Natural History and Taxonomic Notes On

Herpetological Conservation and Biology 9(2):406–416. Submitted: 27 March 2014; Accepted: 25 May 2014; Published: 12 October 2014. NATURAL HISTORY AND TAXONOMIC NOTES ON LIOPHOLIDOPHIS GRANDIDIERI MOCQUARD, AN UPLAND RAIN FOREST SNAKE FROM MADAGASCAR (SERPENTES: LAMPROPHIIDAE: PSEUDOXYRHOPHIINAE) JOHN E. CADLE1, 2 1Centre ValBio, B.P. 33, 312 Ranomafana-Ifanadiana, Madagascar 2Department of Herpetology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA, e-mail: [email protected] Abstract.––Few observations on living specimens of the Malagasy snake Liopholidophis grandidieri Mocquard have been previously reported. New field observations and specimens from Ranomafana National Park amplify knowledge of the natural history of this species. Liopholidophis grandidieri is known from above 1200 m elevation in pristine rain forests with a high diversity of hardwoods and bamboo. In some areas of occurrence, the forests are of short stature (15–18 m) as a result of lying atop well-drained boulder fields with a thin soil layer. Dietary data show that this species consumes relatively small mantellid and microhylid frogs obtained on the ground or in phytotelms close to the ground. A female collected in late December contained four oviductal eggs with leathery shells. One specimen formed a rigid, loose set of coils and body loops, and hid the head as presumed defensive behaviors; otherwise, all individuals were complacent when handled and showed no tendency to bite. I describe coloration and present photographs of living specimens from Ranomafana National Park. The ventral colors of L. grandidieri were recently said to be aposematic, but I discuss other plausible alternatives. Key Words.––conservation; diet; habitat; reproduction; snakes; systematics exhibits the greatest sexual dimorphism in tail length and INTRODUCTION the greatest reported relative tail length of any snake (tails average 18% longer than body length in males; The natural history and systematics of many snake Cadle 2009). Several hypotheses that might explain species from Madagascar remain poorly documented. In these extraordinary characteristics have been considered some cases, morphological characters, geographic (morphological constraint, antipredator defense, sexual ranges, natural history, and colors in life are poorly selection) but no explanation is yet entirely satisfactory documented because observations on living snakes are (Cadle 2009, Glaw et al. 2014). A recent molecular unavailable and few specimens may exist in museum phylogeny of all species of Liopholidophis indicated a collections. Among these is Liopholidophis grandidieri sister relationship between L. grandidieri and the species Mocquard (1904), which was known only from the most similar to it in external characters, L. dolicocercus holotype until the mid-1990s, by which time one (Glaw et al. 2014). additional specimen had been collected and two others At the time of the previous review (Cadle 1996a), no were identified in old museum materials (Cadle 1996a). observations on living specimens of Liopholidophis Liopholidophis grandidieri is still known from few grandidieri were available. Yet detailed natural history specimens and few observations on its behavior or data will ultimately be necessary to fully understand the natural history have been reported. significance of the exceptional morphology of these Eight species of Liopholidophis are currently snakes. The data I report here are based on five recognized (Cadle 1996a; Glaw et al. 2007, 2014). Five specimens of Liopholidophis grandidieri obtained since species (L. dimorphus, L. dolicocercus, L. grandidieri, L. the species was last reviewed (Cadle 1996a). All of the rhadinaea, and L. sexlineatus) are characterized by new observations are from Ranomafana National Park remarkable sexual dimorphism in tail length and (RNP) and vicinity, obtained in January 2001 and exceptionally long tails in males (Cadle 1996a, 2009; December 2003 during 17 days at two field sites. The Glaw et al. 2007, 2014). Liopholidophis varius is less main purpose of this report is to present new data on the exceptional in both characters (Cadle 2009) and these natural history and behavior of L. grandidieri based on features are unknown in L. oligolepis and L. baderi field observations and specimens obtained since the because specimens of only one sex are known for these previous character summaries (Cadle 1996a). I put into species (Glaw et al. 2014). Liopholidophis grandidieri broader context here data on four specimens of L. Copyright © 2014. John Cadle. All Rights Reserved. 406 Herpetological Conservation and Biology grandidieri previously examined (Cadle 1996a: 460), days field work) and the 2003 locality as “Anjamba” update basic taxonomic data for the species, and provide (nine days). for the first time color descriptions and photographs of The following describes habitats at localities within RNP L. grandidieri in life from RNP. where L. grandidieri was observed in 2001 and 2003; local Malagasy names for the trees and bamboos are MATERIALS AND METHODS given. A field team of three logged on average eight hours Amindrabe.––Between the villages of Ambendrana per day during daylight hours and four to six hours at and Amindrabe a trail passes through rice fields and night while making a general faunal inventory secondary forest, with patches of what seem to be (herpetofauna, birds, and mammals); streambeds and relatively undisturbed stunted forest mainly along ridge existing footpaths allowed access to the forest. tops; one specimen of Liopholidophis grandidieri was Liopholidophis grandidieri were encountered found in these ridge-top forests. In the vicinity of the opportunistically and captured by hand (none were Amindrabe field camp (Fig. 1), terrain varies from captured by pitfall trap lines set at each site). I recorded 1200–1250 m elevation and much of it is characterized snout-vent length (SVL, in mm), measured from the tip by ravines separating sharp ridges. A small river, the of the snout to the posterior edge of the anal plate, and Apepina River, occasionally disappears under boulder tail length, measured from the posterior edge of the anal jumbles for tens of meters at a stretch and then reappears plate to the tip of the tail. Relative tail length (RTL) is on the surface. Boulders within the riverbed are covered here defined as tail length divided by SVL. I recorded with a thin to moderately thick layer of moss on exposed mass (g) using Pesola© scales (Pesola AG, Baar, Switzerland). portions. The forest canopy is 15–18 m in higher parts I acquired stomach contents of snakes by regurgitation but patches within the forest are sometimes less, after capture. I prepared vouchers as standard museum especially on ridge tops. I attribute the short stature of specimens by fixation in 10% formalin for the forest to its growth on top of an extensive boulder approximately one month, followed by storage in 70– field. Soil may also be of poor quality and thin. 75% ethanol. Specimens discussed herein were Bamboos are especially dense, including large stands of deposited in the California Academy of Sciences (CAS, the giant species known locally as volotsangana San Francisco), The Field Museum (FMNH, Chicago), (Bambusa madagascariensis; Poaceae). Prominent large the Museum of Comparative Zoology (MCZ, hardwood trees in this forest were tavolo (Cryptocarya Cambridge), the Muséum National d’Histoire Naturelle sp.; Lauraceae), ramy (Carnarium madagascariensis; (MNHN, Paris), The Natural History Museum (BMNH, Burseraceae), and hazondrano (Ilex mitis; London), and the Université d’Antananarivo Aquifoliaceae). Département de Biologie Animale (UADBA, Antananarivo). Anjamba.––The terrain in the vicinity of the field camp was a series of ridges separated by valleys with RESULTS small streams. The streams have substrates of rocks and boulders toward headwaters and sand or gravel Habitats and environments.––I found five new downstream. Uncommonly, the streams have cut to specimens of Liopholidophis grandidieri at three sites bedrock and sometimes flow over steep precipices, from upland rain forests (1200–1250 m; Fig. 1). Details forming small waterfalls. The Iarena River at the of the collection sites are as follows (specimen vouchers campsite has a streambed 3–4 m wide, low current from each locality follow the coordinates): 4 January velocity, and was fairly shallow (300–500 mm) during 2001 on a trail between the villages of Ambendrana and the fieldwork because no substantial rain had fallen for Amindrabe, about 1200 m elevation (21.3833°S, several days. The river had a sand and gravel substrate 47.3600°E; FMNH 261185); 8 and 11 January 2001 near at the camp but contained boulders upstream ~75 m. a field camp approximately 4 km (Euclidean distance) Jumbles of huge boulders were present in some areas, east of Amindrabe village, 1200–1250 m elevation with small caves present among some of these. A trail (21.3833S, 47.4000E; FMNH 261186–87); and 20 and heading east from camp passed over several low ridges 27 December 2003 near a field camp 9 km (Euclidean and arrived at a somewhat larger stream, the distance) east of Anjamba village, 1200 m elevation Ambohibory River. All of the local streams ultimately (21.3255S, 47.4027E; CAS 250594, UADBA–JE flow into the Andranaroa River to the west according to Cadle 15035). The localities are just within the western a local informant. boundary of RNP, or just outside the park near its At Anjamba the forest

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