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Physalaemus Crombiei (Amphibia: Leptodactylidae), a New Frog Species from Espirito Santo, Brazil with Comments on the P

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Physalaemus Crombiei (Amphibia: Leptodactylidae), a New Frog Species from Espirito Santo, Brazil with Comments on the P PROC. BIOL. SOC. WASH. 102(2), 1989, pp. 500-506 PHYSALAEMUS CROMBIEI (AMPHIBIA: LEPTODACTYLIDAE), A NEW FROG SPECIES FROM ESPIRITO SANTO, BRAZIL WITH COMMENTS ON THE P. SIGNIFER GROUP W. Ronald Heyer and Alan J. Wolf Abstract.— Physalaemus crombiei, a new species of the P. signifer group, is described from the State of Espirito Santo, Brazil. Members of this group are most easily distinguished from each other by advertisement call, although morphological differences also exist. The relationships and distributions of members of this species group are not well understood at present. One species group of frogs of the genus lack of parotoid glands, small to large in- Physalaemus breeds in forest puddles and/ guinal glands, slender build and a size range or swamps within the Atlantic Forest sys- of 15-35 mmSVL. tem of Brazil. Members of this group are Physalaemus crombiei differs from P. dei- more easily distinguished from each other maticus Sazima and Caramaschi 1986 by advertisement calls than by their mor- (which apparently does not belong to any phologies. Perhaps because of this, several of Lynch's (1970) species groups) by the fol- species have been described since field re- lowing characteristics: relatively smooth cording equipment has become more avail- skin (granular in deimaticus), slender build able. We describe another new species of (stocky), and inverse V-shaped marks on this group, to be known as: back (absent). Physalaemus crombiei can be distin- guished from other members of the P. sig- Physalaemus crombiei, new species nifer group [P. signifer (Girard, 1853), P. Fig. 1 olfersi (Lichtenstein & Martens, 1856), P. Holotype.—MZVSP 66252, adult male nanus (Boulenger, 1888), P. maculiventris from Brazil: Espirito Santo; adjacent to Re- (A. Lutz, 1925) (Frost 1985:253, is in error serva Biologica Nova Lombardia, near San- in placing maculiventris in the cuvieri group), ta Teresa, approximately 19°55'S, 40°36'W. P. obtectus Bokermann, 1966, P. boker- Collected on 31 Dec 1977 by one of collec- manni Cardoso & Haddad, 1985, and P. tors of paratopotypes. franciscae Heyer, 1985] as follows. Physa- Paratopotypes.-MZUSP 66253-66284, laemus crombiei differs from bokermanni, USNM 285038-285070, taken by Ronald obtectus, olfersi, and signifer by size (P. I. Crombie, Christina Duchene, W. Ronald crombiei 19-22 mm SVL, bokermanni 15- Heyer, and Francisca C. do Val from 28 Dec 17 mm, obtectus 22-27 mm, olfersi 30-35 1977 to 1 Jan 1978. mm, signifer 24-31 mm). Physalaemus Diagnosis.— Physalaemus crombiei is a crombiei also differs from olfersi by the ab- member of the P. signifer group. The sig- sence of a light outline on the ventral border nifer group differs from other Physalaemus of the lateral stripe from the eye to the arm species groups (Lynch 1970) by the follow- (present in olfersi). It is distinguished from ing characteristics: absence of an inner tar- bokermanni, maculiventris and nanus by the sal tubercle, first finger shorter than second, presence of tubercles on the sole of the foot VOLUME 102, NUMBER 2 501 (absent). Physalaemus crombiei differs from foot with four series of small but distinct obtectus by the lack ofnumerous small round fleshy tubercles. markings closely associated with the inverse Color in preservative: Dorsum creamy tan V-shaped marks on the back, by its sharp with indistinct brown markings including canthus rostralis, and pointed snout (nu- an inversed V-shaped mark in shoulder area, merous small round marks associated with an indistinct interorbital mark, a pair of the dorsal pattern, rounded canthus ros- acute marks in sacral area, and a pair of tralis. less pointed snout in obtectus). Phys- very distinct black, white-outlined spots, one alaemus crombiei has a relatively light col- on each upper portion of lumbar glands; ored posterior belly differing from the boldly sides of head mottled brown, distinctly sep- dark and light mottled posterior belly of arated along canthus rostralis from lighter franciscae and maculixentris. dorsal color and leaving a light stripe on Description ofholotype.— Snout subellip- very tip of snout; tympanum brown, distin- tically pointed from above, rounded in pro- guished from surrounding areas by color file; canthus rostralis rather sharp in cross pattern; distinct dark (almost black) supra- section; tympanum barely discernible ex- tympanic stripe extending along flank to- ternally, large, about two-thirds eye diam- wards belly, terminating below mid-lumbar eter; maxillary teeth not visible, but dis- gland level, distinct dorsal dark stripe with cernible by probe: vomerine teeth absent; white outline, brief white outline from pos- vocal slits present, elongate: vocal sac sin- terior tympanum midway to shoulder ven- gle, noticeably but not greatly expanded ex- trally, rest of stripe blending into ventral ternally, extending to level in front of arms; pattern and not distinct ventrally; upper limb finger tips slightly expanded, not disked: fin- surfaces indistinctly cross-banded, fore- gers lacking fringe and web: subarticular tu- arms with darker brown blotch medially, bercles large, globose, pungent; pair of tan each outer shank with series of three black- nuptial pads on inner surface ofeach thumb, ish spots: throat dark brown with small, light rounded-oblong medial pad associated with irregular spots; anterior half ofbelly mottled inner metacarpal tubercle, narrowly sepa- brown and cream, posterior belly cream; rated from elongate distal pad: no prepollex; ventral forearm region dark brown laterally, no ulnar ridge; forearm slightly hypertro- irregularly but distinctly separated from phied; extremely weak supratympanic fold: lighter dorsal color pattern; ventral surface no dorsolateral fold: large ovate lumbar of thighs mottled brown and cream, lighter gland in groin: dorsal skin texture smooth medially, posterior portions with small white in appearance but with scattered small fleshy dots; outer tarsus brown medially with ir- tubercles, especially on upper eyelid; pos- regular dark brown elongate markings cen- terior part of vocal sac wrinkled, rest of trally with lighter dorsal color laterally; sole throat and belly smooth, under surfaces of of foot brown, subarticular and metatarsal thighs areolate; toe tips slightly expanded, tubercles cream: light dorsal pattern dipping toes III and IV most expanded; toes without irregularly but with distinct border onto webs, sides of toes with or without weak posterior surface of each thigh, lowest on ridges; subarticular tubercles globose, medial half of thigh, remainder of lower rounded, pungent; well-developed rounded portions of thigh almost uniform brown, metatarsal tubercle about half the size of with a few ventral white dots. well-developed oblong inner metatarsal tu- Snout-vent length 20.3 mm, greatest bercle; short, weakly developed inner tarsal tympanum diameter 1.7 mm, head length fold on distal one-fourth of tarsus, narrowly 7.0 mm, head width 7.7 mm, thigh length separated from inner metatarsal tubercle, 9.3 mm, shank length 9.3 mm, foot length remainder of tarsus smooth: heel smooth: 9.7 mm. 502 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Holotype of Physalaemus crombiei (MZUSP 66252, SVL = 20.3 mm, male), dorsal and ventral views. Variation. — The dorsal patterns range The following measurements are based from almost uniform brown to distinct dark on 7 adult male (listed first) and 14 adult interorbital bands and two dark, white-out- female (listed second) paratopotypes: SVL lined confluent arrows, one on the mid- 18.9-20.0 mm, 19.2-21.8 mm; head length scapular area, the other on the mid- sacral 33-38% SVL (mean 35%), 32-36% SVL area. In a few specimens the interorbital and (mean 34%); head width 35-39% SVL (mean arrow markings join. In some individuals a 37%), 33-36% SVL (mean 35%); thigh length distinct small mid-scapular light spot is 43-48% SVL (mean 46%), 42-46% SVL present. One individual (USNM 285048) (mean 44%); shank length 45-48% SVL has a faint but distinct light mid-dorsal pin (mean 47%), 43-48% SVL (mean 45%); foot stripe from the tip of the snout to just past length 44-49% SVL (mean 47%), 40-46% the sacrum. Most individuals lack a light SVL (mean 44%). border posterior to the tympanum defining Color in life. —Brief color notes are avail- a portion of the ventral border of the dark able for two individuals. In both, the iris lateral stripe. The stripe is distinct (black) was bronze. In USNM 285051 a slight red in most individuals, and fainter but distinct wash occurred in the groin and front and in a few. The dark spots on the lumbar glands back of the thighs. In USNM 285052 the are distinct in all individuals. The ventral groin and belly had a pinkish cast and the pattern described for the holotype is char- dorsum was red-brown and brown. acteristic of nearly all individuals, although Advertisement call— Data from a series a couple have lighter bellies than does the of calls presumably given by a single un- holotype. In some individuals the dorsal and captured individual; another captured in- posterior thigh patterns are distinct only dividual (USNM 285070) was heard to give medially, near the vent; most have the same a similar call. Call rate approximately one/ pattern as described for the holotype. sec; average call duration 0.376-0.418 sec VOLUME 102, NUMBER 2 503 4 N 3 UJ 2 j » --*-* o .». *T. «*_ _l 1 i 1 1 1 1 1 2 SECONDS Fig. 2. Advertisement call of Physalaemus crombiei. Wave form of second call shown on audiospectrogram, total signal length 0.428 sec. Recorded (no voucher) by W. R. Heyer, 30 December 1977, air temperature 21. 5°C (USNM Tape 7, Cut 5). (mean 0.399 sec); interval between calls (and other species within the genus) or their 0.521-0.651 sec (mean 0.574 sec); 8-10 distributions.
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