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MAR TIENSSENI INTRODUCTION the Montane Rainforests Of HERPETOLOGICAL JOURNAL, Vol. 8, pp. 35-40 (1998) THE MALE NUPTIAL CHARACTERISTICS OF ARTHR OLEPTIDES MA R TIENSSENI NEIDEN, AN ENDEMIC TORRENT FROG FROM TANZANIA'S EASTERN ARC MOUNTAINS MICHAEL W . KLEMENS International Programs, Wildlife Conservation Society, 185th Street at Southern Blvd. , Bronx, New York 10460, USA and Department of Herpetology, American Museum of Natural History, Central Park We st at 79th Street, New York, New York 10024, USA Recent field work in the Udzungwa Mountains of Tanzania has resulted in the discovery of the hitherto unreported nuptial characters of male Arthroleptides martiensseni which are described and illustrated in this contribution. These newly described characters are compared with the nuptial characters of male Arthroleptides dutoiti, a supposedly extinct species from Mount Elgon (Kenya) as well as with members of the West African genus Petropedetes, which exhibit some remarkable morphological and ecological similarities with Arthroleptides martiensseni. INTRODUCTION 1983) from "d' A to lo (region de Mamfe), Cameroon." Six of the eight species of the genus Petropedetes occur The montane rainforests of Cameroon and Tanzania in the western and southern portions of Cameroon, in­ contain a remarkable diversity of anurans, character­ cluding Fernando Po. P. cameronensis has also been ized by a high degree of endemism (Perret, 1966; collected in eastern Nigeria, and P. newtoni in Equato­ Howell, 1993; Lawson, 1993). Two genera, Petrope­ rial Guinea, which indicates a remarkable center of detes (Reichenow, 1874) and Arthrolep tides (Neiden, speciation in the region of the Bight of Biafra (Perret, 1910) contain species that are highly specialized for 1984). living on wet cliff faces, often in and immediately adja­ Neiden (1910) described the genus Arthroleptides cent to waterfalls and cascades. These adaptations based on Arthroleptides martiensseni sp. nov. from include tadpoles that have evolved a distinctive suite of "Amani, Deutschostafrika", now Tanzania. This spe­ morphological characters that allow fortheir develop­ cies has a restricted distribution, endemic to the Eastern ment on mossy, wet rock faces, as well as a distinctive Arc Mountains of Tanzania, reported from the East and "T-shaped" terminal expansion of the digits in adults, West Usambaras, the Ulugurus, and the Udzungwas which allow these frogs to adhere to the slippery rock (Howell, 1993). Neiden considered Arthroleptides to surfaces as described by Boulenger (1905) and Lov­ be intermediate between Petropedetes, with which it eridge (1925). shared the distinctive "T-shaped" terminal phalanges Reichenow (1874) described the genus based on and Arthroleptis, with which it shared the absence of Petropedetes cameronensis sp. nov. from "Bimbia in vomerine teeth and the reduction of webbing between the Cameroon foothills." Boulenger (1887) described the toes. The second member of the genus, A. dutoiti Cornufe r (=Petropedetes) johnstoni from "Rio de! was described by Loveridge ( 1935) from two adults Rey, Cameroons District." This description was based (male and fe male) and an immature specimen from the on a single, sub-adult male (Amiet, 1983). Subse­ eastern slopes of Mt. Elgon, Kenya. This species is con­ quently Boulenger (1900) placed Cornufe r johnstoni in sidered extinct by Baillie & Groombridge (1996). the genus Petropedetes. Bocage (1895) described Petropedetes and Arthroleptides share many morpho­ Ty mp anoceros ( =Petropedetes) newtonii, from logical and ecological similarities. However, certain "Fernao do P6" (=Fernando Po), noting that the large members of the genus Petropedetes possess a suite of tympanum was "surmonte tout pres de son bard morphological characters unique to breeding males. superieur par un tubercule cylindrique. " Boulenger These include enlarged femoralgla nds, the presence of (1905) described P. natator from"Sie rra Leone at 800 a bony, metacarpal projection, tympanic and brachia! feet"and P. palmipes from"Efulen, South Cameroon." hypertrophy, and most distinctive, the presence of a Ah I (1924) described P. obscurus from "Tscharra nipple-like projection on the tympanum. Perret (1984) Dana-Fluss" in Kenya. P. obscurus was synonymized summarized the characters of breeding male with P. cameronensis by Perret (1984) who found the Petropedetes, reporting that the tympanic projection two species morphologically indistinguishable, and occurred in four of the seven recognized species (i.e. considered Ahl's Kenya locality data to be erroneous. perreti, newtoni, parkeri, and johnstoni). Perret also Amiet (1973) described P. perreti from "Nsoung, noted that the location of the projection varied by spe­ 1400-1500 m., Cameroon" and P. parkeri (Amiet, cies, from the centre of the tympanum to the upper 36 M. W. KLEMENS margin. An examination of Perret's data also shows Additional data were collected by conducting time­ that these four species have larger tympani than the constrained transect searches, along trails and three species (i.e. cameronensis, natator and palmipes) river-edge, both during the day and at night; by oppor­ that lack tympanic projections. All four species that tunistic examination of fallentrees and rotten logs; and have tympanic projections also possess a bony, meta­ by making several excavations, to exhume fossorial carpal projection; however, this character is also shared species. with P. palmipes. Specimens were collected to document the diversity Considerable confusion exists in the literature as to of species, and whenever possible, the range in varia­ the structure of the tympanic projection, undoubtedly tion, sexual dimorphism, and life history stages of each the most distinctive sexually dimorphic character of species of amphibian and reptile found. These voucher male Petropedetes. This was illustrated by Noble specimens were anaesthetized, individually labelled, (1931) and by Duellman & Trueb ( 1994 ). These au­ and fixed in formalin. Tissue samples for DNA analysis thors concluded that the projection was the columella were collected prior to fixation, and stored in ethyl al­ thrust through the eardrum, covered by dermis. cohol. Colour photographs were taken of selected Duellman & Trueb (1994) stated that this projection specimens prior to preservation and tape recordings of would diminish the vibratory capacity of the tympa­ frog calls were made. All specimens were exported to num, thereby reducing its sensitivity to high the American Museum of Natural History (AMNH) for frequencies. However, du Toit (1943) in his anatomical follow-up identification and study. Preparation of this monograph on Petropedetes stated that "Petropede tes paper required examining specimens of Petropedetes is unique in possessing a seasonal secondary sexual and Arthroleptides deposited in both the AMNH and characteristic in the form of a so-called tympanic pa­ the Natural History Museum (London). As preserved pilla. The latter, which Noble (1931) mistook for the frogs are quite pliable, all measurements reported on pars externaplectri thrust through the drum is in reality Tables 1 and 2 were made with calipers and rounded to a thickened portion of the outer dermal part of the tym­ the nearest 0.5 mm. Great care was exercised to be con­ panic membrane." He further explains the structure of sistent in measuring technique, i.e. the application of the papilla to be quite complex, consisting of epidermal pressure to the specimen from the caliper's jaws. Un­ and cutis components, with a concentration of glands, less otherwise noted, the sex of all specimens examined collagenous fibres in its basal portion, a subepidermal was verified by internal examination through a slit cut pigment layer that is better developed in the papilla through the abdomen wall. than other portions ofthe tympanum, as well as a super­ RESULTS AND DISCUSSION fi cial cornified layer that is produced into tiny, wart-like asperities. This paper reports the discovery of On 8 December 1995, a series of eleven frogs an equivalent suite of nuptial characteristics in breeding (AMNH 151340-50) was collected at night by the au­ male Arthroleptides martiensseni, and discusses the thor and A. M. Nikundiwe in primary rainforest, taxonomic relationships between Petropedetes and perched on large boulders in cascades on the Arthrolep tides in the light of this new discovery. Nj okomoni River, near where the river drops offof the edge of the Udzungwa Escarpment in Tanzania's MATERIALS AND METHODS Udzungwa Mountains National Park, 7°48'57"S, In December 1995, an intensive, multi-taxa 36°5 1' 15"E, 1100 m. That afternoon the dry weather biodiversity inventory of two sites in the Udzungwa had broken with an exceptionally heavy thunderstorm. Mountains National Park was conducted at the begin­ Two of the largest individuals of this series were ma­ ning of the several-month long rainy season. Nine days ture males (AMNH 151342-43). They were quite were spent at a single site at the foot of the U dzungwa remarkable in that they possessed several of the distinc­ Escarpment, along the Mwaya River at 350 m. This lo­ tive breeding characteristics of male Petropedetes, cality included a mixture of riverine forest, miombo, including the unique tympanic projection which was and disturbed edge and agricultural habitats near located between the center and upper rim of the tympa­ Man'gula village. Nine days were spent at a single up­ num, as well as tympanic and brachia! hypertrophy, and land site (1100-1200 m) along the upper reaches of the a large, metacarpal knob.
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