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CHIARI Et Al Preliminary data on genetic differentiation within the Madagascar spider tortoise SALAMANDRA 41 1/2 35-43 Rheinbach, 20 May 2005 ISSN 0036-3375 Preliminary data on genetic differentiation within the Madagascar spider tortoise, Pyxis arachnoides (BELL, 1827) YLENIA CHIARI, MEIKE THOMAS, MIGUEL PEDRONO & DAVID R. VIEITES Abstract. The spider tortoise, Pyxis arachnoides, is one of the four extant endemic terrestrial chelonian species of Madagascar. Although this species is considered to be vulnerable, little is known about the status and genetic differentiation of its three subspecies. Here we report on field observations made in early 2004 during collection of blood samples of this species for genetic analysis. We investigated the genetic differentiation within the above mentioned subspecies using mtDNA sequences (cytochrome b). This marker showed a low differentiation among the three subspecies of P. arachnoides and an extremely low variation within and among populations of each of the subspecies. In contrast the differentiation to the second species of Pyxis, P. planicauda, was much higher. Our data therefore confirm current taxonomy and suggest that the three subspecies of P. arachnoides should be treated as separate entities for conservation. Key words. Reptilia: Testudinidae: Pyxis arachnoides; subspecies; cytochrome b; Madagascar. Introduction mined. The decline of tortoise populations is the result of many centuries of commercial The high biodiversity and high degree of exploitation (PEDRONO et al. 2000) and it has endemism make Madagascar one of the top been associated with habitat loss and popu- hotspots for biodiversity conservation lation fragmentation (RAXWORTHY & NUSS- worldwide (MYERS et al. 2000). At the species BAUM 2000). Human exploitation is consid- level, endemism in amphibians and reptiles ered to be the cause of the extinction of is greater than 95%. Of the seven endemic Madagascar’s giant tortoises, Dipsochelys chelonian species that historically inhabited grandidieri and D. abrupta (BOUR 1994). the island (six terrestrial and one aquatic) The systematic history of the two Pyxis only five are still present in Madagascar. The species is controversial. BELL (1827) defined four land tortoises (Geochelone radiata, the genus Pyxis for its movable anterior part Geochelone yniphora, Pyxis planicauda and of the plastron. Afterwards, GRANDIDIER (1867) Pyxis arachnoides) share a common ancestor described Testudo planicauda, a species as shown by mitochondrial DNA data (CAC- close to Pyxis (BELL 1827) but without mov- CONE et al. 1999) that probably arrived by able plastron. Based on osteological charac- rafting from Africa (BOUR 1988, 1994). The ters, SIEBENROCK (1902) created the genus Aci- other species of tortoise, Kinixys belliana, nixys for the species described by GRANDIDIER seems to have been introduced from Africa (1867). Within the family Testudinidae, by humans (ANGEL 1941). All endemic spe- BOUR (1979) and OBST (1978, 1980) placed cies are considered to be endangered or vul- Acinixys planicauda within the genus Pyxis, nerable according to IUCN classification designating Acinixys as a subgenus. This (RAXWORTHY & NUSSBAUM 2000, IUCN Red classification was supported from juvenile List), whereas the status of the non-endemic traits shared also with juveniles of other Kinixys belliana on Madagascar is undeter- species. ERNST & BARBOUR (1989) used Pyxis © 2005 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT) http://www.salamandra-journal.com 35 YLENIA CHIARI et al. related to the secretive life style of this spe- cies and the difficulties of access to large parts of its range. These tortoises are active early in the morning, particularly during the rainy season. For the rest of the day they hide mostly underground (JESU & SCHIMMENTI 1995). Morphological examination of speci- mens of P. arachnoides from different locali- ties has revealed the existence of three sepa- rated geographic entities that are presently considered as subspecies (BOUR 1978): Pyxis arachnoides brygooi (VUILLEMIN & DOMERGUE 1972), (Fig. 2a), P. a. arachnoides, BELL Fig. 1. Sampling localities of Pyxis arachnoides. 1827, (Fig. 2b) and P. a. oblonga, GRAY 1869, The different level of grey correspond from North (Fig. 2c). P. a. brygooi occupies the northern- to South to the distribution area of each subspecies most part of the distribution area shown in (lighter grey: P. a. brygooi; medium grey: P. a. Fig. 1, in the south of Morombe; P. a. arach- arachnoides; darker grey: P. a. oblonga). Numbers refer to Tab. 1. noides occurs around Toliary; P. a. oblonga occurs in the extreme southern coast of Ma- dagascar to Fort-Dauphin (BOUR 1987). The and Acinixys as different monotypic genera, main character used to identify the different whereas the results of CACCONE et al. (1999) subspecies is the movable hinge in the ante- agreed with the placement of the two species rior part of the plastron and its coloration in a single genus, as previously proposed by (BOUR 1987). P. a. brygooi presents a rigid BOUR (1979) and OBST (1978, 1980). and uniform yellow colored plastron; P. a. Pyxis arachnoides, the Madagascar spi- arachnoides has a uniform yellow plastron der tortoise, is a small chelonian with a cara- in which the anterior lobe is more or less pace length of up to 15 cm. The carapace is mobile; P. a. oblonga shows a yellow plas- domed and has a starlike pattern with a black tron with black spots on the sides in which rounded edge and yellow center from which the anterior lobe is completely mobile (BOUR several broad rays extend. The plastron var- 1987) (Fig. 2). ies from immaculate yellow to having some Three of the four taxa within the genus dark blotches at the bridge. The head is black Pyxis (P. arachnoides arachnoides, P. a. with some yellow markings (PEDRONO & SMITH brygooi, P. a. oblonga) are currently consid- 2003). ered subspecies, but exceptional sympatric The distribution of P. arachnoides is lim- occurrence has been quoted (BOUR 1987). ited to the south-western region of the island, The taxonomic history of the taxa within the in a semi-arid area with thornbush vegetation genus Pyxis has been quite chaotic. Together from south of Morombe to Fort-Dauphin (see with the above mentioned taxonomic confu- Fig. 1) (BOUR 1978, 1981; KUCHLING 1989). sion between Pyxis arachnoides and P. pla- This species is sympatric with Geochelone nicauda, also P. a. oblonga has been previ- radiata but to date it is not known to be ously considered as P. a. matzi by BOUR heavily exploited or consumed for food by (1979) but the same author later recognised local people as G. radiata, except in places P. a. oblonga as a valid senior synonym of P. where radiated tortoises have disappeared a. matzi (BOUR 1982). Moreover, VUILLEMIN & (PEDRONO et al. 2000). Except a few field DOMERGUE (1972) introduced the genus Py- observations by KUCHLING (1989), little is xoides, characterised by the absence of the known about the ecology, behaviour and movable hinge in the plastron and described status of P. arachnoides. This is probably Pyxoides brygooi. OBST (1978, 1980) and 36 Preliminary data on genetic differentiation within the Madagascar spider tortoise BOUR (1978) agreed with the identification of February 2004. These samples belong to the Pyxoides brygooi as a geographic subspecies three subspecies (Pyxis arachnoides arach- of P. arachnoides. noides, P. a. brygooi, P. a. oblonga) and to We here provide preliminary insights in- Pyxis planicauda (Fig. 3). Blood was taken to genetic differentiation within the species from the forelimbs of the tortoises with insu- P. arachnoides. Our goal is to explore whe- lin syringes. Each tortoise was photographed ther the geographical variants identified as and subsequently released at the place of subspecies are also genetically isolated u- capture. nits, which is of particular importance for Ten populations were sampled and geo- conservation actions. We also want to evalu- graphical coordinates were recorded by GPS ate whether the three subspecies of P. arach- (Tab. 1). The sampling locations extend a- noides are best considered as subspecies ra- long the southern part of Madagascar (Fig. 1) ther than species by comparing the genetic and encompass the known ranges of the three distance between subspecies with the ge- subspecies. netic distance between the distinct species P. We studied differentiation of three sub- arachnoides and P. planicauda. To address species at the population level, using partial these questions we used DNA sequences of sequences of the mitochondrial cytochrome the mitochondrial gene cytochrome b of in- b gene. As outgroup we used a cytochrome b dividuals of each subspecies from different sequence of Geochelone radiata (Genbank localities. In addition, we report on field accession number AF020897). observations made during our sampling trip in south-western Madagascar. Laboratory techniques Materials and Methods DNA was extracted from 170 µl of blood Sampling tissues preserved in modified Queen’s lysis buffer (SEUTIN et al. 1991) using Proteinase K Blood samples of 36 individuals used in this (final concentration 1 mg/ml). DNA was iso- study were collected during fieldwork in lated by a standard salt extraction protocol Madagascar in February 2003 and January/ (BRUFORD et al. 1992). Locality Locality GPS coordinates Subspecies N° of number individuals Manombo Atsimo 0 22°56’44”S; 43°28’04”E P. a. brygooi 1 Manombo Atsimo nr 1 22°57’05”S; 43°28’29”E P. a. brygooi 2 Manombo Atsimo sr 2 22°58’10”S; 43°29’02”E P. a. brygooi 1 Manombo Atsimo- 3 23°00’26”S; 43°30’41”E P. a. brygooi 2 Antsira Ifaty 4 23°07’50”S; 43°36’55”E P. a. brygooi 19 Miary (Toliary) 5 23°18’58”S; 43°44’11”E P. a. brygooi 1 Beheloka 6 23°56’36”S; 43°41’17”E P. a. arachnoides 2 Mangoro- 7 23°78’00”S; 43°66’00”E P. a. arachnoides 3 Andranotohoka Faux Cap 8 25°34’07”S; 45°31’24”E P.
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