The H Rpeto Ogical Jour A
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Volume 13, Number 2 April 2003 ISSN 0268-0130 THE H RPETO OGICAL JOUR A Published by the Indexed in BRITISH HERPETOLOGICAL SOCIETY Current Contents HERPETOLOGICAL JOURNAL, Vol. 13, pp. 69-79 (2003) A REVISION OF THE SCAPHIOPHRYNE MARMORATA COMPLEX OF MARBLED TOADS FROM MADAGASCAR, INCLUDING THE DESCRIPTION OF A NEW SPECIES MIGUEL VENCES1, CHRISTOPHER J. RAXWORTHY2, RONALD A. NUSSBAUM3 AND FRANK GLA W4 'Institute fo r Biodiversity and Ecosystem Dynamics, Zoological Museum, Un iversiteit van Amsterdam, PO Box 94766, 1090 GT Amsterdam, Th e Netherlands :American Museum of Natural Hist01y, Central Park West at 79th Street, New York, NY 10024-5192, USA 3Division of Reptiles and Amphibians, Museum of Zoology, Un iversity of Michigan, Ann Arbor, Michigan 48109-1079, USA 'Zoologische Staatssammlung Miinchen, Miinchhausenstrasse 21, 81247 Miinchen, Germany A revision of the available material hitherto assigned to the endemic Malagasy microhylid toad Scaphiophryne marmorata finds this taxon to be a complex of three species. In this study we resurrect Scaphiophryne sp inasa Steindachner, 1 882 from the synonymy of S. marmorata and describe a new species fromcentral eastern Madagascar. These three Scaphiophryne species are characterized by their distinctly expanded terminal finger discs, a character only shared with S. gottlebei. S. sp inosa is characterized by a highly granular back, with large spiny tubercles above the forelimb insertion and in the tympanic region, while S. marmorata is dorsally covered by less prominent and more regular tubercles. The new species is distinguished from both S. marmorata and S. sp inosa by its large body size (SVL 47-60 mm), a smoother dorsal skin, and reddish terminal fingerdi scs in life. S. sp inosa is distributed in low- and mid-altitude rainforests along the east coast, whereas the new species is only known from mid-altitude rainforest in the Fierenana region. In contrast, records of S. marmorata include eastern mid-altitude rainforests and several more arid western sites. Keywords: Amphibia, Anura, Madagascar, Microhylidae, frog taxonomy INTRODUCTION several species of Scaphiophryneare also known from the mountains and rainforest areas of central and eastern Several of the anuran genera of Madagascar belong Madagascar. According to Blommers-Schlosser & to diverse endemic radiations such as the Mantellidae or Blanc (1991) these comprise Scaphiophryne the microhylid subfamily Cophylinae (Vences & Glaw, madagascariensis, S. marmorata and S. pustulosa. 2001 ). Other genera, in contrast, are Jess speciose and Three further species - S. brevis, S. calcarata and S. have uncertain affinities within their respective higher gottlebei - are known fromwestern and southern Mada taxa. Such genera are often distributed in the xeric re gascar. gions of western Madagascar and show reproductive Vences et al. (2002b) provided evidence that the modes typical for arid environments (Vences et al., Pseudohemisus pustulosus 1945 2000a). name Angel & Guibe, is a junior synonym of Calophrynus madagascariensis One of these groups is the genus Scaphiophryne, Boulenger, 1882. Consequently, they applied the name which - together with the monotypic Paradoxophyla - Scaphiophryne madagascariensis to populations previ is classified in the endemic subfamily Scaphiophryninae ously assigned to S. pustulosa, and verified that no (Blommers-SchJOsser & Blanc, 1991), or even in the available name exists for specimens from the separate family Scaphiophrynidae (Dubois, 1992). Andringitra Massif in south-eastern Madagascar which Scaphiophryne are unique in the morphology of their has thus far been referred to as S. madagascariensis. tadpoles, which are intermediate between the ranoid and Scaphiop hryne madagascariensis and the new form the microhylid type (Wassersug 1984). They reproduce from Andringitra are known from high altitudes along inlentic, often temporarywaters, and Jay a large number the centralmountain chain ofMadagascar, and are char of small eggs (Blommers-Schlosser, 1975; Vences et acterized by lacking greatly expanded terminal discs on al., 2002a). This reproductive mode is typical for sea fingers and toes. In contrast, S. marmorata is known sonal areas such as savannas (e.g. Rodel, 2000), but from forests in eastern Madagascar, and from relict for ests in the arid west (Glaw & Vences, 1994). It is Correspondence: M. Vences, Institute for Biodiversity and characterized by its expanded terminal fingerdiscs (and Ecosystem Dynamics, Zoological Museum, Universiteit van slightly expanded toe discs), a state that so far was Amsterdam, PO Box 94766, 1090 GT Amsterdam, The Netherlands; E-mail: [email protected] thought to be shared by only one further species, S. 70 M. VENCES ET AL. gottlebei from Isalo in south-western Madagascar RESULTS AND SPECIES ACCOUNTS (Busse Bobtne, 1992). Recent fieldwork revealed the & During the revision of the available material thus far presence of additional species with this character in attributed to S. marmorata, we noted the existence of eastern Madagascar. We herein revise the material of two distinct species: a large species with a very granular green-coloured Scaphiophryne with expanded finger and spiny dorsal skin, and a smaller, less granular spe discs available to us, resurrect the name Scaphiophryne cies. Careful examination showed that the less granular sp inosa Steindachner, 1882 and describe one highly dif species is distinguished from the spiny species by the ferentiated new species. following characters: (I) smaller body size (male SVL 32-49 mm vs. 40-48 mm; Table I); (2) presence of a MATERIALS AND METHODS variable numberof regular larger granules on head and The present revision is based on material in the fol body, usually including a symmetrical pair of elongated lowing collections: The Natural History Museum, tubercles in the shoulder region and a smaller pair on the London (formerly British Museum of Natural History) posterior dorsum, these tubercle pairs being especially (BMNH); Museum National d'Histoire Naturelle, Paris distinctin subadults ( vs. a large number of irregular dor (MNHN);Museo Regionale di Scienze Naturali, Torino sal tubercles of different size, and absence of elongated (MRSN); Naturhistorisches Museum Wien (NMW); symmetrical tubercles in the shoulder region); (3) ab University of Michigan, Museum ofZoology (UMMZ); sence oflarge spiny tubercles above forelimb insertion, Zoologisch Museum Amsterdam (ZMA); Zoologisches at posterior end of maxilla, and in tympanic region (vs. Forschungsinstitut und Museum A. Koenig, Bonn presence); (4) tympanum often faintly recognizable, its (ZFMK); Zoologische Staatssammlung Miinchen horizontal diameter larger than 50% of eye diameter (vs. (ZSM). tympanum usually totally concealed, smaller than 50% Specimens were measured withcalip ers to thenearest of eye diameter - if visible); (5) ventral patternvaria ble, 0.1 mm. Measurements taken were SVL (snout-vent but often (especially in juveniles) with contrasting dark length), HW (head width at the maxillary rictus), HL light marbling, the dark colour extending onto the (head length, from the maxillary comrnissure to the posterior belly and the femur (vs. absence of contrasted snout tip), ED (horizontal eye diameter), END ( eye-nos marbling, usually dark on throat, chest and anterior tril distance), NSD (nostril-snout tip distance), NND belly, light on posterior belly and femur). These differ (nostril-nostril distance), TD (horizontal tympanum di ences were already visible in juveniles of both species ameter), HAL (hand length, fromthe carpal-metacarpal (e.g. UMMZ 191157; ZSM 303/2000). We also recog articulations to the tip of the longest finger), FORL nized a third larger species, with a smoother dorsal body (forelimb length, from the axil to the tip of the longest surface and reddish discs on the fingers and toes. The finger), HIL (hindlimb length, fromthe cloaca to the tip three species showed a relevant molecular differentia of the longest toe), FOL (foot length, from the tarsal tion in the sequenced fragment of the 16S rRNA gene metatarsal articulations to the tip of the longest toe), (see comments in the section of S. boriborybelow). FOTL (footlength including tarsus, fromthe tibiotarsal articulation to the tip of the longest toe), IMTL and SCA PHIOPHRYNE MA RMORA TA BOULENGER, 1882 (FIG. 1-2) IMTH (maximum length and height of inner metatarsal Diagnosis. A Scaphiophrynewith highly expanded tubercle), FD4 (maximum width of the terminal disc of terminal discs on the fingers; dorsally green with sym fourth finger). The sex and maturity of preserved speci metrical dark markings, and lacking pink markings; mens were either determined by obvious secondary adult SVL 32-49 mm; typically two symmetricalpairs of sexual characters ofmales in breeding conditions (vocal larger tubercles, an elongated pair in the shoulder region sac) or by dissection and gonad examination. and a smaller pair on the posterior dorsum; absence of Advertisement calls were recorded in the wild or in large spiny tubercles above forelimbinsertion, at poste captivity and were analysed on a PC using the software rior end of maxilla, and in tympanic region; tympanum Cooledit (Syntrillium Corp.). Statistical analyses were typically visible with a horizontal diameter > 50% of eye carried out using SPSS forWindows, version 10. Meas diameter; ventral pattern often with contrasted dark urements are given as range, with mean ± standard light marbling, the dark colour extending onto the deviation