Nematodes Parasites of Testudinidae (Chelonia): List of Species and Biogeographical Distribution

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) redescribed 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 parasite 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. Florida, USA. lepp, 1933, and Pseudoalaeuris quadrilabiata (Ort- lepp, 1933) by Petter (1966); Alaeuris quadrilabiata Alaeuris macroptera (Walton, 1942) quadrilabiata (Ortlepp, 1933), new synonym. Host-type. Psammobates tentorius. Synonymy. Pseudoalaeuris macroptera Wal- Type locality. Niekerk’Hope, Griqualand West, ton, 1942, by Petter (1966). South Africa. 228 S. BOUAMER and S. MORAND

Alaeuris sonorae Petter et Douglass, 1976 zarudnyi and Testudo horsfieldii Gray, 1844 from Iran, by Petter (1966). Testudo hermanni from Alba- Synonymy. Alaeuris kinsellai sonorae Petter et nia, by Baruš and Johnson (1973). Testudo hors- Douglass, 1976, new synonym. fieldii and Testudo graeca from Volga River delta Host-type. Gopherus agassizii. and Caspian Steppe, Dagestan by Markov et al. (1962) Type locality. Sonora, Mexico. and Sharpilo (1976).

Alaeuris venustus (Leidy, 1856) Mehdiella cristata Petter, 1966

Synonymy. Thelastomoides Walton, 1927, by Host-type. Pyxix arachnoides. Petter and Douglass (1976). Type locality. Madagascar. Host-type. Gopherus polyphemus. Record. Astrochelys radiata from Madagascar, Type locality. Georgia, USA. by Petter (1966).

Gopheruris Petter et Douglass, 1976 Mehdiella dollfusi Petter, 1966

Remark. A mono-species genus erected by Petter Synonymy. Mehdiella stylosa dollfusi Petter, and Douglass (1976). 1966, new synonym. Type species. Gopheruris aspicula Petter et Host-type. Pyxix arachnoides. Douglass, 1976. Type locality. Madagascar. Records. Astrochelys radiata from Madagascar by Petter (1966). Astrochelys yniphora (Vaillant, 1885) Gopheruris aspicula Petter et Douglass, 1976 from North-West of Madagascar, by Petter and Brygoo (1972). Host-type. Gopherus agassizii. Type locality. Sonora, Mexico. Mehdiella grassei Petter, 1966 Mehdiella Seurat, 1918 Host-type. Astrochelys radiata. Type locality. Madagascar. Type species. Mehdiella microstoma (Drasche, 1884). Remarks. Petter (1966) split Mehdiella stylosa in Mehdiella longistima Petter, 1966 sub-species. We raise these sub-species to two species. We recognize M. dubinina Skrjabin et Schkhobalova, Host-type. Testudo horsfieldii. 1951 (Bouamer et al. 2001b). Type locality. Iran.

Mehdiella microstoma (Drasche, 1884) Mehdiella petterae Bouamer, Morand et Bourgat, 2001

Synonyms. Oxyuris microstoma Drasche, 1884 Host-type. Testudo hermanni. by Seurat (1918); Tachygonetria microstoma Tha- Type locality. South Catalona, Spain. par, 1925 in part, by Petter (1961); Tachygonetria weissei Seurat, 1918, by Dittmann (1965); Macracis Mehdiella stylosa (Thapar, 1925) papillosa Forstner, 1960, by Petter (1966). Host-type. Testudo graeca. Synonyms. Tachygonetria stylosa females sen- Type locality. Europe, locality unknown. su Thapar 1925, by Petter (1966). Mehdiella hamosa Records. Testudo kleinmanni Lortet, 1833 from Forstner, 1960, by Petter (1966). Mehdiella cordata Egypt by Baylis (1923). Testudo sp. from Afghanistan Forstner, 1960, by Petter (1966). Mehdiella stylosa by Akthar (1937). Testudo graeca ibera by Thapar stylosa (Thapar, 1925), by Bouamer et al. (2003). (1925). Testudo horsfieldii from Tadzhikistan, by Du- Host-type. Testudo graeca ibera. binina (1949). Testudo graeca graeca and Testudo Type locality. Unknown (but turtle lived in Lon- graeca ibera from Europe by Dyk and Dykova (1956), don Zoo). Forstner (1960) and Schad, Kuntz and Wells (1960). Records. Testudo graeca graeca and Testudo Testudo hermanni from Greece by Dittmann (1965). graeca ibera from Palaearctic region, by Thapar Testudo hermanni from Romania, Testudo graeca (1925); Forstner (1960) and Schad, Kuntz and Wells graeca from Algeria and Morocco, Testudo graeca (1960). Testudo graeca graeca from Algeria and NEMATODES PARASITES OF TESTUDINIDAE 229

Morocco, Testudo hermanni from Romania, Testudo Type locality. Madagascar. graeca zarudnyi and Testudo horsfieldii Gray, 1844 from Iran, by Petter (1966). Testudo hermanni Tachygonetria Wedl, 1862 from Albania by Johnson (1973). Testudo horsfieldii and Testudo graeca from Dagestan, Kasakhistan and Type species. of this genus parasite Tachygone- Tadzhikistan, by Sharpilo (1976). tria viviparia Wedl, 1862, of Uromastix spinipes Boulenger, 1885 (Agamidae) from Egypt. Mehdiella uncinata (Drasche, 1884) Synonyms. Pracis Railliet et Henry, 1916 by Wal- ton (1942). Synonyms. Oxyuris inflata Drasche, 1884, by Se- Remarks. Seurat (1918) listed the following species urat (1918). Oxyuris inflata Stossich, 1898, by Seurat names without description: Tachygonetria jugur- (1918). Oxyuris albanica Stossich, 1898, by Seurat thae Seurat, 1918, T. lambdiensis Seurat, 1918, and (1918). Tachygonetria uncinata (Drasche) sensu T. massinissae Seurat, 1918. Petter (1966) considered Thapar (1925), by Petter (1966). Tachygonetria in- this species a Nomina nuda. Petter (1966) splitting flatocervix Akhtar, 1937, by Petter (1966). Tachygo- Tachygonetria dentata (Drasche, 1884), T. longico- netria uncinata (Drasche) sensu Dubinina (1949). lis (Schneider, 1866), T. macrolaimus (Linstow, Mehdiella inflata Forstner, 1960, by Petter (1966). 1899), Bouamer and Morand (2004) raised these sub- Mehdiella uncinata albanica Johnson, 1973, by Bak- species to level of species. Petter (1966) splitting T. co- er (1987). nica (Drasche, 1884) in two sub-species: T. conica Host-type. Testudo graeca. conica (Drasche, 1884) and T. conoca nicollei (Seu- Type locality. Trieste, Italy. rat, 1918), Bouamer and Morand (2002) suppressed Records. Testudo graeca graeca and Testudo T. conica nicollei. Petter (1966) transferred Tachy- graeca ibera by Stossich (1898), Seurat (1918), Tha- gonetria thapari Dubinina, 1949 in the genus Tha- par (1925), Ryšavý (1958) and Forstner (1960). Testu- paria, but Baker (1987) recognised this species in ge- do kleinmanni Lortet, 1833 from Egypt, by Baylis nus Tachygonetria. We agree with Petter’s opinion. (1923). Testudo horsfieldii from India, by Rees Tachygonetria sub-species described only on the (1935), and from Tadzhikistan, by Dubinina (1949). morphology of females by Petter (1966) were consid- Testudo graeca graeca from Algeria and Morocco, ered as Nomina nuda, while waiting for new rede- Testudo hermanni from Romania, Testudo graeca scription. Bouamer and Morand (2002) redescribed the zarudnyi and Testudo horsfieldii Gray, 1844 from type species of the genus and emended the diagnosis. Iran, by Petter (1966). Testudo hermanni from Alba- nia by Johnson (1973). Testudo graeca and Testudo hermanni from Baltic region, by Hristovski (1973). Tachygonetria combesi Bouamer et Morand, 2002 Testudo horsfieldii and Testudo graeca from Volga River delta and Caspian Steppe, Dagestan by Markov Host-type. Testudo hermanni. et al. (1962) and Sharpilo (1976). Type locality. South Catalona, Spain.

Ortleppnema Petter, 1966 Tachygonetria conica (Drasche, 1884)

Type species. Ortleppnema possompesi Petter, Synonyms. Oxyuris conica Drasche, 1884, by Se- 1966. urat (1918). Tachygonetria lobata Dubinina, 1949, by Remark. Petter (1966) erected the genus Ortlepp- Petter (1966). Tachygonetria stylosa Thapar, 1925, nema for two species parasites of Testudunidae from by Petter (1966). Tachygonetria longicollis (Schnei- Madagascar, Pyxix arachnoides and Astrochelys der, 1866) males sensu Thapar (1925) and Dubinina radiata. (1949), by Petter (1966). Tachygonetria conica nicolei (Seurat, 1918) by Bouamer and Morand (2002). Ortleppnema possompesi Petter, 1966 Host-type. Testudo graeca ibera. Type locality. Europe, locality unknown. Host-type. Pyxix arachnoides. Records. Testudo graeca graeca from Tunisia, Type locality. Madagascar. Algeria, by Seurat (1918). Testudo graeca ibera from Record. Astrochelys radiata (Shaw, 1802) from Europe, by Schad, Kuntz and Wells (1960) and Tha- Madagascar, by Petter (1966). par (1925). Testudo horsfieldii from Tadzhikistan, by Dubinina (1949). Testudo graeca graeca from Algeria Ortleppnema radiatum Petter, 1966 and Morocco; Testudo graeca zarudnyi from Iran and Testudo horsfieldii from Caspian region; Testu- Host-type. Astrochelys radiata. do hermanni from Italy, by Petter (1966). Testudo 230 S. BOUAMER and S. MORAND horsfieldii and Testudo graeca from Volga River in part, by Thapar (1925). Tachygonetria massinisse delta and Caspian Steppe, Dagestan by Markov et al. Seurat, 1918, by Thapar (1925). Paracis longicollis (1962) and Sharpilo (1976). Testudo hermanni from Railliet and Henry, 1916 in part, by Thapar (1925). Ta- Albania by Baruš and Johnson (1973). Testudo gra- chygonetria microlaimus female sensu Thapar eca graeca from Morocco, by Bouamer and Morand (1925), by Petter (1966). Tachygonetria testudinis (2002). Forstner (1960), by Petter (1966). Tachygonetria forstneri Skrjabin, Schikholova and lagodovkaja, Tachygonetria dentata (Drasche, 1884) 1967, by Baker (1987). Tachygonetria longicollis longicollis (Seurat, 1918), by Bouamer and Morand Synonyms. Oxyuris dentata Drasche, 1884 by (2004). Host-type. Testudo graeca ibera. Seurat (1918). Tachygonertia expansa Ress, 1935, Type locality. Europe, locality unknown. by Petter (1966). Tachygonertia torticolis Ress, Records. Testudo graeca graeca from Tunisia 1935, by Petter (1966). Tachygonetria dentata and Algeria, by Seurat (1918). Testudo graeca ibera dentata (Drasche, 1884) by Bouamer and Morand from Europe, by Drasche (1884), Thapar (1925) and (2002). Forstner (1960). Testudo graeca graeca from Algeria Host-type. Testudo graeca ibera. and Morocco, Testudo graeca zarudnyi from Iran Type locality. Europe, locality unknown. and Testudo horsfieldii from Caspian region, by Pet- Records. Testudo graeca ibera from Europe, by ter (1966). Testudo hermanni from Albania by Baruš Drasche (1884), Thapar (1925), Ryšavý (1958), Forst- and Johnson (1973). Testudo horsfieldii and Testudo ner (1960) and Schad, Kuntz and Wells (1960) (locality graeca from Volga River delta and Caspian Steppe, unknown). Testudo kleinmanni from Egypt by Baylis Dagestan, by Markov et al. (1962) and Sharpilo (1976). (1923). Testudo graeca graeca from Tunisia and Al- geria, by Seurat (1918). Testudo horsfieldii from Tadzhikistan, by Dubinina (1949). Testudo graeca Tachygonetria longispiculum Petter, 1966 graeca from Algeria and Morocco, Testudo graeca zarudnyi from Iran and Testudo horsfieldii from Host-type. Psammobates tentorius. Caspian region, Testudo hermanni from from Italy, Type locality. Kimberley, South Africa. by Petter (1966). Testudo horsfieldii and Testudo graeca from Dagestan, Kasakhistan and Tadzhikistan, Tachygonetria macrolaimus (Linstow, 1899) by Sharpilo (1976). Testudo graeca graeca from Mo- rocco, by Bouamer and Morand (2002). Synonymy. Tachygonetria macrolaimus macrolaimus (Linstow, 1899) sensu Petter (1966), Ta- Tachygonetria dessetae Petter, 1966 chygonetria macrolaimus macrolaimus, by Bou- amer and Morand (2004). Synonym. Tachygonetria macrolaimus desse- Host-type. Geochelone pardalis. tae Petter, 1966, new synonym. Type locality. Berlier Aquarium, South Africa. Host-type. Pyxix arachnoides. Records. Geochelone pardalis, Zoo of Pretoria, Type locality. Madagascar. South Africa; same host from Swaziland, by Petter (1966). Records. Astrochelys radiata from Madagascar, by Petter 1966). Astrochelys yniphora from Mada- Tachygonetria microlaimus (Linstow, 1899) gascar, by Petter and Brygoo (1972). Synonymy. Oxyuris microlaimus Linstow, 1899, Tachygonetria fitzsimmonsi Petter, 1966 by Seurat (1918). Host-type. Geochelone pardalis. Type locality. South Africa, locality unknown. Synonym. Tachygonetria longicollis fitzsim- Records. Geochelone pardalis; Zoo of Pretoria, monsi Petter, 1966, by Bouamer and Morand (2004) South Africa and Swaziland, by Petter (1966). Testudo Host-type. Geochelone pardalis (Bell, 1828). horsfieldii and Testudo graeca from Dagestan, Ka- Type locality. Zoo of Pretoria, South Africa. zakhstan and Tadzhikistan, by Sharpilo (1976). Records. Geochelone pardalis from Swaziland, by Petter (1966). Tachygonetria nearctica Petter et Douglass (1976) Tachygonetria longicollis (Schneider, 1866) Synonymy. Tachygonetria dentata neartica Synonyms. Oxyuris longicollis Schneider, 1866, Petter et Douglass (1976), by Adamson and Nasher by Seurat (1918). Oxyuris longicollis Drasche, 1884 (1984). NEMATODES PARASITES OF TESTUDINIDAE 231

Host-type. Gopherus agassizii. Tachygonetria quentini Petter, 1966 Type locality. Sonora, Mexico. Synonymy. Tachygonetria dentata quentini Tachygonetria numidica Seurat, 1918 Petter, 1966, by Adamson and Nasher (1984). Host-type. Psammobates tentorius. Synonymy. Tachygonetria macrolaimus (Lin- Type locality. Kimberley, South Africa. stow, 1899) sensu Thapar 1925, by Petter (1966). Record. Geochelone pardalis from Victoria West, Host-type. Testudo graeca graeca. South Africa, by Petter (1966). Type locality. Algeria. Records. Testudo graeca graeca from Tunisia Tachygonetria richardae Petter, 1966 and Algeria, by Seurat (1918). Testudo graeca ibera from Europe, by Thapar (1925) and by Schad, Kuntz Synonymy. Tachygonetria dentata richardae and Wells (1960). Testudo graeca graeca from Alge- Petter, 1966, by Adamson and Nasher (1984). ria and Morocco, by Petter (1966). Host-type. Pyxix arachnoides. Type locality. Madagascar. Tachygonetria palearcticus Seurat, 1918 Records. Astrochelys radiata from Madagascar, by Petter (1966). Synonyms. Oxyuris macrolaimus Linstow, 1899, by Seurat (1918). Tachygonetria macrolai- Tachygonetria robusta (Drasche, 1884) mus (Linstow, 1899) sensu Seurat (1918), by Petter (1966). Tachygonetria pusilla females sensu Thapar Synonyms. Oxyuris Draschei Stossich, 1898, by 1925, by Petter (1966). Tachygonetria pusilla fe- Baylis (1923). O. robusta Drasche, 1884, by Dubinina males sensu Dubinina (1949) by Petter (1966). Tachy- (1949). O. lata Forstner, 1960, by Petter (1966). Meh- gonetria macrolaimus Petter, 1961, by Petter (1966). diella micristoma (Drasche, 1884), sensu Seurat Tachygonetria conica males sensu Thapar 1925, by (1918) in part, by Petter (1966). Tachygonetria mi- Petter (1966). Tachygonetria conica males sensu crostoma (Drasche, 1884), sensu Thapar (1925) in Dubinina (1949), by Petter (1966). Tachygonetria part, by Petter (1966). macrolaimus palearcticus Petter, 1966, new syn- Host-type. Testudo graeca ibera. onym. Type locality. Europe, locality unknown. Host-type. Testudo graeca graeca. Records. Testudo graeca graeca from Tunisia Type locality. Tunisia. and Algeria, by Seurat (1918). Testudo kleinmanni Records. Testudo graeca graeca from Tunisia from Egypt by Baylis (1923). Aspideretes leithii Algeria and Morocco; Testudo graeca zarudnyi (Gray, 1872) (fresh water Turtles) from India by Baylis from Iran and Testudo horsfieldii from Caspian (1936). Testudo graeca ibera from Europe, by Thapar region; Testudo hermanni from Italy, by Petter (1925) and Forstner (1960). Testudo horsfieldii from (1966). Tadzhikistan, by Dubinina (1949). Testudo graeca graeca from Algeria and Morocco; Testudo graeca Tachygonetria pusilla Seurat, 1918 zarudnyi from Iran and Testudo horsfieldii from Caspian region, by Petter (1966). Testudo horsfieldii Synonyms. Oxyuris longicollis Drasche, 1884, and Testudo graeca from Volga River delta and Cas- nec Schneider, 1866, by Seurat (1918). Paracis pian Steppe, Dagestan by Markov et al. (1962) and longicollis (Schneider, 1866), in part by Thapar Sharpilo (1976). Testudo horsfieldii from Afganistan, (1925). Tachygornetria longicollis pusilla (Seurat, by Ryšavý and Johnson (1979). Testudo graeca grae- 1918) sensu Petter (1966), by Bouamer and Morand ca from Morocco, by Bouamer and Morand (2002). (2005). Host-type. Testudo graeca graeca. Tachygonetria rosickyi Ryšavý et Johnson, 1979 Type locality. Tunisia. Records. Testudo graeca ibera from Europe, by Host-type. Testudo horsfieldii. Schneider (1866) and Thapar (1925). Testudo graeca Type locality. Afghanistan. graeca from Algeria and Morocco, Testudo graeca zarudnyi from Iran and Testudo horsfieldii from Tachygonetria seurati Petter, 1966 Caspian region, by Petter (1966). Testudo horsfieldii and Testudo graeca from Volga River delta and Synonyms. Tachygoentria microlaimus males Caspian Steppe, Dagestan, by Markov et al. (1962) and senu Thapar (1925), by Petter (1966), Tachygoentria Sharpilo (1976). microlaimus sensu Petter (1961), by Petter (1966). 232 S. BOUAMER and S. MORAND

Host-type. Testudo graeca graeca. Tachygonetria khallaayounei Bouamer et Morand, 2005 Type locality. Morocco. Records. Testudo graeca graeca from Algeria and Host-type. Testudo graeca graeca. other locality unknown from North Africa, by Petter Type locality. Settat, Morroco. (1961, 1966). Tachygonetria settatensis Bouamer et Morand, 2003 Tachygonetria testudinis Walton, 1942 Host-type. Testudo graeca graeca. Type locality. Settat, Morroco. Host-type. Chelonoidis nigra (Quoy et Graimard, 1824). Type locality. Galapagos Islands. Tachygonetria africana Bouamer et Morand, 2004

Host-type. Geochelone pardalis (Bell, 1828). Tachygonetria tetrapapillata (Caballero, 1944) Type locality. Zoo of Pretoria, South Africa.

Synonymy. Tachygoentria macrolaimus tetra- Tachygonetria pretoriensis Bouamer et Morand, 2004 papillata (Caballero, 1944), new synonym. Host-type. Gopherus polyphemus. Host-type. Geochelone pardalis Type locality. Valle de Mexico, Mexico. Type locality. Zoo of Pretoria, South Africa Records. Gopherus flavomarginatus from Co- ahuila, and Gopherus agassizii from Sonora, Mexico Thaparia Ortlepp, 1933 Gopherus polyphemus from Florida, USA, by Petter and Douglass (1976). Type species. Thaparia macrospiculum Ort- lepp, 1933. Tachygonetria setosa (Seurat, 1918) Remarks. Petter (1966) splitting the species Tha- paria thapari Dubinina, 1949 in two sub-species: Synonyms. Tachygonetria longicollis (Schnei- Thaparia thapari thapari (Dubinina, 1949) and der, 1866) sensu Seurat (1918) in part, by Petter Thaparia thapari australis Petter, 1966; Johnson 1966). Tachygonetria longicolis females sensu Tha- (1973) added a sub-species Thaparia thapari rysavyi par (1925), by Petter (1966). Tachygonetria pusilla Johnson, 1973. Bouamer and Morand (2000) redefined males sensu Thapar (1925), by Patter (1966). Tachy- the genus. gonetria longicollis setosa sensu Patter (1966), by Bouamer and Morand (2005). Thaparia australis Petter, 1966 Host-type. Testudo graeca graeca. Type locality. Tunisia. Synonymy. Thaparia thapari australis Petter, Records. Testudo graeca graeca from North Af- 1966, new synonym. rica; Testudo hermanni from Europe Locality un- Host-type. Geochelone pardalis. known, by Petter (1966). Type locality. Zoo of Pretoria, South Africa.

Thaparia bourgati Bouamer et Morand, 2000 Tachygonetria verreauxi Petter, 1966 Host-type. Testudo hermanni. Host-type. Psammobates tentorius. Type locality. South Catalona, Spain. Type locality. Kimberley, South Africa. Thaparia capensis Fitzsimmons, 1961 Tachygonetria poulini Bouamer et Morand, 2003 Host-type. Chersina angulata (Schweigger, Host-type. Testudo graeca graeca. 1812) Type locality. Morroco. Type locality. Cape Province, South Africa.

Tachygonetria marocana Bouamer et Morand, 2005 Thaparia carlosfelui Bouamer et Morand, 2000

Host-type. Testudo graeca graeca. Host-type. Testudo hermanni. Type locality. Settat, Morroco. Type locality. South Catalona, Spain. NEMATODES PARASITES OF TESTUDINIDAE 233

Thaparia contortospicula Walton, 1942 Thelandros Wedl, 1862

Host-type. Chelonoidis nigra. Synonymy. Avilandros Skrjabin, Schikholova et Type locality. Galapagos Islands. Mozgovoi, 1951. Remarks. The genus Thelandros Wedl, 1862, with Thaparia domerguei Petter, 1966 type species T. atalus Wedl, 1862 from Uromatrix aegyptia (Forskal, 1775) (Agamidae) contain beside Host-type. Pyxix arachnoides. the parasitic of Ethiopian Testudinidae, the parasites Type locality. Madagascar. of herbivores Reptiles (Agamidae, Iguanidae, Scincidae Record. Astrochelys radiata from Madagascar, and Geckonidae) (see Adamson 1981, Adamson and by Petter (1966). Nasher 1984). Petter (1966) erected a sub-genus Archithelandros Thaparia macrocephala Petter et Douglass, 1976 for a species with caudal alae and she split the species parasitic Testudunidae in sub-species. Adamson Host-type. Gopherus flavomarginatus. (1981) and Adamson and Nasher (1984) raised these Type locality. Coahuila, Mexico. species to the level of species. We agree with Adamson (1981) and Adamson and Nasher (1984). Petter (1966) described the females of Oxyuris sp. Thaparia macrospiculum Ortlepp, 1933 as member of a sub-genus Thelandros (Archithelen- dros) Petter, 1966. We considered this species Nomi- Host-type. Psammobates tentorius. Type locality. Niekerk’s Hope, Griqualand West; na Nuda. Adamson and Nasher (1984) suggested that South Africa. this species is probably as an Alaeuris sp. Record. Geochelone pardalis from South Africa, by Petter (1966). Thelandros (Archithelandros) Petter, 1966 Thelandros (Ar.) pyxis Petter, 1966 Thaparia microcephala Petter et Douglass, 1976 Host-type. Pyxix arachnoides. Host-type. Gopherus flavomarginatus. Type locality. Madagascar. Type locality. Coahuila, Mexico.

Thelandros (Ar.) dolichurus Petter, 1966 Thaparia rysavyi Johnson, 1973 Host-type. Astrochelys radiata. Synonymy. Thaparia thapari rysavyi Johnson, Type locality. Madagascar. 1973, new synonym. Record. Pyxix arachnoides from Madagascar, by Host-type. Testudo hermanni. Petter (1966). Type locality. Albania.

Thelandros (Ar.) ortleppii Petter, 1966 Thaparia thapari (Dubinina, 1949)

Host-type. Psammobates tentorius. Synonyms. Oxyuris sp. females sensu Thapar Type locality. Kimberley, South Africa. (1925). Tachygonetria thapari Dubinina, 1949, by Petter (1966). Thaparia thapari thapari (Dubinina, 1949), new synonym. Thelandros (Thelandros) Wedl, 1862 Host-type. Testudo horsfieldii. Thelandros (Thelandros) tcheprakovae Type locality. Tadzhikistan. Petter, 1966 Records. Testudo graeca ibera from Europe (locality unkown), by Thapar (1925) and Schad et al. (1960). Host-type. Psammobates tentorius. Testudo graeca (L.) from Europe by Dyk and Dykova Type locality. Kimberley, South Africa. (1956). Testudo graeca graeca from Algeria and Mo- rocco, Testudo graeca zarudnyi from Iran and Testu- do horsfieldii from Caspian region, Testudo herman- Thelandros (Th.) versterae Petter, 1966 ni from Italy, by Petter (1966). Testudo horsfieldii from Afghanistan, by Johnson (1973). Testudo graeca Host-type. Psammobates tentorius. graeca from Morocco, by Bouamer and Morand (2000). Type locality. Kimberley, South Africa. 234 S. BOUAMER and S. MORAND

Thelandros (Th.) weilleae Petter, 1966 by Sprent (1980), Emys orbicularis (Linnaeus, 1758) (fresh water Turtle) from Europe, by Linstow (1909), Host-type. Psammobates tentorius. from Volga River delta and Caspian Steppe, Dagestan, Type locality. Kimberley, South Africa. by Markov et al. (1962) and Sharpilo (1976). Terra- pene Carolina (Linnaeus, 1758) (wet and marshy land Thelandros (Th.) sexlabiata Ortlepp, 1933 turtle) from Leningrad Museum, by Linstow (1903). Tortoise (species unknown) from North America, by Stiles and Hassall (1894), Pillers (1933) and Jansen Synonymy. Parapharyngodon sexlabiata Fre- and den Broek (1966). itas, 1957, by Petter (1966). Host-type. Psammobates tentorius. Type locality. Niekerk’Hope, Griqualand West; Superfamily Cosmocercoidea South Africa. Records. Geochelone pardalis from South Africa, Remarks. There are two families, Atractidae Rail- by Petter (1966). liet, 1917 and Cosmocercoidae Railliet, 1916, parasites of Testudinidae.

Order Ascaridida Family Atractidae Railliet, 1917 Superfamily Ascaridoidea Remark. This family contain five genera parasites Remark. Only the family of Ascarididae Baird, 1853 of Testudinidae. Petter (1966) splitt Atractis dacty- parasite of Reptiles. luris (Rudolphi, 1819) in sub-species. We raise these sub-species to level of species. Family Ascarididae Baird, 1853 Atractis Dujardin, 1845 Subfamily Ascaridinae Baird, 1853 Type species. Atractis dactyluris (Rudolphi, Remark. There is only one genus and only one spe- 1819). cies parasite of Testudinidae. Synonyms. Cyrtosomum Gedoelst, 1919 and Pseudatractis Yamaguti, 1961, by Baker (1987). Angusticaecum Baylis, 1920 Atractis africana Ortlepp, 1933 Type species. Angusticaecum holopterum (Ru- dolphi, 1819) Host-type. Psammobates tentorius. Type locality. Niekerk’Hope, Griqualand West; Angusticaecum holopterum (Rudolphi, 1819) South Africa.

Synonyms. Ascaris holoptera Rudolphi, 1819, by Atractis baltazardi Petter, 1966 Baylis (1920). Ascaris holoptera Rudolphi, 1819 sensu Dujardin (1845) in part by Sprent (1980). Angusti- Synonymy. Atractis daclyluris baltazardi Pet- caecum brevispiculum Charpin, 1924. ter, 1966, new synonym. Host-type. Testudo graeca. Host-type. Testudo graeca zarudnyi. Type-locality. Europe, locality unknown. Type locality. Iran. Records. Testudo marginata (Schoepff, 1795) from Europe, by Creplin (1846). Testudo horsfieldii Atractis chabaudi Petter, 1966 from Tadzhikistan, by Dubinina (1949). Testudo graeca ibera from Europe, by Forstner (1960) and by Host-type. Pyxix arachnoides. Schad, Kuntz and Wells (1960). Testudo graeca gra- Type-locality. Madagascar. eca from North Africa by Stossich (1898). Testudo hermanni from Petrich region, Bulgaria, by Katalan- Atractis daclyluris (Rudolphi, 1819) Gateeva and Dimitrov (1961) and from Greece, by Dittman (1965). Testudo graeca from Volga River Synonyms. Ascaris dactyluris Rudolphi, 1819 delta and Caspian Steppe, Dagestan, by Markov and and Atractis brevicollis Schneider, 1866, by Thapar Kriuchkov (1961) and Sharpilo (1962, 1976). Chelo- (1925). noidis denticulata from Brazil, by Chaprin (1924), Host-type. Testudo graeca ibera. Kinyxix belliana from Melbourne zoo, South Africa Type-locality. Roma, Italy. NEMATODES PARASITES OF TESTUDINIDAE 235

Records. Testudo graeca graeca from Algeria, Atractis morinae Baer, 1936 by Hallez (1886). Aspideretes leithii from India, by Baylis and Daubney (1922). Chelonoidis denticu- Host-type. Astrochelys radiata. lata from South America (locality unknown), by Tha- Type-locality. Madagascar. par (1925). Testudo graeca graeca from Algeria, Testudo graeca ibera from Roma, Italy, by Petter Atractis perarmata Linstow, 1910 (1966). Testudo hermanni from Balkan, by Hristovski (1973). Host-type. Kinixys belliana. Type-locality. West Africa, locality unknown. Atractis dubininae Petter, 1966 Atractis trematophila Travassos, 1934 Synonyms. Oxyuris sp. sensu Dubinina 1949 by Petter (1966), Atractis daclyluris dubininae Petter, Host-type. Unidentified turtle (specimens found in 1966, new synonym. the digestive tract of a digenetic trematoda (Param- Host-type. Testudo horsfieldii. phistomoidea; Chabaud and Petter (1960) suggested Type locality. Tadzhikistan. that the specimens are parasites of turtle and were Records. Testudo horsfieldii from Tadzhikistan, ingested by a parasitic trematode). by Dubinina (1947). Testudo horsfieldii and Testudo Type-locality. Amazon River, Brazil. graeca from Dagestan, Kasakhistan and Tadzhikistan, by Sharpilo (1976). Testudo horsfieldii from Afghan- Atractis thapari Petter, 1966 istan, by Baruš et al. (1972). Synonymy. Atractis daclyluris, sensu Thaparia (1925), by Petter (1966) Atractis emilii Berenger, 1945 Host-type. Chelonoidis denticulata. Type-locality. America, locality unknown. Host-type. Testudo hermanni hermanni. Type-locality. Balear Islands, Spain. Fitzsimmonsnema Petter, 1966 Atractis fasciolata Gendre, 1909 Type species. Fitzsimmonsnema reptiliae (Fitz- simmons, 1958). Host-type . Kinixys belliana Gray, 1831. Remark. Mono-species genus erected by Petter Type-locality. Guinea. (1966).

Atractis granulosa (Railliet et Henry, 1912) Fitzsimmonsnema reptiliae (Fitzsimmons, 1958)

Synonyms. Atractis granulosa granulosa Rail- Synonymy. Probstmayria reptiliae Fitzsim- liet et Henry, 1912, by Baer (1936). Atractis dacly- mons, 1958 by Petter (1966). luris sensu Baylis and Daubney (1922), by Petter Host-type. Homopus femoralis Boulenger, 1888. (1966). Type-locality. South Africa. Host-type. Manouria emys emys (Schlegel et Müller, 1844). Ibrahima Khalil, 1932 Type-locality. Indomalaysia, locality unknown. Record. Geochelone elegans (Schoepff, 1794) from Type sepcies. Ibrahima ibrahima Khalil, 1932. Bengal, by Baylis and Daubney (1922). Remark. Mono-species genus erected by Khalil (1932). Atractis impura Caballero, 1944 Ibrahima ibrahima Khalil, 1932 Host-type. Gopherus polyphemus. Type-locality. Azacapotzalco, Mexico. Host-type. Tortoise unidentified. Records. Kinosternon scorpioides (Linnaeus, Type-locality. Liberia. 1766) (Kinosternidae, wet land Turtles), from Costa Rica, by Brenes and Bravo Holls (1960). Gopherus Labiduris Schneider, 1866 agassizii from Sonora, Mexico, by Petter and Dou- glass (1976). Type species. Labiduris gulosa (Rudolphi, 1819). 236 S. BOUAMER and S. MORAND

Studies Host species diversity A B

Parasite genus diversity Parasite species diversity C D

Oxyurida species diversity Ascaridida species diversity E F

Palaearctic

Ethiopian

Nearctic

Neotropical

Oriental

Figure 1. (A) Number of studies by biogeographical region; (B) number the hosts by biogeographical region; (C) genus diversity of nematode by biogeographical region; (D) species diversity of nematode by biogeographical region; (E) Oxyurida species diversity biogeographical region; (F) Ascaridida species diversity biogeographical region. NEMATODES PARASITES OF TESTUDINIDAE 237

Labiduris africana Gedoelst, 1916 Family Cosmocercoidae Railliet, 1916

Host-type. Kinixys erosa (Schweigger, 1812). Remark. There is only one genus Raillietnema Type-locality. Leopoldville, Democratic Republic Travassos, 1927, which is a diversified genus of the of Congo. family Cosmocercoidae found in many amphibians. Records. Kinixys australis Hewitt, 1935 from Swaziland and Kinixys erosa from Malawi, by Fitz- Raillietnema Travassos, 1927 simmons (1961). Type species. Raillietnema simples (Travassos, Labiduris bryooi Petter, 1966 1927) in Hyla faber Wied, 1824 from Brazil.

Host-type. Pyxix arachnoides. Raillietnema bainae Petter, 1966 Type-locality. West of Madagascar. Host-type. Kinixys erosa. Labiduris gulosa (Rudolphi, 1819) Type-locality. M’Baïki, Centre Afrique Republic.

Synonymy. Ascaris gulosa Rudolphi, 1819 by Raillietnema kinixys Fitzsimmons, 1964 Schneider (1866). Host-type. Chelonoidis denticulata. Host-type. Kinixys bellina. Type-locality. South America, locality unknown. Type-locality. Nyasaland. Record. Chelonoidis denticulata from Para, Brazil, by Charpin (1924), Chelonoidis denticulata from Bolivar, Venezuela, by Inglis and Diaz-Ungria Superfamily Heterakoidea (1963). Remark. The family Heterakidae Railliet and Hen- ry, 1912 parasite birds, herptiles and rarely mammals. Labiduris ireneata Gonçalves da Costa, 1961

Host-type. Chelonoidis denticulata. Family Heterakidae Railliet et Henry, 1912 Type-locality. Brazil. Record. Baker (1987) cited Inglis and Diaz-Ungria Remark. With only one species and type-species of (1963) reporting Labiduris ireneata parasite of the the genus. “same host from Bolivia and Venezuela”. However, we do not find any mention of this sentence in Inglis and Diaz-Ungria (1963). Africana Travassos, 1920 Africana africana (Gendre, 1909)

Labiduris zshokkei Linstow, 1899 Host-type. Kinixyx belliana. Type locality. Guinea. Host-type. Chelonoidis denticulata. Records. Kinixyx belliana from Malawi, by Type-locality. Brazil. Fitzsimmons (1961). Kinixyx erosa from Congo Record. Chelonoidis denticulata from Brazil and (Baylis, 1939, 1940). Varanus arenarius (Squamata) Venezuela, by Inglis and Diaz-Ungria (1963). from Dahomey, by Gendre (1909). Chamaeleo crista- tus Stuchbury, 1837 (Squamata) from Cameroon, by Rasheed (1965). Varanus niloticus (Linnaeus, 1766) Orientatractis Petter, 1966 (Squamata) from Congo, by Graber (1981). Type species. Orientatractis levanhoai Petter, 1966. DISCUSSION Remark. Mono-species genus erected by Petter (1966). On the basis of host species and biogeographical distribution, we raised the subspecies to level of Orientatractis levanhoai Petter, 1966 species. These changes follow previous studies on this group of Oxyurid (Bouamer and Morand 2002, Host-type. Indotestudo elongata (Blyth, 1853). 2003, 2004, 2005). Moreover, the sequence of 12S gene Type-locality. Saigon, Vietnam. the following species Mehdiella microstoma and 238 S. BOUAMER and S. MORAND

Tachygonetria palearcticus parasites of two differ- (Figs 1C, 1D). A similar observation is noted concern- ent species hosts (Testudo greaca from Morrocco and ing the species diversity of Oxyurids and Ascarids Testudo hermanni from Spain) shows significant dif- (Figs 1E, 1F) ferences among species (Bouamer 2003). We found a good significant relationship between In the genus Alaeuris, subspecies Alaeuris nu- the number of studies and the species diversity of hosts midica numidica and Alaeuris numidica mada- (p < 0.0001); a significant relationship between the gascariensis are parasites of different hosts (Testu- species diversity of parasites and the generic diversity do graeca graeca and Astrochelys radiata) from of the parasites (p = 0.02). We also found a significant distant geographical regions (Neotropical and Mada- relationship between the generic diversity of parasites gascar). In the same manner, he subspecies Alaeuris and the species diversity of hosts (p = 0.02), and gopheri gopheri and Alaeuris gopheri pudica a weak relationship between the species diversity of Gopherus are found on two different hosts (Gopherus parasites and the species diversity of hosts (p = 0.07). flavomarginatus and Gopherus agassizii) also The parasite species diversity of nematodes seems to originated from two geographical regions. be linked to the species diversity of hosts. Five other pairs of subspecies are also raised to the Finally the host specificity does not differ between level of species for the same reasons: Alaeuris kinsel- the Oxyurida (1.6 ± 0.9 host species) and the Ascaridi- lai sonorae and Alaeuris kinsellai kinsellai from da (1.6 ± 1.2 host species). Gopherus flavomarginatus and Gopherus agassizii; Alaeuris quadrilabiata insularis and Alaeuris quadrilabiata quadrilabiata from Pyxix REFERENCES arachnoides and Psammobates tentorius; Mehdiel- Adamson, M. L. 1981. Parapharyngodon osteopili n. sp. la stylosa dollfusi and Mehdiella stylosa stylosa (Pharyngodonidae: Oxyuroidea) and a revision of Para- from Testudo graeca ibera and Pyxix arachnoides; pharyngodon and Thelandros. Systematic Parasitology, Tachygonetria macrolaimus macrolaimus and 3: 105–117. Tachygoentria macrolaimus tetrapapillata from Adamson, M. L. and A. K. Nasher. 1984. Pharyngodonidae Geochelone pardalis and Gopherus polyphemus; (Oxyuroidea; Nematoda) of Agama yemenensis in Saudi Thaparia thapari australis and Thaparia thapari Arabia: hypothesis on the origin of pharyngodonids of her- thapari from Geochelone pardalisi and Testudo bivorous reptiles. Systematic Parasitology, 6: 299–318. horsfieldii. We also raised the two subspecies Tha- Akhtar, S. A. 1937. Report on some nematode parasites of paria thapari rysavyi parasite of Testudo herman- Kabul, with descriptions of new specie. Proceedings of ni from Albania and Atractis daclyluris baltazardi Indian Academic of Science, 6: 263–273. Baker, M. R. 1987. Synopsis of the Nematoda parasitic in parasite of Testudo graeca zarudnyi from Iran to the amphibians and reptiles. Memorial University of New- species level. foundland, Occasional Papers in Biology, 11: 1–325. According to our report, the highest number of tax- Baruš, V. and S. Johnson. 1973. Notes on Mehdiella micros- onomic studies was devoted to the parasites of the toma (Drasche, 1884) from Testudo hermanni Gmelin, Palaearctic region followed by the Ethiopian region. 1788. Folia Parasitologica, 20: 139–140. A lower number of taxonomic studies was devoted to Baruš, V., Kullman, E. and F. Tenora. 1972. Parasitische Ne- the Oriental region, the Neotropical region and the matoden aus Wirbeltieren Afganistans. Acta Scientarium Nearctic region (Fig. 1A). Naturalium Academiae Scientiarum Bohemosiovacae According to Inverson (1992) and Bonin et al. Brno, 12: 1–45. (1998), the highest species diversity of Testudinidae is Baer, J. G. 1936. Une nouvelle espèce d’Oxyure, Atractis morinae n. sp. Chez une Tortue africaine. Revista di Paras- observed in the Ethiopian region (24 species), followed sitologia, 2: 277–288. by the Palaearctic region (6 species) and the Oriental Baylis, H. A. 1920. On the classification of the Ascaridae. (i) region (6 species) (Fig. 1B). The last two regions show The Systematic value of certain characters of the alimen- a low species diversity of Testudinidae with 4 species tary canal. Parasitology, 12: 253–264. in Neotropical region and 4 species in Nearctic region Baylis, H. A. 1923. Report on a collection of parasitic Nema- (Fig. 1B). todes, mainly from Egypt. II. Oxyuridae. Parasitology, 15: We found no relationship between the number of 14–23. taxonomic studies and the species diversity of Baylis, H. A. 1936. The fauna of British India. Nematoda, Vol. described parasites (p = 0.58). There were also no I (Ascaroidea and Stronggyloidea). Taylor and Francis, Londes. XXXVI + 408 pp. relationship between the number of studies and the Baylis, H. A. 1939. Records of some parasitic worms from the generic diversity of parasites (p = 0.58). All of these Belgian Congo. Annals and Magazine of Natural History, suggest that there is no potential bias in the taxonom- 10, 3: 625–629. ic survey of nematode parasites of Testudinidae. Baylis, H. A. 1940. On a further Collection of Parasitic worms Species diversity and genus diversity of nematodes from the Belgian Congo. Annals and Magazine of Natural are high in Ethiopian region and Palaearctic region History, 11, 9: 401–416. NEMATODES PARASITES OF TESTUDINIDAE 239

Baylis, H. A. and R. Daubney. 1922. Report on the parasitic Dittmann, I. 1965. Untersuchungen über Magen-Darm-Nema- Nematodes in the collection of the Zoological survey of toden aus der griechischen Landschildkröte (Testudo India. Memoirs of Indian Museum, 7: 263–284. hermanni L.). Helminthologia, 6: 181–203. Bonin, F., Vevaux, B. and A. Dupré. 1998. Toutes les Tortues Drasche, R. V. 1884. Nematoden aus Testudo graeca. Ver- du Monde. Delachaux et Niestlé, 254 pp. handlungen der Kaiserlich-Königlichen Zoologische- Bouamer, S. 2003. Systematics and phylogeny of Oxyurid Botanischen Gesellschaft in Wien, 33: 325–330. (Nematode) parasite of Paleaerctic Testudinidae (Che- Dubinina, M. H. 1947. Geographitcheskoïe rasprostranenie lonea). PhD. Thesis, Universite de Perpignan. Oxyuridea tcherepach roda Testudo. Dokladi Akademii Bouamer, S. and S. Morand. 2000. Oxyuroids of Palaearctic Nauk SSSR, 58: 1247–1250. (In Russian). Testudinidae: New definition of the genus Thaparia Dubinina, M. H. 1949. [Ecological studies on the parasite fau- Ortlepp, 1933 (Nematoda: Pharyngodonidae), redescrip- na of Testudo horsfieldii Gray from Tadjikistan]. Para- tion of T. thapari thapari, and descriptions of two new zitologicheskij Sbormik Zoologicheskogo Muzeia Akade- species. Comparative Parasitololgy, 67: 169–180. mii Nauk SSSR, 11: 61–97. (In Russian). Bouamer, S. and S. Morand. 2002. Description of T. combesi Dyk, V. and S. Dykova. 1956. Hlistice nalezene v dovezenych n. sp. and redescriptions of four species of the genus zelvach reckych (Testudo graeca L.). Československá Tachygonetria Wedl, 1862 (Nematoda: Pharyngodoni- Parasitologie, 3: 43–48. dae), with a new diagnosis of the genus. Systematic Para- Fitzsimmons, W. M. 1961. Redescription of Labidurus af- sitolgy, 53: 121–139. ricana Gedoest, 1916 with new Host Records and summa- Bouamer, S. and S. Morand. 2003. Descriptions of two new ry of our present Knowledge of the genus Labidurus species of the genus Tachygonetria Weld, 1862 (Nemato- Schneider, 1866. Journal of Helminthology, 35: 91–100. da – Pharyngodonidae) and discussion of the relationships Forstner, M. J. 1960. Ein Beitrag zur Kenntnis parasitischer among the species of the genus. Parasitology Research, Nematoden aus griechischen Landschildkröten. Zeit- 99: 68–73. schrift für Parasitenkunde, 20: 1–22. Bouamer, S. and S. Morand. 2004. Description of Tachygone- Gedoelst, L. 1916. Notes sur la faune parasitaire du Congo tria africana n. sp. and Tachygonetria pretoriensis n. Belge. Revue de Zoologie Africaine, 1: 1–90. sp., and redescription of two species of the genus Tachy- Gendre, E. 1909. Notes d’Helminthologie Africaine (Deuxième gonetria Wedl, 1862 parasite Geochelone pardalis note). Actes de la Société Linéenne de Bordeaux, 63: (Bell, 1828) from South Africa. Systematic Parasitolgy, 58: 33–41. 199–208. Graber, M. 1981. Parasites internes des vertébrés domes- Bouamer, S. and S. Morand. 2005. Descriptions of two new tiques et sauvages, autres que les primates se la Ré- species of Tachygonetria Weld, 1862 (Nematoda – Pharyn- publique Populaires du Congo (d’après la collection Cas- godonidae) and redescriptions of four species parasite of sard-Chambron, 1956–1960). Rôle pathogeneprophylaxie. Palaearctic Testudinidae. Zoosystema, 27 (2): 193–209. Revue d’Elevage et de Medecine Vetirinaire des Pays Bouamer, S., Morand, S. and R. Bourgat. 2001a. Oxyuroids of Tropicaux, 34: 155–167. Palaearctic Testudinidae – New definition of the genus Hallez, P. 1886. Anatomie de l’Atractis dactylura (Duj.). Alaeuris Seurat, 1918 (Nematoda: Pharyngodonidae), Mémoire de la Societe des Sciences. Lille, série 4, 15: 120. redescription of Alaeuris numidica (Seurat, 1918). Jour- Hristovski, N. D. 1973. Nematodes of the Suborder Oxyurata nal of Parasitology, 87: 128–133. Skrjabin, 1923 which parasitize the bodies of poikilother- Bouamer, S., Morand, S. and R. Bourgat. 2001b. Redescription mic animals from the vicinity of Bilota (Macedonia – of Mehdiella microstoma and description of Mehdiella Yugoslavia). Archiv Bioloskih Nauka. Beograd, 25: 85–96. petterae sp. n., with a new definition of the genus Meh- Inglis, X. E. and C. Diaz-ungria. 1963. Sobre genero Labidu- diella Seurat, 1918 (Nematoda: Pharyngodonidae). Folia rus (Ascaridata: Kathlaniidae) con una discusion sobre el Parasitologica, 48: 132–138. desarrollo de la cabeza. Bulletin-Sociedad Venezolana de Bouamer, S., Morand, S. and M. Kara. 2003. Redescription of Ciencias Naturales, 25: 126–154. four species of Mehdiella from Testudinidae with a key to Inverson, J. B. 1992. A revised checklist with distribution the species of the genus. Parasite, 10: 333–42. maps of the turtles of the world. Second printing, 1994: Brenes, R. R. and M. Bravo-Hollis. 1960. Helmintos de la 374 pp. republica de Costa Rica. IX. Nematoda 3 algunos Nema- Jansen, J. and E. van den Broek. 1966. Parasites of zoo-ani- todes de Reptiles, con description de dos nuevos especies: mals in the Netherlands and of exotic animals II. Bijdra- Atractis caballero n. sp. y cyrtosomum longicau- gen tot de Dierkunde, Uitgegeven doo het Koninklijk Zoöl- datum n. sp. Libro Homenaje al Dr. Eduardo Caballero ogisch Genootschap Natura Artis Magistra te Amsterdam, y Caballero, 451–464. 36: 65–68. Chaprin, E. A. 1924. Nematode parasites of the Brazilian land- Johnson, S. 1973. Some oxyurid nematodes of the genera tortoise, Testudo denticulata. Proceedings of the United Mehdiella and Thaparia from the tortoise Testudo her- States National Museum, 65: art. 13, 6 p. manni. Folia Parasitologica, 20: 141–148. Chabaud, A. G. and A. J. Petter. 1960. Sur les Nématodes Khalil, M. 1932. Parasites from Liberia and French Guinea, Atractides. Libro Homenajae al Doctor Eduardo Caballero first part: Nematoda. Zeitschrift für Parasitenkunde, 4: y Caballero. Mexico. D.F. 465–470. 431–458. Creplin, F. C. H. 1846. Nachträge zu Gurlt’s Verzeichniss der Katalan-Gateeva, Sh. and G. Dimitrov. 1961. [Research on Thiere, bei welchen Entozoen gefunden worden sind. nematodes infecting tortoises in the Petric region]. Iz- Archiv für Naturgeschichte, 12: 129–160. vestiia na Tsentralnata Khelminthologichna Laboratoriia. 240 S. BOUAMER and S. MORAND

B’lgarska Akademiia na Naukite, 6: 109–118. (In Bul- Rasheed, S. 1965. Some parasitic nematodes from the Came- garian). roons (W. Africa). Journal of Helminthology, 39: 67–100. Linstow, O. 1903. Entozoa des zoologischen Museums der Ryšavý, B. 1958. [Contribution of the knowledge of nematodes Kaiserlichen Akademie der Wissenschaften zu St. Peters- of imported Testudo graeca L.] Československá Para- burg. 2. Ezhegodnik Zoologicheskago Muzeia Impera- sitologie. 5: 178–183. (In Czech). torskoi Akademii Nauk, 8: 265–294. Ryšavý, B. and S. Johnson. 1979. On some oxyurid nematode Linstow, O. 1909. Parasitische Nematoden. In: Die Süss- of the genus Tachygonetria parasitizing tortoises in wasserfauna Deutschlands. Eine Exkursionsfauna. Heft Afghanistan and Albania. Vestnik Československe Spolec- 15: 47–83. nosti Zoologicke, 43: 148–160. Markov, G. S., Ivanov, V. P., Nikulin, V. P. and V. F. Chernobai. Schad, G. A., Kuntz, R. E. and W. H. Wells. 1960. Nematode 1962. [Helminth parasites of reptiles from Volga-delta and parasites from Turkish vertebrates. Canadian Journal of Caspian steppe]. Trudy Astrakhanskogo Zapovednika, 6: Zoology, 38: 949–963. 145–172. (In Russian). Schneider, A. 1866. Monographie der Nematoden., Berlin. XIII Markov, G. S. and B. P. Kriuchkov. 1961. [Helminth fauna of + 357 pp. the turtle Cryptodira in southern Daghestan]. Uchenye Seurat, L.-G. 1917. Sur les Oxyures des Sauriens de Nord- Zapiski Stalingradskovo Gosudarstvennogo Pedagogi- Africain. Archives de Zoologie Expérimentales et Géné- cheskogo Instituta im A. S. Serafimovicha, 13: 124–133. (In rales, 45: 401–444. Russian). Seurat, L.-G. 1918. Contribution à l’étude de la faune para- Ortlepp, R. J. 1933. On some South African reptilian oxyurids. sitaire de la Tunisie. Nématodes. Archive de l’Institut Pas- Onderstepoort Journal of Veterinary Science and Animal teur de Tunisie, 10: 243–275. Industry, 1: 9–114. Sharpilo, V. P. 1962. [On the study of the Helminth fauna of Petter, A. J. 1961. Redescription et analyse critique de quel- reptiles in the Transcaucasia]. Zbirnik Prats Zoolgichno- ques espèces d’Oxyures de la tortue grecque (Testudo go Museiu. Akademiia Nauk Ukainskoi RSR, 31: 63–69. (In graeca L.). Diversité des structures céphaliques. Annales Russian). de Parasitologie Humaine et Comparée, 10, 648–671. Sharpilo, V. P. 1976. [Parasitic worms of the reptilian fauna of Petter, A. J. 1966. Équilibre des espèces dans les populations the USSR. Systematics, chorology, biology]. Naukova de Nématodes parasites du côlon des tortues terrestres. Dumka, Kiev, 287 pp. (In Russian). Mémoire du Muséum National d’Histoire Naturelle, Paris, Stossich, M. 1898. Saggio di una elmintologica di Trieste. Pro- Nouvelle série A, Zoologie, 39: 1–252. gramma della Civica Scuola Reale Superiore, Trieste. Petter, A. J. and E. R. Brygoo, 1972. Phénomène de gigantisme 162 pp. dans une population de Nématodes parasite de Tortue Travassos, L. 1925. Contributions para o conhecimento da Malgache. Annales de Parasitologie Humaine et Com- fauna helmintolojica dos batraquios do Brasil. Nema- parée, 47: 581–583. todeos intestinais. Sciencias Medicina, 3, 673–687. Petter, A. J. and J. F. Douglass. 1976. Étude des populations Stiles, C. W. and A. Hassall. 1894. A preliminary catalogue of d’Oxyures du côlon des Gopherus (Testudinidae). Bul- the parasites contained in the collections of the United letin du Muséum National d’Histoire Naturelle. série 3, n° States Bureau of the Animal Industry, United States Army 389, Zoologie. 271: 731–768. Medical Museum, Biological Department of the University Petter, A. J. and J. C. Quentin. 1976. Keys to the genera of the of Pennsylvania (Coll. Leidy) and in Coll. Stiles and Coll. Oxyuroidea. CIH Keys to the Nematode Parasites of Verte- Hassall. Veterinary Magazine, 1: 133–354. brates. No 4. R. C. Anderson, A. C. Chabaud and S. Will- Sprent, J. F. A. 1980. Ascadoid nematodes of amphibians and mott (eds.). Commonwealth Agricultural Bureaux Farn- reptiles: Angusticaecum and Krefftascaris n. g. Journal ham Royal. Slough, U.K. 11.2–11.3 p. of Helminthology, 54: 55–73. Pillers, A. W. N. 1933. Notes on parasites in 1932. Veterinary Thapar, G. S., 1925. Studies on the oxyurid parasites of rep- Record. 13: 964–966. tiles. Journal of Helminthology, 3: 83–150. Rees, F. G. 1935. Two new species of Tachygonetria from the Walton, A. C. 1942. Some oxyurids from a Galapagos tortoise. Indian tortoise, Testudo horsfieldii Gray. Proceedings of Proceeding of the Helminthological Society of Washington, the Zoological Society of London, 3: 599–603. 9: 1–17.

Received: May 06, 2005 Accepted: October 03, 2005