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(Geochelone Radiata) in Madagascar

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(Geochelone Radiata) in Madagascar BIOLOGICAL CONSERVATION Biological Conservation 124 (2005) 451–461 www.elsevier.com/locate/biocon Distribution, status, and conservation of radiated tortoises (Geochelone radiata) in Madagascar Thomas E.J. Leuteritz a,*, Trip Lamb b, Jean Claude Limberaza c a Department of Biology, George Mason University, Fairfax, VA 22030, USA b Department of Biology, East Carolina University, Greenville, NC 27858, USA c ANGAP, Cap Sainte Marie Special Reserve, BP 28 Taolagnaro (Fort Dauphin) 614, Madagascar Received 19 April 2004 Abstract The radiated tortoise, Geochelone radiata, one of MadagascarÕs four endemic tortoises, occupies a narrow band of xeric spiny forest along the islandÕs southwest coast. Traditionally avoided by indigenous tribes, these tortoises are now routinely harvested for food. An accurate assessment of human exploitation remains problematic, however, hindered by limited, dated statistics avail- able on tortoise populations. To update the radiated tortoiseÕs status and distribution, we established a series of line transects at seven localities across its range and implemented a mark-recapture study at one of these localities (Cap Sainte Marie). Tortoises currently range from south of Tulear to east of Cap Sainte Marie, at density estimates spanning 27–5744 tortoises/km2. The mark-recapture estimate for Cap Sainte Marie (1905–2105 tortoises/km2) was substantially higher than its transect estimate (654 tortoises/km2) though comparable to actual tortoise captures (1438) there. Thus, our transect density values probably err as under- estimates, and from these data, we calculate a conservative total population size of 12 million radiated tortoises. We also examined mitochondrial DNA sequences (ND4 gene) for two individuals/locality in a preliminary assessment of genetic variation across the speciesÕ range. Only two ND4 haplotypes were recovered, the more common haplotype representing 13 of the 14 individuals. We offer several conservation recommendations in light of our survey results. Ó 2005 Elsevier Ltd. All rights reserved. Keywords: Radiated tortoises; Geochelone radiata; Distribution; Density; mtDNA variation; Madagascar 1. Introduction 1996; Nussbaum and Raxworthy, 2000). The tortoiseÕs geographic range is roughly coincident with regions The radiated tortoise or sokatra (Geochelone radiata) occupied by the Mahafaly and Antandroy, local peoples is one of four tortoise species endemic to Madagascar who have a taboo (fady) against touching or eating tor- (Ernst and Barbour, 1989; Juvik, 1975). The tortoiseÕs toises (Brown and Mbola Versene, 1997; Lewis, 1995; natural distribution is limited to xeric spiny forests on Nussbaum and Raxworthy, 2000). However, recent the Mahafaly and Karimbola plateaus of southwestern immigrants from other regions do exploit radiated Madagascar (Iverson, 1992). G. radiata is classified as tortoises, often in large quantities. For example, the ‘‘Vulnerable’’ on the IUCN Red List (Hilton-Taylor, Madagascar big-headed turtle (Erymnochelys madaga- 2000), due to threats posed by overcollecting and habitat scariensis) a freshwater turtle of lowland Western Mad- loss (Durrell et al., 1989b; Baillie and Groombridge, agascar is commonly used as a food source by the Sakalava peoples (Kuchling, 1988). Cart and truckloads * Corresponding author. Tel.: +1 909 335 5383; fax: +1 909 307 6952. of tortoises have been observed in the larger cities, where E-mail address: [email protected] (T.E.J. Leuteritz). tortoise meat is especially popular around Christmas 0006-3207/$ - see front matter Ó 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2005.02.003 452 T.E.J. Leuteritz et al. / Biological Conservation 124 (2005) 451–461 (Lewis, 1995; Nussbaum and Raxworthy, 2000). Many the Manombo River (60 km north of Tulear) and found Malagasy also keep tortoises penned with their chickens populations in the southeast to be highly fragmented and ducks as a means to ward off poultry diseases (Dur- east of Cap Sainte Marie. OÕBrien (2002) corroborated rell et al., 1989b; personal observation). Moreover, radi- these patterns of local extirpation in both northern ated tortoises face standard but significant sources of and eastern termini of the speciesÕ historic range. habitat degradation through deforestation, overgrazing, Prior to 1995, efforts to estimate tortoise populations and charcoal production (Nussbaum and Raxworthy, were limited to incidental counts taken by car, i.e., the 2000). number of tortoises seen along a stretch of road. Lewis Historically, the range of G. radiata encompassed a (1995) conducted a two-month field survey to determine 50 km band along the countryÕs southwestern coast the status of radiated tortoise populations throughout (Decary, 1950), from southeast of Morombe to the their range. His density estimates, derived by transect Bay of Ranofatsy (40 km east of Ambovombe). Offering sampling across five sites, varied from 262.2 to 1076.7 minor revision, Juvik (1975) extended the range north to tortoises/km2, from which he extrapolated a conserva- Morombe but observed that populations were ‘‘severely tive total population size of 1.6–4 million. However, Le- depleted’’ north of the Onilahy River (south of Tulear), wisÕ transects were established along existing trails or and further noted that animals were ‘‘rarely encoun- paths, which can result in nonrandom sampling of hab- tered’’ east of Ambondro or Antaritarika along the itat and thereby generate biased counts. southeastern range terminus (Fig. 1). In a subsequent The purpose of this study was to (1) delimit the cur- survey, Lewis (1995) did not observe tortoises north of rent range of G. radiata; (2) establish a series of transects Fig. 1. Map of southwestern Madagascar depicting the core range (shaded) of Geochelone radiata. Transect sites, indicated by [], are as follows: 1 – Cap Sainte Marie, 2 – Lavanono, 3 – Ankirikirika, 4 – Nisoa-Ambony, 5 – Lavavolo, 6 – Vohombe, and 7 – Lake Tsimanampetsotsa. T.E.J. Leuteritz et al. / Biological Conservation 124 (2005) 451–461 453 across different sites to assess tortoise densities; and (3) At each chosen site, two transects, at least 1 km use these data to estimate population size, both region- apart, were cut through dry spiny forest habitat at a ally and range-wide. Our survey provides a much- bearing of 45° NW. Transects were 1 m wide and 809– needed update and detailed account on the distribution 2244 m in length, depending on accessibility. General and status of G. radiata, complementing OÕBrien et al.Õs qualitative assessments of floristic composition and hab- (2003) recent view on overexploitation of this species. In itat condition were noted for each site. To insure sight- addition, we conducted a preliminary survey of genetic ing consistency, a local guide, an Association Nationale variation to determine whether G. radiata exhibits any Pour La Gestation Des Aires Protegees (ANGAP) ran- degree of phylogeographic structure. Although the radi- ger, and the primary author walked each transect to- ated tortoise has a restricted distribution, its range is gether twice daily on sunny days during peak tortoise roughly contiguous with another endemic tortoise, activity times, 06:30–10:00 h and 15:30–18:00 h. Surveys Pyxis arachnoides (spider tortoise), for which four sub- were conducted in January and February, during the re- species are recognized. Radiated tortoises do not exhibit gionÕs rainy season and period of greatest tortoise move- distinct morphological variation geographically (Love- ment (Durrell et al., 1989b; Lewis, 1995). ridge and Williams, 1957; Bour, 1978). Nonetheless, Perpendicular distances (from transect line to the ani- their congruent distribution with the polytypic P. arach- mal), sighting distances (from observer to animal at mo- noides suggests some potential for historical population ment of detection), and sighting angles (between transect isolation. This possibility, coupled with the tortoiseÕs line and line of sight of the animal) were recorded using ‘‘Vulnerable’’ IUCN status, underscores the prudence a 100 m tape measure (±0.01 m) and an azimuth com- of a range-wide genetic survey. pass for each animal detected along a transect. Time of capture, size, weight, age, sex, behaviour, and habitat were also recorded. Given the time constraints of com- 2. Methods pleting transect runs, tortoises were simply marked with paint to avoid multiple captures. Data were pooled per 2.1. Line transect surveys activity period per site. Transect data were analyzed using DISTANCE 3.5 Ten potential localities, spanning the speciesÕ range software (Buckland et al., 1993; Thomas et al., 1998) (from Tule´ar to Andohahela National Park), were se- to determine tortoise density and population structure. lected and searched for tortoise sign as well as suitabil- The statistical methodology used for line transect ity for transect sampling (Lewis, 1995; WWF, 1998, analysis is based on Fourier analysis, the accuracy of 1999). These localities were chosen for their accessibil- which depends on four assumptions: (1) objects directly ity, minimally disturbed habitat, and, in certain cases, on the line will not be missed; (2) objects are fixed at the their use in previous transect surveys. Three sites – initial sighting position (i.e., they do not move and are Beheloka and Mahaleotse (SE of Tule´ar), and Ando- not counted more than once); (3) distances and angles hahela National Park (20 km NW of Ft Dauphin, are measured exactly; and (4) all sightings are indepen- 100 km NE of Ambovombe)
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