Junker_27-36.indd 27 Onderstepoort Journal of Veterinary Research, 73:27–36 (2006) A check-list of the pentastomid parasites of crocodilians and freshwater chelonians K. JUNKER and J. BOOMKER* Department of Veterinary Tropical Diseases, University of Pretoria Private Bag X04, Onderstepoort, 0110 South Africa ABSTRACT JUNKER, K. & BOOMKER, J. 2006. A check-list of the pentastomid parasites of crocodilians and fresh- water chelonians. Onderstepoort Journal of Veterinary Research, 73:27–36 Based on published records and own data a summary is given of the geographical distribution of the currently known species of pentastomid parasites infecting crocodiles and alligators, as well as fresh- water chelonians. A brief generic diagnosis is provided for each genus. Fourteen out of the currently 23 living crocodilian species have been recorded as being host to one or more pentastomes. Out of the 32 pentastome species six are considered species inquirendae. Pres- ently, six genera of crocodilian pentastomes, Agema, Alofia, Leiperia, Sebekia, Selfia and Subtriquetra are recognized. African crocodiles harbour eight pentastome species, six of which have been recorded from the Nile crocodile, Crocodylus niloticus. Three species belong to the genus Sebekia, Alofia being represented by two and Leiperia by only one species. Two species, Alofia parva and Agema silvae- palustris, occur in the dwarf crocodile, Osteolaemus tetraspis, and the slender-snouted crocodile, Crocodylus cataphractus, exclusively, but a single Sebekia species is shared with the Nile crocodile. The genus Agema is endemic to the African region. Infective stages of the pentastome Sub triquetra rileyi, thought to utilize Nile crocodiles as final hosts, have been recovered only from fishes. The larg- est number of pentastome species is found in the Australasian region. Of these, the Indo-Pacific croc odile, Crocodylus porosus, harbours seven, representing the genera Alofia, Sebekia, Lei peria and Selfia. Selfia is exclusive to the latter host. The genus Subtriquetra has been reported from “Indian crocodiles”, a term possibly referring to either Crocodylus palustris, Crocodylus porosus or Gavi alis gangeticus. Ten species of pentastomes parasitizing the crocodilian genera Alligator, Cai- man, Crocodylus and Melanosuchus have been recorded from the Neotropical region including the southern states of the North American continent. The two most wide-spread pentastome genera, Alofia and Sebekia, have been recorded together with representatives of the genus Sub triquetra and immature and larval forms of Leiperia. To date the two monospecific genera, Pelonia, from two terrapin species, Pelusios sinuatus and Pelo- medusa subrufa, in South Africa, and Diesingia from Hydraspis geoffroyana and Hydromedusa tectif- era in South America, are the only chelonian pentastomes recovered world-wide. A possible excep- tion is the crocodilian pentastome Sebekia mississippiensis which can reach maturity in exper imentally infected terrapins. Keywords: Agema, Alligator, Alofia, Caiman, Crocodylus, Diesingia, Gavialis, Hydraspis, Hydro- medusa, Leiperia, Melanosuchus, Pelomedusa, Pelonia, Pentastomida, Phrynops, Sebe- kia, Selfia, Subtriquetra, terrapins * Author to whom correspondence is to be directed. E-mail: jboomker@op.up.ac.za Accepted for publication 1 August 2005—Editor 28 check-list of pentastomid parasites of crocodilians and freshwater chelonians INTRODUCTION The pentastomid parasites of chelonians and croco- dilians are currently divided into the family Sebekidae and Subtriquetridae. The former comprises seven gen era, namely, Agema, Alofia, Selfia, Sebekia, Lei- peria, Diesingia and Pelonia. While the first four genera inhabit the lungs and bronchioles of the croco dilian host, Leiperia occurs in the trachea and bronchi (Riley, Spratt & Winch 1990; Riley 1994; Riley & Huch zermeyer 1996; Riley, Hill & Huch zer- meyer 1997). Diesingia and Pelonia parasitize the lungs of chelonian final hosts (Junker & Boomker 2002; Junker, Riley & Boomker 2003). A single member of Sebekia, Sebekia mississippi- ensis, might be able to reach maturity in chelonians, too, but as yet no mature specimens have been col- lected from naturally infected hosts (Dukes, Shealy & Rogers 1971). Members of the monogeneric fam- ily Sub triquetridae inhabit the nasopharynx of their crocodilian final hosts, but Subtriquetra rileyi, of which currently only infective larvae have been re- covered, needs verification (Winch & Riley 1986a; Junker, Boomker & Booyse 1998). During the past 10 years renewed progress has been made as regards the taxonomy of crocodilian pentastomes. The older genera, Alofia, Sebekia and Leiperia, have been revised, and examination of new material has led to the description of several new genera and species (Riley 1994; Riley et al. 1990, 1997; Riley & Huchzermeyer 1996; Junker et al. 1998). However, there is a dearth of data concern- ing the chelonian pentastomids. In order to provide a quick reference tool, this check- list consists of two parts, following the example of Sambon (1922). The first part lists the parasites un- der their scientific names, their synonyms and their authorities. A short generic diagnosis precedes each genus and the parasites are grouped according to the geographic distribution of their respective hosts. The list starts with Africa, followed by Australasia. South and North America are listed last. The second part of the check-list alphabetically lists the hosts and their synonyms, and, also in alpha- betical order, their respective parasites. The no- menclature and synonyms of the crocodilian and chelonian hosts are according to Getz (2002). Only references dealing with mature pentastomes are included in the check-list, but for completeness’ sake the intermediate hosts of pentastome species of which only larval forms are known are listed. PARASITE/HOST CHECK-LIST OF THE PENTASTOMIDA FAMILY SEBEKIDAE SAMBON, 1922 Genus Agema Riley, Hill & Huchzermeyer, 1997 GENERIC DIAGNOSIS: Overall shape typical of smaller members of the Sebekidae; males claviform, fe- males with more uniform diameter and conical pos- terior terminating in a small blunt point; adult hooks smooth; blades smoothly curved without abrupt right-angled bend near to base; blade on anterior hook pair larger than that of posterior pair; fulcrum without cowl; mouth ovoid and sides of cadre united anteriorly and posteriorly by segments of chitin which appear as two crescents; copulatory spicules deli- cate and elongate; basal section without a hooked collar (Riley et al. 1997). AFRICA 1. Agema silvaepalustris Riley, Hill & Huchzermeyer, 1997 Crocodylus cataphractus Riley, Hill & Huchzermeyer (1997), Repub- lic of the Congo Osteolaemus tetraspis Riley, Hill & Huchzermeyer (1997), Repub- lic of the Congo Genus Alofia Giglioli, 1922 GENERIC DIAGNOSIS: Size small; body banana-shaped; hooks large with long, narrow, slightly curved blade and a slender base; absence of chitinous formation at the base of the anterior hook; chitinous buccal cadre large and U-shaped; intestine as in Sebekia; found in crocodiles (Fain 1961 in Riley 1994); caudal extremity of female bluntly rounded, often swollen into a bulb (Riley 1994); hooks usually smooth (rare- ly with patches of minute spines [see Alofia nilotici and Alofia parva (Riley & Huchzermeyer 1995a, b)], blades finely canaliculated, bent through almost a right angle at the base; peg-like extension of oral cadre projects into oesophagus; copulatory spicules with double-hooked collar on the shorter of the two anterior extensions (Riley & Huchzermeyer 1995a, b; Junker, Boomker & Bolton 1999). AFRICA 1. Alofia nilotici Riley & Huchzermeyer, 1995 Crocodylus niloticus 29 K. JUNKER & J.BOOMKER Riley & Huchzermeyer (1995a), Bo t swana Junker, Boomker & Bolton (1999), South Africa 2. Alofia parva Riley & Huchzermeyer, 1995 Crocodylus cataphractus Riley & Huchzermeyer (1995b, 2000), Re- pub lic of the Congo Osteolaemus tetraspis osborni Riley & Huchzermeyer (1995b, 2000), Re- public of the Congo 3. Alofia simpsoni Riley, 1994 Unknown crocodilian Riley (1994), Ghana Crocodylus niloticus Riley & Huchzermeyer (1995a), Botswana Junker, Boomker & Bolton (1999), South Africa AUSTRALASIAN REGION 4. Alofia ginae Giglioli, 1922 Unknown crocodilian Sambon (1922), Samoa Most probably Crocodylus porosus, as it is the only crocodilian whose range ex- tends as far as Fiji in the Pacific Ocean and it is known to have colonized many small islands as far as nearly 1 000 km from land (Ross 1989). Crocodylus porosus (?) Riley (1994), Philippines The collector did not specify the host, but Riley (1994) concludes it to be C. porosus, as the only other Philippinian crocodile, Crocodylus mindorensis, does not occur in the region from which the parasites were recovered. 5. Alofia merki Giglioli, 1922 Sebekia merki Heymons, 1941 Unknown crocodilian Sambon (1922), Samoa Most probably Crocodylus porosus, as it is the only crocodilian whose range ex- tends as far as Fiji in the Pacific Ocean and it is known to have colonized many small islands as far as nearly 1 000 km from land (Ross 1989). Crocodylus porosus Riley (1994), Northern Territory, Australia Riley (1994), Philippines 6. Alofia indica (Von Linstow, 1906) Hett, 1924, spe- cies inquirenda Gavialis gangeticus Hett (1924), India SOUTH & NORTH AMERICAN REGION 7. Alofia platycephala (Lohrmann, 1889) Giglioli, 1922 Pentastomum platycephalum Lohrmann, 1889; Porocephalus platycephalus Shipley, 1898; Rei- ghardia platycephala Sambon, 1910 Unknown crocodilian Lohrmann (1889), South America Caiman crocodilus Self & Rego (1985), Brazil Caiman latirostris Heymons (1941), Paraguay Comment: Hirst (1922) described Alofia adriatica (Hirst, 1922) Giglioli, 1922 from an unknown host from the Adriatic. As crocodilians do not occur in the Adriatic this species will have to remain species in- quirenda until further material becomes available. Genus Leiperia Sambon, 1922 GENERIC DIAGNOSIS: Female with spirally coiled ab- domen; broad bands of chloride cells; hooks smooth, flat-topped with sharply curved blade; oral cadre V- shaped with large anterior flanges, numerous pores around the pharynx; copulatory spicules heavily chit- inized with complex internal supporting structures, shorter of the two anterior extensions forms a smooth collar (double in L. australiensis) around the longer spatulate extension; cirrus tip modified into a flat- tened trumpet of longitudinally-striated chitin (Riley & Huchzermeyer 1996; Junker, Boomker, Swa ne- poel & Taraschewski 2000). AFRICA 1. Leiperia cincinnalis (Vaney & Sambon, 1910) Sambon, 1922 Reighardia cincinnalis Vaney & Sambon, 1910; 30 check-list of pentastomid parasites of crocodilians and freshwater chelonians Porocephalus nematoides De Beauchamp, 1918 Crocodylus cataphractus Fain (1961), Central Africa: infective larva Crocodylus niloticus Vaney & Sambon (1910), Uganda Sambon (1922), Zimbabwe Rodhain & Vuylsteke (1932), Democratic Republic of the Congo Junker, Boomker, Swanepoel & Tara schew- ski (2000), South Africa AUSTRALASIAN REGION 2. Leiperia australiensis Riley & Huchzermeyer, 1996 Crocodylus johnsoni Riley & Huchzermeyer (1996), Northern Ter ritory, Australia Crocodylus porosus Riley & Huchzermeyer (1996), Northern Territory, Australia SOUTH & NORTH AMERICAN REGION 3. Leiperia gracilis Diesing, 1936, species inquiren- da Pentastoma gracile Diesing, 1836 (partim); Pen- ta stomum gracile Leidy, 1856; Pentastoma gra- cilis Parona, 1891; Porocephalus gracilis Shipley, 1898; Porocephalus crocodili Wheeler, 1915 (par- tim); Leiperia neotropica Heymons & Vitz thum, 1935 Immature and larval forms were recovered from the following hosts, but adults have not been collected. Alligator mississippiensis Leidy (1856, in Sambon 1922), locality un- known, North America Crocodylus acutus Heymons (1935), South America Caiman crocodilus Heymons (1935), Brazil Genus Sebekia Sambon, 1922 GENERIC DIAGNOSIS: Body short and squat with 58– 94 compressed annuli; lateral lines conspicuous; cephalothorax small, wedge-shaped and projecting nipple-like from the abdomen, ventral surface con- tinuous with that of the abdomen; mouth subtermi- nal and shaped like an inverted ‘U’; oral cadre oval to elongate, highly variable in shape, without long, parallel sides and generally united anteriorly; hooks small, equal or subequal, claw-shaped, with convex or flat dorsal surface; all hooks spiny (rarely only the anterior pair); all fulcra often with spinous anterior extension (rarely only the anterior pair); hook barb curved, strongly united and continuous with shank; spicules generally obpyriform, with one or two fine sclerotized rods supporting membranous region distally. Parasites of the lungs of crocodilians, rarely of chelonians (Riley et al. 1990). AFRICA 1. Sebekia cesarisi Giglioli, 1922 Crocodylus sp. Sambon (1922), Africa Crocodylus niloticus Riley & Huchzermeyer (1995a), Botswana Junker, Boomker & Bolton (1999), South Africa 2. Sebekia okavangoensis Riley & Huchzermeyer, 1995 Sebekia cesarisi Riley, Spratt & Winch, 1990 Crocodylus cataphractus Riley & Huchzermeyer (2000), Republic of the Congo Crocodylus niloticus Riley & Huchzermeyer (1995a), Botswana Junker, Boomker & Bolton (1999), South Africa Osteolaemus tetraspis Riley & Huchzermeyer (2000), Republic of the Congo 3. Sebekia wedli Giglioli, 1922 Pentastoma oxycephalum var. minor Wedli, 1861; Sebekia oxycephala Self & Rego, 1985 Crocodylus niloticus Devos (1939), Democratic Republic of the Congo Riley, Spratt & Winch (1990), Uganda Riley & Huchzermeyer (1995a), Botswana Junker, Boomker & Bolton (1999), South Africa 31 K. JUNKER & J.BOOMKER AUSTRALASIAN REGION 4. Sebekia johnstoni Riley, Spratt & Winch, 1990 Crocodylus johnsoni Riley, Spratt & Winch (1990), Northern Territory, Australia Crocodylus porosus Riley, Spratt & Winch (1990), Northern Territory, Australia 5. Sebekia multiannulata Riley, Spratt & Winch, 1990 Crocodylus johnsoni Riley, Spratt & Winch (1990), Northern Territory, Australia Crocodylus porosus Riley, Spratt & Winch (1990), Northern Territory, Australia 6. Sebekia purdieae Riley, Spratt & Winch, 1990 Crocodylus porosus Riley, Spratt & Winch (1990), Northern Ter- ritory, Australia 7. Sebekia jubini (Vaney & Sambon, 1910) Sambon, 1922, species inquirenda Porocephalus jubini Vaney & Sambon, 1910 Crocodylus siamensis Sambon (1922), locality unknown, south- east Asia 8. Sebekia novaeguineae Riley, Spratt & Winch, 1990 Crocodylus novaeguineae Riley, Spratt & Winch (1990), Papua New Guinea SOUTH & NORTH AMERICAN REGION 9. Sebekia acuminata Travassos, 1924, species in- quirenda Unknown crocodilian Travassos (1924), Brazil 10. Sebekia divestei Giglioli, 1922 Crocodylus acutus Sambon (1922), locality unknown, Neo- tropical region 11. Sebekia microhamus Self & Rego, 1985 Caiman crocodilus Self & Rego (1985), Brazil 12. Sebekia mississippiensis Overstreet, Self & Vliet, 1985 Pentastoma oxycephalum Diesing, 1836 (par- tim); Pentastomum gracile (syn. Leiperia graci- lis) Leidy, 1856 Alligator mississippiensis Deakins (1971), USA Hazen, Aho, Murphy, Esch & Schmidt (1978), USA Overstreet, Self & Vliet (1985), USA 13. Sebekia oxycephala (Diesing, 1836) Sambon, 1922 Pentastoma proboscideum Rudolphi, 1819 (par- tim); Pentastoma oxycephalum Diesing, 1836 (partim); Pentastoma gracile Diesing, 1836 (par- tim); Pentastomum oxycephalum Diesing, 1850 (partim); Pentastomum gracile Diesing, 1850 (par tim); Pentastomum heterodontis Leuckart, 1860; Pentastomum oxycephalum Chatin, 1882; Porocephalus oxycephalus Stiles, 1893; Penta- stoma proboscideum crocodili scleropis Rudolphi (Shipley in Sambon 1922); Reighardia oxyceph- ala Vaney & Sambon, 1910; Porocephalus croco- dili Wheeler, 1913 (partim); Sebekia oxycephala Sambon, 1922 (partim); Bdukus ichthyius Holl, 1929; Leiperia heterodontis Heymons & Vitzthum, 1935; Sebekia crocodili Heymons & Vitzthum, 1935 Alligator mississippiensis Sambon (1922), locality unknown Caiman crocodilus Sambon (1922), locality unknown Winch & Riley (1986b), Trinidad, South America Caiman latirostris Heymons (1941), locality unknown Crocodylus acutus Sambon (1922), locality unknown 14. Sebekia samboni Travassos, 1924, species in- quirenda Unknown crocodilian Travassos (1924), Brazil 15. Sebekia trinitatis Riley, Spratt & Winch, 1990 32 check-list of pentastomid parasites of crocodilians and freshwater chelonians Caiman crocodilus Riley, Spratt & Winch (1990), Trinidad, South America Genus Selfia Riley, 1994 GENERIC DIAGNOSIS: Size small, cephalothorax mi- nute in comparison with diameter of the abdomen; 78–82 well defined annuli; abdomen strongly curled ventrally; caudal extremity of female abruptly tapered to blunt point; hooks very small, with tiny blade only slightly offset from transversely creased and folded shank; rear of anterior hooks enveloped by soft, spinous cowl which forms an extension of the ful- crum; buccal cadre somewhat variable in shape, being oval to more U-shaped, but lacking parallel sides; copulatory spicule of male like that of Alofia (Riley 1994). AUSTRALASIAN REGION 1. Selfia porosus Riley, 1994 Crocodylus porosus Riley (1994), Northern Territory, Australia Genus Diesingia Sambon, 1922 GENERIC DIAGNOSIS: Hooks smooth, flat-topped, with sharply curved blades; fulcra with anterior cowl-like extension, extension smooth in posterior and spiny in anterior fulcra; oral cadre open anteriorly with an oesphageal peg similar to that in Alofia; copulatory spicule with cowry shell-shaped base, the short, ven- tral extension is transformed into a structure resem- bling the collembolan fulcrum, and is connected to the base by a joint (Junker, Riley & Boomker 2003). SOUTH & NORTH AMERICAN REGION 1. Diesingia megastoma (Diesing, 1836) Sambon, 1922 Pentastoma megastomum Diesing, 1836; Pen ta- stomum megastomum Leuckart, 1860; Poro - cephalus megastomus Shipley, 1898; Sebekia mega stoma Travassos, 1923; Sebekia crocodili Hey mons & Vitzthum, 1935; Diesingia megasto- ma Heymons; 1941; Butantanella megastoma Da Fonseca & Ruiz, 1956; Sebekia megastoma Self & Rego, 1985 Phrynops geoffroanus Diesing (1836), Brazil Hydromedusa tectifera Da Fonseca & Ruiz (1956), Brazil Genus Pelonia Junker & Boomker, 2002 GENERIC DIAGNOSIS: Hooks smooth, dorsally convex, with sharply bent blades, fulcra without extensions; oral cadre more or less U-shaped, closed anteriorly by delicate chitinous fibres; copulatory spicules al- most identical to those of Sebekia wedli, with cowry shell-shaped base and the short anterior extension ending in a smooth collar, the long spatulate exten- sion carries small chitinous teeth (Junker & Boomker 2002). AFRICA 1. Pelonia africana Junker & Boomker, 2002 Pelomedusa subrufa Junker & Boomker (2002), South Africa Pelusios sinuatus Junker & Boomker (2002), South Africa FAMILY SUBTRIQUETRIDAE FAIN, 1961 Genus Subtriquetra Sambon, 1922 GENERIC DIAGNOSIS: Body elliptical, ventrally flat- tened and dorsally dome shaped with flattened mar- gins; hooks simple, slender and sharply pointed, dis- posed in a curved line; oral cadre rounded (Fain 1961; Winch & Riley 1986a; Junker et al. 1998). AFRICA 1. Subtriquetra rileyi Junker, Boomker & Booyse, 1998 Infective larvae: Tilapia rendalli swierstrae Junker, Boomker & Booyse (1998), South Africa Oreochromis mossambicus Junker, Boomker & Booyse (1998), South Africa AUSTRALASIAN REGION 2. Subtriquetra megacephala (Baird, 1853) Sam- bon, 1922 Pentastoma megacephalum Baird, 1853; Poro- cephalus megacephalus Shipley, 1898 Crocodylus palustris Sambon (1922), Sunderbunds, India Crocodylus palustris, Crocodylus porosus or 33 K. JUNKER & J.BOOMKER Gavialis gangeticus? (“Sangor crocodile”) Sambon (1922), Bengal, India 3. Subtriquetra shipleyi Hett, 1924 Crocodylus palustris Crocodylus porosus or Gavialis gangeticus ? (“Indian crocodile”) Hett (1924), India SOUTH & NORTH AMERICAN REGION 4. Subtriquetra subtriquetra (Diesing, 1836) Pentastoma proboscideum Bresmer, 1824 (par- tim); Pentastoma subtriquetrum Diesing, 1836; Pen tastomum subtriquetrum Diesing, 1850; Pen- ta stomum pusillum Diesing, 1856; Linguatula sub- triquetra Raillet, 1883; Linguatula pusilla Ship ley, 1898 Caiman crocodilus Sambon (1922), South America Winch & Riley (1986a), Trinidad, South America Melanosuchus niger Sambon (1922), South America HOST/PARASITE CHECK-LIST OF THE PENTASTOMIDA Crocodylia FAMILY ALLIGATORIDAE (CUVIER, 1807) (Alligators and caimans) Genus Alligator Cuvier, 1807 1. Alligator mississippiensis (Daudin, 1801) Daudin, 1802 (American alligator) Crocodilus mississipiensis Daudin, 1801 Leiperia gracilis, species inquirenda, larval forms only Sebekia mississippiensis Sebekia oxycephala Genus Caiman Spix, 1825 1. Caiman crocodilus (Linnaeus, 1758) (Common or Spectacled caiman) Lacerta crocodilus Linnaeus, 1758; Caiman scler- ops Schneider, 1801 (fide Medem 1981); Pero- suchus fuscus Cope, 1868; Alligator (Jacare) chi apasius Bocourt, 1876 Alofia platycephala Leiperia gracilis, species inquirenda, larval forms Sebekia microhamus Sebekia oxycephala Sebekia trinitatis Subtriquetra subtriquetra 2. Caiman latirostris (Daudin, 1801) (Broad-snout- ed caiman) Crocodilus latirostris Daudin, 1801; Caiman fis- sipes Spix, 1825; Champsa fissipes Wagler, 1828 (fide Hoogmoed & Gruber, 1983); Alligator cynocephalus Duméril & Bibron, 1836; Jacare la- tirostris Gray, 1862; Alligator latirostris Boulanger, 1886; Jacaretinga latirostris Vaillant, 1898 Alofia platycephala Sebekia oxycephala Genus Melanosuchus Gray, 1862 1. Melanosuchus niger (Spix, 1825) (Black caiman) Caiman niger Spix, 1825 Subtriquetra subtriquetra FAMILY CROCODYLIDAE (CUVIER, 1807) (Crocodiles) SUBFAMILY CROCODYLINAE (CUVIER, 1807) Genus Crocodylus Laurenti, 1768 1. Crocodylus acutus (Cuvier, 1807) (American crocodile) Crocodilus acutus Cuvier, 1807 Leiperia gracilis, species inquirenda, larval forms only Sebekia divestei Sebekia oxycephala 2. Crocodylus cataphractus Cuvier, 1825 (Slender- snouted crocodile) Crocodilus cataphractus Falconer, 1846 Agema silvaepalustris Alofia parva Leiperia cincinnalis, infective larva Sebekia okavangoensis 34 check-list of pentastomid parasites of crocodilians and freshwater chelonians 3. Crocodylus johnsoni Krefft, 1873 (Australian freshwater crocodile) Tomistoma kreffti Gray in Krefft, 1873 (nomen nudum); Crocodilus (Philas) johnstoni Gray, 1874; Crocodylus johnstoni Cogger, 2000 Leiperia australiensis Sebekia johnstoni Sebekia multiannulata 4. Crocodylus niloticus Laurenti, 1768 (Nile croco- dile) Crocodilus vulgaris Cuvier, 1807; Crocodilus mul- ti scutatus Rüppell in Cretzschmar, 1826; Croco - di lus marginatus Geoffroy, 1827; Crocodilus mad- a gascariensis Grandidier, 1872; Crocodilus vul- garis var. madagascariensis Boettger, 1877 Alofia nilotici Alofia simpsoni Leiperia cincinnalis Sebekia cesarisi Sebekia okavangoensis Sebekia wedli 5. Crocodylus novaeguineae Schmidt, 1928 (New Guinea crocodile) Sebekia novaeguineae 6. Crocodylus palustris Lesson, 1831 (Mugger, Marsh crocodile) Subtriquetra megacephala Subtriquetra shipleyi (“Indian crocodile”) 7. Crocodylus porosus Schneider, 1801 (Indo- Pacific or Saltwater crocodile) Crocodylus natans Meyer, 1795; Crocodilus po- rosus Schneider, 1801; Crocodilus oopholis Schneider, 1801; Crocodilus biporcatus Cuvier, 1807; Crocodilus biporcatus raninus Müller & Schlegel, 1844; Oopholis pondicherianus Gray, 1862; Crocodylus porosus australis Deraniyaga- la, 1953; Crocodylus porosus minikanna Deran i- ya gala, 1953 Alofia ginae (possibly Crocodylus mindoren- sis, but distributionally unlikely) Alofia merki Leiperia australiensis Sebekia johnstoni Sebekia multiannulata Sebekia purdieae Selfia porosus Subtriquetra shipleyi (“Indian crocodile”) 8. Crocodylus siamensis Schneider, 1801 (Siamese crocodile) Crocodilus galeatus Cuvier, 1807 Sebekia jubini, species inquirenda Genus Osteolaemus Cope, 1861 1. Osteolaemus tetraspis Cope, 1861 (Dwarf croc- odile) Crocodilus frontatus Murray, 1862; Halcrosia af- zelii Lilljeborg, 1867; Halcrosia nigra Gray, 1867; Halcrosia nigra Gray, 1870; Osteoblepharon os- borni Schmidt, 1919; Osteolaemus tetraspis tet- raspis Wermuth & Mertens, 1961 Agema silvaepalustris Alofia parva Sebekia okavangoensis FAMILY GAVIALIDAE ADAMS, 1854 (Gharials) Genus Gavialis Oppel, 1811 1. Gavialis gangeticus (Gmelin, 1789) (Gharial) Lacerta gangetica Gmelin, 1789 Subtriquetra megacephala (“Indian croco- dile”) Subtriquetra shipleyi (“Indian crocodile”) CHELONIA Suborder Pleurodira (Side-necked turtles) FAMILY PELOMEDUSIDAE COPE, 1868 Genus Pelomedusa Wagler, 1830 1. Pelomedusa subrufa (Lacépède, 1788) (Cape ter ra pin) Testudo subrufa Lacépède, 1788; Testudo gale- ata Schoepff, 1792; Testudo badia Donndorf, 1798; Testudo rubicunda Suckow, 1798; Emys olivacea Schweigger, 1812 (non Emys olivacea Gray, 1855); Pentonyx capensis Duméril & Bib- ron, 1835; Pentonyx gehafie Rüppell, 1835; Pen- tonix americana Cornalia, 1849; Pelomedusa mozambica Peters (nomen nudum) in Gray 1855 (?); Pelomedusa mossambicensis Peters (no- men nudum) in Lichtenstein 1856; Pelomedusa 35 K. JUNKER & J.BOOMKER nigra Gray, 1863; Pelomedusa gasconi Roche- brune, 1884; Pelomedusa galeata Boulanger, 1889; Pelomedusa galeata var. disjuncta Vaillant & Grandidier, 1910; Pelomedusa galeata orangen- sis Hewitt, 1935; Pelomedusa galeata devilliersi Hewitt, 1935; Pelomedusa galeata dama ren sis Hewitt, 1935; Pelomedusa subrufa wett steini Mertens, 1937; Testudo emys arabica N.-Ehren- berg in Stresemann 1954 Pelonia africana Genus Pelusios Wagler, 1830 1. Pelusios sinuatus (Smith, 1838) (Serrated hinged terrapin, African serrated mud turtle) Sternotherus sinuatus Smith, 1838; Sternotherus dentatus Peters, 1848 (nomen nudum); Sterno- thaerus sinuatus Boulanger, 1889; Sternothaerus bottegi Boulanger, 1895; Pelusios sinuatus zulu- ensis Hewitt, 1927; Pelusios sinuatus leptus Hewitt, 1927 Pelonia africana FAMILY CHELIDAE GRAY, 1825 (Snake-necked turtles) Genus Hydromedusa Wagler, 1830 1. Hydromedusa tectifera Cope, 1869 [1870] (South American snake-necked turtle, Uruguay snake- necked turtle) Hydromedusa platanensis Gray, 1873; Hydro - med usa wagleri Günther, 1884 Diesingia megastoma Genus Phrynops Wagler, 1830 1. Phrynops geoffroanus (Schweigger, 1812) Gor- zula & Señaris, 1999 (Geoffroy’s side-necked turtle) Emys geoffroana Schweigger, 1812; Emys geof- freana Schweigger, 1812 (fide Boulanger, 1886); Emys depressa Merrem, 1820 (non Emys de- pressa Spix, 1824); Emys viridis Spix, 1824 (?); Emys geoffroyana Gray, 1831; Platemys geoffre- ana Duméril & Bibron, 1835; Platemys neu wiedii Duméril & Bibron, 1835; Platemys waglerii Du- méril & Bibron, 1835; Platemys tuberosa Peters, 1870; Platemys geoffroyana Boulanger, 1886; Hydraspis geoffroyana Boulanger, 1889; Hydras- pis wagleri Boulanger, 1889; Hydraspis tuberosa Boulanger, 1889; Hydraspis boulangeri Bohls, 1895; Phrynops geoffroana Mertens et al., 1934; Phrynops geoffroana geoffroana Müller, 1939; Phrynops tuberosa Mertens et al., 1934; Phrynops geoffroana tuberosa Müller, 1939 Diesingia megastoma REFERENCES DA FONSECA, F. & RUIZ, J.M. 1956. Was ist eigentlich Penta- stoma megastomum Diesing, 1836? (Porocephalida, Poro- cephalidae). Senckenbergiana Biologica, 37:469–485. DEAKINS, D.E. 1971. Pentastomes from Blackbeard Island, Georgia, with notes on North American pentastomes. Journal of Parasitology, 57:1197. DEVOS, R. 1939. Deux porocephales des crocodiles de Luapula. Annales de la Société Belge de Médecine Tropicale, 19:193– 195. DIESING, K.M. 1836. Versuch der Monographie der Gattung Pentastoma. Annalen des Wiener Museums der Natur ge- schich te, 1:1–32. DUKES, G.H., SHEALY, R.M. & ROGERS, W.A. 1971. Sebekia oxycephala (Pentastomida) in largemouth bass from Lake St. John, Concordia Parish, Louisiana. Journal of Para sitol- ogy, 57:1028. FAIN, A. 1961. Pentastomides de l’ Afrique Centrale. Annales du Musée Royale de l’ Afrique Centrale, Série 8, 92:1–115. GETZ, W. 2002. Reptilia. http://srs.embl-heidelberg.de:8000/ HAZEN, T.C., AHO, J.M., MURPHY, T.M., ESCH, G.W. & SCHMIDT, G.D. 1978. The parasite fauna of the American alligator (Alligator mississippiensis) in South Carolina. Jour- nal of Wildlife Diseases, 14:435–439. HETT, M.L. 1924. On the family Linguatulidae. Proceedings of the Zoological Society of London, 1924: 107–159. HEYMONS, R. 1935. Pentastomida, in Bronns Klassen und Ord- nungen des Tierreichs, Vol. 5. Leipzig: Akademische Verlags- gesellschaft. HEYMONS, R. 1941. Beiträge zur Systematik der Pentastomiden VI—Die Arten der Gattung Alofia im Vergleich mit Sebekia. Zeitschrift für Parasitenkunde, 12:419–432. HIRST, S. 1922. On a new linguatulid from the Adriatic. Annals and Magazine of Natural History, 10:289. JUNKER, K. & BOOMKER, J. 2002. Description of Pelonia afri- cana n. g., n. sp. (Pentastomida: Sebekidae) from the lungs of Pelomedusa subrufa and Pelusios sinuatus (Chelonia) in South Africa. Onderstepoort Journal of Veterinary Research, 69:53–59. JUNKER, K., BOOMKER, J. & BOLTON, L. 1999. Pentastomid infections in the Nile crocodile (Crocodylus niloticus) in the Kruger National Park, South Africa, with a description of the males of Alofia simpsoni. Onderstepoort Journal of Veterinary Research, 66:65–71. JUNKER, K., BOOMKER, J. & BOOYSE, D.G. 1998. Pentastomid infections in cichlid fishes in the Kruger National Park and the description of the infective larva of Subtriquetra rileyi n. sp.. Onderstepoort Journal of Veterinary Research, 65:159–167. JUNKER, K., BOOMKER, J., SWANEPOEL, D. & TARA SCHEW- SKI, H. 2000. Leiperia cincinnalis Sambon, 1922 from Nile crocodiles Crocodylus niloticus in the Kruger National Park, South Africa, with a description of the male. Systematic Para- sitology, 47:29–41. 36 check-list of pentastomid parasites of crocodilians and freshwater chelonians JUNKER, K., RILEY, J. & BOOMKER, J. 2003. Redescription of Diesingia megastoma (Diesing, 1836) Sambon, 1922, a pen- ta stomid parasite from the South American terrapin Hydro- medusa tectifera. Systematic Parasitology, 56:211–218. LOHRMANN, E. 1889. Untersuchungen über den anatomischen Bau der Pentastomen. Archiv für Naturgeschichte, 1:303– 337. OVERSTREET, R.M., SELF, J.T. & VLIET, K.A. 1985. The pen- tastomid Sebekia mississippiensis sp. n. in the American al- ligator and other hosts. Proceedings of the Helminthological Society of Washington, 52:266–277. RILEY, J. 1994. A revision of the genus Alofia Giglioli, 1922 and a description of a new monotypic genus, Selfia: two genera of pentastomid parasites (Porocephalida: Sebekidae) inhab- iting the bronchioles of the marine crocodile Crocodylus po- rosus and other crocodilians. Systematic Parasitology, 29: 23–41. RILEY, J., HILL, G.F. & HUCHZERMEYER, F.W. 1997. A de- scription of Agema, a new monotypic pentastomid genus from the lungs of the African dwarf and slender-snouted croco diles. Systematic Parasitology, 37:207–217. RILEY, J. & HUCHZERMEYER, F.W. 1995a. Description of four species of pentastomid parasites belonging to the genera Alofia Giglioli, 1922 and Sebekia Sambon, 1922, from a sin- gle Nile crocodile Cocodylus niloticus from Botswana. Sys- tematic Parasitology, 31:221–238. RILEY, J. & HUCHZERMEYER, F.W. 1995b. Pentastomid para- sites of the family Sebekidae Fain, 1961 in West African dwarf crocodiles Osteolaemus tetraspis Cope, 1851 from the Congo, with a description of Alofia parva n. sp. Onderstepoort Journal of Veterinary Research, 62:151–162. RILEY, J. & HUCHZERMEYER, F.W. 1996. A reassessment of the pentastomid genus Leiperia Sambon, 1922 with a de- scription of a new species from both the Indopacific crocodile Crocodylus porosus and Johnston’s crocodile C. johnstoni in Australia. Systematic Parasitology, 34:53–66. RILEY, J. & HUCHZERMEYER, F.W. 2000. Diet and lung para- sites of Swamp forest dwarf crocodiles (Osteolaemus tet- raspis osborni) in the Northern Congo Republic. Copeia, 2: 582–586. RILEY, J., SPRATT, D.M. & WINCH, J.M. 1990. A revision of the genus Sebekia Sambon, 1922 (Pentastomida) from crocodil- ians with descriptions of five new species. Systematic Para- sitology, 16:1–25. RODHAIN, J. & VUYLSTEKE, C. 1932. Contribution à l’ étude des porocéphales des crocodiles africains. Révue de Zoolo- gie et de Botanique Africaines, 23:1–11. ROSS, C.A (Ed.). 1989. Crocodiles and alligators, London: Meere hurst Press. SAMBON, L.W. 1922. A synopsis of the family Linguatulidae. Journal of Tropical Medicine and Hygiene, 25:188–206; 391–428. SELF, J.T. & REGO, A.A. 1985. Reassessments and revisions of certain genera and species of the family Sebekidae (Pen ta- stomida) including a description of Sebekia microhamus n. sp.. Systematic Parasitology, 7:33–41. TRAVASSOS, L. 1924. Sebekia du poumon des crocodiles d’Amerique. Comptes rendus hebdomaires des séances et mémoires de la Société de Biologie, 90:239–240. VANEY, C. & SAMBON, L.W. 1910. Preliminary notes on three new species of tongue worms (Linguatulidae) in the collec- tion of the “Museum d’Histoire Naturelle”, Paris. Transactions of the Royal Society of Tropical Medicine and Hygiene, 3: 129–134. WINCH, J.M. & RILEY, J. 1986a. Studies on the behaviour, and development in fish, of Subtriquetra subtriquetra: a uniquely free-living pentastomid larva from a crocodilian. Parasitology, 93:81–98. WINCH, J.M. & RILEY, J. 1986b. Morphogenesis of larval Se- bekia oxycephala (Pentastomida) from a South American crocodilian (Caiman sclerops) in experimentally infected fish. 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