Described from the Rainforests of Northeastern Madagascar (Microhylidae: Cophylinae)

Home , Cophylinae, Mantellidae, Plethodontohyla

Two Unknown Arboreal Frogs (genus Platypelis) Described from the Rainforests of Northeastern Madagascar (Microhylidae: Cophylinae)

FRANCO ANDREONE1*, DANTE B. FENOLIO2, AND MARK E. WALVOORD2

1Museo Regionale di Scienze Naturali , Sezione di Zoologia, Via G. Giolitti, 36 ,I-10123 Torino, ITALY 2University of Oklahoma , Department of Zoology, 730 Van Vleet Oval, Richards Hall, Norman, Oklahoma, USA

Abstract: Two new arboreal microhylid frogs are described from the rainforests of northeastern Madagascar. Platypelis tetra is a very small frog, reaching about 20mm SVL, and inhabits phytotelms in screw Pandanus pines. It was found in some low and mid-altitude rainforests, such as Anjanaharibe-Sud, Besariaka, Tsararano, and Masoala Peninsula. This species diverges from the other Platypelis by its small size and colouration, with a series of whitish spots on the back, of which four are more evident. Furthermore, its advertisement call is composed by a long series of discrete notes at about 3.5-4KHz of frequency, and a repetition rate of about 3.0notes/s. The second species, Platypelis mavomavo, reaches a larger size (about 30mm SVL), and is characterized by a yellowish colouration of the ventral surface, and a dorsal surface with a network of dark spots on a beige-yellow background. It is currently known from Anjanaharibe-Sud and Ambolokopatrika, around the Andapa water-basin, but is expected to have a wider distribution.

Key words: Amphibia; Anura; Microhylidae; Platypelis; New species; Madagascar

this landmass belong to a few families, of INTRODUCTION which they represent an offshoot radiation. Madagascar is well known for its biodiversity According to recent studies (Vences and and high endemism (Glaw and Vences, 1994). Glaw, 2001) they are included in four families: Amphibians are no exception with more than Ranidae, Mantellidae, Microhylidae, and Hyper- 210 endemic species (Andreone and Luiselli, oliidae. Of these, the microhylids are still the 2003) and many others still in wait for most enigmatic Glade, and their phylogenetic description. Like other vertebrates of Mada- and taxonomic relations are largely unstudied. gascar (e. g., fish and mammals, see Benstead Of the three recognized Malagasy subfamilies et al,, 2000; Yoder et al., 2003), the frogs of (Dyscophinae, Scaphiophryninae, and Cophyli- nae), the cophylines account for a high number *Corresponding author. Tel: +39-011-4320- of species inhabiting rainforests and secondary 7310; Fax: +39-011-4323-331. altitude savannahs (Andreone,1999). Indeed, E-mail address:[email protected] little is known of their ecology and distribution 92 Current Herpetol. 22 (2) 2003 except that they are specialized to a variety deforestation outpaces description. Written of microhabitats: e. g., the enigmatic Rhom- records and museum specimens might become bophryne testudo and some Plethodontohyla all we have for future studies of the most species (such as P. ocellata, P. alluaudi, and sensitive species, which are rapidly vanishing. P. serratopalpebrosa) are fossorial or terres- The present paper is just a step in this trial, the small Stumpffia are adapted to leaf direction and deals with the description of two litter, and most of the Anodonthyla, Platype- new Platypelis that we found in northeastern lis, Cophyla, and some Plethodontohyla are Madagascar. One of these frogs is a Pandanus- arboreal or semiarboreal. The conditions and specialized Platypelis, while the second is an evolutionary pressures that led to such a arboreal generalist. They are sufficiently dramatic ecological radiation in these frogs are divergent from all known Platypelis, in both unknown. Further, whether an ecologically morphology and bioacoustics, to warrant specialised group is indeed monophyletic is the description here. subject of a separate work (Andreone et al., in press). MATERIALS AND METHODS One reason for the lack of knowledge about cophyline microhylids is that they often lead a We searched for frogs during the night with cryptic life, which makes their study difficult. the help of flashlights. Vocalizing males were In fact, many taxa are known from a few located by following calls at sunset or during specimens and are only occasionally re- rainfall. One species described here is a micro- discovered (e.g., Plethodontohyla coudreaui; habitat specialist, living within phytotelmata Andreone, in press a). This was confirmed by of screw-pines, Pandanus sp., and we spent our own experience: during field surveys we daytime hours searching the leaf axils of these often found specimens that could be grouped plants. After capture and photography of live with one another representing the same eco- colouration, specimens were sacrificed after logical and morphological cluster, but were anaesthesia in chlorobutanol, fixed in 4% difficult to assign to any known taxon. This formalin or 90% ethanol, and preserved in leads us to the idea that, especially in micro- 70% ethanol. They are currently deposited in hylids, many more species await description, the Museo Regionale di Scienze Naturali, and it is not an exaggeration to affirm that Torino (MRSN). most likely the final number of species will sta- Measurements follow standard methods, bilize around the double that currently known. using digital callipers or the dissecting micro- Many of these specimens still await description scope's micrometer (to the nearest 0.1mm): and will be the object of further contributions, SVL (snout-vent length), HW (head width), using both traditional (e.g., morphological HL (head length, from the maxillary commis- analysis and comparisons of museum sam- sure to the snout tip), ED (horizontal eye ples), and other methods (e.g., acoustics, kary- diameter), END (eye-nostril distance), NSD ology, DNA). While it is important to collect (nostril-snout tip distance), NND (inter-narial ecological and natural history information on distance), TD (horizontal tympanum diameter), unknown species, the discovery and rapid HAL (hand length, from the carpal-metacarpal description of new frogs from Madagascar is articulations to the tip of the longest finger), important for two reasons: (1) it accelerates FORL (forelimb length, from the axilla to the the cataloguing of the whole Malagasy batra- tip of the longest finger), HIL (hindlimb chofauna, providing powerful tools to draw length, from the cloaca to the tip of the biogeographic relationships for conservation longest toe), FOL (foot length, from the actions (see Andreone and Luiselli, 2003), and tarsal-metatarsal articulations to the tip of the (2) it helps clarify the taxonomy and phylog- longest toe), FOTL (foot length including eny of this subfamily in a region where tarsus, from the tibiotarsal articulation to the ANDREONE ET AL. -TWO NEW FROGS FROM MADAGASCAR 93 tip of the longest toe), TIBL (tibia length). A2164-2165, one juvenile and two adult When available, type specimens and/or other males, Ambohidroina Forest, Masoala Penin- representative specimens of presumably related sula, Mahalevona Fivondronana, Toamasina taxa from zoological collections were exam- Faritany (Tamatave Province), 15°26.00'S, ined, as well as the original descriptions and 49°57.56'E, 840m asl, collected by J. E. subsequent works (e. g., Blommers-Schlosser Randrianirina, 26 January 2002; MRSN A2166, and Blanc, 1991; Glaw and Vences, 1994). The one juvenile, same date and collector, 28 advertisement calls were recorded with a SONY January 2002; MRSN A2167, an adult male, TC-D3 tape recorder with external micro- same date and collector, 29 January 2002; phone and analyzed with a VOXYS 3.1 sound MRSN A2169, one juvenile, same date and system (Andreone et al., 2003). The proposed collector, 30 January 2002; MRSN A2170, common names for the new species are provided in the authors' original languages (Italian and English) and in Malagasy.

Platypelis tetra sp. nov. Four-spotted tree cophyline frog (English), cofilino degli alberi dalle quattro macchie (Ital- ian), sahonkely anatihazo misy tebok'efatra (Malagasy). Figs. 1-3

Type series FIG. 1. Platypelis tetra. Holotype (male, MRSN Holotype: MRSN A2174, adult male, Camp- A2174) from Analabe Valley, Anjanaharibe-Sud, site W2, Anjanaharibe-Sud Massif, Analabe western slope, NE Madagascar. Valley, Befandriana Fivondronana, Mahajanga Faritany (Majunga Province), 14°46'S, 49°26'E, 1,250m asl, collected by F. Andreone, H. Randriamahazo, and J. E. Randrianirina, 5 February 1996. Paratypes: MRSN A2171-2173, three adult females, same data as the holotype; MRSN A1952-1953, an adult male and an adult female, Tsararano Forest, Andatony Anivo, Andapa Fivondronana, Antsiranana Faritany (Diego Suarez Province), 14°54.8'S, 49°42.6'E, 600m asl, collected by F. Andreone and J. E. Randrianirina, 5 and 10 December 1996; MRSN A2175, one juvenile, Besariaka Forest, Andapa Fivondronana, Antsiranana Faritany (Diego Suarez Province), between 14°49'S-14°50'S, 49°35'E-49°36'E, about 700m asl, collected by J. E. Randrianirina, 1 May 1996; MRSN A646.1-4, three adult males and one juvenile, Masomihenjina Forest, Masoala Peninsula, Maroantsetra Fivondro- nana, Toamasina Faritany (Tamatave Prov- FIG . 2. Particular of the head (A), hand (B), ince), 15°25'S, 49°46'E, collected by R. and foot (C) of the holotype of Platypelis tetra Nincheri, 24 July 1993; MRSN A2161, (male, MRSN A2174). 94 Current Herpetol. 22 (2) 2003

at the free margin, no groove or notch, free behind for about two thirds of its length. Pupil horizontal. Vocal sac moderate in size, single, and in gular position. Skin on dorsum of head, body, belly, and limbs smooth. Fingers moderate in length bearing circular, moderately sized discs compared with phalanges; disc on first digit smaller than others: 1<2<4<3; subarticular tubercles faint, circular, not elevated; super- FIG. 3. Sonagram of part of a call (two notes numerary tubercles absent; palmar tubercle out of a longer note series) of Platypelis tetra, semi-distinct, not elevated. No webbing between recorded at Anjanaharibe-Sud, NE Madagascar. fingers. Feet with semi-distinct supernumerary Recording temperature: 18C. tubercles; vestiges of metatarsal tubercle; no other distinct tubercles. Small pads on toes, same date and collector, 9 February 2002. circular in shape. No webbing between toes. Toe length: 1<2<5<3<4. Diagnosis An arboreal Platypelis characterized by the Coloration following combination of characters: small After five-seven years of preservation in adult body size, canthus rostralis indistinct, toe ethanol, all the specimens still show contrast- and finger pads more circular than ovoid and ing colouration. Dorsum of head, body, and moderate in size; hands and feet without limbs are brown or tan with vague darker webbing, tarso-metatarsal articulation reach- markings, and legs have irregular tan spots. ing the eye, dorsal surface smooth, dorsal Four bigger dorsal whitish spots are present on colour shading from brown to tan, sometimes the back (two just behind the head, and two with irregular markings and at least four other on the sacral bone). Other smaller scat- oblong white spots on the dorsal surface. tered light points are also present of the back and superior parts of legs in some individuals. Description In some cases the four main spots form a The SVL in the examined males ranged sort of hourglass figure on the dorsum. Flanks from 15.7 to 18.2mm. In the females from with a brown dorsolateral band beginning 17.5 to 19.4mm. Other measurements of the posterior to the eye, passing through the union type series are given in Table I. Head as long of the arms with the body and down each as wide; HW 32-41% of SVL; HL 31-41% flank, ending at the groin; brown bar is edged SVL; snout short, slightly protruding beyond above and below by a thin cream line. The margin of lip; rounded and blunt in dorsal anterior surface of the arm with a brown line view and in profile; END less than ED; END extending from the union of the arms with the 20-26% of HL; eye moderate in size; ED body and terminating at or before the elbow. 28-36% of HL. Top of head flat; cranial Iris brownish shading to copper, with periph- crests absent; canthus rostralis indistinct; eral fine spots and dark shadings. Ventral internarial area not depressed; nostril more surface grey to creamish with a few brown circular than ovoid; protruding laterally. Weak flecks, sometimes forming dark spots. supratympanic fold usually visible, tympanum round, TD 34-67% of ED. Choanae small, Variation round, separated, partially obscured by pala- The paratypes come from Anjanaharibe, tal shelf of maxillary arch. Vomerine teeth Besariaka, Tsararano, and Masoala, spanning present and posterior to choanae. Tongue therefore about 200km. These specimens trapezoid in shape with rounded edges, widest agree with the holotype in colouration. The ANDREONE ET AL. -TWO NEW FROGS FROM MADAGASCAR 95

TABLE 1. Morphometric measurements (in mm) of type specimens of Platypelis tetra sp. nov. and P. mavomavo sp. nov. M=male, F=female, J=juvenile, holotype marked with an asterisk. For abbreviations see the text.

individuals from Masomihenjina (Masoala) Natural history and distribution show a paler dorsal colouration, and a The holotype and some of the paratypes contrasted lateral band. Colouration in life were collected at Anjanaharibe-Sud, western was basically similar to that described for slope (Analabe Forest), Campsite W2 (around preserved animals, although much more vivid. 1200m asl), where the habitat is comprised of a mid-high altitude transitional rainforest with Etymology tall trees and abundant lichens, mosses, and The word "tetra" is derived from the Greek ferns (Raxworthy et al., 1998; Andreone et al., "tetras" means "four" and is attributed to the 2000). The specimens were found within a fact that most specimens show four lighter plant of the genus Pandanus. A single speci- spots on the back. This name is used as a men was found at Besariaka Forest. This noun with the same meaning. forest at about 60km south of Andapa is rather degraded, especially in areas far from streams, due to cattle, cutting of trees by 96 Current Herpetol. 22 (2) 2003 villagers, and use of path systems. The speci- dorsal surface, in the possession of white men was found within a Pandanus, and did dorsal spots, and brown dorsolateral bars; (3) not call. Other paratypes came from Tsa- from P. occultans in lacking webbing between rarano. The mountain chain and forest of the the toes, a triangular marking on the back, in same name lie midway between the Anjanaha- a tibio-tarsal articulation extending to the ribe-Sud Massif and the Masoala Peninsula, eye, in the possession of dorsal spots, and in and is formed by several hills. The forest was the presence of the dorsolateral bands. at that time (1996) mostly intact, perhaps due Within the wealth of Platypelis species, P. to its distance from large-sized villages. The tetra is morphologically similar to P, tuber- remainder of specimens were found at ifera, although the latter reaches a larger size Masoala at two low altitude sites (Masomi- (up to 40mm). Platypelis tetra could be henjina and Ambohidroina), always within misidentified with juveniles of this species, Pandanus. This species is evidently special- from which it differs in lacking toe webbing, ized to this plant, and is a phytotelm the absence of a dorsal median line, the frequenter (Lehtinen, 2003). Concerning the possession of white dorsal spots, and in the distribution, P. tetra was until now found only possession of a brown dorsolateral bar in the northeastern sector of Madagascar, an bordered in cream. area which is interesting in terms of endemicity and species diversity (Andreone, in press Platypelis mavomavo sp. nov. b). We do not have evidence of a wider Yellowish tree cophyline frog (English), distribution, although we strongly suspect that cofilino arboreo giallastro (Italian), sahonkely its presence runs southward, and possibly, mavomavo (Malagasy) northwards. Figs. 4-6

Advertisement calls Type series Calls were recorded on 5 February 1996 Holotype. MRSN A2435, adult male, between 1830 and 1900h, at an air tempera- Ambolokopatrika, Antsinj orano, 14°32.6'S, ture of 18.0-18.5C. They consisted of an 49°25.8'E, 975m asl, Andapa Fivondronana, almost endless series of chirping metallic Antsiranana Faritany (Diego Suarez Province), notes. Temporal call parameters were: note collected by F. Andreone, G. Aprea, and J. E. duration 171-318ms (x±SD=266±41ms, Randrianirina, 10 December 1997; Paratypes. N=17), duration of inter-note intervals 148- MRSN A2433, an adult female, same data and 398ms (226±59ms, N=13), note repetition collectors, 18 December 1997; MRSN A2431, rate 3.0/s. The frequency showed a simple adult female, Ambolokopatrika, Andemakat- pattern, with a dominant frequency between sara, 14°31.8'S, 49°26.5'E, 875m asl, collected 3500-4000Hz. by F. Andreone and J. E. Randrianirina, 2 June 1997; MRSN A2432, adult male, Amboloko-

Comparisons with other species patrika, Andranomadio, 14°32.4'S, 49°26.3'E, Of the other Platypelis species possibly 890m asl, collected by F. Andreone, G. Aprea, sympatric and similarly sized, P. tetra differs: and J. E. Randrianirina, 1 December 1997; (1) from P. barbouri in lacking webbing MRSN A2434, adult male (dissected for kary- between the toes and fingers, in lacking the ological analyses), Ambolokopatrika, Antsin- red or green markings on the dorsal surface of jorano, collected by F. Andreone, G. Aprea, the groin and thighs, in the presence of white and J. E. Randrianirina, 1 December 1997. dorsal spots, and of a brown dorsolateral bar bordered in cream; (2) from P. cowani in Other specimens lacking webbing between the toes, in the Platypelis mavomavo was first discovered in absence of a beige triangular marking on the January 1996 at Anjanaharibe-Sud, western ANDREONE ET AL. -TWO NEW FROGS FROM MADAGASCAR 97

FIG. 4. Platypelis mavomavo. Holotype (male, MRSN A2435), dorsal view, from Amboloko- patrika, NE Madagascar.

FIG. 6. Platypelis mavomavo. Holotype (MRSN A2435), ventral view.

beyond margin of lip; rounded in dorsal view (in profile, lightly truncated); END less than FIG. 5. Particular of the head (A), hand (B), ED; END 23-30% of HL; eye moderate in and foot (C) of the holotype of Platypelis size; ED 34-42% of HL. Top of head flat; mavomavo (male, MRSN A2435). cranial crests absent; canthus rostralis indistinct; internarial area not depressed; nostrils slope. The specimen discovered was photo- protruding dorsally and laterally. Supratym- graphed but not captured. panic fold visible, tympanum round, TD 41- 47% of ED. Choanae small, round, and sepa- Diagnosis rated. Vomerine teeth present and posterior An arboreal species of Platypelis character- to choanae. Tongue trapezoid in shape with ized by the following combination of charac- rounded edges, widest at the free margin, with ters: medium adult body size (SVL=35mm), a slight frontal notch. Vocal sac moderate in yellowish colouration, with extended brownish size, single, and in gular position. Skin on marblings, digital expansions triangular in the dorsum of head, body, and limbs quite smooth, fingers and ovoid on toes, vestigial webbing although in some specimens elongated warts between the toes, dorsal surface from smooth are visible. Ventral surfaces smooth. Forearm to slightly warty. broad, fingers moderate in length bearing subtriangular, large discs; disc on first digit Morphological description smaller than others: 1<2=4<3; subarticular The three males ranged in size from 24.4 to tubercles faint, circular; supernumerary tuber- 28.6mm; the two females were respectively cles absent; palmar tubercle semi-distinct, 26.8 and 27.9mm long. Other measurements not elevated. No webbing between digits of of the type series are given in Table I. Head as hands. Feet with semi-distinct supernumerary long as wide; HW 34-40% of SVL; HL 33- tubercles. Small pads on digits of feet, ovoid in 35% SVL; snout short, slightly protruding shape. Vestigial webbing between third and 98 Current Herpetol. 22 (2) 2003

fourth, and between fourth and fifth toes. Toe likely that, similarly to other species (e. g., P. length: 1<2≦5<3<4. pollicaris) it may hide during the day and dry periods in trunk-holes, under barks, and at the Coloration basis of epiphytic plants. Given its presence at After about six years in preservative solu- Ambolokopatrika and Anjanaharibe-Sud, we tion, the dorsum of this new Platypelis has expect the species' distribution to include become light brownish, with scattered lighter other areas of northeastern Madagascar. spots and reticulations. The legs have approx- imately the same colouration as the back, with Comparisons with other species some pigmented transverse areas or spots. Platypelis mavomavo is most similar to Most of the hands and feet are brown or black. Platypelis cowani and P. pollicaris. They are Pupil horizontal, iris yellowish with dark spots. different from P. mavomavo especially in The ventral surface is cream/yellow with colouration. In particular, both have a brown some faint dark marbling, more visible on the dorsum, with light spots, and their belly is throat. In life, the colouration is similar in typically white. Platypelis tsaratananaensis pattern, although more distinct. The yellow is (at least the specimens from Anjanaharibe- conspicuous and bright in live animals (espe- Sud) has a more slender body, and it hides in cially on the belly), but disappeared after bamboo internodes. preservation in formalin and/or ethanol. DISCUSSION Variation The type series consists of four specimens The discovery of the new Platypelis species from Ambolokopatrika, quite similar in mor- described herein, at some localities of north- phology and size to the holotype. eastern Madagascar highlights the incomplete knowledge of the microhylid fauna of Mada- Advertisement calls gascar. Platypelis tetra is likely the smallest Unknown. arboreal frog of Madagascar, reaching a SVL of only 18mm. It is smaller than P. barbouri Etymology (23mm), Anodonthyla boulengeri (23mm), The specific name mavomavo (pronounce: and A. nigrigularis (24mm) (records from "mow-woo mow-woo") is a Malagasy term: Glaw and Vences,1994). This small size cou- "mavo" means yellow , and the duplicate pled with sticky skin and toe pad shape makes "mavomavo" adjective yellowish . It makes Platypelis tetra a good climber of the leaves obvious reference to the typical species' colou- beneath which it lives. At Anjanaharibe-Sud ration and is used as an invariable noun stand- we found Platypelis tetra in syntopy with ing in apposition to the generic name. another Platypelis of small size (P. occultans), P. tuberifera, Mantidactylus cf. bicalcaratus Natural history and distribution and M pulcher. The type series was collected at Amboloko- Platypelis tetra lives in close relationship patrika, where the habitat is a mid-high with the Pandanus microhabitat, and, there- altitude transitional rainforest, with tall trees fore, needs a pristine or mature secondary and an abundance of epiphytic plants. The rainforest. Platypelis mavomavo was found specimens were found during rainy nights only at some intact forest fragments of Anjan- climbing well above the ground (about 2-4m) aharibe-Sud and Ambolokopatrika forests. In on small tree trunks. A similar habitat was this sense, both these cophylines are both observed for one specimen (not captured) at closely tied to the existence of an intact rain- Anjanaharibe-Sud. This species is apparently forest and to high humidity levels. For these not a phytotelm specialist, although it is reasons they are good candidates for being ANDREONE ET AL. -TWO NEW FROGS FROM MADAGASCAR 99

ecologically sensitive species, as stressed by LITERATURE CITED Vallan (2000) and Andreone and Luiselli (2003). ANDREONE,F. 1999. Chapter 6. Madagascan Given the increasing deforestation occurring amphibians. p. 213-261. In: S. Yamagishi (ed.), in the Andapa water basin (Berner, 1995), the Madagasukaru no Dobutsu [The Animals of protection of the remnant forest fragments Madagascar-Its wonderful adaptive radiation]. corresponds to the protection of the Panda- Shoukabo, Tokyo. (in Japanese) nus screw pines where P. tetra and other frogs ANDREONE, F. In press a. Geographic distribution. live. The sites where the new species were Plethodontohyla coudreaui. Herpetol. Rev. found are either partly included in a reserve ANDREONE, F. In press b. Crossroad of herpetolog- network or are in places where logging is still ical diversity: survey work for an integrated conducted at a low rate. As outlined by Vallan conservation of the amphibian and reptile fauna et al. (in press) this kind of forest exploitation in Northern Madagascar. Ital. J. Zool. does not have an immediate negative effect on ANDREONE, E; G. APREA, M. VENCES, AND G. the batrachofauna. The long-term effects are ODIERNA. 2003. A new frog of the genus Manti- still largely unknown, but it is obvious that dactylus from the rainforests of north-eastern the physical reduction of the rainforest will Madagascar, and its karyological affinities. coincide with the disappearance of many frog Amphibia-Reptilia 24: 285-303. species. ANDREONE, F. AND L. M. LUISELLI. 2003. Conser- vation priorities and potential threats influencing the hyper-diverse amphibians of Madagascar. ACKNOWLEDGEMENTS Ital. J. Zool. 70: 53-63. We thank G. Aprea, H. Randriamahazo and ANDREONE, F., M. VENCES,D. R. VIEITES, F. J. E. Randrianirina for field assistance, and the CLAW, AND A. MEYER. In press. Recurrent Malagasy authorities for collection and export ecological adaptations revealed through a molec- permits. R. Nincheri helped with material, ular analysis of the secretive cophyline frogs of photographs and personal observations. P. Madagascar. Mol. Phyl. Evol. Bergo and V. Mercurio assisted with the speci- BENSTEAD,J. P., M. L. J. STIASSNY,P. V. LOISELLE, mens' setting. The research of FA was made in K. J. RISENG,AND N. RAMINOSOA.2000. River cooperation between the MRSN, and the conservation in Madagascar, p. 205-231. In: P. J. Parc Botanique et Zoologique de Tsimbazaza, Boon, B. R. Davies and G. E. Petts (eds.), Global Antananarivo, and supported by WWF, MRSN, Perspectives of River Conservation: Science, and WCS. Further support to DF and MEW Policy and Practice. John Wiley and Sons, was provided by "Radeau des Cimes", the Chichester. Oklahoma City Zoo and Zoological Society, P. BERNER, P. O. 1995. Preparation d'un volet Stout, the University of Oklahoma Depart- forestier d'un projet de conservation et devel- ment of Zoology, B. and H. Fenolio, G. and B. oppement integre de la region est de Madagas- Walvoord, and K. Walvoord. Equipment and car: le cas du complexe des reserves de Marojejy other support was provided by the Sam Noble et d'Anjanaharibe-Sud. Akon'ny Ala 17: 28-39. Oklahoma Museum of Natural History, A. BLOMMERS-SCHLOSSER,R. M. A., ANDC. P. BLANC. Estep, C. Leary, M. Mandica, M. Hoefnagels, 1991. Amphibiens (premiere partie). Faune de V. Hutchison, and J. Caldwell. Special thanks Madagascar 75 (1): 1-379. to our climbing technician in Madagascar, N. GLAW,F. AND M. VENCES.1994. A Fieldguide to Baiben. M. Matsui and an anonymous referee the Amphibians and Reptiles of Madagascar, reviewed a first version of this paper. 2nd Edition. Vences and Glaw Verlag, Koln. LEHTINEN,R. M. 2003. The use of screw pines (Pandanus spp.) by amphibians and reptiles in Madagascar. Herpetol. Bull. 82: 20-25. 100 Current Herpetol. 22 (2) 2003

RAXWORTHY,C. J., F. ANDREONE,R. A. NUSSBAUM, R. DOLCH, in press. Consequences of low, selec- N. RABIBISOA,AND H. RANDRIAMAHAZO. 1998. tive wood exploitation on Amphibians in the Amphibians and reptiles of the Anjanaharibe- tropical rain forest of An'ala in East Madagascar. Sud Massif, Madagascar: Elevational distribu- Oryx. tion and regional endemicity. p. 79-92. In: S. M. VENCES, M. AND F. CLAW. 2001. When molecules Goodman (ed.), A Floral and Faunal Inventory claim for taxonomic change: New proposals on of the Eastern Slopes of the Reserve Speciale the classification of Old World treefrogs. Spixi- d'Anjanaharibe-Sud, Madagascar, with Refer- ana 24: 85-92. ence to Elevational Variation. Fieldiana Zoology YODER, A. D., M. M. BURNS, S. ZEHR, T. (n. s.) 90, Field Museum of Natural History, DELEFOSSE, G. VERON, S. M. GOODMAN, AND J. Chicago. J. FLYNN. 2003. Single origin of Malagasy VALLAN, D. 2000. Influence of forest fragmentation Carnivora from an African ancestor. Nature on amphibian diversity in the nature reserve of 421: 734-737. Ambohitantely, highland Madagascar. Biol. Conserv. 96: 31-43. VALLAN, D., F. ANDREONE, V. H. RAHERISOA, AND Accepted: 11 December 2003