A N N A L E S Z O O L O G I C I (Warszawa), 2006, 56(2): 225-240 NEMATODES PARASITES OF TESTUDINIDAE (CHELONIA): LIST OF SPECIES AND BIOGEOGRAPHICAL DISTRIBUTION SALAH BOUAMER* and SERGE MORAND CBGP (Centre de Biologie et de Gestion des Populations) Campus International de Baillarguet, CS 30 016, 34988 Montferrier sur Lez Cédex, France e-mail: [email protected]. *Corresponding author Abstract.— One hundred and four Nematodes parasites of Testudinidae have been recorded. A list of all valid species within orders, superfamily, family and genus are given. For each species the host-type, locality-type and other records are presented. The sub- species are raised to level of species. The highest number of taxonomic studies concerned the Palaearctic region and the Ethiopian region; but a lower number of studies were devoted to the other regions: Oriental, Neotropical and the Nearctic. The parasite species diversity of nematodes seems to be linked to the species diversity of hosts. A good significant relationship between the number of taxonomic studies and the species diversity of hosts was noted and a significant relationship between the species diversity of parasites and the generic diversity of the parasites was also observed. The host specificity does not differ between the oxyurida species and the ascaridida species. ± Key words.— Nematoda, Oxyurida, Ascaridida, Testudinidae, biogeographical distribution, relationship. INTRODUCTION genera and redescribed the type-species. Bouamer and Morand (2003, 2005) redescribed five species, raised The parasitic nematodes of terrestrial tortoises four sub-species to level of species, and described four belong to the orders Oxyurida and Ascaridida, with species parasite of Testudo graeca Linneaus, 1758 the majority of the species belonging to the Oxy- from Morocco. urida. Studies concerning nematodes from Ethiopian The studies of the parasitic nematodes from Palae- hosts were realised by four authors: Gendre (1909) and arctic were initiated by five authors: Drasche (1884), Gedoelst (1916) who gave the first descriptions of par- Stossich (1898), Seurat (1917, 1918), Baylis (1923) and asitic nematodes of terrestrial tortoises of West Africa, Thapar (1925), focusing on Testudinidae of Mediter- Ortlepp (1933) who gave the first description of para- ranean area. Dubinina (1947, 1949) in a study of the sitic species of Southern African tortoises and Petter parasites of Testudo horsfieldii Gray, 1844, gave the (1966) who redescribed and described a certain num- information of the nematodes of the East region of ber of species and also gave the first descriptions of Palaearctic. Petter (1966) tried to rectify taxonomic Malagasy species. Bouamer and Morand (2004) re- errors in the precedent studies, redescribed and de- described two species, raised two sub-species to level scribing a great number of species. Bouamer and Mo- of species, and described two species parasite of Geo- rand (2000), Bouamer et al. (2001a, 2001b), Bouamer chelone pardalis (Bell, 1828) from Zoo of Pretoria, and Morand (2002) emended the diagnosis of four South Africa. 226 S. BOUAMER and S. MORAND Walton (1942) was the first investigator of the para- Remarks. Baker (1987) cited 34 species and sub- sitic nematodes of tortoises from the Galapagos Islands species that are parasites of Testudinidae, Iguanidae (Neotropical region). and Agamidae, with 16 species and 9 sub-species para- Petter and Douglass (1976) provided a study on the sites of Testudinidae. Bouamer et al. (2001a) revised nematodes of Nearctic tortoises, with description and the diagnosis of this genus. redescription of several species. The parasitic nematodes of Oriental tortoises are Alaeuris numidica (Seurat, 1918) not well known. Baylis and Daubney (1922) redescrib- ed one species and Petter (1966) gave the description Synonyms. Alaeuris alaeuris Thapar, 1925 and of a second species. A. forcipiformis Forstner, 1960, by Petter (1966); Until now 104 parasite species of Testudinidae with Alaeuris numidica numidica (Seurat, 1918), new 79 Oxyurida and 25 Ascaridida can be considered as synonym. valid species with 29 species from Palaearctic region Host-type. Testudo graeca graeca Linneaus, (24 Oxyrida and 5 Ascaridida), 50 species from Ethi- 1758. opian region (34 Oxyrida and 16 Ascaridida), 17 spe- Type locality. Tunisia and Algeria, (Seurat (1918) cies from Nearctic region (16 Oxyrida and 1 Ascaridi- did not indicate precisely it locality-type exactly). da), 11 species from Neotropical region (5 Oxyurida Records. Testudo graeca ibera Palad, 1814 in and 6 Ascaridida) and 2 species from Oriental region London Zoo, by Thapar (1925); Tortoise, locality un- (2 Ascaridida). known, by Forstner (1960). Testudo graeca zarudnyi In the present study, we give the complete data of Nikolsky, 1896 from Iran; Testudo graeca graeca and the parasitic nematodes of Testudinidae. We give syn- Testudo horsfieldii Gray, 1844 locality unknown by onymies of each species, host-type, locality-type and Petter (1966). Testudo graeca, Kazakhstan by Sharpi- other records (if available). We then give the list of all lo (1976). Testudo graeca graeca from Settat, Moroc- valid species and on the basis of host species and bio- co and Testudo hermanni from South Catalonia, geographical distribution, we raised the subspecies to Spain, by Bouamer et al. (2001a). level of species. These changes follow previous studies on this group of Oxyurid (Bouamer and Morand, 2002, 2003, 2004, 2005). Alaeuris auricularis (Walton, 1942) This data set allows the investigation of the para- site species diversity in relation to host species diversi- Synonymy. Pseudoalaeuris auricularis Wal- ty and their biogeographical distribution. ton, 1942, by Petter (1966). Host-type. Chelonoidis nigra (Quoy et Graimard, 1824). TAXONOMY Type locality. Galapagos Islands. Alaeuris brevicollis (Walton, 1927) Order Oxyurida Synonymy. Thelastomoides Walton, 1927, by In the superfamilly Oxyuridea, the Pharyngodo- Petter and Douglass (1976). nidae is the only family parasitizing Testudinidae, Host-type. Gopherus polyphemus (Daudin, Only seven genera occur in this host group; the other 1802). genera of Pharyngodonidae occur in amphibians, fish Type locality. Georgia, USA. and mammals. Alaeuris caballeroi Petter et Douglass, 1976 Pharyngodonidae Travassos, 1919 Host-type. Gopherus flavomarginatus from There are six genera parasites of Testudinidae. Coahuila, Mexico. Type locality. Coahuila, Mexico. Alaeuris Thapar, 1925 Record. Gopherus agassizii from Sonora, Mexico, by Petter and Douglass (1976). Type-species. Alaeuris numidica (Seurat, 1918). Alaeuris conspicua Ortlepp, 1933 Synonyms. Pseudalaeuris Walton, 1942 by Petter (1966); Thelastomoides Walton, 1927 by Petter and Host-type. Psammobates tentorius (Bell, 1828). Douglass (1976); Travassolaimus Vigueras, 1938 by Type locality. Niekerk’Hope, Griqualand West; Petter and Quentin (1976). South Africa. NEMATODES PARASITES OF TESTUDINIDAE 227 Record. Geochelone pardalis (Bell, 1828) by Pet- Host-type. Chelonoidis nigra. ter (1966), South Africa. Type locality. Galapagos Islands. Alaeuris dupuisi Petter, 1966 Alaeuris madagascariensis Petter, 1966 Host-type. Pyxix arachnoides Bell, 1827. Synonymy. Alaeuris numidica madagascar- Type locality. Madagascar. iensis Petter (1966), new synonym. Record. Astrochelys radiata (Shaw, 1802) from Host-type. Astrochelys radiata (Shaw, 1802). Madagascar, by Petter (1966). Type locality. Madagascar. Alaeuris gopheri Petter et Douglass, 1976 Alaeuris mazzottii Petter et Douglass, 1976 Synonymy. Alaeuris gopheri gopheri Petter et Host-type. Gopherus flavomarginatus. Douglass, 1976, new synonym. Type locality. Coahuila, Mexico. Host-type. Gopherus flavomarginatus Legler, 1959. Type locality. Coahuila, Mexico. Alaeuris paramazzottii Petter et Douglass, 1976 Host-type. Gopherus polyphemus. Alaeuris kinsellai Petter et Douglass, 1976 Type locality. Florida, USA. Record. Gopherus agassizii from Sonora, Mexico, Synonymy. Alaeuris kinsellai kinsellai Petter by Petter and Douglass (1976). et Douglass, 1976, new synonym. Host-type. Gopherus flavomarginatus. Type locality. Coahuila, Mexico. Alaeuris pharyngodentata (Walton, 1942) Alaeuris insularis Petter, 1966 Synonymy. Pseudoalaeuris pharyngodentata Walton, 1942, by Petter (1966). Synonymy. Alaeuris quadrilabiata insularis Host-type. Chelonoidis nigra. Petter, 1966, new synonym. Type locality. Galapagos Islands. Host-type. Pyxix arachnoides. Type locality. Madagascar. Alaeuris poweri (Ortlepp, 1933) Records. Astrochelys radiata (Shaw, 1802) from Madagascar, by Petter (1966). Synonyms. Tachygonetria poweri Ortlepp, 1933, by Petter (1966); Pseudoalaeuris poweri (Ort- lepp, 1933), by Petter (1966). Alaeuris longicollis (Walton, 1927) Host-type. Psammobates tentorius. Type locality. Niekerk’Hope, Griqualand West; Synonymy. Thelastomoides Walton, 1927, by South Africa. Petter and Douglass (1976). Host-type. Gopherus polyphemus. Type locality. Georgia, USA. Alaeuris pudica Petter et Douglass, 1976 Records. Gopherus flavomarginatus from Coa- huila, and Gopherus agassizii from Sonora, Mexico, Synonymy. Alaeuris gopheri pudica Petter et by Petter and Douglasss (1976). Douglass, 1976, new synonym. Host-type. Gopherus agassizii (Copper, 1863). Alaeuris macrolabiata Petter et Douglass, 1976 Type locality. Sonora, Mexico. Synonymy. Alaeuris gopheri macrolabiata Pet- Alaeuris quadrilabiata (Ortlepp, 1933) ter et Douglass, 1976, new synonym. Host-type. Gopherus polyphemus. Synonyms. Tachygonetria quadrilabiata Ort- Type locality.