken with an ocular micrometer fitted in a Wild dissecting microscope; larger measurements were made with digital calipers read to the nearest 0.1 mm or with a ruler read to the nearest 1 mm. For anuran webbing formulae, we use the notational device of Savage and Heyer (1967) as modified for general use by Myers and Duellman (1982: 6). Tadpoles were staged according to Gosner (1960), which has been long accepted by many workers as a standard for taxonomic descriptions, and further championed in McDiarmid and Altig's (1999: 8–11) masterful Tadpoles. We use the larval terminology in chapter 3 (Altig and McDiarmid, 1999) of Tadpoles, although we are perplexed why the term “jaw sheath” was proposed as a replacement for the more familiar and visually descriptive “beak”.3

Vocalizations

Tape recordings of frog calls were converted to electronic format through a Kay CSL 4500 16-bit analog-to-digital converter (sampling rate 22050 Hz) and stored on a computer hard drive and optical discs. Waveforms and sound spectrographs were analyzed and printed using the CSL software (Kay Elemetrics Co.). Sound spectrographs are similar to older analog standards: wideband, with an effective bandwidth filter of 323.00 Hz, and/or narrowband, with an effective bandwidth filter of 63.09 Hz. It is important to note that time measurements of notes were made from wideband spectrograms and/or from the waveforms, whereas statements concerning frequency were derived from narrowband analysis. Failure to distinguish between time and frequency correlated aspects of changing bandwidth has caused ambiguity and confusion in the bioacoustical literature (Zweifel and Myers, 1989: 11).

We cannot improve on Zweifel's (1985: 272) concise definitions for the sometimes confused terms “call” and “note”:

Notes may be nonpulsatile (single pulsed) or, depending on degree of amplitude modulation of the waveform, weakly to strongly pulsed. Variation is tremendous (e.g., see Heyer et al., 1990: 387–404) and there are borderline cases of ambiguity between notes and pulses (e.g., Myers and Donnelly, 1997: 23).

Species Synonymies

In addition to senior and some junior synonyms, the synonymies may include references that mention Auyantepui specimens or that otherwise are pertinent to an understanding of the listed taxon. There is no intention of completeness. Additional references for the rapidly growing anuran literature can be found in Barrio Amorós (2004), Faivovich et al. (2005), Frost et al. (2006), Grant et al. (2006), and Señaris and Ayarzagüena (2005).

Taxonomic accounts of genera and species are arranged alphabetically within families. Lack of strict alphabetical order of listed frog families reflects recent taxonomic changes, which require corroboration as a matter of course.

Material

Stream below Camp 2, <1750 m: EBRG 2956 (holotype), MHNLS 12916 (paratype). See Señaris (“1993” [1995]) for description.

Material

1.1 km NNE Camp 1, 1650 m: AMNH A-164834–164839 (larvae), EBRG 2649 (7 larvae). Camp 1, 1700 m: AMNH A-164817–164822, EBRG 2693–2699. Camp 3, 1850 m: AMNH A-164823, 164840–164843 (larvae), EBRG 2650 (8 larvae), 2700. Camp 4, 1600 m: AMNH A-164824–164833, 164844–164848 (larvae), EBRG 2651 (15 larvae), 2701–2709. All from the 1994 AMNH–Terramar Expedition to Auyantepui.

Diagnosis

A small frog (♂ about 21–24 mm, ♀ about 24–28 mm SVL) with an elongated median lingual process; no cloacal tubercles; fingers with keel-like lateral folds, broadest and best developed on fingers II and III; feet moderately webbed, toes with folded flaplike fringing; dorsal blotched pattern; an incomplete oblique pale lateral line either present or absent. See Discussion and Other Comparisons.

Morphology

Adult male Anomaloglossus tepuyensis about 21–24 mm SVL, adult females larger at about 24–28 mm SVL (table 1). Sexual maturity was judged by the presence of open vocal slits in males (open only on right in two specimens of 22 mm SVL), and by the presence of enlarged pigmented ova and convoluted oviducts in females.

Table 1

Size (in mm) and Proportions of Anomaloglossus tepuyensis La Marca, from the Summit of Auyantepui^a

Dorsal skin granular in life, becoming somewhat tuberculate on hind limbs (fig. 13), but skin much smoother in preservative; ventral skin virtually smooth in preservative, weakly granular on throats of some males. Dorsal skin forming a well-defined rounded, posteriorly projecting flap well above vent, which opens at upper level of thighs. Proximal thighs coarsely granular below vent, but no pronounced, symmetrically arranged tubercles in cloacal region.

Head slightly wider than long; greatest head width (between angles of jaws) 33%–39% of SVL. Snout sloping, rounded to bluntly pointed in profile, truncate to broadly rounded or bluntly pointed in dorsal and ventral view. Nares situated near tip of snout and directed slightly posterolaterally; nares visible from front, barely visible from above or below; posterior rim of naris bordered behind by a crescent-shaped, finely incised groove; posterior rim not raised but usually bearing a small tuberclelike prominence posterodorsally to naris. Canthus rostralis slightly rounded; loreal region weakly concave, sloping slightly outward to lip. Interorbital region much wider than upper eyelid. Snout subequal to eye length; distance from center of naris to edge of eye/eye length  =  0.55–0.74. Usually two low but inconspicuous postrictal tubercles. Tympanum usually inconspicuous, concealed posterodorsally by a diffuse supratympanic bulge; tympanum less than half of eye length.

Hand moderate, its length 24%–29% of SVL, 62%–82% of greatest head width. Relative lengths of appressed fingers III > IV > II > I; disc of finger I either failing to reach or else overlapping disc of finger II. Discs of all fingers moderately expanded; third finger disc in adults 1.4–2.0 times wider than distal end of adjacent phalanx, average width greater in females. Third finger broadly keeled or fringed but not swollen. Keel-like lateral folds on fingers usually low and inconspicuous except on finger III and, to a lesser extent, finger II; these keel-like structures appear to be tightly folded ventromedially around the fingers—with the edge of the fold recurved in a groove along the ventral side of the finger, usually best developed on medial side of finger III (figs. 14, 21); no obvious sexual dimorphism in finger keeling, which is also detectable but inconspicuous in small juveniles.

Base of palm with large median metacarpal tubercle, generally distally flattened, but in some specimens slightly cordiform or rounded; smaller elliptical inner metacarpal tubercle on base of first finger; one or two low, rounded subarticular tubercles (one each on fingers I and II, two each on fingers III and IV, with distal tubercle tending to be inconspicuous on IV). A faint outer metacarpal fold extending from lateral side of fourth finger to the large palmar tubercle, sometimes bearing a small tubercle. No ulnar tubercles or fold. Often a small tubercle situated below palmar tubercle toward lateral side of wrist (fig. 14). No fleshy supracarpal pad atop wrist.

Hind legs moderately long, with heel of adpressed limb reaching from between eye and snout to beyond snout; tibia 49%–55% of adult SVL, relatively shorter in females (table 1). Relative lengths of appressed toes IV > III ≥ V > II > I; first toe short, reaching or failing to reach base of subarticular tubercle of second toe. Toe discs moderately expanded (sometimes only weakly expanded on first toe), subequal or larger than finger discs. Feet moderately webbed, the webbing distally continuous with distinct lateral fringing on all toes; the lateral fringing is folded downward along sides of toes (“folded flaplike fringes”, see Discussion and fig. 14). Modal webbing formula of adults: I 1½–2½ II 1½–3 III 2½–3½ IV 4–2½ V. One specimen has a short first toe with only the first phalanx (disc) free of web; otherwise there is slight variation only between the third to fifth toes: III (2–3)–(3½–4) IV (2–4)–(2–2½) V.

A small, usually elliptical inner metatarsal tubercle, and a small, usually round outer metatarsal tubercle subequal in size (either may be equal to or larger than the other). One to three nonprotuberant subarticular tubercles (one each on toes I, II, two each on III and V, three on IV). The fringe along outer edge of the fifth toe is continuous with a weak outer metatarsal fold that terminates at the outer metatarsal tubercle. Distinct fringe along free edge of first toe is continuous with an oblique tarsal fold extending proximolaterad on the distal half of the tarsus (fig. 14); the nearly straight tarsal fold terminates abruptly in a curved tuberclelike structure (similar to condition shown in fig. 20A [A. tamacuarensis]).

Fig. 20

Hind foot structure of two species of Anomaloglossus characterized by folded flaplike fringes on toes. A. A. tamacuarensis, with well-developed flaplike fringes (AMNH A-131342, paratopotype). B. A. parkerae, with relatively weak flaplike fringes (KU 167328, paratype). See also figure 14 (A. tepuyensis).

Fig. 21

Anomaloglossus tepuyensis. Finger III (right hand) enlarged from figure 14, showing keel-like lateral fold in greater detail, including diagrammatic section of a short transverse cut across fringe. An underlying layer of melanophores shows through the somewhat translucent lateral fold, giving it the appearance of being irregularly pigmented. Scale line  =  1 mm.

Teeth present on maxillary arch. Tongue longer than wide; free posteriorly, with rounded margin; a usually prominent median lingual process is either pointed or distally rounded and is present in all specimens including juveniles. Vocal slits large, each extending from edge of tongue nearly to angle of jaw.

Coloration

In life (fig. 13), the dorsal ground color of Anomaloglossus tepuyensis ranged from grayish brown to orangish brown and bronzy brown in adults, mostly orange-brown in subadults, and brown or gray-brown with conspicuous white flecking in small juveniles—all with irregularly pigmented dark brown or blackish brown markings (obscure or indefinite only in a few specimens) usually arranged in the following pattern (fig. 15): (1) a dark interorbital bar, often with a slightly concave (sometimes straight) anterior margin, with or without a posteriorly produced apex; (2) a V-shaped to cordiform (transversely straight in a few) median dark blotch between arms, sometimes connected with the following; (3) a transverse pair of smaller paravertebral blotches at midbody—these markings sometimes fused with the anterior blotch to form a vaguely X-shaped dorsal mark; (4) usually a single small median posterior blotch near end of body.

The dorsal dark markings listed above are formed from irregular distributions of melanophores and usually contain pale areas of brown ground color; the paired paravertebral blotches and posterior median blotch often are circular with pale centers (ocellus-like). The snout anterior to the interorbial marking is brown like the rest of the dorsum, often with one or a few small dark marks or blotches, but the top of the snout is unmarked in about 27% of the sample and is never darkened overall. The eyelids occasionally have a few unpigmented dots but are usually unicolor except where crossed by the interorbital bar.

An incomplete pale oblique lateral line (in life, pale brown to blue) present or absent; extending from groin and terminating dorsolaterally behind arms when distinct, but often poorly defined and broken, or completely absent (frequently present on one side but absent on the other). A blackish stripe or face mask along side of snout passing through eye and widening above shoulder, tending to continue posteriad just below dorsum but changing along its lower edge on flanks to indefinite brown mottling or reticulum with irregular or ill-defined pale spots of bluish white, pale blue, or pale bronze, this area interrupted or not by a pale oblique lateral line of similar color. Pale spotted pattern of flanks extending to ventrolateral edge of venter (no hint of a pale ventrolateral stripe).

Black face stripe sometimes continuous around tip of snout, which otherwise varies from uniform brown to pale spotted. Upper lip pale to dark brownish, variously and indefinitely marked with small pale areas or spots (sometimes bluish white) or irregular dark blotching. Often a bronze (white to pale brownish in preservative) stripe extending obliquely from eye to forearm, just below the postocular widening of the black face mask; this pale postocular stripe distinct in some specimens, virtually absent in others in which it may lack contrast with a pale lip or else be interrupted and obscured by infusion of dark pigment.

Forearms and hind legs brown with distinct to vague dark bars or irregular spots, and with a tendency toward orange suffusion on the thighs. Often a dark brown stripe along anterior edge of upper arm, sometimes extending forward to corner of mouth, with a similar horizontal brown stripe varying from distinct to absent along anterior surface of thigh. Posterior thigh brown with indefinite pattern of pale flecks or small spots, only rarely with an ill-defined horizontal brown line or stripe on one or both sides (EBRG 2695, 2696, 2701). Some specimens have conspicuous pale spots or crossbands on the joints of the digits of hands and feet, but this pattern usually vague or digits uniformly pigmented. Digit tips noted as being “white marked” in life, but in preservative all digital scutes tending to be completely dark above, only occasionally with a pair of small inconspicuous pale spots.

Ventral color varies ontogenetically in life: (1) small juveniles are green or greenish gray, with silvery white flecking; (2) larger juveniles had all ventral surfaces orange except greenish on underside of thigh and shank and gray on palms and soles; (3) a few adult males at camp 1 were greenish gray ventrally becoming yellowish green under the limbs, whereas an adult male at camp 3 had the throat and chest gray with conspicuous white flecking that continued along the sides of the belly, the belly and undersides of limbs otherwise bright greenish yellow; ventral color at camp 4 (including adult males and females) was mostly gray with white flecking on throat and/or chest, turning bright greenish yellow or even (in a few) golden yellow posteriorly and under limbs, but a few adults had even the throat and chest bright colored like the belly and undersides of limbs. (Some degree of sexual dimorphism in ventral coloring is probable, but specimens could not be reliably sexed in the field.)

In preservative, ventral surfaces (fig. 16) vary from pale tan with a weak suffusion of melanophores, most dense on throat, to overall darker brown owing to dense suffusion of melanophores, which sometimes form a reticulum around pale spots. The axillary region tends to be weakly pigmented or unpigmented (but not forming in life a flash mark differently colored from ventral surfaces), this area being continuous ventrally with a small unpigmented area proximal to the arm insertion, which stands out as a small distinct spot (except in specimens with the palest venters). Undersides of hands and feet dark brown.

Iris pale bronze or orange-bronze, darkened with black suffusion, sometimes paler below pupil than above; pale bronze or orange-bronze pupillary ring usually interrupted by a black tick mark below pupil. Nictitating membrane translucent, with brown pigment along free edge.

Habitus and Proportions

Head and body elliptical and rounded at both ends in dorsal view; midbody width 70%–75% of head-body length. Head-body depressed (midbody depth 76%–80% of midbody width), somewhat flattened dorsally and ventrally. Snout rounded in dorsal and lateral view. Eyes dorsal, directed laterally; interorbital distance (between centers of pupils) 3.9 mm. Nares dorsal, directed laterally; internarial distance (between centers of nares) 2.6 mm. Nares approximately equidistant to tip of snout and anterior edge of eye. Spiracle sinistral, directed posterodorsally, positioned low on body, a tube (1.5–2.1 mm long) with the medial wall attached to the body; opening 6.9–7.6 mm from snout, at 55%–58% of head-body length. Vent tube dextral to caudal fin.

Lateral line system with a supraorbital and infraorbital branch originating from near upper labium and connecting behind eye. Angular branch originating ventral to the infraorbital branch near the upper lip and continuing posteriorly as the middle body branch; a branch connecting the angular branch to the infraorbital branch posterior to the eye, this branch continuing ventrally onto the ventral surface of the body anterior to the spiracle. Superior trunk branch continuing onto tail in the dorsal fin just above the musculature, for about 25% of tail length.

Tail 67%–68% of total length, with low fins, maximal height approximately equal to body depth; dorsal fin not extending onto body; dorsal fin higher than ventral fin; tail tip rounded, extending slightly past tip of musculature.

Pigmentation

In preservative, head-body brown dorsally and laterally; ventral body clear, flecked with melanophores and some light brown blotches; tail musculature tan with irregularly shaped dark brown blotches and flecks; ventral fin lacking pigment near body, with dark blotches and flecks posteriorly; dorsal fin dark near body, with flecks and blotches posteriorly.

Mouth Parts

Mouth ventral. Oral disc laterally emarginate. Labial teeth in 2/3 rows; anterior rows nearly equal in length, A2 row with median gap; posterior rows 2 and 3 equal in length, slightly longer than P1; P3 not as heavily keratinized as P1 and P2. Upper jaw sheath a broad arch with slender lateral processes and blunt serrations; lower jaw sheath V-shaped with blunt serrations like those on upper jaw. Marginal papillae from lateral edges of upper labium in single row above emargination; in weak double row below emargination and then single across middle of lower labium; papillae on lower labium more pointed than those on upper labium.

Ontogenetic Changes

Changes in body size are summarized in table 2. The ratio of head-body length to total length decreases from 38% in stage-25 tadpoles to 31% in stage-36 specimens. Body width increases during development relative to oral disc width; the ratio of oral disc width to body width changes from 55% in stage-25 tadpoles to 42% in the single stage 40-specimen. The lateral line system is not visible in several stage-25 tadpoles, but is distinct by stage 29.

In several stage-25 tadpoles, labial tooth row P3 is lightly keratinized (this row commonly is the last to be keratinized in dendrobatoid larvae [e.g., Myers et al., 1978; Donnelly et al., 1990; Myers and Donnelly, 1997]).

There is some variation in the marginal papillae on the lower labium (a single row or double rows), but we did not detect ontogenetic change in the character. (Ontogenetic change from one to two rows seems to be common among dendrobatoids.)

Holotype

EBRG 2733 (field no. CWM 19318), an adult male from summit of Auyantepui at 5°58′N, 62°33′W (AMNH–Terramar Camp 4), 1600 m, Bolívar, Venezuela; collected February 19–21, 1994, AMNH–Terramar Expedition. Figure 22.

Etymology

The specific name is a Latin adjective pertaining to celestial angels, including, perhaps, legendary pilot Jimmy Angel of Auyantepui fame.

Diagnosis

This species belongs to the former Hyla geographica group (as defined by Duellman, 1973), as indicated by the external characters of calcars, prepollical spines not projecting through the skin, basal webbing between fingers, and general habitus. Presence of a flat mental gland and close resemblance with Hypsiboas roraima places it in the benitezi group of Hypsiboas (Faivovich et al., 2005: 86–87).

It is further distinguished by the following suite of characters (in the format of Duellman and Hoogmoed, 1992): (1) body slender, with head distinct from body; (2) skin on dorsal surfaces including upper eyelids weakly granular (in life) or smooth (in preservative); head skin not co-ossified; (3) tympanum distinct; (4) fingers with basal webbing; (5) toes about half webbed; (6) fringes absent on limbs; a short conical calcar present; (7) axillary membrane absent; (8) dorsum brown with pale lichenlike spots and irregular black spotting; broken black vertebral stripe present (at least in holotype); flanks grayish with small black spots; limbs brown with gray crossbanding; rear of thigh gray; venter pale (gray in life), unmarked; foot webbing brown like dorsum; upper half of palpebrum clear except for brown upper edge and brown flecking, lower half brown; (9) vomerine odontophores angular.

Hypsiboas angelicus is remarkably similar to Hypsiboas roraima (Duellman and Hoogmoed) from Cerro Roraima, differing most conspicuously in lacking pale reticulation on the palpebral membrane.

Morphology

An adult male 38.4 mm SVL, with large vocal slits; a shallow subgular vocal sac, irregularly folded against posterior half of throat and chest. A thin, loosely organized mental gland on anterior half of gular region, with a convexly rounded posterior edge (fig. 25, ventral view). Body slender, head approximately as wide as long, prominent, much wider than body (head width/greatest body width  =  1.25); snout slightly sloping, short, truncate in dorsal and lateral views, not projecting beyond margin of upper lip; canthus rostralis rounded; loreal region concave, inclined outward to rounded lip; nostrils protuberant laterally; internarial area slightly depressed; top of head flat; upper eyelid as wide as interorbital distance; eye length greater than its distance to naris (eye/naris-eye  =  1.12), nearly as long as snout; tympanum distinct except for upper edge, circular, deflected dorsolaterally; separated from eye by a distance slightly greater than its length; supratympanic fold weak, but covering upper edge of tympanic annulus.

Axillary membrane absent; forearm slender, with row of large, ill-defined, closely spaced low ulnar tubercles situated along ventrolateral edge of arm; fingers slender, with rounded terminal discs; third finger disc subequal with length of tympanum; relative finger length III > IV > II > I; subarticular tubercles rounded, low, barely protuberant; supernumerary tubercles few, indistinct proximally; large palmar tubercle rounded, low and very indistinct; thenar tubercle elongate, not well defined, with median projection distally; basal webbing between fingers II–IV.

Hind limb long and slender; tibia 53.4% of SVL; heel bearing short conical tubercular calcar; toes with rounded terminal discs subequal in size to those on fingers; relative toe length IV > V > III > II > I; subarticular tubercles low, rounded, only slightly protuberant; supernumerary tubercles few, indistinct proximally; inner metatarsal tubercle oval; outer metatarsal tubercle small and indistinct; no tarsal fold; toes moderately webbed, with formula: I 2–2½ II 1½–2½ III 1½–2½ IV 2–1½ V (fig. 23).

Vomerine odontophores oblique, weakly arched, situated between large choanae and separated by a distinct medial gap, each with about a dozen teeth. Tongue large, covering floor of mouth, only lateral edges free, fully attached behind.

Skin on dorsal surfaces, including upper eyelids, weakly granular in life (fig. 22), becoming virtually smooth in preservative; skin on throat and under arms smooth in preservative, coarsely granular (areolate) on venter; proximal posteroventral surfaces of thighs crowded with smooth tubercles; ventral thigh surfaces covered with large, probably glandular tubercles, low and smooth in preservative. Supracloacal flap short, slightly overlapping vent opening, which is directed posteriorly at upper level of thighs; vent situated immediately above a smooth vertical channel situated between two large, vertically elongate tubercles, which in turn are flanked by large, smoothly rounded tubercles that grade into those on proximal posteroventral surfaces of thighs (fig. 24).

Measurements (in Mm)

Snout to vent 38.4; tibia 20.5; hand from proximal edge palmar tubercle to tip of longest finger 10.5; foot from proximal edge inner metatarsal tubercle to tip of longest toe 14.1; head length on the diagonal from tip of snout to angle of jaw 13.6; greatest head width (near angle of jaws) 13.8; width of upper eyelid 4.0; greatest body width 11.0; width of interorbital area 4.0; internarial distance 3.0; rear edge naris to anterior corner of eye 4.2; eye length from anterior to posterior corner 4.7; eye to tympanum 2.5; horizontal length of tympanum 2.2; width of third finger disc 2.1; width of fourth toe disc 1.9.

Coloration

In life (fig. 22), grayish brown, with lichenlike spots of pale rose and irregular black spotting on dorsum; a narrow, broken black vertebral stripe from snout to midbody; flanks gray with small black spots; limbs brown with inconspicuous gray crossbanding; rear of thigh gray; foot webbing brown like limbs. Dull grayish yellow under chin, turning uniformly light gray over all remaining ventral surfaces, including palms and soles. Iris pale golden yellow, with very fine, scarcely noticeable, black venation. Upper half of palpebral membrane clear in life (except for brown upper edge and brown flecking, as noted in preservative), lower half brown.

In preservative (fig. 25), pattern well retained, with rose coloring faded from lichenous dorsal spots, and with throat and venter turned pale yellowish and undersides of limbs pale brownish.

Material

Camp 2, 1750 m: AMNH A-164998 (1 larva), from the 1994 AMNH–Terramar Expedition to Auyantepui.

This curious little tadpole was found in a pool of water at the bottom of a deep (> 10 m) sandstone crevice, into which our colleague Petia Alcócer had descended by rope. Assuming that it does not represent a frog otherwise unknown from the Auyán summit, we tentatively associate it with Hypsiboas angelicus.

Habitus and Proportions

Head-body length 12.6 mm, body width 7.0 mm, body depth 6.0 mm, total length 35.7 mm. Body slightly depressed (body width/body depth  =  1.17); snout acutely rounded in dorsal view, rounded in profile. Nares dorsolateral, directed laterally, 1.5 mm from tip of snout; distance between centers of nares 2.1 mm. Small eyes dorsal, directed laterally, diameter 0.4 mm; interocular plane 2.6 mm from tip of snout. Spiracle a 0.7 mm long tube adhered to body wall, sinistral. Large vent opening dextral to caudal fin; vent 1.5 mm long, with inner wall free of body.

Lateral line system (not illustrated) not pronounced, including: a pair of supraorbital lines diverging in straight lines from center of snout to top of eye; high and middle lateral lines on body; an angular branch with a short anteriorly extending loreal branch; a gular branch extending from the middle lateral line and ventrally converging toward the angular branch.

A single unpigmented bump (gland?), 0.4 mm in diameter, near end of body on each side.

Tail length 64.7% of total length. Maximum tail height 16.8% of total length. Fins unequal in height; dorsal fin 1.8 mm and ventral fin 1.6 mm at maximum tail height. Tail distally forked.

Pigmentation

In preservative, head-body uniformly covered by rodular melanophores, brown in color. Brown rodular melanophores also overlying the pale yellowish brown tail musculature and extending proximally along both fins, becoming sparse and disappearing before the end of the tail. Well-defined pale yellow chromatophores are sparsely distributed, occurring mainly on the sides of the body and along the base of the upper tail fin.

Mouthparts

Mouth ventral. Oral disc width 2.2 mm, not emarginate. Labial teeth in two anterior and three posterior rows, which are subequal in length except for last posterior row being shortest. Anterior tooth rows medially broken. Edges of upper and lower jaw sheaths with pointed serrations; laterally, the lower jaw sheath is only narrowly keratinized. Oral disc nearly surrounded by pointed marginal papillae, except for an anterior median break in the row. The marginal papillae are disposed in a single row (looking like a double row where folded posteriorly, fig. 28).

Material

BMNH 1975.1356 from Auyantepui, 3200 ft. [975 m], collected by Adrian Warren (field no. 112).

Material

TADPOLES. 1.1 km NNE Camp 1, 1650 m: EBRG 2645. Camp 4, 1600 m: AMNH A-164852–164857, EBRG 2646–2647, 2731–2732. Camp 5, 2100 m: AMNH A-164858–164864, EBRG 2648. All larvae from the 1994 AMNH–Terramar Expedition to Auyantepui.

FROGS. Auyantepui summit, 1700 m, 5°53′N, 62°35′W: AMNH A-169282, EBRG 3750, 3751, collected by Ricardo Guerrero, February 6–8, 1988.

The last three specimens are adult males 28.1–32.4 mm SVL (x ¯  =  30.70 mm, S.D.  =  2.29 mm, N  =  3); tibia length 52%–59% of SVL; head width 37%–39% of SVL, 95%–102% of diagonal head length; eye length greater than distance from its anterior edge to center of naris (EN/eye  =  0.88–0.92); upper eyelid more than half the width of interorbital space (IOD/eyelid  =  1.38–1.56); disc on finger III wider than discs on toes III–IV. Prepollical spine large, lacking an orifice (not protrusile, fig. 30).

Coloration

Coloration is somewhat variable in preservative (originally fixed in 70% ethanol according to the collector), but all have a black canthal line and a variable dorsal pattern of small black spots (fig. 31). There are about 3–6 spots along the outer side of the lower arm, and conspicuous spotting on dorsal surfaces of the hind limbs, especially on the tibia and tarsus. Several black spots scattered over the dorsum are conspicuous in two specimens but reduced to a few black specks in another. One specimen (EBRG 3751) is dull brown dorsally owing to dispersal of pigment from the melanophores; the other two specimens are very pale orangish or (in AMNH A-169282) pale orange over all dorsal surfaces, with minute melanophores visible under magnification. Rears of thighs are pale orangish; ventral surfaces, including palms and soles, are immaculate, the color varying from very pale orangish (nearly colorless) to pale orange. Two specimens have occasional silvery iridophores associated with the melanophores; the iridophores are relatively numerous in AMNH A-169282, appearing as silvery flecking under magnification.

Based on living specimens from the type locality on the southern slope of Auyantepui (970 m), Hypsiboas jimenezi has “a green dorsum with a dense matrix of [brown or reddish brown] chromatophores and with large spots on limbs and forelimbs”; white dorsolateral lines were present in topotypic specimens but absent in others (Señaris and Ayarzagüena, 2006: 310, fig. 1). In preservative, the green specimens faded either to cream or to reddish brown in specimens that had a very dense matrix of chromatophores. The bright yellow venter became cream in preservative (Señaris and Ayarzagüena, 2006: 312, 313).14

Coloration

We noted in the field that some tadpoles had a noticeably bright red iris. The red color was explicitly noted for one large larva from Camp 4 (AMNH A-164853, stage 37), and for some (probably the larger ones) in one lot of 11 from Camp 5 (EBRG 2648, stages 25–33; AMNH A-164858–164864, stages 24–40). However, Duellman (1997: 20) described two small (stage-25) Hypsiboas sibleszi tadpoles as having a pale bronze iris. Although ontogenetic change or some other confounding factor cannot be ruled out at this time, red eye color seems diagnostic at least for larger larvae of the Auyantepui sample.

Hoogmoed (1979a: 29) said that H. sibleszi tadpoles in stages 42 and 43 have white interorbital and lateral lines, which are precursors of the pattern found in some but not all adults. None of our larvae show such markings. However, our largest tadpoles (stages 37 and 40) have conspicuous dark brown spots on the dorsum, which appear to be precursors of the adult spotted pattern of Hypsiboas jimenezi (compare figs. 31 and 33). Rodular melanophores are present among larger variably shaped melanophores.

Mouthparts

We here compare Hoogmoed's (1979a: 27–29) description of the mouth parts of Hypsiboas sibleszi larvae. Hoogmoed described sibleszi as having 2/4 labial tooth rows, with the P4 row poorly developed and consisting of several isolated patches of small denticles. In our sample of Hypsiboas jimenezi, some tadpoles in stages 25, 27, 28/29, and 32 have 2/4 tooth rows, but the most common formula is 3/5 rows (observed in stages 27, 28, 29, 31, 33, 37, and 40). A labial tooth-row formula of 2/5 is present in some tadpoles in stages 25, 27, and 40, and the formula 3/4 rows occurs in a single stage-33 larva.

The A3 and P5 rows, when present, are often obscured by the folds of the oral disc. The A1, A2, A3, P1, and P5 rows in our sample may have median gaps (in those tadpoles with 2/4 or 2/5 labial tooth rows, the A2 row has the median gap and the A1 lacks it; in tadpoles with 3/4 or 3/5 labial tooth rows, the A1 row has a very wide gap, A2 lacks one, and the A3 row has a narrow gap).

As shown by Hoogmoed (1979a: fig. 8d) for Hypsiboas sibleszi and in our figure 32 for Hypsiboas jimenezi, the oral disc is anteromedially emarginate on the upper labium and has a posterolateral fold or indentation on each side of the lower labium. Hypsiboas jimenezi additionally shows a median fold in the lower labium; the median fold varies from a weak median indentation (easily overlooked or considered artifactual) to a pronounced tuck (as in fig. 32) and would seem to be species specific.

The upper jaw sheath is a broad arch with blunt serrations and slender lateral processes; the lower jaw sheath is V-shaped with pointed serrations. Hoogmoed (1979a) illustrated and described a single row of marginal papillae around the oral disc in Hypsiboas sibleszi, but there are two rows in our sample of Hypsiboas jimenezi. As also shown by Hoogmoed (1979a: fig. 8d), the marginal papillae are broken by a median gap on the upper labium, at the point of anterior emargination (fig. 32). There are a few blunt submarginal papillae on the lower labium in some of our larvae (fig. 32).

Hoogmoed (1979a: 29) mentioned that there may be “connections between the third and fourth lower denticle rows” in H. sibleszi larvae, and we observed anomalies in various tooth (denticle) rows in several tadpoles of Hypsiboas jimenezi. In one larva in stage 25 (EBRG 2646) and in another in stage 27 (AMNH A-164859), the A2 row curves upward to occupy a position normally occupied by A1 on the left. This results in a short A2 on the left and a short A1 on the right. In some cases, the tooth row consists of two or three ridges (short lateral segments and long medial segment), whereas in others there are short rows that are perpendicular to the tooth row (located near the submarginal papillae), and in one case there is a short lateral ridge between the P2 and P3 rows. The A3 and P5 rows in our samples are made up of several short ridges that are often adjacent to the marginal papillae.

Lateral Line System

The infraorbital and supraorbital branches originate near the upper labium and join behind the eye to form a loop. There are scattered neuromasts inside the orbital loop. The middle trunk branch originates in the middle of the body slightly anterior to the spiracle, then curves ventrally posterior to the spiracle, to terminate at the level of the hind limb. There are two superior trunk branches, originating as parallel branches just posterior to the orbital loop; one branch curves medially and continues posteriorly in the dorsal tail fin, and the other branch curves ventrally and continues along the middle of the tail musculature. Hoogmoed's (1979a: fig. 8) drawing of H. sibleszi shows some resemblances to the above description.

There is an inconspicuous small unpigmented bump near the end of the body on each side (fig. 32).

Material

Camp 4, 1600 m: AMNH A-164849–164851. All from the 1994 AMNH–Terramar Expedition to Auyantepui.

These frogs were found only at Camp 4, where conditions were much wetter than at the other camps. Five adult males were collected. These males are 30.6–33.1 mm SVL (x ¯  =  31.46 mm, S.D.  =  1.02 mm, N  =  5); tibia length 53%–54% of SVL; head width 39%–41% of SVL, 84%–91% of diagonal head length; eye length slightly shorter than distance from its anterior edge to center of naris (EN/eye  =  1.06–1.13); upper eyelid less than half the width of interorbital space (IOD/eyelid  =  2.00–2.50); disc on finger III wider than discs on toes III–IV, which are subequal.

Coloration

In life (fig. 35), the ground color of the five adult males varied either from light green to yellowish green, or from pale to medium blue-green.15 A variable number of small pale yellow dots sparsely scattered on dorsum, conspicuous or not; some specimens with dorsal surfaces also densely dotted with brown melanophores overall, this dotting inconspicuous or not evident in others (but conspicuously present in all specimens in preservative). Digits distally light orange above and below. Pale interorbital bars and dorsolateral stripes are lacking in this sample.

Vocal sac light yellow-green; rest of ventral surfaces light blue-green. Venter somewhat transparent, with heart, liver (including gall bladder), and intestinal tract tightly bound in white peritoneum; linea masculina visible. Bones green. Iris pale bright yellow, with suffusion of light red-brown around pupil and with fine black venation.

Material

Camp 1, 1700 m: AMNH A-164865–164878, 164879–164880 (larvae), EBRG 2596–2609, 2628 (larvae). Camp 2, 1750 m: AMNH A-164881–164899, 164900–164909 (larvae), EBRG 2610–2619, 2629–2631 (larvae). Camp 3, 1850 m: AMNH A-164910–164917, 164918–164923 (larvae), EBRG 2620–2627, 2632–2634 (larvae). Camp 4, 1600 m: AMNH A-164924–164926, EBRG 2637–2639. Camp 5, 2100 m: AMNH A-164927–164931, 164932–164935 (larvae), EBRG 2640–2644, 2635–2636 (larvae). All from the 1994 AMNH–Terramar Expedition to Auyantepui.

Additional Material

Auyantepui Plateau, 2200 m (7218 ft): AMNH A-46033–46035, 146339 (larvae), from the 1937–1938 AMNH–Phelps Venezuelan Expedition.

Females are larger than males (x ¯ female SVL/x ¯ male SVL  =  1.24) and there is little overlap in size between sexually mature males and females. The following measurements were obtained by combining all adult males and all adult females from the 1994 Expedition:

There is no obvious variation in size across sites (data not shown); analysis of variance for males from camps 1–5 shows no statistically significant difference between the mean SVL at the 95% confidence level (F  =  2.30, P  =  0.0731).

All adult males have a well-developed nuptial pad on the base of the thumb; the pad is a pale brown color owing to minute centers of cornification (visible with a dissecting microscope). The vocal slits of males are relatively small apertures situated near the corner of the mouth; it is difficult to find these openings on some specimens, and a few individuals of adult size with nuptial pads seem to lack them. There is a degree of sexual dimorphism in skin texture, with males well covered with small spicules and with females tending to have smoother or slightly pustulate skin, as seen in figure 38 (males above, female below); some adult females, however, do have sparse spicules (especially anteriorly) additional to smoother pustules. Adult males have enormous white testes that fill most of the body cavity; females in the sample have convoluted oviducts and numerous small pigmented ova.

Coloration

Based on a large sample of 95 specimens, Tepuihyla edelcae was diagnosed as having a uniformly dark dorsal color (Ayarzagüena et al., 1992a: 122), with some limited variation being mentioned in the description (p. 126). However, in life this is a colorfully variable frog, at least on Auyantepui (figs. 37–38), with a basic dorsal color of brown, orange-brown, red-brown, or gray; a few specimens had the dorsolateral areas slightly lighter than the darker middorsum and one individual (AMNH A-164916) has distinct pale dorsolateral stripes that are retained in preservative. Some frogs had orange or yellowish spots or blotches over the head and dorsum and also on the throat. Limbs colored like the dorsum, rarely with indication of dark crossbanding; rear of thigh orange-brown to gray, sometimes with small areas of silvery gray frosting. Upper lip white or variably suffused with dark pigment. Throat and venter dirty white, turning gray under the limbs. Bones white. The iris varies from light bronze to dark brown or gray, with variable black venation and a tendency for a dark horizontal stripe through the pupil.

In preservative the frogs are darker, but any light markings are well retained in our material. Two specimens were noted as having either faint or distinct dark crossbanding on the limbs (AMNH A-164869, 164916, respectively); the rear of the thigh is light to dark orange. Some specimens retain a distinct pale labial line in preservative; Ayarzagüena et al. (1992a: 126) seemed to suggest that this is a juvenile character retained in some adults. We see no ontogenetic linkage in the Auyantepui material, inasmuch as the upper lip is dark in some small specimens such as AMNH A-164872 (a metamorph 16 mm SVL), EBRG 2639 (20 mm), and EBRG 2641 (25 mm).

Habitus and Proportions

Head and body elliptical and rounded at both ends in dorsal view; maximum body width 53.8%–54.6% of head-body length. Body higher than wide (maximum body depth about 117.6%–118.2% of maximum body width); body flattened dorsally and somewhat wedge-shaped in profile, increasing in depth from anterior to posterior. Eyes dorsal, directed laterally, not visible from below. Nares dorsal, directed anterolaterally, approximately equidistant between tip of snout and eye. Spiracle sinistral, low on body, a short tube (1.4–2.2 mm long) directed posteriorly with medial wall attached to body; spiracle opening at 58.2%–59.5% of head-body length. Vent tube attached to ventral fin, opening dextral to fin.

Lateral line system somewhat indistinct. Supraorbital and infraorbital branches forming a loop around eye; superior and middle trunk branches originate behind eye; superior branch continues posteriorly in dorsal fin above musculature; middle trunk branch continues posteriorly in the middle of tail musculature; inferior body branch originates on body near spiracle and curves ventrally to the ventral body.

Tail 64.7%–66.5% of total length; maximum tail height approximately equal to body depth; dorsal fin not extending onto body, higher than ventral fin; musculature height greater than fin height; margins of dorsal fin slightly scalloped; tail tip rounded.

Pigmentation

In preservative, head-body brown dorsally and laterally; ventral skin clear, with underlying tissue having an iridescent blue sheen; tail musculature tan, flecked with melanophores, becoming vaguely blotched posteriorly. Dorsal and ventral fins brown (heavy melanophore deposition), paler at tail tip and adjacent to body on lower fin.

Mouth Parts

Mouth ventral. Oral disc not emarginate. Labial teeth in 2/3 (AMNH A-164907) or 2/4 rows (AMNH A-164902); A1 and A2 rows subequal in length, A2 row with distinct median gap; P1 slightly shorter than P2; P3 either shorter than P1 or equal to P2; P1 with or without a short gap in median curvature; P4 row (in AMNH A-164902) small and barely keratinized. Upper jaw sheath a broad arch bearing pointed serrations and with slender lateral processes, its surface with a minutely pebbled texture that is highly reflective under certain light conditions. Lower jaw sheath V-shaped with pointed serrations. Marginal papillae with a broad anteromedial gap on upper labium; one row of marginal papillae on upper labium and along lateral margins of disc; two rows of slightly blunter papillae on lower labium; several pointed to blunt submarginal papillae laterally.

Ontogenetic Changes

In three available tadpoles of Tepuihyla edelcae in stage 24/25, maximal tail height exceeds body depth, and body depth and width are approximately equal. The body depth exceeds tail height and body width in all other stages. Changes in body size are summarized in table 4.

The three larvae in stage 24/25 have paired patches of raised, pigmented skin posterior and lateral to the oral disc. These raised patches may be adhesive glands; they are reduced to pigmented spots in early stage-25 larvae, with the spots disappearing by late stage 25.16

The stage 24/25 larvae have 2/2 tooth rows that are barely keratinized; jaw sheaths are keratinized in this stage and pointed serrations are present; marginal papillae are also present in these tadpoles but submarginal papillae are lacking. Early stage-25 tadpoles have 2/3 labial tooth rows; the P3 row is the smallest and lightly keratinized; these tadpoles also lack submarginal papillae. The posteriormost tooth row (P4) is the last to appear and it is often not well developed. The submarginal papillae also appear relatively late in development.

Variation in the tooth-row formula is as follows: The three available tadpoles in stage 24/25 have 2/2 tooth rows; some tadpoles in stages 25 (N  =  6), 26 (1), 27 (2), 33 (1), and 37 (1) have 2/3 tooth rows; all others (stages 25–40) have 2/4 tooth rows.

Some unusual configurations of mouthparts are not associated with ontogeny and may be environmentally caused (see Natural History). In one stage-33 tadpole (AMNH A-164902), the P4 row is short, is situated to the right of the midline, and is positioned in the marginal row of papillae. In some cases, the tooth-row ridges are comprised of short and long sections (i.e., A1 in three sections—two short lateral and one long medial section; A1, A2, or P2 in two sections—one short and one long). In some tadpoles, the tooth-row ridges are unusually short (P1 and P2 missing on the left, A1 and A2 short on right, P3 short on right). In many tadpoles, the P4 row is small, difficult to discern, and mixed with the marginal papillae. In one late stage-25 larva, there are large pointed papillae between tooth rows), and in one there is a single papilla in the median gap on the upper labium.

Material

Camp 2, 1750 m: AMNH A-164992–164997, EBRG 2686–2691, all topotypes. Camp 5, 2100 m: EBRG 2692. All from the 1994 AMNH–Terramar Expedition to Auyantepui.

Coloration

In life (fig. 41), Stefania schuberti was olive to olive-brown over the dorsal surfaces, with ill-defined black spotting and limb bars. The rear thigh surfaces were grayish brown with small vague spots of yellow or olive. Ventral surfaces were gray, with vague white mottling that is most evident on the chest. Iris olive above, below dark bronze greatly darkened by black venation.

Juvenile frogs of about 21–23 mm SVL were similar to adults, with some color differences. They were gray above, with a suffusion of reddish brown in the blackish spots and limb bars. The rear of thigh was black with gray flecking. Ventral surfaces were gray with whitish flecking, especially on the throat and chest. Iris pale gray across the upper third, very dark brown on the lower two thirds, with slight black venation (which did not seem to contribute much to the darkening of the lower two thirds).

Natural History

Most specimens were found at night at a series of low waterfalls below Camp 2 (fig. 7 top), where they were discovered mostly by eyeshine while they were looking out of crevices around the falls or sitting on wet rock. One adult female (EBRG 2691) was found by day on a sandstone wall in the dark of a crevice cave that had a floor of water. Two juveniles (21–23 mm SVL) were found by day under rocks on a sandstone ledge well above the waterfalls below Camp 2 (fig. 7, bottom). Another juvenile (30 mm SVL) was found under a rock near the south rim of Auyantepui, near Camp 5.

Stefania appeared to be reproductively inactive during our work in February. Calling was not heard (although vocal slits are lacking in male Stefania, at least a few species are believed to call [see Duellman and Hoogmoed, 1984: 8]). The two females judged as sexually mature at 62 and 85 mm SVL have convoluted oviducts and maturing ova of about 1.3 to 1.5 mm in diameter. Three immature females of 50–52 mm SVL have nonconvoluted oviducts and largest ova of about 0.6–0.8 mm. The two adult males are 72 and 74 mm SVL.

Holotype

EBRG 2725 (field no. CWM 19213), an adult female from summit of Auyantepui at 5°54′N, 62°29′W (AMNH–Terramar Camp 2), 1750 m elevation, Bolívar, Venezuela, collected February 9–11, 1994, AMNH–Terramar Expedition.

Paratopotypes

AMNH A-164951–164953, EBRG 2724, 2726–2727, with same collection data as holotype.

Other Paratypes

Camp 1, 1700 m: AMNH A-164936–164950, EBRG 2709–2723, collected 2–4 February, 1994. Camp 4, 1600 m: AMNH A-164954, EBRG 2728–2729, collected 23 February 1994. All from the 1994 AMNH–Terramar Expedition.

Etymology

The specific name auricarens (“lacking ears”) is an adjective of common gender derived from the Latin auris (“ear”) + carens (the present participle of careo, “to be without”), alluding to the absence of the tympanic apparatus.

Definition and Diagnosis

A small Eleutherodactylus belonging to the unistrigatus species group and characterized by the following combination of characters (in the format of Lynch and Duellman, 1980, 1997): (1) dorsal skin tubercular, ventral skin areolate; (2) tympanic membrane and annulus absent (annulus possibly vestigial in a few specimens, see Remarks); (3) snout truncate to bluntly pointed in dorsal view, rounded in profile; canthus rostralis gently concave or nearly straight, rounded; (4) upper eyelid weakly tubercular (tubercles low, subequal); (5) vomerine odontophores small but distinct, with detectable teeth, oblique, well separated and situated between and behind choanae; (6) males lacking vocal slits/vocal sac; weak nuptial pads seemingly present; (7) finger I shorter than II; broad discs on fingers II–IV; (8) weak lateral keeling but no fringes on fingers; (9) ulnar tubercles absent or low and inconspicuous (not forming a distinct line); (10) no pronounced calcar tubercle; (11) inner metatarsal tubercle oval, much larger than small outer metatarsal tubercle; (12) toes weakly keeled, webless; toe discs broad, subequal with finger discs; fifth toe much longer than third; (13) varying shades of brown above, with interorbital bar and often a vague undulating longitudinal dark marking on dorsum; polymorphism limited, but occasionally with a pale postocular stripe posteriorly widening to cover flanks, or with a few oblique dark stripes extending posteroventrad from indefinitely patterned dorsum; (14) four males about 16–18 mm, nine females 20–25 mm SVL.

Eleutherodactylus auricarens is distinguished from sympatric E. pulvinatus by absence of a distinct tympanic apparatus, which is externally well defined in E. pulvinatus (see following species account for other differences).18 The only other Eleutherodactylus known to lack the tympanic apparatus in eastern and southern Venezuela is E. yaviensis from the northwestern tepuis, although specimens in one population of yaviensis (Cerro Yutajé) have a vestigial annulus and membrane concealed under the skin (see Remarks). E. yaviensis is a larger species with a higher degree of pattern polymorphism, although it lacks the vague undulating middorsal marking that is present in many specimens of auricarens (see Remarks for further comparisons between E. auricarens and E. yaviensis).

Measurements of Holotype (in Mm)

The holotype (figs. 44, 45) is an adult female as revealed by presence of large ova and convoluted oviducts. SVL 23.6, tibia length 13.6, foot length from proximal edge inner metatarsal tubercle to tip of toe IV 11.2, head width 9.2, head length on the diagonal from tip of snout to angle of jaw 8.0, upper eyelid width 2.7, interorbital distance 3.2, internarial distance 2.3, center of nostril to eye 3.0, eye length 3.1, tympanum absent, width of third finger disc and width of fourth toe disc both 1.3.

Morphology

Females are larger than males (x ¯ female SVL/x ¯ male SVL  =  1.33). Head little wider than long, usually wider than body (body as wide as head in a few froglets, wider than head in some gravid females); head width 36%–40% of SVL in adults, 34%–39% in juveniles. Snout truncate to bluntly pointed in dorsal view, rounded in profile; eye-nostril distance 74%–97% of eye length in adults, 72%–94% in juveniles; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis gently concave or nearly straight, edge rounded; loreal region concave, sloping outward to lip. Upper eyelid bearing low, more-or-less subequal small tubercles; upper eyelid width usually narrower than interorbital distance (82%–95% of interorbital distance in adults, 72%–108% in juveniles). Tympanic apparatus absent (as judged from dissection of all adults and selected juveniles, but see Remarks); a usually pronounced glandular ridge (“supratympanic” fold) sloping from eye nearly to forearm, the skin below this fold varying from rugose to nearly smooth (see fn. 18 and Remarks). Two or three sometimes inconspicuous postrictal tubercles between corner of mouth and forearm. Choanae small to moderate, oval to round, not concealed by palatal shelf of maxillary arch; vomerine odontophores distinct, rounded to slightly elongate and slanted posteromedially, distinctly separated medially, situated posterior and median to choanae, each bearing several teeth. Tongue longer than wide, posterior edge varying from straight to rounded, with or without a slight notch, posterior half free. Adult males lacking vocal slits (hence there is no vocal sac); adult males appearing to have a weakly developed, unpigmented, nuptial pad along edge of thenar tubercle.

Dorsal skin finely granular in life (fig. 42) and weakly tuberculate, somewhat more strongly tuberculate on limbs; middorsal raphe present, although only anterior portion visible in some juveniles; throat smooth, venter areolate; a broad triangular area of glandular skin present on posteroventral thigh surfaces; discoidal fold (inconspicuous on most specimens) situated anterior to groin; no ulnar ridge or defined line of tubercles.

Relative lengths of appressed fingers III > IV > II > I; tip of first finger sometimes reaching disc of second; males seeming to have a weakly developed unpigmented nuptial pad along edge of thenar tubercle; fingers with lateral keeling absent or weak, but downward, tightly folded flaplike fringing developed in some adults on medial sides of fingers II–III and lateral side of V (e.g., AMNH A-164938♀, 164954♀). Finger discs broader than long, slightly rounded to nearly truncate, with subdigital pads much wider than long; disc on thumb slightly expanded. Palmar tubercle large, cordiform; thenar tubercle oval, large; subarticular tubercles low to somewhat protuberant, either rounded or conical; several supernumerary palmar tubercles (fig. 43), either low or slightly protuberant, sometimes nearly as large as subarticular tubercles.

Hind limbs relatively long; heels overlap when held at right angles to sagittal plane; tibia 50%–60% of SVL (table 5). Relative lengths of appressed toes IV > V > III > II > I; tip of toe I not reaching disc of toe II; tip of toe V normally extending to the distal edge of, or well past, the ultimate subarticular tubercle of toe IV; tip of toe III extending to or slightly past the penultimate subarticular tubercle of toe IV.19 Toe discs broad, usually as broad or slightly broader than those on fingers. Toes with lateral keeling weak or absent, but downward, tightly folded flaplike fringing developed in some adults on medial sides of toes III–IV and lateral side of toe V (e.g., AMNH A-164938♀, 164954♀). Basal webbing essentially absent.20 Inner metatarsal tubercle large, an elongated ovoid, much larger than small round outer metatarsal tubercle; several or many small to moderate supernumerary plantar tubercles (fig. 43), low or protuberant; subarticular tubercles larger, moderately protuberant, rounded to nearly conical. Calcar tubercles low and indistinct, none prominent. Tarsus somewhat tuberculate, lacking a tarsal fold or constantly positioned tarsal tubercle (a small tubercle sometimes distinguishable on distal part of tarsus near inner metatarsal tubercle).

Coloration

In life (fig. 42), dorsal coloration some shade of brown—gray-brown, brown, blackish brown, greenish brown, orange-brown—with darker interorbital bar and often with a vague, ill-defined, middorsal longitudinal dark marking that usually has a few transverse or sometimes chevronlike lateral extensions or undulations (fig. 44). The longitudinal dorsal mark is lacking in some specimens, including the following: one male and two females each have a dark dorsum and a contrasting pale (orange or gray in life) postocular stripe that extends ventrolaterally and expands on the flank; and a few females have indefinite dark dorsal coloration, from which a few dark stripes extend obliquely (posteroventrally) onto the flank.

Some specimens with an ill-defined dark stripe below the canthus rostralis from eye to snout, often broken or incomplete, with only the anterior half remaining as a longitudinal dark blotch enclosing the nostril, absent in some. Often a brown postocular stripe with or without conspicuously darkened upper and lower edges, sometimes represented only by its upper edge—a narrow black line superimposed on the postocular (“supratympanic”) fold. Frequently, one to several dark bars radiating from eye to lip, with pigmented but slightly lighter interspaces, but lip sometimes moderately pigmented without obvious pattern.

Dark crossbanding on limbs sometimes indistinct, but forearms usually with one distinct band, thighs usually with two, shanks usually with three. Groin suffused with orange, which color often occurs also on concealed part of shank, anterior side of thigh and/or top and rear of thigh. In other specimens, the posterior thigh and concealed part of the shank is uniformly gray or brown in life, uniform or with pale dots.

There is some ontogenetic change in ventral coloration. In life, the ventral surfaces are nearly black in the smallest juveniles, then becoming gray or greenish gray in larger specimens. Ventral coloration tends to be darkest (blackish) under the head, where an occasional specimen has a weakly developed pale midgular line (which is usually absent, see fig. 45); there may be patches or dots of bronze on the gray chest and sometimes on the throat as well. Some sexual dimorphism was suggested by two amplectant males, whose ventral surfaces and posterior thighs (as well as dorsal surfaces, see fig. 42) were much darker than in their respective females, but in preservative there are males and females showing a range of ventral pigmentation from pale to dark. In preservative, the ventral surfaces are pale yellowish, sparsely to densely punctuated with brown, the throat usually being somewhat darker to much darker than the belly (fig. 45). Palms and soles lightly pigmented, palmar and plantar tubercles tending to be somewhat darker, some subarticular tubercles on feet also may be pigmented.

There is individual variation in the living color of the iris, which may be light to dark bronze, golden bronze, or greenish bronze, tending to be paler above than below pupil, and with a red or reddish brown horizontal stripe (which may be ill-defined) through pupil.

Material

Camp 1, 1700 m: AMNH A-164955, juv. ♀, collected 2–4 February, 1994. Camp 4, 1600 m: AMNH A-164956♂, EBRG 2730♂, collected 19–22 February 1994. All from the 1994 AMNH–Terramar Expedition to Auyantepui.

Eleutherodactylus pulvinatus is readily distinguished from sympatric E. auricarens by presence of an externally distinct, relatively large tympanum (fig. 47A). The two adult males in our collection measured 22.8 and 23.0 mm SVL, with the following proportions: tibia/SVL  =  0.583 in both; head width/SVL  =  0.387–0.399; eye-naris/eye length  =  0.765–0.906; tympanum/eye  =  0.313–0.324; eyelid/IOD  =  0.852–0.920. The juvenile female is 13.4 mm; tibia/SVL  = 0.575; head width/SVL  =  0.373; eye-naris/eye length  =  0.850 tympanum/eye  =  0.350; eyelid/IOD  =  1.063. The males have two slightly whitish nuptial pads, one along edge of thenar tubercle and the other on posterodorsal side of thumb; these are best developed in AMNH A-164956, the second pad only weakly indicated in EBRG 2730. Vomerine odontophores are small but easily seen, situated between and slightly posterior to the choanae.

Coloration

In life, gray-brown over the dorsal surfaces, with obscure darker markings and crossbanding on the limbs; a poorly defined dark interorbital bar and what seem to be remnants of a dark W-shaped scapular mark are perhaps somewhat more evident in preservative (fig. 48) than in life; the thighs appeared virtually uniform gray in life (pale orangish brown in preservative, with or without pale flecking). EBRG 2750 has several pale oblique bars crossing the flanks, but these markings are lacking in AMNH A-164956. A dark brown stripe below the canthus rostralis, from eye to nostril, is evident in one (EBRG 2750) of the adult males but vague in the other. Both have an oblique postocular brown stripe with a blackish upper edge, and several dark bars radiating from eye to lip (most distinct in EBRG 2750, obscure in the other).

The ventral surfaces were medium gray overall, with inconspicuous pale gray flecking on throat and venter (pale brown in preservative, with unpigmented flecks and small spots). There was a horizontal red-brown stripe through the pupil; the iris was bright bronze above the pupil, with minimal black venation, whereas below the pupil it was darkened by black venation in a more-or-less vertically aligned pattern.

The juvenile female (AMNH A-164955) lacks a distinct dorsal pattern, lacks oblique pale bars on the flanks, and lacks the brown canthal stripe, although the dorsally black-edged postocular stripe is present; the upper lip has dark bars along its length. The rear of thigh is brown with pale flecking, and the ventral surfaces are like those of the adults except a darker brown.

Material

Camp 1, 1700 m: AMNH A-164957–164963, EBRG 2653–2659. Camp 3, 1850 m: AMNH A-164964–164985, EBRG 2660–2680. 4 km NE Camp 3, 1430 m: EBRG 2681. Camp 4, 1600 m: AMNH A-164986–164989, A-164854–164857 (4 larvae), EBRG 2652 (larva), 2682–2685. All from the 1994 AMNH–Terramar Expedition to Auyantepui.

Color and pattern are somewhat variable (figs. 49–50), as is size. Combining all adult males and all adult females gives the following measurements:

Size overlap between the sexes is considerable, but females are significantly larger than males (t  =  −2.7482, P  =  0.0117). Sexual maturity in males on Auyantepui seems to be attained between about 48 and 54 mm SVL21; four subadults in the range 49–54 mm SVL had unopened vocal slits, unpigmented or pale gray vocal sacs, and small and/or noncornified thumb spines; two males, coded as adult at 52 mm SVL, each had only one open vocal slit; the vocal sacs were dark and the thumb spine weakly cornified in one of these specimens, but the other had pale vocal sacs and a noncornified thumb spine.

Most adult males have a single, externally evident thumb spine (fig. 51) in various stages of cornification, but a specimen each from camps 1 and 3 have two thumb spines. Less than half the adult males have 1–2 small spines on each side of the chest. Some have several minute spinules clustered around the spines, and one specimen (AMNH A-164979) from Camp 3 has a band of small spinules or tubercles extending across the chest (fig. 51). Adult male rugosus are characterized by paired chest spines but not by such a “distinctive band of black tubercles across the chest in larger males”, which is a property of Leptodactylus lithonaetes (Heyer, 1995: 699, 701, 708). A patch of black skin tubercles on the anterior throat is diagnostic of L. lithonaetes (Heyer, 1955: 708) but absent in L. rugosus, including all specimens from Auyantepui.

Color in Life

See figures 49–50 and table 6, which summarizes color and pattern by locality.

Table 6

Leptodactylus rugosus Noble. Color and Pattern Variation (in Life) in Three Populations on the Summit of Auyantepui

Holotype

EBRG 2905 (field no. CWM 19354), an adult female from summit of Auyantepui at 5°58′N, 62°33′W (AMNH–Terramar Camp 4), 1600 m elevation, Bolívar, Venezuela, collected February 22, 1994, AMNH–Terramar Expedition.

Paratype

AMNH R-140230 (CWM 19165), a male from summit of Auyantepui at 5°51′N, 62°32′W (AMNH–Terramar Camp 1), 1700 m, Bolívar, Venezuela, collected February 7, 1994, AMNH–Terramar Expedition. Figure 53.

Etymology

The specific name montigena is an adjective meaning “mountain born”. It is derived from the Latin noun mons (“mountain”) + the adjective-forming suffix -genus (“born or produced in a certain place”).

Diagnosis

A small highland species of Arthrosaura readily distinguished from its nearest geographic neighbor (A. testigensis, Los Testigos range, east of Auyantepui) in having three (vs. four) supraoculars and in having a pale dorsolateral line on each side and an unmarked venter (vs. four narrow yellowish stripes on dorsum and brown spots on venter). A presumably unnamed species from the Chimantá massif (southeast of Auyantepui) evidently differs from A. montigena in having a dark brown middorsal stripe (Gorzula and Señaris, 1999: photo 89).

Arthrosaura montigena shares many characteristics of A. versteegii (French Guiana to extreme eastern Venezuela25) but differs in having a more complete pale dorsolateral line and in not having the ventral or ventrolateral scales sharply demarcated by a zone of small scales. A key to the named species of Arthrosaura is provided at the end of this section.

Coloration

In life (fig. 53), dorsum brown with variably distinct tan dorsolateral lines; flanks blackish brown, with some small pale spots. Palms and soles black; other ventral surfaces either sexually dimorphic or individually variable as follows: Male—all ventral surfaces except palms and soles uniformly bright orange. Female—underside of head creamy white, turning light yellow under throat and over venter and base of tail, becoming orange under distal three fourths of tail. Iris orange. Tongue black.

Color and Pattern in Preservative

More detailed examination of the preserved specimens (fig. 56) shows the female to have a distinct pale dorsolateral line extending from the postorbitals onto the base of the tail. The male has this line extending weakly from the eye and along the outer edge of the parietal, then becoming distinct on the neck, present but poorly defined on the body, and then again distinct from above the groin onto the base of the tail; the dorsolateral line is continued posteriorly from the proximal part of the tail as a series of pale brown spots that disappear before the end of the tail in the female (tail incomplete in male). A parallel row of white spots on lower side of tail, also disappearing before the end of tail.

The hexagonal body scales—inconspicuously speckled with brown on a lighter brown ground and with a slight darkening along the keels—are sharply bicolored, with the free ends grayish brown. The grayish distal ends overlap irregular dark brown pigmentation across the bases of the scales in each succeeding row, forming a series of dark transverse lines across the dorsum and flanks, but there is an increase in the amount of dark brown pigmentation on the flanks of both specimens, and some of the scale tips of the lateral scales are white in the male, accounting for its pale-spotted blackish sides in life (fig. 53). The pattern of dark transverse lines extends only onto base of the tail, which becomes marked with small irregular dark spots and darker keels.

Top and sides of head brown, with some slightly darker brown pigmentation but no definite patterning. Supralabials appear either brown with white spots in the male, or white with brown bars in the female. Several elongated whitish spots with darker borders on side of neck. Ventral surfaces and ventrolateral scales on lower flanks white, with some light to dark brown pigmentation on the ventrolaterals and lateral row of ventrals; no pigment on genials, but some irregular dark brown markings at base of forelimb insertion and an occasional very small dark mark in preanal area or elsewhere; ventral surfaces otherwise immaculate.

Method

The genitalia of the paratype of Arthrosaura montigena were everted in the field, and the left organ was later removed and inflated with carmine-dyed petroleum jelly. Because of a puzzling, well-defined aperture between the lobes of the organ and because the apices of the lobes seemed not to fully evert, the jelly was squeezed out and the organ soaked overnight in a solution of 3% KOH (also injected into the lumen) before being reinflated. The right organ was then removed, injected with 3% KOH and soaked in the KOH solution for about 10 hours before being inflated. Results were comparable, however, and the two organs are essentially identical. The tips of the lobes could not be completely extended manually, even by internal probing with the round head of a no. 000 insect pin. The following description and illustrations are of the left organ, showing maximally attained inflation.

Description

The inflated hemipenis (fig. 57) is 5.5 mm wide by about 7 mm in overall length; it is bilobed for about 30% of total length. The broad middle part of the sulcate side of the organ is nude, with the sulcus spermaticus being a deep, well-defined groove running a medial course to the crotch; the sulcus bifurcates as it enters the crotch, with the branches extending centripetally probably to the apices of the lobes. There is a well defined aperture (the “orificium”) about 0.2 mm wide between the sulcate branches in the middle of the nude crotch, opening into a tunnel extending proximally into the hemipenis to a depth of perhaps 1 mm (which would  =  14% of hemipenial length).

The median part of the asulcate side of the organ is nude from the base to the crotch. Plicae arise on either side of the nude midline in a weak chevron pattern; the basal six rows of plicae terminate laterally on each side, at a nude area on the lower side of the hemipenis; a small spine rises at the lateral terminus of rows 2–5. Starting with row 7, the asulcate plicae meet a matching set of plicae on the sides of the lobes. At magnifications exceeding 20 times, the edges of all the plicae are lined with minute, presumably mineralized spinules.

Comments

The puzzling orifice (here termed orificium) in the hemipenial crotch opens into a relatively deep tunnel of unknown function. The possibility was considered that the orificium might open into the lumen of a long, uneverted process. However, the opening and the shaft are equally well defined in both left and right hemipenes and there is no visible indication that the structure might be eversible. Although the apical tips of the lobes resisted complete eversion, the hemipenes overall were maximally inflated without apparent deformation of their distal, proximally extending tunnels. The orificium is a heretofore unknown hemipenial structure.

The hemipenis of Arthrosaura montigena is somewhat similar to that of Arthrosaura synaptolepis (fig. 3 in Donnelly et al., 1992), which differs most notably in having the hemipenial lobes terminally flattened and in having a bumplike “pronounced elevation of tissue” instead of an orificium. Clearly, there is considerable taxonomic potential in the hemipenial morphology of Arthrosaura.

Material

1.1 km NNE Camp 1, 1650 m: AMNH R-140200–140201, EBRG 2906. Camp 2, 1750 m: AMNH R-140202–140205, EBRG 2907–2909. Camp 3, 1850 m: AMNH R-140206–140208, EBRG 2910–2918. 4 km NE Camp 3, 1430 m: AMNH R-140209–140210. Camp 4, 1600 m: AMNH R-140211–140212, EBRG 2919. Camp 5, 2100 m: AMNH R-140213, EBRG 2920. All from the 1994 AMNH–Terramar Expedition to Auyantepui.

Coloration in Life

A juvenile (AMNH R-140201, 38 mm SVL) was bicolored, with a brown dorsum and black sides (fig. 58 top); ventral surfaces whitish. Color becomes variable in larger specimens.

One subadult (EBRG 2906, 61 mm SVL) had a dark greenish brown dorsum; the throat and venter were pale yellow with suffusions of orange-brown beneath neck and orange-brown spotting ventrolaterally, becoming pale greenish beneath limbs and tail.

Another subadult (AMNH R-140202, 60 mm SVL) had a black-checkered, dark olive-brown dorsum; ventral and ventrolateral surfaces as in specimen above; iris brown around pupil, with dense gray suffusion toward periphery.

An adult female (AMNH R-140200, 87 mm SVL) was dorsally brown when photographed (fig. 58 bottom), but, at time of preservation, it was dorsally olive, with paler areas and some blackish scales; ventral surfaces pale yellowish; iris olive green turning gray toward periphery. A large male (AMNH R-140206, 85 mm SVL) was black above with conspicuously yellow-speckled sides.

Hemipenis

The hemipenes of AMNH R-140208 (73 mm SVL) were everted in the field. Both organs were later removed, immersed for 1 hr in 3% KOH solution, and inflated with carmine-dyed petroleum jelly. Attempts to obtain maximum inflation (usually a desirable thing) caused an apparent overinflation and swollen distortion of one lobe of each organ, but all structures are completely everted. The following description and illustration (fig. 59) are based primarily on the left hemipenis, with some interpretation provided by comparison with the right organ; the apparent distortion of the swollen lobe is not illustrated, being replaced by a mirror-image drawing of the opposite lobe.

The everted hemipenis is robust, measuring 5.5 mm long and 4.2 mm across the midsection. The organ is very weakly bilobed, with the lobes strongly inclined to the sulcate side (fig. 59). The nude sulcate faces of the lobes are nearly flattened and are confluent with a pair of pronounced knobs (2 mm long, 1.5 mm wide) extending distally. There are four additional, much smaller protuberances, including a small round knob situated at the sulcate-side base of each large knob and a pair of protuberances in the lobular crotch. The sulcus spermaticus curves mediad from the base of the organ (coming from the left in the left organ [fig. 59], from the right in the right organ); the sulcus then extends centrolineally to bifurcate just beyond the base of the lobular area, with each branch extending between a large knob and the adjacent medial protuberance. The midsection of the hemipenis is nearly encircled by eight angulate tissue ridges that are interrupted by the sulcus spermaticus on one side and a narrow median nude space on the other. On the asulcate side the ridges are chevron-shaped and pointed distally, whereas the chevrons point proximally on the sides of the hemipenis. Comblike rows of minute, presumably mineralized spinules are embedded in the tissue folds; there are no other spines. The base of the organ is nude.

Material

Camp 1, 1700 m: AMNH R-140214–140217, EBRG 2902–2903. Camp 2, 1750 m: EBRG 2904. From the 1994 AMNH–Terramar Expedition to Auyantepui. Additional Material: Auyantepui plateau, 2200 m (7218 ft.): AMNH R-61013, from the 1937–1938 AMNH–Phelps Venezuelan Expedition.

Coloration

Dorsal color and pattern variable in life, but pattern usually including a dark brown interorbital bar, pale diagonal line on shank, and a pair of brown sacral markings. Two juvenile females had tan middorsal stripes on brown (fig. 60 middle) or greenish gray dorsa; another juvenile was frosted gray over a complex darker pattern (fig. 60 top). Adults were gray or slightly greenish gray with indefinite darker gray markings, especially on the flanks (fig. 60 bottom). Ventral surfaces dirty white, grayish white, or chin white with venter pale yellow. Pale labial stripe and stripe through ear pure white, grayish white, or pale greenish gray (this marking tending to become obscure in preservative).

Dewlap of two males orange, with white or grayish white scales in basal rows, scales darker gray or blackish gray in distal rows. One adult female with a smaller but similarly colored dewlap. The small juvenile dewlaps seemed more variable based on four specimens from Camp 1: One juvenile (♂) had a red dewlap with yellow scales basally and dark gray scales peripherally; two others had either a red (♂) or orange (♀) dewlap with a mixture of pale and dark scales; a fourth juvenile (♀) had a large bluish black basal spot on the dewlap, which had a bright orange periphery and mostly white scales (only a few dark scales).

Iris of adults dark bronze heavily suffused with black and with a pale blue pupillary ring; iris of two juveniles appeared dark brown with a bluish white area above the pupil. Tongue pale yellowish or orangish; mouth lining pale grayish, throat lining not pigmented.

Natural History

Six of our seven specimens were collected in the forest (fig. 4 bottom) near Camp 1; one was sleeping on a leaf 1 m aboveground at night and the others were on the forest floor by day. An additional specimen was found by day in a patch of forest near Camp 2.

Characters Consistent with A. chrysolepis

A conspicuous sheen is emphasized in the original description of nitens and in the name,30 suggestive of the metallic sheen seen in some anoles, including occasional specimens of chrysolepis (see below). The pair of dark sacral spots seems to be the only character really suggestive of the chrysolepis complex, but their presence is variable and they are not confined to A. chrysolepis. The indistinct thigh pattern seems relatively uninformative.

Characters Inconsistent with A. chrysolepis

The fact that Wagler took the trouble to mention the sacral markings and the obsolete31 thigh banding indicates that the animal was otherwise patternless. If so, the absence of all head markings, tail banding, vertebral stripe or dorsal chevrons or rhomboids, lateral marbling, or other such pattern elements is inconsistent with what is known of the normal variation of Anolis chrysolepis. (An occasional old, poorly preserved and faded specimen of chrysolepis might conceivably lack describable pattern, but neither should such a specimen show the bright colors attributed to nitens.)

As previously stated (Myers and Donnelly, 1997: 59), the green dorsal coloring ascribed to nitens is inconsistent with Anolis chrysolepis, which is a variably patterned, basically brown or gray lizard. This requires additional elaboration.

“green” in the Color Variation of Anolis chrysolepis

Beebe (1944) indicated the great range of colors and color patterns of his composite A. chrysolepis/A. nitens at study sites in Venezuela and Guyana, including a specimen of chrysolepis having a general ground color of “olive with a strong greenish yellow tinge” and another one “grayish olive”. A search of Myers' field catalog reveals a few specimens having a slight greenish cast in the basic ground colors in life, including a “greenish gray” juvenile (EBRG 2902) and a “greenish gray” adult (AMNH R-140217) from Auyantepui and a “pale greenish brown” adult female from Cerro Guanay.

Such pale greenish casts, washes, or suffusions are not rare in the normal variation of brown or gray-colored reptiles and amphibians, but they disappear more quickly than normal greens in preservative and often are photographically elusive as well (e.g., Lynch and Myers, 1983: 522, 540). Nonetheless, under certain light, a pale bluish gray or pale greenish gray ground color can be seen on the lower sides or venters of some preserved Auyantepui specimens (AMNH R-140214–140217), although, in life, a greenish gray color was evident in only one of these (AMNH R-140217).

However, it is difficult imagining that anyone would describe such subtle hues as “green” without qualification.32 We earlier puzzled over the green in Wagler's description:

A slight metallic bronzy or golden sheen sometimes is evident in living A. chrysolepis (fig. 229 in Ávila-Pires [1995] seems to show this) and especially in preserved specimens. However, reasons for presence of a sheen, sometimes in association with a greenish hue, are not evident to us. Duméril and Bibron (1837: 91, 93–94) described the holotype of Anolis refulgens as having a ground color of “gris verdâtre doré [golden greenish gray, or ‘greyish…with metallic gloss’ as rendered by Boulenger, 1885: 92]” above and “blanche, glacée de verdâtre [white with a greenish glaze]” below, with dark head marbling, oblique bars, and crossbands on limbs and tail. Perusal of AMNH material revealed a specimen of A. chrysolepis scypheus (AMNH R-97359) having a tan ground color replaced by irregular areas of pale metallic blue-green on the posterior dorsum and left side, but oblique blackish brown lateral bars are well developed even in the blue-green area. The metallic shine in these few specimens is reminiscent of Wagler's description of nitens, which, however, seems to have had a patternless body and tail.

Summary Comments on Anolis nitens

No author has argued that Wagler's (1830) description of nitens demonstrably applies to chrysolepis, only that the name has priority and has been applied to that species. However, despite perceived shortcomings of Wagler's description, his portrayal of color is so vivid that the name nitens probably could be assigned if a virtually patternless, green lizard were found matching the description—but would the hypothetical animal prove to be a rare color-pattern variant of a known species (e.g., A. chrysolepis) or a representative of a rare or geographically restricted species not currently recognized (i.e., A. nitens sensu stricto)?

The name nitens is a nomen dubium that cannot unambiguously be assigned to any known lizard. Article 23.3.6 allows authors who reject the synonymy to use the junior name chrysolepis as valid. However, although never in prevailing usage historically, nitens remains in competing use with chrysolepis because of prior (and arguable) interpretation of Article 23.

Until Ávila-Pires (1995), the body of work comprising the continuous usage of nitens was not impressive, inasmuch as the name had been mentioned mostly in lists and had been applied in print only to a few dozen specimens by 1970, compared with 761 specimens later assigned by Vanzolini and Williams (1970: 13) to the better known name chrysolepis. Studies using the name nitens subsequent to Ávila-Pires (1995) include Gorzula and Señaris (1999), Glor et al. (2001), and McDiarmid and Donnelly (2005).

Because of the confusion that has been engendered, we now agree with Hoogmoed (1973) that stability will best be served by petitioning the International Commission on Zoological Nomenclature to suppress nitens. A proposal for suppression was submitted by the first author in December 2007. Meanwhile we use the name Anolis chrysolepis as allowed by Article 23.3.6 (ICZN, 1999).

Material

Camp 1, 1700 m: AMNH R-140218. Camp 2, 1750 m: AMNH R-140219, EBRG 2921–2924. Camp 3, 1850 m: AMNH R-140220–140227, 140228 (eggs), EBRG 2925–2928, 2929 (eggs). Camp 5, 2100 m: AMNH R-140229, EBRG 2930–2932. All from the 1994 AMNH–Terramar Expedition to Auyantepui. Additional Material: Auyantepui, 1100 m (3609 ft.) [elevation may be in error, see Remarks], AMNH R-61011 (holotype). Auyantepui Plateau, 2200 m (7218 ft.): AMNH R-61012, 61014–61015, 61017 (paratypes). Type specimens from the 1937–1938 AMNH–Phelps Venezuelan Expedition. (Another paratype, AMNH R-61016, was exchanged to the MCZ in 1958; four additional paratypes  =  MBUCV 3053–3056 fide Roze, 1958a).

Based on dissection only of the AMNH share of specimens (N  =  11) from the 1994 expedition, females are appreciably smaller than the males. Five females, all reproductively active (see Natural History below), average 70.2 mm (67–75 mm) SVL, whereas four adult males average 88.3 mm (80–92 mm) SVL. Two additional specimens are immature males of 56 mm and 74 SVL; the last individual (AMNH R-140218) has faint indications of black patches starting to appear under the thighs and in the cloacal area.

Coloration

In life (fig. 63), overall black above, with light orange spots that are formed from small clusters of orange granular scales; white or tan scales are interspersed among the orange spots in smaller clusters of scales or as individual scales. Ventral surfaces spotted or mottled bluish white or pale blue on black (ventral pattern in fig. 64). Venter in juveniles bronzy white with vague black reticulum. Adult males have a large black patch under each thigh and smaller pre- and postcloacal patches. Tongue pinkish gray or whitish, with gray tip. Iris turquoise blue, except noted as green in one adult male.

Hemipenis

The hemipenes are tiny (≤ 6.5 mm long) relative to the size of the males. The organ lacks spines of any size and is divided for nearly half its length (fig. 65). The basal stalk of the hemipenis is nude, without ornamentation; there is a pronounced swelling below the crotch, on the distal third of the asulcate side of the stalk. The hemipenial lobes are calyculate, with the calyces being confined to the separate lobes and not confluent across the hemipenial crotch. The sulcus spermaticus bifurcates proximal to the crotch; the sulcus branches extend in centrolineal orientation onto the apices of the lobes. Each sulcus branch terminates as a nude channel between dense lines of columnar papillae, which rise above the calyces on the tips of the hemipenial lobes (fig. 65C).

Holotype

EBRG 3403, an adult male from summit of Auyantepui at 5°46′N, 62°32′W, 2100 m elevation, Bolívar, Venezuela; collected by Ricardo Guerrero, February 6–8, 1988. (The type locality is the same as the later-established Camp 5 of the 1994 AMNH-Terramar Expedition; see map 1.) Figure 67.

Etymology

This snake is named for our colleague and field companion Dr. Ricardo Guerrero, who collected the only known specimen several years before the AMNH–Terramar Expedition.

Diagnosis

An Atractus of small adult size (1 ♂ 250 mm total length), 17 dorsal scale rows, with a dark vertebral stripe and ill-defined lateral stripes; a prominent midventral line of large black spots; supralabials 7, with labials 2–3 touching short loreal and 3–4 touching eye; maxillary teeth 6; hemipenis noncapitate, feebly bilobed, completely spiny, with a long basal naked pocket on asulcate side.

Atractus guerreroi most closely resembles its geographic neighbor Atractus steyermarki Roze (Chimantá massif), which is relatively melanistic in color (dark stripes not evident or inconspicuous in life) and which has more conspicuous scale organs on some cephalic plates. Atractus guerreroi has a smaller head and shorter snout than A. steyermarki, with disproportionately smaller frontal, internasal, and loreal plates but with comparable sized parietals and prefrontals (as shown in fig. 72). See Remarks and Comparisons.

Proportions and Scutellation

Total length 250 mm, tail length 35 mm (14.0% of total). A small slender snake, with body slightly wider than high, rounded ventrolaterally; greatest head width 70% of head length from snout to end of parietals and 63% of length from snout to end of mandible; greatest body width 2.7% of SVL. Dorsal scales smooth, lacking apical pits, in 17-17-17 rows; anal ridges present. Ventrals 166, anal plate undivided, subcaudals in 41 pairs.

Head slightly wider than neck; snout bluntly rounded in dorsal view, rounded in profile; rostral wider than high, visible from above; internasals small, slightly wider than long, half (50%) the length of prefrontal suture; prefrontals large, longer than wide (greatest prefrontal width 80% of greatest length); prefrontal suture asymmetrical (dextral) to the internasal suture; prefrontal suture 89% length of frontal plate; supraoculars longer than wide; frontal wider than long, pentagonal; interparietal suture much longer than prefrontal suture and longer than frontal plate (interparietal suture 1.28 × frontal length). See figure 68 for cephalic scutellation.

Eye moderate, longer than loreal, contained 2.0 times in snout length (sagittal); eye length 1.4 times longer than its distance to lip; eye protuberant to level of lip and visible from below. Pupil subcircular (sensu Myers, 1984). Nasal divided above and below naris, its greatest length 2 times loreal length; loreal short (73% of eye length), 1.6 times longer than its greatest height, broadly separated from internasal, entering eye; no preoculars; supralabials 7, 2nd–3rd touching loreal, 3rd–4th touching eye; postoculars 2, the upper 1.5 times larger than lower; temporals 1 + 2, the upper one in row 2 very elongated, reaching past end of parietal.

Infralabials 7, first pair in contact behind mental, first three in contact with genials; single pair of genials, each genial 2.7 times longer than wide; four large median gulars (including two preventrals) between genials and first ventral. Head-plate tubercles tiny, inconspicuous, most concentrated on snout and chin, otherwise sparse. Some larger tubercles are discernible in a few places, particularly along the posterior edges of the prefrontals and anterolateral edges of the frontal.

Color Pattern

The specimen is medium brown, with a generally poorly defined pattern of narrow brown stripes and a low lateral pale stripe (fig. 67). There is most prominently a middorsal stripe (vertebral scale row and adjacent edges of paravertebrals) extending from the parietal plates to the tail tip. And there are indications of several ill-defined lateral stripes—on scale-row 6, on the adjacent edges of rows 3–4, and on the adjacent edges of rows 2–3. The lower 1½ scale rows are white in preservative, forming a low lateral pale stripe. The lateral brown striping and the low pale stripe are anteriorly vague, becoming somewhat more pronounced posteriorly.

The white ventral surface bears a median line of prominent blackish brown markings, starting anteriorly as a small spot on the first ventral and gradually enlarging posteriorly as close-set half-moon markings that fuse to a solid stripe under the tail. The lateral edges of the ventral plates are brown (paler than the blackish midventral markings), setting off the pale lower lateral stripe.

The head is brown, with a pale area on the outer edge of each parietal anteriorly and a poorly defined pair of large pale spots posteriorly on the ends of the parietals. The brown pigmentation extends onto the upper edges of the supralabials, which otherwise are immaculate white. The anterior face of the rostral plate is dark brown, with a white lower edge. There is a pair of brown spots on the chin, each spot occupying a first infralabial and part of the adjacent genial; underside of head and neck otherwise white. A dorsad extension of white extending onto side of neck behind mouth.

Maxillary Teeth

Examined in situ on right side. There are six well-spaced, recurved teeth, the first three large and subequal, the last three smaller and decreasing in size, with the ultimate tooth being very small. The teeth are firmly ankylosed and there are no empty sockets.

Hemipenis

The noncapitate, noncalyculate hemipenis is spinose over all except the spinulate basal section. Weak bilobation, most evident in the uneverted organ, is discernible but inconspicuous after eversion. The right retracted organ was 10.5 mm long from base to the ends of very short lobes. Bilobation started at the level of subcaudal 14. The lobes were 0.3 mm long; the two slips of retractor muscle merged 1.2 mm posterior to insertion, with the major retractor originating at the level of subcaudal 27. This organ was removed, soaked in a solution of 3% KOH for 10 hours and manually everted.

The everted hemipenis (fig. 69) is 11 mm long. The sulcus spermaticus forks more than halfway (64%) from the base, the two branches extending in centrolineal orientation to the tips of the lobes; bilobation is negligible (< 3% total length of organ). The distal 73% of the organ is densely covered with spines all around, mostly medium-sized but smaller near the sulcus spermaticus; the space between the sulcate branches is sparsely covered in small spines.

The basal 27% of the hemipenis is spinulate. There is a long basal naked pocket on the asulcate side, extending distad nearly halfway (45%) from the base, well into the zone of spines.

Notes on Atractus steyermarki

Roze (1958b) named Atractus steyermarki based on two specimens from elevations of 1430 m and 2160 on Chimantá-tepui. Later, Roze (1961: 116; 1966: 87) mentioned that the species had been rediscovered by Steyermark “en la región de El Dorado”. The lowland town of El Dorado, about 150 km NE Chimantá, lies at an elevation of 130 m, but elevations above 1200 m occur well to the south along the “El Dorado–Santa Elena de Uairén road” (see the regional description in Duellman, 1997). Roze took the locality at face value and assumed a broad distribution “en la Gran Sabana en general”, but he provided no descriptive data for the specimen. The El Dorado snake, cited by Roze (1966: 87) as an unnumbered MBUCV specimen (Museo de Biología de la Universidad Central de Venezuela, Caracas), apparently has not been seen by other Venezuelan workers (e.g., Lancini and Kornacker, 1989; Gorzula and Señaris, 1999).

Gorzula (1992: 272) reported an additional specimen of Atractus steyermarki from Chimantá Camp VIII ( =  2600 m fide Huber, 1992: 31). Gorzula and Señaris (1999: 161–162) also provided data for this specimen and mentioned the El Dorado record as unverified, concluding that “the available evidence indicates that this species is endemic to the highland summits of the Chimantá Massif”. The known elevation range for Atractus steyermarki on Chimantá is 1430–2600 m.

The holotype of Atractus steyermarki (figs. 70, 71) has been examined; see table 9 for detailed measurements. The specimen is in fair condition and pliable, although somewhat desiccated; it evidently has faded sufficiently in preservative to reveal a basic striped pattern (see below). There are six maxillary teeth, which appear much as described for A. guerreroi above, and the hemipenis also is similar to that of A. guerreroi. Unfortunately, both hemipenes are damaged and can not be easily illustrated. The left organ had been opened irregularly from the medial side and is torn. The proximal section of the right organ is mutilated, although the distal part (opened midventrally) is in good condition; the right organ extends to subcaudal 13. The hemipenes are spinose and very feebly bilobed; the attachment slips of retractor muscle are correspondingly very short.

Fig. 70

Atractus steyermarki Roze. The holotype (FMNH 69920♂) from the Chimantá massif, in dorsal and ventral view, ×1.3.

Fig. 71

Atractus steyermarki, showing enlarged lateral view of head of holotype (FMNH 69920♂) from the Chimantá massif, about ×7.5. The dark spots along edges of some scales anterior and posterior to eye are pigmented scale organs, which also are conspicuous along the edges of some plates on top the head.

Fig. 72

Upper: Atractus guerreroi, new species from Auyantepui (holotype, 215 mm SVL). Lower: A. steyermarki from Chimantá (holotype, 219 mm SVL). Heads of holotypes shown at same scale. The noticeably smaller head and shorter snout of A. guerreroi is not explainable by the slight differences in snout-vent length; it has disproportionately smaller frontal, internasal, and loreal plates, although prefrontals and parietals are comparable to those of A. steyermarki.

The male holotype (“270” [257] mm total length) and female paratype (420 mm) of Atractus steyermarki were described by Roze (1958b) as patternless dark gray, turning paler on the lower 1–2 scale rows. The midventral dark spots in the male cover most of the ventrals in the larger, more melanistic female. Gorzula (1992: 272) described the living colors of his intermediate-sized specimen (a male, 373 mm total length) as dark brown above, yellow below, with dark brown pigment midventrally.

The aforesaid descriptions document Atractus steyermarki as having melanistic tendencies that evidently cause the dorsum to appear essentially unicolor in life. Nonetheless, after a half century of preservation, the holotype has faded sufficiently for a striped pattern to appear (especially when immersed in alcohol, see fig. 70). Three prominent dark brown stripes are now visible—a middorsal stripe confined to the scales of the vertebral row, a high lateral stripe centered on row 6 and overlapping onto adjacent edges of rows 5 and 7, and a lateral stripe on row 3 and adjacent edge of row 2. Scales in rows 1 and 2 are mostly pale gray (with brown pigment at the scale base), appearing as a low pale lateral stripe set off by the brown stripe above and black ventral tips below. The midventral marking starts as small spots on ventral 13, enlarging posteriorly as black half-moons that nearly fuse to form a median stripe.

Comparison of Atractus steyermarki and A. guerreroi

As mentioned above, the maxillary teeth and hemipenes of Atractus steyermarki appear to be essentially the same as in A. guerreroi. The midventral dark pattern also is essentially the same in Atractus guerreroi and A. steyermarki. The dark dorsal stripes and ill-defined pale “stripe” on the first two scale rows of A. guerreroi are similar to the above-described pattern on the faded holotype of A. steyermarki, but there are differences.

The middorsal stripe of A. guerreroi is microscopically less dense and overlaps onto the paravertebral rows. Both have a stripe on row 6, but in steyermarki it is better defined and wider, overlapping onto the adjacent rows. Atractus guerreroi lacks the vivid lateral dark stripe (row 3 and edge of row 2) of steyermarki, having instead two very ill-defined lateral stripes or dark suffusions along the adjacent edges of rows 3–4 and rows 2–3. These differences in lateral stripes might be diagnostic, but, at this time, there is no way of assessing variation within populations. In A. steyermarki, the conspicuous stripes on the faded holotype seem likely to reflect a juvenile pattern that becomes ontogenetically obscured by melanism.

The holotypes of both Atractus guerreroi and A. steyermarki have the very minute, inconspicuous cephalic sensory tubercles that are widespread among colubrids. Both also have a larger kind of tubercle, which, in A. guerreroi, is discernible especially along the posterior edges of the prefrontals and anterolateral edges of the frontal. These organs are blackish and more numerous—or at least much more conspicuous—in A. steyermarki, where they cause the snake to look as if some of its head plates had been “riveted on”.33 in A. steyermarki, the structures are prominent along the posterior edges of the rostral, internasals, and prefrontals, somewhat less prominent along the lateral edges of the frontal, and conspicuously present laterally on the top edge of the loreal, posterodorsal edge of the nasal, and on the upper parts of supralabials 2–3 (fig. 71).

The male holotypes of Atractus steyermarki and A. guerreroi are of comparable size, the former only 4 mm longer in SVL (3 mm longer in tail length), but A. guerreroi appears to have a disproportionately smaller head, as shown by measurements (table 9) and as readily visualized in side-by-side comparison (fig. 72). The parietals and prefrontals appear to be of comparable size in A. guerreroi and A. steyermarki, but the internasals and frontal are disproportionately smaller in A. guerreroi. Atractus guerreroi has a relatively shorter snout, as also reflected in a shorter loreal, which is 64% of eye length in guerreroi vs. 92% in steyermarki. Both have 7 supralabials, but there is a slight difference in the configuration. In A. guerreroi, the first supralabial is far enough removed from the loreal that contact might seem unlikely in intraspecific variation, whereas in A. steyermarki the first supralabial is in point contact with the loreal on the left side and nearly in contact on the right side.

In summary, Atractus guerreroi most closely resembles A. steyermarki. The limited data suggest to us that the evolutionary course of A. steyermarki has been toward melanism and that of A. guerreroi toward reduced head size—hypotheses needing larger sample sizes.

Chironius fuscus (Linnaeus)

Figure 73

Fig. 73

Chironius fuscus (Linnaeus), ×0.7; the species is broadly distributed in the lowlands of northern South America. This specimen (BMNH 1976.235, subadult ♀) was caught on the summit of Auyantepui in August 1974; a shed skin found in 1994 indicates a possibly persistent presence of this racer on the Auyán summit.

Coluber fuscus Linnaeus, 1758: 222–223. “Habitat in Asia” in error.

Chironius fuscus: Roze, 1966: 97. Peters and Orejas-Miranda, 1970: 60. Dixon et al., 1993: 113, 261 (generic revision; BMNH specimen listed from “south side Auyantepui, 2283 m [in error for 6850 ft.  =  2088 m]”).

Material

Southern side of Auyantepui (“below final cliff face”), 6850 ft. [2088 m]: BMNH 1976.235, collected by Adrian Warren, August 17, 1974. Camp 3, 1850 m: AMNH R-155717 (fragment of shed skin, tentatively identified), from the 1994 AMNH–Terramar Expedition to Auyantepui.

The one whole summit specimen (fig. 73) is a female, probably subadult, 937 mm total length, 279 mm tail length (29.8% of total length). Dorsal scales completely smooth except for a few weak anal ridges laterally above cloacal region, in 10-10-10 rows; apical pits absent. Ventrals 158, subcaudals 103, paired; anal plate entire. Supralabials 9/8, 2nd–3rd touching loreal, 4th–6th in eye; infralabials 8, first five touching anterior genials. Preoculars 1, postoculars 2. Temporals 1 + 1. About 41 maxillary teeth on right. Color in preservative indistinctly marked with dark gray transverse bands; the bands are separated by white middorsal scales anteriorly on the body and by whitish transverse lines posteriorly. These markings along with black scale edges and small spots give an overall mottled appearance of gray, black, and white (fig. 73 top).34 Supralabials mostly yellowish white, with labials 2–3 having dark posterior edges. An ill-defined black postocular stripe extending from eye across lower part of primary temporal and upper part of ultimate supralabial. Underside of head immaculate yellowish white; black transverse lines across bases of anterior ventrals, these lines becoming heavier, with intervening black spots posteriorly (fig. 73 bottom).

The fragment of shed skin is from a small snake of juvenile size. The scales are relatively large, in 10 rows anteriorly, eight rows posteriorly; no keeled scales are discernible; apical pits absent over most of body, but relatively large pits are present on some scales posteriorly on the neck, with paired pits being more common than single ones. The eyes are relatively large; there are two postoculars and 1 + 2 temporals; both postocular scales are in contact with the primary temporal. Anal plate torn.

Material

Camp 2, 1750 m: AMNH R-140231, from the 1994 AMNH–Terramar Expedition to Auyantepui. Cabanayén, 1230 m, (about 70 km SE Auyantepui): AMNH R-104803, collected by S. Gorzula in 1975. Marco de Fronteira BV8, Roraima, Brazil: MZUSP 9235–9236, collected by C. M. Carvalho in 1986.

The single specimen (figs. 74–Fig. 7576) from our expedition to Autantepui is a juvenile female, 221 mm total length, 35 mm tail length (15.8% of total length); dorsal scales lacking apical pits, smooth, in 17-17-15 rows, with reduction occurring by fusion of rows 3 + 4 at ventrals 107 on left side/104 on right. Ventrals 174, subcaudals 54, paired; anal plate divided. Supralabials 8, 2nd–3rd touching loreal, 4th–5th in eye; infralabials 8, first four touching anterior genials. Maxillary teeth 16 + 2 on right side.

Coloration

In life (fig. 74), AMNH R-140231 was black with a thin, yellow nuchal bar followed by 40 ringlike yellow markings on body—the first 16 of these narrow crossbands being mostly unbroken above, the posterior ones broken dorsally, often with the two halves in alternating sequence. The ringlike crossbands are greenish yellow laterally and a deeper golden yellow dorsally. These markings are narrow, mostly about one-scale wide dorsally; they become wider on the lower sides, most conspicuously on the anterior body where they are about two-scales wide; markings are indistinct atop tail. Ventral and subcaudal surfaces transversely black-banded and checkered on pale greenish white. Some areas of the pale ventral color are transversely continuous with the much narrower yellow crossbands above, but the overall aspect is not of a “ringed” pattern (fig. 75).

Head black, with dull olive-yellow small spots on parietals and adjacent temporal region and with olive suffusions on snout. Supralabials heavily edged in black on light yellow (fig. 76); infralabials pale greenish yellow with black edging; genial plates and gular region white. Iris brown with a small, ill-defined patch of bronze above pupil. Tongue, including tips, black.

Venezuelan Specimens

Some previous assignments of specimens to this taxon are in error (see under Comparisons below). As documented herein, we know of Liophis trebbaui from six specimens—four from Venezuela and two from northern Brazil. The Venezuelan material includes our one specimen taken on the summit of Auyantepui at 1750 m, and the two type specimens collected on its south slope at about 1000–1100 m (see Comments on Type Specimens and Type Locality). A fourth Venezuelan specimen (AMNH R-104803) comes from Cabanayén, 1230 m elevation, about 70 km southeastward from Auyantepui (5°36′N, 61°44′W, fide Gorzula and Señaris, 1999: 246). According to the AMNH catalog, this specimen was collected at Cabanayén by Stefan Gorzula in 1975.

The specimen AMNH R-104803 (fig. 77) is a juvenile female that compares well with the one from the Auyán summit (compare figs. 74 and 76 with fig. 77). It is 200 mm total length, 35 mm tail length (17.5% of total length); dorsal scales lacking apical pits, smooth, in 17-17-15 rows. Ventrals 170, subcaudals 52, paired; anal plate divided. Supralabials 8, 2nd–3rd touching loreal, 4th–5th in eye; infralabials 10, first five touching anterior genials. Maxillary teeth 17 + 2 on right side. There is a narrow nuchal ring (a shorter nuchal bar in the Auyán specimen) and 40 pale ringlike markings anterior to the tail (as in the Auyán specimen). Many of the pale markings are medially broken, with the two halves sometimes offset; they are noticeably wider on the lower sides (fig. 77). The labials and parts of the genials are edged with black; pale ventral surfaces are banded and checkered with black (but less heavily marked than in the Auyán specimen).

This specimen was one of a small collection donated to the American Museum by Gorzula in 1977, for collaborative study with then Research Associate Janis Roze. The unidentified specimen was assigned a catalog number and tagged, but it was held aside until Roze's research materials were dispersed into the main collection in the late 1990s. The specimen is listed as Liophis sp. by Gorzula and Señaris (1999: 172, 250), who, however, gave the locality as “Parupa, CVG Agricultural Station”, 1220 m at 5°41′N, 61°31′W. The locality “Parupa” is shown as “Campo Parupa” on Huber and Berry's topographic map of the Venezuelan Guayana (issued in Steyermark et al., 1995), about 20 km NE Kavanayén. We assume Cabanayén ( = Kavanayén) to be correct, since this name was provided for AMNH cataloging, although it makes little difference for present purposes. Cabanayén and Campo Parupa are in reasonable proximity on the Gran Sabana, roughly 70–90 km SE–ESE of Auyantepui (some AMNH specimens collected by Gorzula are catalogued as “Parupa, nr. Cabanayen”). Gorzula and Señaris (1999: 246, 250) described Cabanayén as being in a “mixture of open savannas, scrub, and partially burned forest” and Parupa as “open savanna [and] riverine forest”.

Brazilian Specimens

Two specimens were collected by Dr. Celso Morato de Carvalho in February 1986, in Roraima, near Vila Pacaraima, border marker BV (Brazil-Venezuela) 8, about 4°29′N, 61°09′W. Myers visited the locality in June 1993, in company with Drs. Carvalho and W. R. Heyer. The site is a rocky stream east of town, at about 830 m elevation in a very dense and humid evergreen forest. The locality is some 200 km SE Auyantepui, suggesting that Liophis trebbaui may be distributed fairly extensively in the uplands east of the Río Caroní.

Both specimens are juvenile males. Following data are for MZUSP 9235 and 9236, respectively: Total length (tail length) 203 (35) and 222 (42) mm; tail length 17.2% and 18.9% of total length. Smooth dorsal scales lacking pits in 17-17-15 rows. Ventrals (subcaudals) 169 (53) and 173 (57); anal plate divided. Supralabials 8, 2nd–3rd touching loreal, 4th–5th in eye; infralabials 10/10 and 11/10, 1st–5th touching anterior genials. Maxillary teeth on right side 17 + 2 and 18 + 2. A narrow nuchal ring (as in fig. 77, not a dorsal bar as in fig. 75); 39 and 38 pale ringlike markings anterior to tail, the rings relatively complete in MZUSP 9236, more extensively broken in MZUSP 9235; some rings medially broken with the two halves slightly offset; all rings except first (nuchal ring) noticeably wider on lower sides. Pale markings in temporal region and on frontal plate similar to specimen in figure 77. The supralabials, infralabials, and genials are dark edged, relatively lightly so in MZUSP 9235, blacker in MZUSP 9236. Venter with transverse black bars (narrower than in fig. 75), many broken and offset to give a checkered appearance.

Natural History

Our specimen of Liophis trebbaui seems to be the first one from the Auyán summit. It is a juvenile found during the afternoon, in a shallow, rock-bottom pool containing Tepuihyla tadpoles (fig. 7 bottom). The diet of the similar Liophis torrenicola includes fish at 1000 m on neighboring Cerro Guaiquinima, but fish are absent from the higher summit of Auyantepui (although doubtless present on the lower slopes), where anurans and their tadpoles are the likely food source. No other specimens were seen despite intensive search.

Gran Sabana Specimens (L. miliaris sensu lato)

The “km marker 144” locality cited above for “Liophis cobella trebbaui” was referenced by Dixon (1983a: 164) to KU 167584, a snake collected by W. E. Duellman in 1974, at an elevation of 1210 m. Although the specimen (fig. 78) is here assigned to Liophis miliaris sensu lato, it may represent an unnamed species. It is a female 695 mm in total length, including 122 mm tail length (17.6% of total); dorsal scales lacking scale pits, smooth, in 17-17-15 rows; ventrals 176; paired subcaudals 55; anal plate divided; supralabials 8, 2nd–3rd touch loreal, 4th–5th in eye; infralabials 10/9, first five touching anterior genials; maxillary teeth 16 + 2 on right side. In life (Duellman's fieldnotes and transparency), the dorsum was black with a greenish yellow spot on each scale; venter cream with black markings; iris dark reddish brown; tongue black. The top of the head is black, with dense pale spotting; the supralabials, infralabials, and genials have heavy black margins (fig. 78).

Fig. 78

Liophis miliaris sensu lato, ×0.6. Dorsal and ventral views of KU 167584, from 1210 m elevation at km marker 144 on road from El Dorado to Santa Elena de Uairén, Bolivar, Venezuela. This specimen has been cited in the literature as “Liophis trebbaui”, a smaller, nonspotted snake (figs. 74–Fig. 75Fig. 7677).

A second, larger miliaris-like snake with a color pattern and scutellation virtually identical to the above is AMNH R-114801, collected by Gorzula in 1975 at Cabanayén (elevation 1230 m fide Gorzula and Señaris, 1999: 246). It is a female 860 mm in total length, 143 mm tail length (16.6% of total); ventrals 178, subcaudals 52. Head scales as above, except 10 infralabials on both sides; 15 + 2 maxillary teeth. A juvenile specimen of Liophis trebbaui (fig. 77) was collected at the same locality.

With total lengths of 695 and 860 mm, the aforesaid specimens (KU 167584, AMNH R-114801) here assigned to Liophis miliaris sensu lato are appreciably larger and more muscular than the adult holotypes of Liophis torrenicola (457 mm), L. trebbaui (570 mm), and L. ingeri (630 mm).

Liophis miliaris had not been explicitly recorded from Venezuela when Dixon (1983b) revised the species, although KU 167584 was the source for Hoogmoed's (1979b: 276) indication of the maximum elevation (1210 m) attained by L. miliaris and its placement in “Western Guiana” (M. S. Hoogmoed, personal commun.).37 Dixon's (1983a) mistaken assignment of that specimen (KU 167584) to “L. cobella trebbaui” presumably was based on meristic data, similarity in head color pattern, and knowledge that a pale-spotted color pattern occurs in several nominal species of Liophis. However, L. trebbaui does not have a spotted pattern—Dixon (1983a: 157) seems to have extrapolated this from Roze's (1958b: 304) description of L. ingeri, which Dixon regarded as a synonym of L. trebbaui. Roze mentioned una apariencia reticulada produced by pale areas on the head plates and dorsal scales on the anterior body of L. ingeri. This weakly spotted anterior pattern may prove diagnostic of L. ingeri, but it is not the vivid “salt and pepper” pattern that characterizes the larger Venezuelan specimens here assigned to L. miliaris (e.g., compare fig. 78 with photos of L. ingeri in Donnelly and Myers [1991: fig. 30] and especially in Gorzula and Señaris [1999: photo 114]).

The two specimens of Liophis miliaris s.l. mentioned above come from the northern part of Venezuela's Gran Sabana (southeastward of Auyantepui). They do not match descriptions of any of the seven subspecies recognized by Dixon (1983b, 1989). Liophis miliaris chrysostomus, of lowland Amazonia, is similar in having a heavily checkered black venter but, unlike the present specimens, the underside of the tail tends to be unmarked (see Dixon, 1983b: fig. 3C). Furthermore the black ventral checkering in chrysostomus is mostly straight-edged with sharp corners (as is commonplace among snakes with such patterns), whereas in the present specimens the heavy markings have mostly nonlinear edges giving a different aspect from normal checkering (compare fig. 78 with fig. 3C in Dixon, 1983b).

Other Specimens (L. m. miliaris)

We are aware of two additional Venezuelan specimens of Liophis miliaris. (1) AMNH R-114755 comes from Santa Elena de Uairén (Estado Bolívar), near the Brazilian border (some 130 km SE of the Cabanayén locality mentioned above). It is a female (preserved in a hardened state and not accurately measurable) that resembles the two Gran Sabana specimens in having dark-edged labials and genials, and a spotted (salt-and-pepper) color pattern, but which differs from them as follows: a lower number of ventrals (ventrals 159, subcaudals 51); head dark brown without pale spots; pale venter inconspicuously marked with thin, pale gray streaks across the ventral plates; subcaudal surfaces unmarked; pale spots on scales of the first dorsal row enlarged to cover most of the scales. The last character causes the pale color of the first-row scales to nearly fuse with the pale venter. In these aspects of color pattern, AMNH R-114755 resembles photographs of the Linnaean holotype of Liophis miliaris (Gans, 1964, fig. 23 [a composite of 5 photos]). Thus, the AMNH specimen from Santa Elena (misidentified as L. cobella in Gorzula and Señaris, 1999: 170) fits Dixon's concept of Liophis miliaris miliaris, which occurs nearby in northern Guyana (Dixon, 1983b: fig. 1).

(2) The second specimen is MHNLS 1634 from km 67 on the El Dorado–Santa Elena de Uairén road (Estado Bolívar); Kornacker et al. (2002: 32) give the coordinates as 06°18′N, 61°18′W. The locality is some 180 km (airline) N of Santa Elena de Uairén, roughly 130 km ENE of Auyantepui, and nearly 100 km NNE of Cabanayén. It was identified as Liophis miliaris miliaris by Kornacker et al. (2002), who provided a photograph in dorsal view and scale data including 156 ventrals and 52 subcaudals (sex not stated). The numbers of ventrals and subcaudals of MHNLS 1634 from km 67 are close to the 159 and 51 counts given above for AMNH R-114755 from Santa Elena de Uairén. These two specimens resemble Gran Sabana specimens (e.g., fig. 78) in the spotted salt-and-pepper dorsal pattern, but head and ventral patterns are quite different.

The above specimens of L. m. miliaris resemble each other in lacking conspicuous pale spotting atop the head and in ventral pattern. Gilson Rivas Fuenmayor kindly provided a photograph of MHNLS 1634 in preservative that shows an orange-tinged venter with markings virtually the same as those of AMNH R-114755—the venters are marked inconspicuously with thin gray streaks across some ventral plates, the subcaudal surfaces are immaculate, and pale spots on scales of the first dorsal row are enlarged and nearly fused with the ventral color.

Two Kinds of Liophis “miliaris” in Venezuela

Gans (1964) surveyed the variation and synonymy of Liophis miliaris (Linnaeus) in southeastern South America; he suggested that it is a polytypic species but did not formally recognize subspecies. Dixon (1983b) enlarged the study to include northern populations of other miliaris-like snakes. Dixon associated pre-existing names to seven subspecies, of which the two northernmost ones (miliaris and chrysostomus) appear to be widely allopatric (map in Dixon, 1983b: fig. 1); see Giraudo et al. (2006) for comments on southern populations. Dixon's (1983b: 796, 799) conclusion that the type specimen of Linnaeus' miliaris came from Suriname is particularly noteworthy, since it defined the nominotypical subspecies—L. m. miliaris—as the allopatric population in northern Guyana, Suriname, and French Guiana. The nominotypical form is now known in southeastern Venezuela from Santa Elena de Uairén (AMNH R-114755) and from km 67, about 180 km to the north (MHNLS 1634)—at elevations apparently well below 1000 m.

Liophis miliaris miliaris seems to be replaced at higher elevations (> 1000 m) in the Gran Sabana by an unnamed miliaris-like snake of very different aspect (KU 167584, AMNH R-114801). This form seems to differ from lower-elevation L. m. miliaris in having a higher ventral count, in having a conspicuously spotted head and, especially, in having a distinctively patterned ventral surface (fig. 78). The photograph of a living specimen from “Kavac, Gran Sabana” probably belongs to this population, judged from its vividly patterned head (Kornacker, 1995 [as L. r. zweifeli]; Kornacker et al., 2002: fig. 2).

On Liophis zweifeli

As demonstrated by Kornacker (1995) and Kornacker et al. (2002), specimens of Venezuelan Liophis “miliaris” can easily be confused with Liophis zweifeli (Roze) ( = Liophis reginae zweifeli sensu Dixon, 1983c), a snake with a similar salt-and-pepper dorsal pattern (Roze, 1966: fig. 39); the similarity is real and collections should be checked with that in mind. The taxon zweifeli can be distinguished from Venezuelan “miliaris” in having apical pits on the dorsal scales (although these may be few and hard to find), in a lower number of ventrals (140–148 fide Roze [1966: 165], 132–148 fide Dixon [1983c: 115), and in a longer tail with higher number of subcaudals (78–86 fide Roze [1966: 165], 69–88 fide Dixon [1983c: 127]).

Also, Liophis reginae zweifeli sensu Dixon possibly differs from Venezuelan L. “miliaris” in lacking black markings on the underside of head and lower part of the supralabials. The ventral color pattern seems to be variable in zweifeli; the holotype was described as having ventral scales heavily mottled with black on the posterior two thirds of the body (Roze, 1959: 5), but in an available paratype (AMNH R-59430) the venter is mostly pale, with minimal black edging laterally over most of the venter (subcaudals immaculate). A concise description of the taxon Liophis reginae zweifeli and re-evaluation of its relationships are needed. The extraordinarily variable and wide-ranging Liophis reginae (see map in Dixon, 1983c: 118) is likely a composite species; one species (Liophis oligolepis Boulenger) already has been resurrected from its synonymy, as summarized by Cunha and Nascimento (1993: 73–74).

Mastigodryas boddaerti (Sentzen)

Figure 79

Fig. 79

Mastigodryas boddaerti, ×0.87. The first specimen (AMNH R-61018) collected on the summit of Auyantepui, by the 1937–1938 AMNH–Phelps Venezuelan Expedition. This snake, an invader from the lowlands, appears to have maintained a persistent presence on the Auyán summit for many years.

Coluber boddaerti Sentzen, 1796: 59. Holotype and type locality unknown.

Eudryas b. boddaertii: Stuart, 1933: 2 (the double -ii termination used by various authors is an incorrect subsequent spelling).

Dryadophis boddaerti: Stuart, 1941: 66 (in generic revision).

Dryadophis boddaertii: Roze, 1958a: 264; 1966: 120 (Auyantepui specimens).

Mastigodryas boddaerti: Peters and Orejas-Miranda, 1970: 192 (new generic combination).

Material

Auyantepui Plateau, 2200 m (7218 ft.): AMNH R-61018, from the 1937–1938 AMNH–Phelps Venezuelan Expedition. Camp 2, 1750 m: AMNH R-155716 (shed skin), from the 1994 AMNH–Terramar Expedition to Auyantepui.

The one whole summit specimen (fig. 79) examined is an adult male 1036 mm total length, 286 mm tail length (27.6% of total length); dorsal scales with paired apical pits, smooth, in 17-17-15 rows. Ventrals 183, subcaudals 105, paired; anal plate divided. Supralabials 9, 2nd–3rd touching loreal, 4th–6th in eye; infralabials 10, first five touching anterior genials. Temporals 2/1 + 2. Maxillary teeth 17 + 2 on right. Color gray, presumably brown in life, with a faint remnant of a pale lateral stripe on scale row 4. (The juvenile pattern of M. boddaerti is very different, consisting of squarish dorsal blotches alternating with lateral ones.)

The shed skin from Camp 2 is fragmentary but identifiable. It has smooth scales with paired apical pits, in 17 dorsal scale rows anteriorly. Shapes of the dorsal head plates correspond to those of the whole summit specimen and another specimen (AMNH R-61035) from the base of Auyantepui, and the configuration of 2/1 + 2 temporals (on both sides of the head) is identical in all three.

Nomenclatural Comment

The available name boddaerti Sentzen (1796) is affixed to this species with complete arbitrariness—under a philosophy “often at a considerable stretch of the imagination … to use as many old names as possible” (Stuart, 1933: 1). The name is practically a nomen dubium and could well have been discarded as such. The fate of the original specimen is unknown, the type locality is unknown (the type locality cited in Kornacker [1999: 115] is that of a presumed synonym), and the original “description” could apply to many snakes (some of which are listed in the long synonymy in Boulenger, 1894: 11–12). The only data provided by Sentzen (1796: 59) were a count of ventrals and caudals (“Scut. 181 – 108  =  289”) and total length and tail measurements in old German feet (“Ganze Länge 2 Fuss, 8 Zoll; Schwanzlänge 7 Zoll, 7 Lin.”), for which there was more than one system.

Stuart (1941: 66) restricted the name to this species simply because Merrem (1820: facing pages 110) had early used Boddaertische and Boddaerti for a second specimen whose habitat was “vorzüglich [especially] Surinam”, which, in Stuart's imagination, suggested “Dutch Guiana as the type locality”. In its present sense, the specific name boddaerti has become well established through the influence especially of Stuart (1933, 1941), Roze (1966), and Peters and Orejas-Miranda (1970). A neotype should not be designated until there is actual need for defining the taxon more objectively.

Auyantepui Records

The snake known as Mastigodryas boddaerti has a broad geographic range and appears to be an invader from the lowlands with an established presence on the Auyán plateau. Records are few, but they span many years.

Roze (1958a: 265; 1966: 122) reported four specimens from Auyantepui, including AMNH R-61018 described above, and AMNH R-61035. The AMNH catalog shows the first as collected by W. H. Phelps on the summit at 2200 m elevation, in December 1937 or January 1938; the second was taken at at 460 m (base of Auyantepui) in February 1938. A third specimen (MBUCV 3073) was taken by P. Trebbau at 1020 m (Guayaraca) on the southern slope in April 1956. Two additional localities were given by Roze with identical catalog numbers (MBUCV 3072) and identical data except for one listed as “Auyantepui” and one as “El Oso, Auyantepui”, both said to have been collected at 1800 m by P. Trebbau on April 24, 1956. Presumably these two entries are a duplication and apply to a single specimen. Aside from being on the southern end of Auyantepui, the location of Trebbau's 1800 m “El Oso” locality is uncertain. Dunsterville (1965: 170, foldout map) mapped a “Bosque del Oso” but gave an elevation of about 2050 m and equated it roughly with the “2200-meter camp” of the 1937–1938 expedition.

In May 1989, Fabián Michelangeli photographed a cazadora identifiable as Mastigodryas boddaerti on the northern end of the western branch of Auyantepui (inaccurately cited as “Sima Aonda” in A. Michelangeli, 2005: 135, fig. 7 [F. Michelangeli, personal commun.]). During the 1994 expedition, we found only the shed skin at Camp 2, on the eastern branch of the Auyán plateau.

The above records, from 1937 to 1994, suggest that Mastigodryas boddaerti maintains a permanent presence on the summit of Auyantepui.

Material

West side of Auyantepui summit, 5197 ft. [1584 m], at 5°55′56.9″N, 62°39′32.2″W fide R. Guerrero (personal communication). Specimen not preserved, photographed March 22, 2006 by José Miguel Pérez G. (see fig. 80).

The photograph provides the only record of a fer-de-lance from the summit of any tepui.

“widespread” Guayana Highland Endemics

Hyalinobatrachium crurifasciatum, Hypsiboas sibleszi, Eleutherodactylus pulvinatus, Tepuihyla edelcae, Liophis trebbaui. The first three frogs are upland species found occasionally on tepuis, but Tepuihyla edelcae is known only from tepuis. The snake Liophis trebbaui, once thought to be endemic to Auyantepui, appears to be widely distributed in uplands east of the Río Caroní, from Auyantepui to the Brazilian frontier.

Endemic to Auyantepui Region

Anomaloglossus tepuyensis, Hypsiboas jimenezi. These occur on the Auyán slopes and in adjacent lowlands as well as on the summit.

Possible Auyán Summit Endemics

Oreophrynella cryptica, Centrolene gorzulai, Hypsiboas angelicus, Stefania schuberti, Eleutherodactylus auricarens, Anadia sp., Arthrosaura montigena, Tropidurus bogerti, Atractus guerreroi. These nine species are known at present only from the Auyán summit, but some (e.g., C. gorzulai, E. auricarens) seem likely to inhabit forest at lower elevations and might prove to be endemic to the Auyán massif broadly.41

4. Various authors have noted that a few genera of amphibians and reptiles are represented by obviously related species on different tepuis, a fact of obvious biogeographic significance. However, biogeographic scenarios for the highland herpetofauna are uncomfortably speculative, given the virtual lack of testable phylogenetic hypotheses and absence of molecular studies. Myers and Donnelly (2001: 80) simply observed that “tepui endemics are likely to have counterparts” on other tepuis—defining “counterparts” as a purposely vague term used for congeneric species judged to be phenotypically and ecologically similar, with phylogenetic closeness being probable.

With one notable exception, the Auyán endemics do have endemic congeneric counterparts on other tepuis (or, in the case of Centrolene, at least in other parts of the Guayana highlands), although degree of relatedness is unknown. The exception is the common lizard Tropidurus bogerti, whose phylogenetically closest relatives may occur south of the Amazon. Interestingly, T. bogerti does have a phenotypically and ecologically similar “counterpart” in the form of Plica lumaria on Cerro Guaiquinima.

Among the Auyán summit taxa, at least Oreophrynella (and perhaps Stefania and Tepuihyla) may represent part of a relatively old and widespread highland fauna in Pantepui. Certain other elements postulated as part of this fauna are absent from Auyantepui. The missing taxa include the genus Riolama, species of the Phenacosaurus neblininus complex, and species of the dwarf Thamnodynastes complex, although members of the last two groups occur in the neighboring Chimantá massif. Such speculation assumes (1) that the involved species comprise monophyletic highland groups rather than separate invasions from lower elevations, and (2) that spotty distribution patterns are the result of random extinction events as well incomplete collecting.

5. Lowland species of amphibians and reptiles “doubtless invade all tepuis, although they may not necessarily reach the highest summits … or, if they do, they may not establish permanent populations there [and although] lowland invaders are expected they can seldom be successfully predicted” (Myers and Donnelly, 2001: 81). The summit of Auyantepui has been invaded by at least the following eight species from lower elevations: Hyalinobatrachium taylori, Leptodactylus rugosus, Neusticurus rudis, Anolis chrysolepis planiceps, Chironius fuscus, Leptodeira annulata ashmeadii, Mastigodryas boddaerti, Bothrops atrox.

Of these only the Anolis seems to be a regular invader of tepuis, although there are tepui records for a few of the other genera and species as well (for records, see McDiarmid and Donnelly, 2005; Señaris and Ayarzagüena, 2005: 230, for H. taylori).

Venomous snakes have rarely been found on tepuis. The present report of Bothrops atrox is only the second record of a viper from a tepui summit, the other being the holotype of Bothrops lichenosa ( =  Bothriopsis taeniata) collected in 1955 on the summit of Chimantá Tepui. Coral snakes occur as rarely; Myers (2005) pictured a specimen of Micrurus psyches from the lower part (1400 m) of the summit of Cerro Guanay.

Whether lowland invaders can maintain permanency on tepui summits is information hard to come by. However, documentation provided herein for Mastigodryas boddaerti indicates that this lowland racer has maintained a presence on the Auyán summit for nearly 60 years at least. And, judged from the number of records, the lowland snake Leptodeira annulata may be a permanent resident of the Chimantá massif, although only a single specimen is known from Auyantepui.