Zoological Journal of the Linnean Society (2001), 133: 495–529. With 29 ﬁgures doi:10.1006/zjls.2001.0301, available online at http://www.idealibrary.com on Revision of the Boophis majori group (Amphibia: Mantellidae) from Madagascar, with descriptions of ﬁve new species FRANK GLAW 1 *, MIGUEL VENCES 2 , FRANCO ANDREONE 3 and DENIS VALLAN 4 1 Zoologische Staatssammlung, Mu ¨nchhausenstr. 21, 81247 Mu ¨ nchen, Germany 2 Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany 3 Museo Regionale di Scienze Naturali, Sezione di Zoologia, Via G. Giolitti, 36, 10123 Torino, Italy and 4 Universita ¨t Bern, Zoologisches Institut, Baltzerstr. 6, 3012 Bern, Switzerland Received July 2000; accepted for publication February 2001 A systematic revision of the Malagasy frogs hitherto attributed to the Boophis difﬁcilis group is presented. The difﬁcilis holotype is not conspeciﬁc with other specimens hitherto named Boophis difﬁcilis but belongs to the B. tephraeomystax group based on lack of webbing between ﬁngers and the presence of heterogeneously granular ventral skin. Rhacophorus difﬁcilis Boettger, 1892 is considered as junior synonym of Boophis tephraeomystax. The Boophis difﬁcilis group is dissolved and its species are transferred to a new species group named after Boophis majori, the oldest of the included taxa. All species of the B. majori group, their type specimens, and their geographical distributions are revised based on new morphological, bioacoustic and ecological data. Five new species of the group from the eastern rainforests of Madagascar are described. Morphological differentiation within the B. majori group is low, although some species can be distinguished by characters such as snout–vent length, relative tympanum size, or coloration. The most reliable character for species identiﬁcation are advertisement calls which are strikingly different between most species of the group. Species diversity in Boophis is highest in central eastern Madagascar and gets lower at the northern and southern borders of the island. Available data do not allow a comprehensive phylogenetic analysis of the B. majori group, but a northern subgroup of small species without red coloration and a southern subgroup of larger species with distinct red pigments can be distinguished. Osteological data for B. miniatus are provided. Within Boophis, representatives of all species groups except the B. tephraeomystax group are characterized by a synapomorphic reduction of the anterolateral hyoidal process.  2001 The Linnean Society of London ADDITIONAL KEY WORDS: Madagascar – biogeography – biodiversity – bioacoustics – taxonomy. (1994) recognized seven Boophis species groups. Ac- INTRODUCTION cording to these authors the Boophis difﬁcilis group The genus Boophis Tschudi, 1838 is endemic to Ma- consists of ﬁve named species: Boophis difﬁcilis dagascar and adjacent islands. Boophis was considered (Boettger, 1892), Boophis majori (Boulenger, 1896), a synonym of the Oriental genus Rhacophorus by Guibe ´ Boophis miniatus (Mocquard, 1902), Boophis blom- (1978), but was re-established by Blommers-Schlo ¨sser mersae Glaw & Vences, 1994 and Boophis marojezensis (1979b) who distinguished eight phenetic species Glaw & Vences, 1994. These species are characterized groups. Most Boophis are arboreal frogs with a typical by small size (snout–vent length of males 21–33 mm), treefrog habitus (enlarged ﬁnger discs, broad and an- a predominantly brownish (never green) back, often teriorly rounded head, large eyes, no dorsolateral with hourglass shaped markings, a whitish (not trans- ridge). parent) venter, and the lack of a distinct canthus In the most recent classiﬁcation Glaw & Vences rostralis. Many species have reddish colour on the legs, feet, webbing or other skin areas of the body (sometimes only visible when preserved). As far as is known, these species live alongside brooks. The * Corresponding author. E-mail: Frank.Glaw@zsm.mwn.de 495 0024–4082/01/120495+35 $35.00/0  2001 The Linnean Society of London