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Home , Alopoglossus angulatus, Alopoglossus atriventris, Amphisbaena (lizard), Amphisbaena fuliginosa, Anisolepis, Anisolepis undulatus

The Journal of Venomous and Toxins including Tropical Diseases ISSN 1678-9199 | 2010 | volume 16 | issue 4 | pages 543-572

Checklist of helminths from and amphisbaenians (Reptilia, ) of ticle r A

Ávila RW (1), Silva RJ (1) eview R

(1) Department of Parasitology, Botucatu Biosciences Institute, São Paulo State University (UNESP – Univ Estadual Paulista), Botucatu, São Paulo State, .

Abstract: A comprehensive and up to date summary of the literature on the helminth parasites of lizards and amphisbaenians from South America is herein presented. One-hundred eighteen from twelve countries were reported in the literature harboring a total of 155 helminth species, being none acanthocephalans, 15 cestodes, 20 trematodes and 111 . Of these, one record was from Chile and , three from , three from Uruguay, eight from , nine from Surinam, 13 from Paraguay, 12 from , 27 from , 17 from Argentina, 39 from and 103 from Brazil. The present list provides host, geographical distribution (with the respective biome, when possible), site of infection and references from the parasites. A systematic parasite-host list is also provided.

Key words: , Nematoda, Trematoda, Squamata, neotropical.

INTRODUCTION The present checklist summarizes the diversity of helminths from lizards and amphisbaenians Parasitological studies on helminths that of South America, providing a host-parasite list infect squamates (particularly lizards) in South with localities and biomes. America had recent increased in the past few years, with many new records of hosts and/or STUDIED REGIONS localities and description of several new species (1-3). Such studies, however, date back to 1920, This checklist was prepared based on records when Dr. Lauro Travassos and his colleagues of nematodes, acanthocephalans, cestodes, presented important contributions to the field and trematodes published from 1914 to 2010, of helminth systematics (4-6). excluding material from the so called “grey The first comprehensive efforts to literature”, that is, records of unpublished theses summarize the knowledge about helminths of or scientific meetings. The of helminths squamates are those from Baker (7), who listed follows: nematodes, cestodes, trematodes and all available published data on the occurrence of acanthocephalans (6, 8, 11-15). Whenever nematodes infecting and amphibians. possible, taxonomy of hosts is reviewed with Other studies have a regional scope and treat recent publications. each helminth group separately, for example, Each record is constituted by the class, order, nematodes collected from reptiles of Brazil, superfamily, family and species of the helminth; as digenetic trematodes of amphibians and well as the name of the authority who described it reptiles from Argentina and acanthocephalans and year; besides its host(s), geographical record in amphibians and reptiles from Brazil and with country and state/province (when available) Paraguay (8-10). and information on biome of host. Comments Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America include taxonomy, synonyms and also changes in Zamora-Chinchipe (ZC), Sucumbíos (SU), the host taxonomy. and Orellana (OR). Abbreviations for all political units cited in • Peru: Amazonas (AM), Ancash (AN), this paper are: Apurímac (AP), Arequipa (AR), Ayacucho • Brazil: Acre (AC), Alagoas (AL), Amapá (AP), (AY), Cajamarca (CA), Callao (CL), Cusco Amazonas (AM), Bahia (BA), Ceará (CE), (CU), Huancavelica (HU), Huánuco (HA), Distrito Federal (DF), Goiás (GO), Espírito Ica (IC), Junin (JU), La Libertad (LL), Santo (ES), Maranhão (MA), Mato Grosso Lambayeque (LA), Lima (LI), Loreto (LO), (MT), Mato Grosso do Sul (MS), Minas Gerais Madre de Dios (MD), Moquegua (MO), Pasco (MG), Pará (PA), Paraíba (PB), Paraná (PR), (PA), Piura (PI), Puno (PU), San Martín (SM), Pernambuco (PE), Piauí (PI), Rio de Janeiro Tacna (TA), Tumbes (TU), and Ucayali (UC). (RJ), Rio Grande do Norte (RN), Rio Grande • Colombia: Amazonas (AM), Antioquia (AN), do Sul (RS), Rondônia (RO), Roraima (RR), Arauca (AR), Atlántico (AT), Bolívar (BO), São Paulo (SP), Santa Catarina (SC), Sergipe Boyacá (BY), Caldas (CA), Caquetá (CQ), (SE), and Tocantins (TO). Casanare (CS), Cauca (CU), Cesar (CE), • Argentina: Buenos Aires (BA), Catamarca Chocó (CH), Córdoba (CO), Cundinamarca (CA), Córdoba (CR), Corrientes (CO), (CN), Guainía (GU), Guaviare (GA), Huila Chaco (CH), Chubut (CB), Entre Rios (ER), (HU), La Guajira (LG), Magdalena (MA), Meta Formosa (FO), Jujuy (JU), La Pampa (LP), La (ME), Nariño (NA), Norte de Santander (NS), Rioja (LR), Mendoza (ME), Misiones (MI), Putumayo (PU), Quindío (QU), Risaralda Neuquén (NE), Rio Negro (RN), Salta (SA), (RI), San Andrés (SA), Santander (SN), Sucre San Juan (SJ), San Luis (SL), Santa Cruz (SC), (SU), Tolima (TO), Valle del Cauca (VC), Santa Fe (SF), Santiago del Estero (SE), Tierra Vaupés (VA), and Vichada (VI). del Fuego (TF), and Tucumán (TU). • Chile: Aisén (AI), Antofagasta (AN), • Paraguay: Alto Paraguay (AP), Alto Paraná Araucanía (AU), Arica (AR), Atacama (AT), (AA), Amambay (AM), Distrito Capital (DC), Biobío (BI), Coquimbo (CO), Los Rios (LR), Boquerón (BO), Caaguazú (CA), Caazapá Los Lagos (LL), Magallanes (MG), Maule (CZ), Canindeyú (CN), Central (CE), (MA), O´Higgins (OH), Santiago (AS), Concepción (CO), Cordillera (CR), Guairá Tarapacá (TA) and Valparaiso (VA). (GU), Itapúa (IT), Misiones (MI), Ñeembucú • Uruguay: Artigas (AR), Canelones (CA), (NE), Paraguarí (PA), Presidente Hayes (PH), Cerro Largo (CL), Colônia (CO), Durazno and San Pedro (SP). (DU), Flores (FL), Florida (FO), Lavalleja • Bolivia: Chuquisaca (CH), Cochabamba (CO), (LA), Maldonado (MA), Montevidéu (MO), Beni (BE), La Paz (LP), Oruro (OR), Pando Paysandú (PA), Río Negro (RN), Rivera (RI), (PA), Potosí (PO), Santa Cruz (SC), and Tarija Rocha (RO), Salto (SA), San José (SJ), Soriano (TA). (SO), Tacuarembó (TA), and Treinta y Tres • Venezuela: Amazonas (AM), Anzoátegui (TT). (AN), Apure (AP), Aragua (AR), Barinas (BA), • Surinam: Brokopondo (BR), Commewijne Bolívar (BO), Carabobo (CA), Cojedes (CO), (CO), Coronie (CR), Marowijne (MA), Delta Amacuro (DA), Falcón (FA), Guárico Nickerie (NI), Para (PA), Paramaribo (PR), (GU), Lara (LA), Mérida (ME), Miranda Saramacca (SA), Sipaliwini (SI), and Wanica (MI), Monagas (MO), Nueva Esparta (NE), (WA). Portuguesa (PO), Sucre (SU), Táchira (TA), • French Guiana: Awala-Yalimapo (AY), Trujillo (TR), Yaracuy (YA), Vargas (VA), and Apatou (AP), Camopi (CA), Cayenne Zulia (ZU). (CY), Grand-Santi (GS), Iracoubo (IR), • Ecuador: Azuay (AZ), Bolívar (BO), Kourou (KO), Macouria (MC), Mana Cañar (CA), Carchi (CR), Cotopaxi (CO), (MA), Maripasoula (MR), Matoury (MT), Chimborazo (CH), Galápagos (GA), Imbabura Montsinéry-Tonnegrande (MO), Ouanary (IM), Loja (LO), Pichincha (PI), Tungurahua (OU), Papaïchton (PA), Régina (RE), Remire- (TU), El Oro (EO), Esmeraldas (ES), Guayas Montjoly (RM), Roura (RO), Saint-Élie (SE), (GU), Los Ríos (LR), Manabí (MA), Morona- Saint-Georges (SG), Saint-Laurent-du-Maroni Santiago (MS), Napo (NA), Pastaza (PA), (SL), Saül (SA), and Sinnamary (SI).

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 544 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

• : Barima-Waini (BW), Pomeroon- Hosts: torquatus Supenaam (PS), Essequibo Islands-West Site of infection: Small intestine Demerara (EI), Demerara-Mahaica (DM), Distribution: Brazil (19). Mahaica-Berbice (MB), East Berbice- Corentyne (EC), Cuyuni-Mazaruni (CM), Polymorphida Potaro-Siparuni (PS), Upper Takutu-Upper Centrorhynchidae Essequibo (TE), and Upper Demerara-Berbice Hosts: Mabuya macrorhyncha; M. agilis Raddi, (DB). 1823, Hemidactylus mabouia Moreau de Jonnés, 1818, Enyalius bilineatus Duméril & Bibron, LIST OF HELMINTHS 1837 Site of infection: Peritonium, liver; mesentery Acanthocephala Distribution: Brazil – Queimada Grande island Hosts: Tropidurus torquatus Wied-Neuwied, 1820 (Atlantic ) and Valinhos (Cerrado), SP; Site of infection: Stomach wall Grussaí (Restinga) and Nova Iguaçu (Atlantic Distribution: Argentina – CO (Chaco). Forest), RJ; Praia das Neves (Restinga) and Comments: No indication of family and/or Marechal Floriano (Atlantic Forest), ES. , but probably belong to Centrorhynchidae, Comments: All citations above consist of since several cystacanths of this family have been cystacanth occurrence and are, thus, difficult to found in South American lizards (16). identify to species level, because reproductive structures are required (20-24). Archiacanthocephala Oligacanthorhynchida Centrorhynchus Lühe, 1911 Oligacanthorhynchidae Centrorhynchus tumidulus Rudolphi, 1919 Oligacanthorhynchus Travassos, 1915 Hosts: Ameiva ameiva; Tropidurus torquatus; Oligacanthorhynchus sp. Tupinambis teguixin Linnaeus, 1758 Hosts: Ameiva ameiva Linnaeus, 1758 Site of infection: Body cavity Distribution: Paraguay – Rio Negro (Chaco), PH Comments: Tupinambis teguixin found in Rio de (10). Janeiro and in open in South America may actually correspond to T. merianae. Palaeacanthocephala Distribution: Brazil – Rio de Janeiro (Atlantic Echinorhynchida Forest), RJ (25, 26). Echinorhynchidae Hosts: Mabuya macrorhyncha Hoge, 1946 Centrorhynchus sp. Site of infection: Stomach wall Hosts: Tupinambis teguixin Distribution: Brazil – Trancoso (Restinga), BA. Site of infection: Not informed Comments: No indication of genus (17). Distribution: Paraguay – AP (Chaco) (10).

Acanthocephalus Koelreuther, 1771 Nematoda Acanthocephalus saurius Bursey & Goldberg, Adenophorea 2003 Enoplida Hosts: oshaugnessyi Boulenger, 1885 Capilariidae Site of infection: Small intestine Capillaria (Zeder, 1800) Bell & Beverley Burton Distribution: Brazil – AC (Amazon) (18). 1981 Capillaria (= Amphibiocapillaria) freitaslenti Acanthocephalus sp. Araujo & Gandra, 1941 Hosts: Enyalius perditus Jackson, 1978 Hosts: Tropidurus torquatus; Ameiva ameiva Site of infection: Stomach Site of infection: Intestine Distribution: Brazil – São Sebastião (Atlantic Distribution: Brazil – GO (Cerrado); Peru – CU Forest), SP (3). (Amazon) (1, 8).

Echinorhynchus Zoega in Müller, 1776 Ichthyocapillaria Moravec, 1982 Echinorhynchus sp. Ichthyocapillaria (= Pseudocapillaria) maricaensis

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 545 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

Rodrigues, 1992 it is considered a valid species by some authors Hosts: lutzae Mertens, 1938 (8, 37-39). Site of infection: Small intestine Distribution: Brazil – Maricá (Restinga), RJ (27). Cyrtosomum Gedoelst, 1919 Cyrtosomum longicaudatum Brenes and Bravo Ascaridida Hollis, 1960 Anisakidae Hosts: Anolis punctatus Daudin, 1802; Anolis Contracaecum Railliet & Henry, 1912 transversalis Duméril, 1851 Contracaecum sp. Site of infection: Large intestine Hosts: Tupinambis teguixin Distribution: Peru – CU (Amazon); Brazil – AM Site of infection: Not informed (Amazon) (1, 40). Distribution: Surinam – PR (Amazon) (28). Cyrtosomum sp. Ascarididae Hosts: Ophiodes striatus Dujardinascaris Baylis, 1927 Site of infection: Large intestine Dujardinascaris sp. Distribution: Brazil – MS (Cerrado) (41). Hosts: Kentropyx pelviceps Cope, 1868, Tupinambis teguixin Maracaya Díaz-Ungría, 1964 Site of infection: Stomach Maracaya belemensis Adamson & Baccam, 1988 Distribution: Peru – CU (Amazon) (1). Hosts: alba Linnaeus, 1758 Site of infection: Large intestine Freitasascaris Sprent, 1983 Distribution: Brazil – Belém (Amazon), PA (42). Freitasascaris alata Baylis, 1947 Hosts: Tupinambis teguixin Maracaya graciai Díaz-Ungría, 1969 Site of infection: Intestine Hosts: Distribution: Surinam – PR (Amazon); Brazil – Site of infection: Intestine AM (Amazon); Uruguay – MO Distribution: Venezuela – Maracay, AR (7, 42). Comments: Tupinambis teguixin from Uruguay may actually correspond to T. merianae. (28-30). Maracaya pusilla (Miranda, 1924) Adamson & Baccam, 1988 Hexametra Travassos, 1919 Hosts: Amphisbaena sp. Hexametra boddaertii Baird, 1860 Site of infection: Intestine Hosts: Cnemidophorus littoralis Rocha, Araujo, Distribution: Brazil – BA. Vrcibradic & Costa, 2000; C. ocellifer Spix, Remarks: The former Aplectana pusilla was 1825; Mabuya agilis; M. frenata Cope, 1862; M. transferred to Maracaya (42), according to a macrorhyncha; Tropidurus torquatus revision of the genus Maracaya (43), but it is cited Site of infection: Body cavity in the genus Aplectana by other authors (42-44). Distribution: Brazil – Jurubatiba, Grussaí and Marica (Restinga), RJ; Salvador and Trancoso Cosmocercidae (Restinga), BA; Valinhos (Cerrado), SP (17, 23, gen. sp. 31-35). Hosts: Cercosaura argulus Peters, 1863 Site of infection: Small intestine Atractidae Distribution: Peru – Rio Nanay (Amazon), LO Atractis Dujardin, 1945 (45). Atractis cruciata Linstow, 1902 Hosts: Tupinambis teguixin, Stenocercus Aplectana Railliet & Henry, 1916 roseiventris Aplectana albae Adamson & Baccam, 1988 Site of infection: Intestine Hosts: Amphisbaena alba; Amphisbaena ridleyi Distribution: Brazil – Belém (Amazon), PA; Boulenger, 1890 Ecuador – PA (Amazon). Site of infection: Intestine Comments: Atractis cruciata is considered a Distribution: Brazil – Belém (Amazon), PA; junior of A. opeatura (7, 36). However, Fernando de Noronha Island, PE (42, 46).

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 546 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

Aplectana raillieti Travassos, 1925 Paradollfusnema Baker, 1982 Hosts: Amphisbaena alba Paradollfusnema Baker, 1981 Site of infection: Intestine Hosts: Leposternon microcephalum Distribution: Brazil – Angra dos Reis (Insular Site of infection: Intestine Atlantic Forest), RJ. Distribution: Brazil – Recife, PE (8, 53). Comments: The host Amphisbaena fusca was originally cited as Amphisbaena alba (5, 7, 8, 47). Raillietnema Travassos, 1927 Raillietnema brachyspiculatum Bursey, Goldberg, Aplectana tucumanensis Ramallo, Bursey & Salgado-Maldonado, and Mendez-de-la-Cruz, Goldberg, 2008 1998 Hosts: Amphisbaena bolivica Mertens, 1929 Hosts: annularis (O’Shaughnessy, Site of infection: Large intestine 1881) Distribution: Argentina – San Miguel de Tucumán Site of infection: Large intestine (Chaco), TU (44). Distribution: Ecuador – PA (Amazon) (39).

Aplectana unguiculata Rudolphi, 1819 Heterakidae Hosts: Amphisbaena sp. Africana Travassos, 1920 Site of infection: Not informed Africana chabaudi Baker, 1981 Distribution: Brazil. Hosts: Uranoscodon superciliosus Comments: The species is actually considered as Site of infection: Intestine species inquirenda (7). Distribution: Brazil – PA, RO and RR (Amazon) (50, 54). Aplectana vellardi Travassos, 1926 Hosts: Enyalius perditus Africana dardanelosi Ávila & Silva, 2009 Site of infection: Intestine Hosts: Hoplocercus spinosus Distribution: Brazil – Ibitipoca (Cerrado), MG Site of infection: Intestine (48). Distribution: Brazil – MT (Amazon) (55).

Cosmocerca Diesing, 1861 Moaciria Freitas, 1956 Cosmocerca rara Freitas & Vicente, 1966 Moaciria alvarengai Freitas, 1956 Hosts: Leposternon microcephalum Wagler, 1824 Hosts: Trachylepis atlantica Schmidt, 1945 Site of infection: Large intestine Site of infection: Large intestine Distribution: Brazil – Rio de Janeiro (Atlantic Distribution: Brazil – Fernando de Noronha forest), RJ (8, 49). Island, PE (8, 46, 56).

Cosmocerca vrcibradici Bursey & Goldberg, 2004 Spinicauda Travassos, 1920 Hosts: angulatus Linnaeus, 1758; Spinicauda spinicauda Olfers, 1819 Alopoglossus atriventris Duellman, 1973; Anolis Hosts: Ameiva ameiva, Tupinambis teguixin, fuscoauratus D’Orbignyi, 1837; Cercosaura Tupinambis merianae, Trachylepis atlantica, eigenmanni Griffin, 1917; C. oshaugnessyi Enyalioides praestabilis (O’Shaughnessey, 1881) Boulenger, 1885; Uranoscodon superciliosus Site of infection: Intestine Linnaeus, 1758 Distribution: Brazil – MT (Pantanal), Rio de Site of infection: Intestine Janeiro, RJ, São Gonçalo, RN and Fortaleza, CE, Distribution: Brazil – AC, AM, PA, RO (Amazon); Fernando de Noronha Islands, PE; Peru – CU Ecuador – SU (Amazon) (18, 50-52). (Amazon); Surinam – PR (Amazon); Venezuela – AR; Ecuador – PA (Amazon) (4, 7, 28, 39, 46, Cosmocerca sp. 50, 57-60). Hosts: Enyalius perditus Site of infection: Stomach Strongyluris Mueller, 1894 Distribution: Brazil – São Sebastião (Insular Strongyluris oscari Travassos, 1923 Atlantic forest), SP (3). Hosts: Ameiva ameiva; Anolis fuscoauratus; A.

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 547 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America punctatus; A. transversalis; Enyalius iheringii Falcaustra belemensis Baker & Bain, 1981 Boulenger, 1885; E. perditus; Eurolophosaurus Hosts: Neusticurus bicarinatus Linnaeus, 1758; nanuzae Rodrigues, 1981; Mabuya agilis; ecpleopus Cope, 1876; P. strangulatus plica Linnaeus, 1758; Plica umbra Linnaeus, 1758; Cope, 1868 Stenocercus caducus; S. roseiventris D’Orbigny in Site of infection: Rectum Duméril & Bibron 1837; Tropidurus sp.; T. guarani Distribution: Ecuador – PA (Amazon) (39). Alvarez, Cei & Scolaro, 1994; T. spinulosus Cope, 1862; T. torquatus; T. melanopleurus Falcaustra sp. Site of infection: Stomach, intestine Hosts: Anolis punctatus Distribution: Brazil – AC, AM, BA, CE, DF, ES, Site of infection: Large intestine GO, MG, MS, PA, PB, RJ, RO, SP (Pantanal, Distribution: Brazil – AM (Amazon) (40). Restinga, Amazon, Caatinga, Atlantic Forest); Argentina (Chaco); Paraguay (Chaco); Ecuador Seuratidae – SU and PA (Amazon); Peru – CU (Amazon); Skrjabinelazia Sypliaxov, 1930 Bolivia – Florida province, SC. Skrjabinelazia galliardi Chabaud, 1973 Comments: Currently, 31 species are recognized Hosts: Gonatodes humeralis Guichenot, 1855 within the genus, and based on morphological Site of infection: Stomach characteristics, only four occurrs in neotropical Distribution: Brazil – Belém (Amazon), PA (8, regions (61). Strongyluris oscari seems to be 74, 75). extremely polymorphic, since the species S. freitasi, S. travassosi and S. sai have been Skrjabinellazia intermedia Freitas, 1940 synonimized (62). The records of S. oscari Hosts: Anolis puncatus; Cnemidophorus nativo infecting T. spinulosus in the region of Salobra, Rocha, Bergallo & Peccinini-Seale, 1997; MS, Brazil correspond to T. guarani (1, 3, 32, 35, Tropidurus guarani; T. torquatus; Stenocercus 39-41, 48, 51, 60, 62-69). caducus (Cope, 1862) Site of infection: Stomach, intestine Kathlaniidae Distribution: Brazil – Salobra (Pantanal) and Cruzia Travassos, 1917 Bodoquena (Cerrado), MS; Salvador and Cruzia fulleborni Khalil & Vogelsang, 1930 Guaratiba (Restinga), BA; Jurubatiba (Restinga), Hosts: Tupinambis teguixin RJ; Paraguay – (Chaco); Peru – CU (Amazon). Site of infection: Intestine Comments: Originally described as Salobrella Distribution: Argentina; Paraguay (7, 70-72). intermedia found in Tropidurus spinulosus from Salobra, MS. See comments on host taxonomy at Cruzia rudolphii Ruiz, 1947 Strongyluris oscari (1, 35, 41, 62, 65, 76, 77). Hosts: Hoplocercus spinosus Fitzinger, 1843 Site of infection: Large intestine Subuluridae Distribution: Brazil – Dois Irmãos do Buriti Subulura Molin, 1860 (Cerrado), MS (71, 73). Subulura lacertilia Vicente, Van-Sluys, Fontes & Kiefer, 2000 Cruzia tentaculata Rudolphi, 1819 Hosts: Eurolophosaurus nanuzae, Cnemidophorus Hosts: Tupinambis teguixin nativo Site of infection: Intestine Site of infection: Intestine Distribution: Uruguay – MO (29, 71). Distribution: Brazil – Serra do Cipó (Cerrado), MG; Guaratiba (Restinga), BA (66, 77, 78). Cruzia travassosi Kalil & Vogelsangi, 1932 Hosts: Tupinambis merianae Subulura sp. Site of infection: Intestine Hosts: Tropidurus torquatus Distribution: Brazil – Salobra (Pantanal) and Site of infection: Small intestine Bodoquena (Cerrado) MS; Bolivia; Argentina (7, Distribution: Brazil – Jurubatiba (Restinga), RJ 41, 71). (35).

Falcaustra Lane, 1915 Oxyurida

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 548 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

Oxyuridae Distribution: Brazil – Santa Luzia (Caatinga), PB; Gynaecometra Araújo, 1976 Peru; Venezuela – La Puerta, GU, Blanquilla, Los Gynaecometra bahiensis Araújo, 1976 Frailes, Los Testigos and Margarita; Colombia – Hosts: Polychrus acutirostris Spix, 1825 LG; Surinam – PR and Marienburg, CO (8, 29, Site of infection: Intestine 82, 83). Distribution: Brazil – Xique-xique (Caatinga), BA; Campo Grande (Cerrado), MS (41, 79). Ozolaimus megatyphlon Rudolphi, 1819 Hosts: Iguana iguana Typhlonema Kreis, 1938 Site of infection: Intestine Typhlonema sp. Distribution: Brazil – Santa Luzia (Caatinga), PB; Hosts: Tropidurus guarani Peru; Venezuela – La Puerta, GU, Blanquilla, Los Site of infection: Not informed Frailes, Los Testigos and Margarita; Colombia – Distribution: Brazil – Salobra (Pantanal), MS (8). LG; Surinam – PR and Marienburg, CO (8, 29, 82, 83). Pharyngodonidae Alaeuris Thapar, 1925 Paralaeuris Cuckler, 1938 Alaeuris caudatus Lent & Freitas, 1948 Paralaeuris dorochila Cuckler, 1938 Hosts: Iguana iguana Hosts: Conolophus subcristatus Site of infection: Intestine Site of infection: Intestine Distribution: Brazil – Exu (Caatinga), PE; Distribution: Ecuador – Galapagos Islands (7). Venezuela – La Puerta, GU; Peru – TU (8, 29, 80). Parapharyngodon Chatterji, 1933 Alaeuris conolophi Cuckler, 1938 Parapharyngodon alvarengai Freitas, 1957 Hosts: Conolophus subcristatus Gray, 1831 Hosts: Trachylepis atlantica, Ameiva ameiva, Site of infection: Intestine Amphisbaena ridleyi Distribution: Ecuador – Galapagos Islands (7). Site of infection: Large intestine Distribution: Brazil – Fernando de Noronha Alaeuris galapagensis Cuckler, 1938 Island, PE and Itaguaí (Restinga), RJ. Hosts: Conolophus subcristatus Gray, 1831 Comments: after the synonym it was cited as Site of infection: Intestine Thelandros alvarengai(46, 84, 85). Distribution: Ecuador – Galapagos Islands (7). Parapharyngodon arequipensis Calisaya & Alaeuris labicula Cuckler, 1938 Córdova, 1997 Hosts: Conolophus subcristatus Hosts: peruvianus (Lesson, 1826) Site of infection: Intestine Site of infection: Large intestine Distribution: Ecuador – Galapagos Islands (7). Distribution: Peru – Omate, MO. Comments: This species is apparently ignored by Alaeuris longispicula Cuckler, 1938 severeal authors (86-90). Hosts: Conolophus subcristatus Site of infection: Intestine Parapharyngodon largitor Alho & Rodrigues, Distribution: Ecuador – Galapagos Islands (7). 1963 Hosts: Ameiva ameiva, Mabuya agilis, Alaeuris vogelsangi Lent & Freitas, 1948 Hemidactylus mabouia, guttulatum Hosts: Iguana iguana Site of infection: Intestine Site of infection: Intestine Distribution: Brazil – Manguinhos and Grumari Distribution: Brazil – Exu (Caatinga), PE, CE; (Restinga), RJ and Valinhos (Cerrado), SP; PERU Venezuela – La Puerta, GU (8, 29, 81). – LA (8, 22, 33, 91, 92).

Ozolaimus Dujardin, 1845 Parapharyngodon moqueguensis Calisaya & Ozolaimus cirratus Linstow, 1906 Córdova, 1997 Hosts: Iguana iguana Hosts: Microlophus peruvianus, Dicrodon Site of infection: Intestine guttulatum

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 549 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

Site of infection: Large intestine Parapharyngodon verrucosus Freitas & Dobbin, Distribution: Peru – Moquegua, MO, LI, TU, PI, 1959 LA, LL, AN. Hosts: Diploglossus lessonae Comments: check P. arequipensis (89, 90, 92). Site of infection: Intestine Distribution: Brazil – João Alfredo (Caatinga), Parapharyngodon riojensis Ramallo, Bursey & PE; CE (Caatinga) (8, 81, 99). Goldberg, 2002 Hosts: punae Cei, Etheridge & Parapharyngodon yurensis Calisaya & Córdova, Videla, 1985; P. palluma Molina, 1782; Liolaemus 1997 buergeri Werner, 1907 Hosts: Microlophus peruvianus Site of infection: Large intestine Site of infection: Large intestine Distribution: Argentina – LR (2, 86). Distribution: Peru – Yura, AR. Comments: check P. arequipensis (89, 90). Parapharyngodon sceleratus Travassos, 1923 Hosts: Ameiva ameiva; Cnemidophorus littoralis; Parapharyngodon sp Eurolophosaurus nanuzae; Hemidactylus Hosts: Ameiva ameiva, Cnemidophorus nativo, mabouia; Kentropyx pelviceps; Liolaemus lutzae; Hemidactylus mabouia, Tropidurus torquatus, T. Mabuya agilis; M. bistriata; M. caissara Rebouças- etheridgei, Spieker, 1974; M. frenata; M. macrorhyncha; Site of infection: Intestine Microlophus albermalensis Baur, 1890; P. Distribution: Brazil – Salobra, MS; Cabo Frio, johnwrighti; Phyllopezus pollicaris; T. guarani; T. Nova Iguaçu and Maricá, RJ, Salvador and hispidus; T. itambere; T. semitaeniatus Spix, 1825; Guaratiba, BA; Argentina – CO, SA; Peru – TU, T. torquatus; T. melanopleurus and Tropidurus sp. LA, LI (8, 16, 21, 77, 92, 100-102). Site of infection: Intestine Distribution: Brazil – Abrolhos, Trancoso Pharyngodon Diesing, 1861 (Restinga) and Canudos, BA; CE; Linhares and Pharyngodon cesarpintoi Pereira, 1935 Praia das Neves, ES; Serra do Cipó (Cerrado), Hosts: Cnemidophorus sp., Ameiva ameiva, MG; Salobra (Pantanal), MS; Xavantina, GO; Liolaemus lutzae, Tropidurus itambere Cachimbo, PA; Mogeiro, Lagoa Remígio, Site of infection: Large intestine Umbuseiro and João Pessoa, PB; Garanhuns, Distribution: Brazil – Juazeiro and Mogeiro PE; Grumari, Rio de Janeiro, Arraial do Cabo, (Caatinga), PB, BA, RN; Maricá (Restinga), RJ; Grussaí, Manguinhos and Maricá, RJ; Cruzeta, Bodoquena (Cerrado), MS. Currais Novos and Ceará Mirim (Caatinga), RN; Comments: The type of host is apparently Queimada Grande, Caraguatatuba and Valinhos, misidentified in the original description (60). SP; Bolivia: Florida province, SC and El Carmen; However, C. lemniscatus does not occur in the Paraguay; Peru – CU; Ecuador – Galapagos Caatinga biome, but C. occelifer and at least three Islands (1, 7, 8, 17, 20, 22, 27, 32-34, 62, 65, 66, undescribed species occur (41, 60, 103, 104). 69, 81, 91, 93-97). Comments: This species has been widely reported Pharyngodon micrurus Freitas & Ibañez, 1963 as P. scleratus (1, 86). However, according to the Hosts: , D. guttulatum original description and the latter redescription, Site of infection: Intestine the proper spelling of the specific name issceleratus Distribution: Peru – Trujillo, LL; TU, PI, LA, AN, (59, 93). Cnemidophorus occelifer, cited by Ribas et LI (7, 92, 105). al. (94), actually corresponds to C. littoralis (31). Pharyngodon travassosi Pereira, 1935 Parapharyngodon senisfaciecaudus Freitas, 1957 Hosts: Ameiva ameiva Hosts: Liolaemus signifer Duméril & Bibron, 1837 Site of infection: Large intestine Site of infection: Large intestine Distribution: Brazil – Areia (Caatinga), PB (60). Distribution: Bolivia – LP. Comments: It was originally described infecting Pharyngodon sp. Liolaemus lenzi, which actually corresponds to L. Hosts: Tropidurus torquatus, Tropidurus hispidus signifier (98). Spix, 1825

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 550 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

Site of infection: Intestine Distribution: Peru – Pumarongo, CA; Argentina; Distribution: Brazil – Salvador, BA; CE (8, 81). Chile (2, 111, 112).

Pseudostrongyluris Guerrero, 1971 Spauligodon oxkutzcabiensis (Chitwood, 1938) Pseudostrongyluris polychrus Guerrero, 1971 Hosts: Thecadactylus solimoensis Bergmann Hosts: Polychrus marmoratus & Russell, 2007; Phyllodatylus reissii Peters, Site of infection: Not informed 1862; P. inaequalis Cope, 1876; P. johnwrighti; Distribution: Venezuela – Federal District (7). P. microphyllus Cope, 1876; Tropidurus guarani; Bogertia lutzae (Loveridge, 1941). Skrjabinodon Inglis, 1968 Site of infection: Intestine Skrjabinodon dixoni Bursey & Goldberg, 2007 Distribution: Peru – CU, (Amazon); Paraguay Hosts: flaviceps(Guichenot, 1855) – Arroyo Corrientes (Chaco), PA; Brazil – AL Site of infection: Large intestine (Atlantic Forest). Distribution: Peru – LO (Amazon); Ecuador – SU Comments: Thecadactylus was recently subject (106). to revision, and T. solimoensis was described by Bergmann & Russel in localities that encompasses Skrjabinodon heliocostai Vicente, Vrcibradic, the Cuzco, Peru (1, 65, 97, 113-115). Muniz-Pereira & Pinto, 2000 Hosts: Mabuya frenata Spauligodon viracochai Freitas, Vicente & Ibañez, Site of infection: Large intestine 1968 Distribution: Brazil – Valinhos (Cerrado), SP Hosts: Phyllodactylus gerrhopygus Wiegmann, (107). 1834; P. angustidigitus Dixon & Huey, 1970 Site of infection: Intestine Skrjabinodon spinulosus Vicente, Vrcibradic, Distribution: Peru – Trujilo, LL and IC (116, 117). Rocha & Pinto, 2002 Hosts: Mabuya dorsivittata Thelandros Wedl, 1862 Site of infection: Intestine Thelandros bulbosus Salas & Campos, 1974 Distribution: Brazil – Itatiaia (Atlantic Forest), Hosts: Microlophus peruvianus RJ; Itirapina (Cerrado), SP (108,109). Site of infection: Not informed Distribution: Peru. Spauligodon Skrjabin, Schikhobalova & Comments: Despite the citation by Pérez et al. Lagodovsk., 1960 (117), this oxyurid was incorrect described in Spauligodon loboi Ramallo, Bursey & Goldberg “Libro de Resumenes del IV Congreso Nacional (2002) de Biologia”, a meeting abstract. Due to an Hosts: Liolaemus capillitas Hulse, 1979; L. inaccordance with articles 8.1 and 9.9 of the huacahuasicus Laurent, 1985; L. quilmes International Code for Zoological Nomenclature, Etheridge, 1993; L. ornatus Koslowsky, 1898; L. it was therefore considered a species inquirenda ramirezae Lobo, 1999 (107, 118). Site of infection: Large intestine Distribution: Argentina – Tafídel Valle and Thelandros capacyupanquii Freitas, Vicente & Amaicha del Valle, TU; El Cerrillo, Andalgalá, Ibañez, 1968 Belén, and Santa María, CA; San Antonio de los Hosts: Dicrodon holmbergi Schmidt, 1957; Cobres, Cachi, and La Poma, SA; and Tilcara and Dicrodon guttulatum Humahuaca, JU (110). Site of infection: Intestine Distribution: Peru – Trujilo, LL, IC, AN (92, 107, Spauligodon maytacapaci Vicente & Ibañez (1968) 118). Hosts: Leiocephalus sp.; Liolaemus andinus Koslowsky, 1895; L. chilensis Lesson, 1830; Thelandros sp. L. elongatus Koslowsky, 1896; L. lemniscatus Hosts: Tropidurus etheridgei Gravenhorst, 1838; L. pictus Duméril & Bibron, Site of infection: Intestine 1837; L. tenuis Duméril & Bibron, 1837 Distribution: Argentina – Departamento Anta Site of infection: Intestine (Chaco), SA (102).

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 551 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

Rhabditida Jurubatiba and Grussaí (Restinga), RJ; Marechal Rhabdiasidae Floriano (Atlantic Forest) and Praia das Neves Chabirenia Lhermitte Vallarino, Bain, Deharo, (Restinga), ES; Peru – Rio Nanay, LO. Bertani, Voza, Attout & Gaucher, 2005 Comments: all aforementioned citations consist Chabirenia cayennensis Lhermitte Vallarino, Bain, of cysts and thus difficult the identification to Deharo, Bertani, Voza, Attout & Gaucher, 2005 species level since reproductive structures are Hosts: Ameiva ameiva required (20, 22-24, 35, 45). Site of infection: Mouth Distribution: French Guiana – CY (119). Diplotriaenidae Hastospiculum Skrjabin, 1923 Rhabdias Stiles & Hassall, 1905 Hastospiculum sp. Rhabdias anolis Bursey, Goldberg & Telford, 2003 Hosts: Plica umbra Hosts: Anolis punctatus Site of infection: Peritoneum Site of infection: Lungs Distribution: Peru – CU (Amazon) (1). Distribution: Peru – CU (Amazon) (1). Filariidae Rhabdias elegans Gutierrez, 1945 Filaria Mueller, 1787 Hosts: Anolis punctatus Filaria multipapilla Molin, Site of infection: Lungs Hosts: Dracaena guianensis Daudin, 1802 Distribution: Ecuador – PA (Amazon) (1). Site of infection: Body cavity Distribution: Brazil – Belém (Amazon), PA. Rhabdias sp. Comments: The species is actually considered Hosts: undulatus Wiegmann, 1834; species inquirenda, because these first records Anolis fuscoauratus; A. punctatus; Enyalius were lost and no new information about the iheringii; E. bilineatus; E. perditus; Tropidurus species has been presented since the description hispidus (123-125). Site of infection: Lungs, stomach Distribution: Brazil – Marechal Floriano (Atlantic Onchocercidae Forest), ES; PA, AM (Amazon); São Sebastião Macdonaldius Khanna, 1933 (Atlantic Forest), SP, Chapada do Araripe Macdonaldius grassi (Caballero, 1954) (Caatinga), CE; Ecuador – SU. Hosts: Phyllopezus pollicaris (Spix, 1825) Comments: Of the neotropical lizards, only Site of infection: Underneath the epidermis of the Rhabdias anolis has been described, despite throat several reports in the same region (3, 24, 40, 51, Distribution: Paraguay – CO (Chaco) (126). 120, 121). OswaldofilariaTravassos, 1933 Strongyloididae Oswaldofilaria azevedoi Bain, 1974 Strongyloides Grassi, 1897 Hosts: Polychrus marmoratus Linnaeus, 1758; Strongyloides cruzi Rodrigues, 1968 Stenocercus roseiventris Hosts: Hemidactylus mabouia Site of infection: Body cavity Site of infection: Small intestine Distribution: Brazil – Belém (Amazon), PA; Peru Distribution: Brazil – Manguinhos, RJ (8, 122). – CU (Amazon) (1, 127).

Spirurida Oswaldofilaria belemensis Bain and Dulahian, Acuariidae 1974 Hosts: Cercosaura argulus; Hemidactylus Hosts: Dracaena guianensis mabouya; Mabuya agilis; M. macrorhyncha; Site of infection: Heart, aorta e vena cava Tropidurus torquatus; Enyalius bilineatus; Distribution: Brazil – Belém (Amazon), PA (8). Phyllodactylus lepidopygus Tschudi, 1845 Site of infection: Body cavity Oswaldofilaria brevicaudata Rodhain and Distribution: Brazil – Queimada Grande Island Vuylsteke, 1937 (Atlantic Forest) and Valinhos (Cerrado), SP; Hosts: Iguana iguana, Anolis punctatus

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 552 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

Site of infection: Body cavity 1974 Distribution: Brazil – Marajó (Amazon), PA, AM Hosts: Mabuya bistriata Spix, 1825 (Amazon), Exu (Caatinga), PE; Venezuela – ZU Site of infection: Body cavity, large intestine (7, 40, 125, 127-129). Distribution: Brazil – Belém (Amazon), PA; Peru – CU (Amazon) (1, 127). Oswaldofilaria petersi Bain and Sulahian, 1974 Hosts: Tupinambis teguixin, Tropidurus hispidus Physalopteridae Site of infection: Body cavity Abbreviata Travassos, 1920 Distribution: Brazil – Belém (Amazon), PA, Abbreviata spiralis (Schneider, 1866) Chabaud, Ibiraba (Caatinga), BA (8, 130). 1956 Hosts: Amphisbaena sp. Oswaldofilaria spinosa Bain and Sulahian, 1974 Site of infection: Not informed Hosts: Mabuya bistriata Distribution: Brazil Site of infection: Body cavity Comments: This species is considered a species Distribution: Brazil – Belém (Amazon) (8). inquirenda (7).

Oswaldofilaria sp. Physaloptera Rudolphi, 1819 Hosts: Tupinambis teguixin, Mabuya frenata Physaloptera lutzi Cristofaro, Guimarães & Site of infection: Body cavity Rodrigues, 1976 Distribution: Surinam – PR; Brazil – Valinhos Hosts: Ameiva ameiva; Cnemidophorus abaetensis (Cerrado), SP (28, 34). Reis, Dias, Rocha & Vrcibradic, 2002; C. littoralis; Enyalius bilineatus; Eurolophosaurus nanuzae; Piratuba Freitas & Lent, 1947 Liolaemus alticolor Barbour 1909; L. ornatus; Piratuba digiticauda Lent & Freitas, 1941 L. quilmes; Tropidurus guarani; T. itambere; T. Hosts: Tropidurus torquatus, Tropidurus guarani, torquatus Plica umbra, Kentropyx calcarata Site of infection: Stomach Site of infection: Body cavity, intestine Distribution: Argentina – SA, TU; Bolivia – Distribution: Brazil – Salvador, BA; Piratuba Roboré (Pantanal); Brazil – Salvador, Serrinha (Amazon), PA; Salobra (Pantanal), MS; Juara and Canudos (Caatinga), BA; Linhares and (Amazon), MT; Peru – CU; Paraguay – Chaco. Conceição da Barra (Restinga), Marechal Floriano Comments: Tropidurus spinulosus found in (Atlantic Forest), ES; Serra do Cipó (Cerrado), Salobra actually correspond to T. guarani (1, 62, MG; Porto Esperança and Salobra (Pantanal), 63, 65, 125, 131). MS; Xavantina, MT; Cachimbo (Amazon), PA; Maricá and Jurubatiba (Restinga), RJ; Valinhos Piratuba lainsoni Bain, 1974 (Cerrado), SP; Paraguay – Chaco (8, 24, 31, 32, Hosts: Anolis punctatus, Polychrus marmoratus 35, 62, 65, 66, 94-96, 132, 133). Site of infection: Body cavity, large intestine Comments: Cnemidophorus occelifer, cited by Distribution: Brazil – Belém (Amazon), PA; Peru Ribas et al. (94), actually corresponds to C. – CU (Amazon) (1, 127). littoralis (31).

Piratuba scaffi Bain, 1974 Physaloptera retusa Rudolphi, 1819 Hosts: Ameiva ameiva Hosts: Ameiva ameiva; Ameiva festiva Site of infection: Body cavity Lichtenstein, 1856; Amphisbaena alba; Anolis Distribution: Brazil – Belém (Amazon), PA (127). fuscoauratus; Anolis punctatus; Cnemidophorus abaetensis; C. lemniscatus; C. littoralis; C. native; Piratuba shawi Bain, 1974 C. ocellifer; Enyalius bilineatus; Iguana iguana; Hosts: Kentropyx calcarata Spix, 1825 Kentropyx altamazonica; K. calcarata; K. pelviceps; Site of infection: Body cavity Leiosaurus bellii Duméril & Bibron, 1837; L. Distribution: Brazil – Belém (Amazon), PA (127). catamarcensis Koslowsky, 1898; Liolaemus lutzae; L. neuquensis Cei & Videla, 2003; Mabuya agilis; Piratuboides Bain & Sulahian, 1974 M. bistriata; M. dorsivittata Cope, 1862; Ophiodes Piratuboides zeae (Bain, 1974) Bain and Sulahian, striatus Spix, 1824; Plica plica; P. umbra; Potamites

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 553 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America ecpleopus; Stenocercus roseiventris; Thecadacytlus MS corresponds to T. merianae (personal solimoensis; Tropidurus guarani; T. hispidus; T. observation) (8, 16, 22, 31, 33, 35, 41, 45, 92, 96, torquatus; Tupinambis rufescens Günther, 1871; 101, 102, 143). T. longilineus Ávila-Pires, 1995; T. teguixin; ; Tropidurus melanopleurus Physalopteroides Wu & Liu, 1940 Boulenger, 1902 Physalopteroides venancioi Lent, Freitas & Site of infection: Stomach, intestine Proença, 1946 Distribution: Brazil – Salvador and Guaratiba Hosts: Alopoglossus atriventris Duellman, 1973; (Restinga), BA; Conceição da Barra (Restinga) Ameiva ameiva; Cercosaura ocellata Wagler, 1830; and Marechal Floraino (Atlantic Forest), ES; Cnemidophorus native; Kentropyx altamazonica; Salobra (Pantanal), MS; Juara (Amazon), MT; K. calcarata; K. pelviceps; Mabuya agilis; M. Altamira, Belém, Cachimbo, Novo Progresso bistriata; M. macrorhyncha; Thecadactylus and Santarém (Amazon), PA; Itatiaia (Atlantic solimoensis; Tropidurus torquatus; Tupinambis Forest), Maricá and Jurubatiba (Restinga), RJ; Ilha teguixin Seca (Atlantic Forest) and Itirapina (Cerrado), Site of infection: Stomach SP; Paraguay – Chaco; Bolivia – El Carmen and Distribution: Brazil – AC (Amazon); Trancoso Florida, SC; Argentina; Peru – CU (Amazon); and Guaratiba (Restinga), BA; Juara (Amazon), Surinam – PR (Amazon); Uruguay – Montevideo; MT; Magé, Grussaí and Jurubatiba (Restinga), Venezuela – Colón, ZU; Atures, AM; Salamanca, RJ; Praia das Neves (Restinga), ES; Peru – CU NE; Cumuná (Arid zone) and La Orchila, Mar (Amazon) (1, 17, 23, 33, 35, 52, 77, 131, 144). Caribe; Colombia – CH. Comments: Baker (7) cited several species of lizards Thubunaea Seurat, 1914 that does not occur in Brazil, such as Tupinambis Thubunaea iguanae (Telford, 1965) rufescens, Cnemidophorus lateristrigus, Euprepis Hosts: Microlophus peruvianus; M. thoracicus spixii, Podinema graphica and P. scripta. According (Tschudi, 1845) to the same author, Pygopus gronovii actually Site of infection: Stomach corresponds to Ophiodes striatus and Podinema Distribution: Peru – R. N. Paracas, IC (117). corresponds to Tupinambis, but no species are currently recognized as synonyms of P. scripta Thubunaea parkeri Baylis, 1926 and P. graphica (7). Cnemidophorus occelifer, Hosts: Microlophus occipitalis Peters, 1871; cited by Ribas et al. (94), actually corresponds to Dicrodon heterolepis C. littoralis (1, 8, 24, 28, 29, 31, 32, 35, 51, 57, 62, Site of infection: Stomach 69, 77, 94, 108, 126, 131, 134-142). Distribution: Peru (7, 145).

Physaloptera sp. Spiruridae Hosts: Ameiva ameiva; Cercosaura argulus; Spirurinae Cnemidophorus litoralis; C. ocellifer; Hemidactylus Hosts: Hemidactylus mabouia mabouia; Hoplocercus spinosus; Mabuya agilis; M. Site of infection: Stomach, lungs, and small macrorhyncha; Polychrus acutirostris; Tropidurus intestine etheridgei; T. torquatus; Tupinambis merianae Distribution: Brazil – Rio de Janeiro, RJ (146). Duméril & Bibron, 1839; Tupinambis teguixin; Dicrodon guttulatum. Micropleuridae Site of infection: Stomach Micropleura Linstow, 1906 Distribution: Brazil – Abrolhos and Salvador Hosts: Tupinambis meriana (Restinga), BA; Linhares and Praia das Neves Site of infection: Body cavity (Restinga) and Sooretama (Cerrado), ES; Salobra Distribution: Brazil – Salobra (Pantanal), MS. (Pantanal) and Bodoquena (Cerrado), MS; Comments: This record must be discharged, Ilha Grande, Cabo Frio, Grussaí, Grumari and because no mention of Micropleura was found in Jurubatiba (Restinga), RJ; Valinhos (Cerrado), Travassos and Freitas (147) as cited by Vicente et SP; Alegria (Pampas), RS; Argentina – SA and FO al. (8). Moreover, Micropleura may be found in (Chaco); Peru – Rio Nanay, LO, LI. the body cavity of crocodiles and turtles (148). Comments: Tupinambis teguixin of Salobra,

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 554 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America

Diaphanocephalidae Distribution: Ecuador – San Pablo (Amazon), LR Diaphanocephalus Diesing, 1851 (154). Diaphanocephalus diesingi Freitas & Lent, 1938 Hosts: Tupinambis teguixin Oswaldocruzia benslimanei Durette-Desset, Site of infection: Small intestine Anjos & Vrcibradic, 2006 Distribution: Brazil – Belém (Amazon), PA (149). Hosts: Enyalius bilineatus Duméril & Bibron, 1837 Diaphanocephalus galeatus Rudolphi, 1819 Site of infection: Small intestine Hosts: Tupinambis merianae, T. rufescens, T. Distribution: Brazil – Marechal Floriano, ES (24, teguixin 155). Site of infection: Intestine Distribution: Brazil – Salobra (Pantanal) and Oswaldocruzia brasiliensis Lent & Freitas, 1935 Bodoquena (Cerrado), MS; Manguinhos, Angra Hosts: Hemidactylus mabouia dos Reis e Rio de Janeiro (Atlantic Forest), RJ; Site of infection: Small intestine Pedras Altas, RS; Ilha Grande and São Paulo, SP, Distribution: Brasil – Rio de Janeiro, RJ (8, 156). Fernando de Noronha, PE; Argentina; Bolivia; Surinam – PR (8, 28, 46, 149, 150). Oswaldocruzia burseyi Durette-Desset, Anjos & Vrcibradic, 2006 Diaphanocephalus jacuruxi Alho, 1965 Hosts: Enyalius perditus Hosts: Dracaena guianensis Site of infection: Stomach Site of infection: Small intestine Distribution: Brazil – São Sebastião (Atlantic Distribution: Brazil – Ilha das Onças (Amazon), Forest), SP (3, 155). PA (8, 151). Oswaldocruzia fredi Durette-Desset, Anjos & Kalicephalus Molin, 1861 Vrcibradic, 2006 Kalicephalus sp. Hosts: Enyalius iheringii Hosts: Tupinambis teguixin Site of infection: Stomach Site of infection: Intestine Distribution: Brazil – São Sebastião (Atlantic Distribution: Brazil – Sooretama, ES; Alegria Forest), SP (3, 155). (Pampas), RS. Comments: Species of Kalicephalus spp. have Oswaldocruzia peruensis Ben-Slimane, Verhaag been reported in several species, and & Durette-Desset, 1995 seems to be restricted to these animals, likewise Hosts: Stenocercus roseiventris, Cercosaura Diaphanocephalus spp. (a sister taxon) in argulus, Anolis punctatus Tupinambinae. Published records could be a Site of infection: Stomach misidentification or a case of incidental infection Distribution: Peru – HA and CU (Amazon) (1, in a Tupinambis teguixin (8, 143, 152). 45, 157).

Molineidae Oswaldocruzia vitti Bursey & Goldberg, 2004 Kentropyxia Baker, 1982 Hosts: Alopoglossus angulatus, A. atriventris, Kentropyxia sauria Baker, 1982 Anolis fuscoauratus, A. punctatus, Cercosaura Hosts: Kentropyx calcarata eigenmanni, C. oshaugnessyi Site of infection: Small intestine Site of infection: Intestine Distribution: Brazil – Belém and Novo Progresso Distribution: Brazil – AC, AM, PA, RO (Amazon); (Amazon), PA (136, 153). Ecuador – SU (Amazon); Peru – CU (Amazon) (1, 18, 40, 51, 52). Oswaldocruzia Travassos, 1917 Oswaldocruzia bainae Ben-Slimane & Durette- Oswaldocruzia sp. Desset, 1996 Hosts: Tropidurus torquatus, Enyalius catenatus Hosts: Anolis chrysolepis Duméril & Bibron, 1837; (Wied, 1821), E. perditus, K. calcarata A. fuscoauratus Site of infection: Intestine Site of infection: Small intestine Distribution: Brazil – Bodoquena (Cerrado)

J Venom Anim Toxins incl Trop Dis | 2010 | volume 16 | issue 4 555 Ávila RW, Silva RJ. Checklist of helminths lizards and amphisbaenians (Reptilia, Squamata) of South America and Salobra (Pantanal), MS; Juara (Amazon), Oochoristica travassosi Rego & Ibañez, 1965 MT; Cachimbo (Amazon), PA; Rio de Janeiro, Hosts: Leiocephalus sp.; Liolaemus vallecurensis RJ; Ibitipoca, MG; Argentina – SA, JU and TA Pereira, 1992; Phyllodactylus johnwrigthi Dixon (Chaco); Paraguay – Assuncion and Chaco. & Huey, 1970 Comments: The aforementioned records of Site of infection: Small intestine T. torquatus are those of O. mazzai and that of Distribution: Peru – Moche; Argentina – SJ (2, Enyalius spp. are from O. subauricularis, both 97, 167). considered species inquirenda (48, 131, 155, 158). Oochoristica insulamargaritae López-Neyra and Cestoda Diaz-Ungría, 1957 Hosts: Ameiva ameiva Linstowiidae Site of infection: Small intestine Oochoristica Lühe, 1898 Distribution: Venezuela – Isla Margarita (163, Oochoristica ameivae (Beddard, 1914) 168). Hosts: Ameiva ameiva, Cnemidophorus nativo, Mabuya agilis, M. macrorhyncha Oochoristica vanzolinii Rego & Rodrigues, 1965 Site of infection: Intestine Hosts: Hemidactylus mabouia, Eurolophosaurus Distribution: Brazil – Cachimbo (Amazon), nanuzae PA; Serra do Navio (Amazon), AP; Manaus Site of infection: Small intestine (Amazon), AM; Guaratiba and Trancoso Distribution: Brazil – Rio de Janeiro, RJ; Serra do (Restinga), BA; Praia das Neves (Restinga), ES; Cipó, MG (66, 162). Grussaí (Restinga), RJ; Peru – CU (Amazon) (1, 17, 23, 77, 159-161). Oochoristica sp. Hosts: Alopoglossus atriventris, Amphisbaena Oochoristica bressalui Fürhmann, 1927 ridleyi, Mabuya frenata, Tupinambis merianae, Hosts: Tropidurus guarani, T. hispidus, T. torquatus Trachylepis atlantica Site of infection: Small intestine Site of infection: Small intestine Distribution: Brazil – Salvador and Serrinha, BA; Distribution: Ecuador – SU; Brazil – Valinhos, Cachimbo (Amazon), Arraial do Cabo, Cabo SP; Fernando de Noronha Islands, PE (34, 46, Frio and Rio de Janeiro, RJ; Bolivia – El Carmen; 52). Paraguay (19, 65, 160, 162). Semenoviella Spasskii, 1951 Oochoristica freitasi Rego & Ibañez, 1965 Semenoviella amphisbaenae Rudolphi, 1819 Hosts: Dicrodon heterolepis; D. guttulatum Hosts: Linnaeus, 1758; A. Duméril & Bibron, 1893 alba Site of infection: Small intestine Site of infection: Intestine Distribution: Peru – Trujillo, LL; TU, PI. Distribution: Brazil – Belém (Amazon), PA; Comments: Bursey and Goldberg (163) and Pirassununga (Cerrado), SP (169). Arizmendi-Espinosa et al. (164) apparently ignored the presence of O. freitasi in their Mesocestoididae findings of Oochoristica spp. infecting lizards in Mesocestoides Vaillant, 1863 neotropical regions. The species is not included Mesocestoides sp. in any synonymy of the revisions made by Bursey Hosts: Anolis transversalis et al. (165). Moreover, the species was cited in Site of infection: Not informed Guillén-Hernández et al. (166). Distribution: Brazil – RO (Amazon) (40).

Oochoristica iguanae (Baylis, 1919) Bursey & Proteocephalidea Goldberg, 1996 Proteocephalidae Hosts: Iguana iguana, Dicrodon guttulatum Cairaella Coquille & De Chambrier, 2008 Site of infection: Small intestine Cairaella henrii Coquille & De Chambrier, 2008 Distribution: Venezuela – Isla Margarita; Peru – Hosts: Anolis trachyderma Cope, 1876 LA, PI (92,163). Site of infection: Intestine

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Distribution: Ecuador – San Pablo de Kantesyia, Brachycoelium (Dujardin, 1845) SU (170). Brachycoelium salamandrae (Frolich, 1789) Dujardin, 1845 Ophiotaenia La Rue, 1911 Hosts: Leposoma osvaldoi Ávila-Pires, 1995; Ophiotaenia flava Rudin, 1917 Anolis nitens Wagler, 1830 Hosts: Kentropyx pelviceps Site of infection: Small intestine Site of infection: Not informed Distribution: Brazil – Novo Progresso (Amazon), Distribution: Peru – CU (Amazon) (1). PA. Comments: Cited as B. mesocoeliiformis when Ophiotaenia nicoleae Coquille & De Chambrier, found infecting Anolis scypheus (A. nitens) (6, 2008 137). Hosts: Thecadactylus solimoensis Site of infection: Intestine Platynosomum Distribution: Ecuador – San Pablo de Kantesyia, Platynosomum sp. Looss, 1906 SU. Hosts: Amphisbaena ridleyi, Trachylepis atlantica Comments: Thecadactylus was recently subjected Site of infection: Gallbladder, bile duct to revision, and a T. solimoensis was found in from Distribution: Brazil – Fernando de Noronha localities that encompass San Pablo de Kantesyia, Island, PE (46). Ecuador (113, 170). Harmotrematidae Ophiotaenia sp. Helicotrema Odhner, 1912 Hosts: Anolis fuscoauratus Helicotrema asymetricum (Travassos, 1922) Site of infection: Small intestine Viana, 1924 Distribution: Brazil – Santarém (Amazon), PA Hosts: Iguana iguana (51). Site of infection: Intestine Distribution: Brazil – Pantanal (174, 175). Tejidotaenia Freze, 1965 Tejidotaenia appendiculata (Baylis, 1947) Helicotrema magniovatum Odhner, 1912 Hosts: Tupinambis teguixin Hosts: Iguana iguana Site of infection: Small intestine Site of infection: Intestine Distribution: Surinam – PR (Amazon); Brazil – Distribution: Brazil (6). Serra do Navio (Amazon), AP; Linhares, ES (28, 171). Helicotrema spirale (Diesing, 1850) Odhner, 1912 Hosts: Iguana iguana Trematoda Site of infection: Intestine Distribution: Brazil (6). Echinostomata Echinostomida Pronocephalidae Superfamily Echinostomatoidea Looss, 1899 Iguanacola Gilbert, 1938 Iguanacola navicularis Gilbert, 1938 Cathaemasiidae Hosts: Amblyrhynchus cristatus Bell, 1825 Pulchrossomoides Freitas & Lent, 1937 Site of infection: Not informed Pulchrosomoides elegans Freitas & Lent, 1937 Distribution: Ecuador – Galapagos Islands (14). Hosts: Tupinambis teguixin, Iguana iguana, Mabuya macrorhyncha Myosaccus Gilbert, 1938 Site of infection: Stomach Myosaccus amblyrhynchi Gilbert, 1938 Distribution: Brazil – Porto Esperança (Pantanal), Hosts: Amblyrhynchus cristatus MS; Praia das Neves (Restinga), ES (6, 23, 172, Site of infection: Not informed 173). Distribution: Ecuador – Galapagos Islands (14).

Plagiorchiida Cetiosaccus Gilbert, 1938 Dicrocoeliidae Cetiosaccus galapagensis Gilbert, 1938

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Hosts: Amblyrhynchus cristatus Site of infection: Small intestine Site of infection: Not informed Distribution: Brazil – Conceição da Barra, ES Distribution: Ecuador – Galapagos Islands (14). (19).

Paradistomum Kossack, 1910 Plagiorchis vicentei Rodrigues, 1994 Paradistomum parvissimum (Travassos, 1918) Hosts: Hemidactylus mabouia Hosts: Ameiva ameiva, Hemidactylus mabouia, Site of infection: Small intestine Iguana iguana, Liolaemus lutzae, Mabuya Distribution: Brazil – Teresópolis, RJ (179). macrorhyncha, M. agilis, Tropidurus torquatus, Tupinambis teguixin Family Brachycoeliidae Site of infection: Gallbladder, liver, small intestine Mesocoelium Odhner, 1910 Distribution: Brazil – Arraial do Cabo, Grussaí, Mesocoelium monas (Rudolphi, 1819) Freitas, Nova Iguaçu, Manguinhos, Maricá, RJ; Praia das 1958 Neves (Restinga) and Santa Teresa, ES; Salvador Hosts: Alopoglossus angulatus, Amphisbaena sp., and Trancoso (Restinga), BA (17, 19, 21, 23, 27, Amphisbaena ridleyi, Cercosaura eigenmanni, 100, 146, 176, 177). Diploglossus lessonae, Trachylepis atlantica, Leposternon microcephalum, Tropidurus Paradistomum rabusculum Kossack, 1910 torquatus, Uranoscodon superciliosus Hosts and records: Gymnodactylus geckoides Spix, Site of infection: Intestine 1825 Distribution: Brazil – PA, RO, RR (Amazon); João Site of infection: Not informed Alfredo (Caatinga) and Fernando de Noronha, Distribution: Brazil (6). PE; Maricá and Rio de Janeiro (Restinga), RJ; Ecuador – SU (Amazon) (6, 18, 46, 50, 52, 100). Macroderidae Pneumotrema Bhalerao, 1937 Family Urotrematidae Pneumotrema travassosi Bhalerao, 1937 Urotrema Braun, 1900 Hosts and records: Amphisbaena alba Urotrema shirleyae Zamparo, Brooks & Tkach, Site of infection: Kidney, ureter, intestine. 2005 Distribution: Brazil (172). Hosts: Anolis fuscoauratus Site of infection: Small intestine Family Plagiorchiidae Distribution: Brazil – Santarém (Amazon), PA Allopharynx (Strom, 1928) (51). Allopharynx daileyi Bursey, Goldberg & Vitt, 2005 Hosts: Uranoscodon superciliosus HOST-PARASITE LIST Site of infection: Small intestine Distribution: Brazil – PA, RO, RR (Amazon) (50). Family Amphisbaena alba Dasymetra Nicoll, 1911 Maracaya belemensis Dasymetra tupinambis Nasir & Diaz, 1971 Maracaya graciai Hosts: Tupinambis teguixin Aplectana albae Site of infection: Intestine Aplectana raillieti Distribution: Venezuela – Sucre (178). Physaloptera retusa Semenoviella amphisbaenae Styphlodora Looss, 1899 Pneumotrema travassosi Styphlodora condita Faria, 1911 Hosts: Tupinambis rufescens (Günther, 1871) Amphisbaena bolivica Site of infection: Kidney, ureter, intestine Aplectana tucumanensis Distribution: Argentina (9). Amphisbaena fuliginosa Plagiorchis Lühe, 1899 Semenoviella amphisbaenae Plagiorchis freitasi Vicente, 1978 Hosts: Tropidurus torquatus Amphisbaena ridleyi

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Aplectana albae Raillietnema brachyspiculatum Parapharyngodon alvarengai Mesocoelium monas Family Platynosomum sp. Oochoristica sp. Rhabdias sp.

Amphisbaena sp. Anolis chrysolepis Maracaya pusilla Oswaldocruzia bainae Mesocoelium monas Anolis fuscoauratus Leposternon microcephalum Cosmocerca vrcibradici Cosmocerca rara Strongyluris oscari Mesocoelium monas Rhabdias sp. Paradollfusnema amphisbaenia Physaloptera retusa Oswaldocruzia bainae Family Oswaldocruzia vitti Amblyrhynchus cristatus Ophiotaenia sp. Iguanacola navicularis Urotrema shirleyae Myosaccus amblyrhynchi Cetiosaccus galapagensis Anolis nitens Brachycoelium salamandrae Iguana iguana Alaeuris caudatus Anolis punctatus Alaeuris vogelsangi Cyrtosomum longicaudatum Ozolaimus cirratus Strongyluris oscari Ozolaimus megatyphlon Falcaustra sp. Oswaldofilaria brevicaudata Skrjabinellazia intermedia Physaloptera retusa Rhabdias anolis Oochoristica iguanae Rhabdias elegans Pulchrosomoides elegans Rhabdias sp. Paradistomum parvissimun Oswaldofilaria brevicaudata Helicotrema asymetricum Piratuba lainsoni Helicotrema magniovatum Physaloptera retusa Helicotrema spirale Oswaldocruzia peruensis Oswaldocruzia vitti Conolophus subcristatus Alaeuris conolophi Anolis trachyderma Alaeuris galapagensis Cairaella henrii Alaeuris labicula Alaeuris longispicula Anolis transversalis Paralaeuris dorochila Cyrtosomum longicaudatum Strongyluris oscari Family Mesocestoides sp. Enyalioides praestabilis Spinicauda spinicauda Polychrus acutirostris Gynaecometra bahiensis Hoplocercus spinosus Physaloptera sp. Africana dardanelosi Cruzia rudolphii Polychrus marmoratus Physaloptera sp. Pseudostrongyluris polychrus Oswaldofilaria azevedoi Morunasaurus annularis Piratuba lainsoni

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Family Spauligodon loboi Enyalius bilineatus Centrorhynchidae Rhabdias sp. Spauligodon maytacapaci Acuariidae Physaloptera lutzi Liolaemus lutzae Physaloptera retusa Ichthyocapillaria (= Pseudocapillaria) maricaensis Oswaldocruzia benslimanei Parapharyngodon sceleratus Pharyngodon cesarpintoi Enyalius catenatus Physaloptera retusa Oswaldocruzia sp. Paradistomum parvissimum

Enyalius iheringii Liolaemus neuquensis Strongyluris oscari Physaloptera retusa Rhabdias sp. Oswaldocruzia fredi Liolaemus ornatus Spauligodon loboi Enyalius perditus Physaloptera lutzi Acanthocephalus sp. Aplectana vellardi Cosmocerca sp. Spauligodon maytacapaci Strongyluris oscari Rhabdias sp. Liolaemus quilmes Oswaldocruzia burseyi Spauligodon loboi Oswaldocruzia sp. Physaloptera lutzi

Leiosaurus belli Liolaemus ramirezae Physaloptera retusa Spauligodon loboi

Leiosaurus catamarcensis Liolaemus signifer Physaloptera retusa Parapharyngodon senisfaciecaudus

Family Liolaemidae Liolaemus alticolor Spauligodon maytacapaci Physaloptera lutzi Liolaemus vallecurensis Liolaemus andinus Oochoristica travassosi Spauligodon maytacapaci Phymaturus palluma Liolaemus buergeri Parapharyngodon riojensis Parapharyngodon riojensis Phymaturus punae Liolaemus capillitas Parapharyngodon riojensis Spauligodon loboi Family Liolaemus chilensis Eurolophosaurus nanuzae Spauligodon maytacapaci Strongyluris oscari Subulura lacertília Liolaemus elongatus Parapharyngodon sceleratus Spauligodon maytacapaci Physaloptera lutzi Oochoristica vanzolinii Liolaemus huacahuasicus

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Leiocephalus sp. Physaloptera lutzi Spauligodon maytacapaci Physaloptera retusa Oochoristica travassosi Oochoristica bressalui

Microlophus albermalensis Tropidurus hispidus Parapharyngodon sceleratus Parapharyngodon sceleratus Pharyngodon sp. Microlophus occipitalis Rhabdias sp. Thubunaea parkeri Oswaldofilaria petersi Physaloptera retusa Microlophus peruvianus Oochoristica bressalui Parapharyngodon arequipensis Parapharyngodon moqueguensis Tropidurus itambere Parapharyngodon yurensis Parapharyngodon sceleratus Thubunaea iguanae Pharyngodon cesarpintoi Physaloptera lutzi Microlophus thoracicus Thubunaea iguanae Tropidurus melanopleurus Strongyluris oscari Plica plica Parapharyngodon sceleratus Strongyluris oscari Physaloptera retusa Physaloptera retusa Tropidurus semitaeniatus Plica umbra Parapharyngodon sceleratus Strongyluris oscari Hastospiculum sp. Tropidurus spinulosus Piratuba digiticauda Strongyluris oscari Physaloptera retusa Tropidurus torquatus Stenocercus caducus Acanthocephala Skrjabinellazia intermedia Echinorhynchus sp. Strongyluris oscari Centrorhynchus tumidulus Capillaria (= Amphibiocapillaria) freitaslenti Stenocercus roseiventris Hexametra boddaertii Atractis cruciata Strongyluris oscari Strongyluris oscari Skrjabinellazia intermedia Oswaldofilaria azevedoi Subulura sp. Physaloptera retusa Parapharyngodon sceleratus Oswaldocruzia peruensis Parapharyngodon sp. Pharyngodon sp. Tropidurus etheridgei Acuariidae Parapharyngodon sp. Piratuba digiticauda Thelandros sp. Physaloptera lutzi Physaloptera sp. Physaloptera retusa Physaloptera sp. Tropidurus guarani Physalopteroides venancioi Strongyluris oscari Oswaldocruzia sp. Skrjabinellazia intermedia Oochoristica bressalui Typhlonema sp. Paradistomum parvissimum Parapharyngodon sceleratus Plagiorchis freitasi Spauligodon oxkutzcabiensis Mesocoelium monas Piratuba digiticauda

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Tropidurus sp. Phyllodactylus microphyllus Strongyluris oscari Spauligodon oxkutzcabiensis Parapharyngodon sceleratus Phyllodactylus reissii Uracentron flaviceps Spauligodon oxkutzcabiensis Skrjabinodon dixoni Physaloptera retusa Phyllopezus pollicaris Macdonaldius grassi Uranoscodon superciliosus Parapharyngodon sceleratus Cosmocerca vrcibradici Africana chabaudi Thecadactylus solimoensis Allopharynx daileyi Spauligodon oxkutzcabiensis Mesocoelium monas Physaloptera retusa Physalopteroides venancioi Family Ophiotaenia nicoleae Hemidactylus mabouia Centrorhynchidae Family Parapharyngodon largitor Gonatodes humeralis Parapharyngodon sceleratus Skrjabinelazia galliardi Parapharyngodon sp Strongyloides cruzi Family Acuariidae Diploglossus lessonae Physaloptera sp. Parapharyngodon verrucosus Spirurinae Mesocoelium monas Oswaldocruzia brasiliensis Oochoristica vanzolinii Ophiodes striatus Paradistomum parvissimum Cyrtosomum sp. Plagiorchis vicentei Physaloptera retusa

Family Family Bogertia lutzae Ameiva ameiva Spauligodon oxkutzcabiensis Oligacanthorhynchus sp. Centrorhynchus tumidulus Gymnodactylus geckoides Capillaria (= Amphibiocapillaria) freitaslenti Paradistomum rabusculum Spinicauda spinicauda Strongyluris oscari Phyllodactylus angustidigitus Parapharyngodon alvarengai Spauligodon viracochai Parapharyngodon largitor Parapharyngodon sceleratus Phyllodactylus inaequalis Parapharyngodon sp. Spauligodon oxkutzcabiensis Pharyngodon cesarpintoi Pharyngodon travassosi Phyllodactylus gerrhopygus Chabirenia cayennensis Spauligodon viracochai Piratuba scaffi Physaloptera lutzi Phyllodactylus johnwrighti Physaloptera retusa Oochoristica travassosi Physaloptera sp. Parapharyngodon sceleratus Physalopteroides venancioi Spauligodon oxkutzcabiensis Oochoristica ameivae Oochoristica insulamargaritae Phyllodactylus lepidopygus Paradistomum parvissimum Acuariidae

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Ameiva festiva Diaphanocephalus jacuruxi Physaloptera retusa Kentropyx altamazonica Cnemidophorus abaetensis Physaloptera retusa Physaloptera lutzi Physalopteroides venancioi Physaloptera retusa Kentropyx calcarata Cnemidophorus lemniscatus Kentropyxia sauria Physaloptera retusa Piratuba digiticauda Piratuba shawi Cnemidophorus littoralis Physaloptera retusa Hexametra boddaertii Physalopteroides venancioi Parapharyngodon sceleratus Oswaldocruzia sp. Physaloptera lutzPhysaloptera retusa Physaloptera sp. Kentropyx pelviceps Dujardinascaris sp. Cnemidophorus nativo Parapharyngodon sceleratus Skrjabinellazia intermedia Physaloptera retusa Subulura lacertilia Physalopteroides venancioi Parapharyngodon sp. Ophiotaenia flava Physaloptera retusa Physalopteroides venancioi Tupinambis longilineus Oochoristica ameivae Physaloptera retusa

Cnemidophorus ocellifer Tupinambis merianae Hexametra boddaertii Cruzia travassosi Physaloptera retusa Physaloptera sp. Physaloptera sp. Diaphanocephalus galeatus Spinicauda spinicauda Cnemidophorus sp. Oochoristica sp. Pharyngodon cesarpintoi Tupinambis rufescens Dicrodon heterolepis Physaloptera retusa Pharyngodon micrurus Diaphanocephalus galeatus Thubunaea parkeri Styphlodora condita Oochoristica freitasi Tupinambis teguixin Dicrodon guttulatum Centrorhynchus tumidulus Oochoristica freitasi Centrorhynchus sp. Oochoristica iguanae Contracaecum sp. Pharyngodon micrurus Dujardinascaris sp. Parapharyngodon moqueguensis Freitasascaris alata Parapharyngodon lagitor Atractis cruciata Parapharyngodon sp. Spinicauda spinicauda Thelandros capacyupanquii Cruzia fulleborni Physaloptera sp. Cruzia tentaculata Oswaldofilaria petersi Dicrodon holmbergi Oswaldofilaria sp. Thelandros capacyupanquii Physaloptera retusa Physaloptera sp. Dracaena guianensis Physalopteroides venancioi Oswaldofilaria belemensis Diaphanocephalus diesingi

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Diaphanocephalus galeatus Centrorhynchidae Tejidotaenia appendiculata Hexametra boddaertii Pulchrosomoides elegans Strongyluris oscari Paradistomum parvissimum Parapharyngodon largitor Dasymetra tupinambis Parapharyngodon sceleratus Acuariidae Family Physaloptera retusa Alopoglossus angulatus Physaloptera sp. Cosmocerca vrcibradici Physalopteroides venancioi Oswaldocruzia vitti Oochoristica ameivae Mesocoelium monas Paradistomum parvissimum

Alopoglossus atriventris Trachylepis atlantica Cosmocerca vrcibradici Moaciria alvarengai Physalopteroides venancioi Parapharyngodon alvarengai Oswaldocruzia vitti Mesocoelium monas Oochoristica sp. Spinicauda spinicauda Platynosomum sp. Cercosaura argulus Oochoristica sp. Cosmocercidae Acuariidae Mabuya bistriata Physaloptera sp. Parapharyngodon sceleratus Oswaldocruzia peruensis Oswaldofilaria spinosa Piratuboides zeae Cercosaura eigenmanni Physaloptera retusa Cosmocerca vrcibradici Physalopteroides venancioi Oswaldocruzia vitti Mesocoelium monas Mabuya caissara Parapharyngodon sceleratus Cercosaura ocellata Physalopteroides venancioi Mabuya dorsivittata Skrjabinodon spinulosus Cercosaura oshaugnessyi Physaloptera retusa Acanthocephalus saurius Cosmocerca vrcibradici Mabuya frenata Oswaldocruzia vitti Hexametra boddaertii Parapharyngodon sceleratus Leposoma osvaldoi Skrjabinodon heliocostai Brachycoelium salamandrae Oswaldofilaria sp. Oochoristica sp. Neusticurus bicarinatus Falcaustra belemensis Mabuya macrorhyncha Echinorhynchidae Potamites ecpleopus Centrorhynchidae Falcaustra belemensis Hexametra boddaertii Physaloptera retusa Acuariidae Physaloptera sp. Potamites strangulatus Physalopteroides venancioi Falcaustra belemensis Oochoristica ameivae Pulchrosomoides elegans Family Scincidae Paradistomum parvissimum Mabuya agilis

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FINAL REMARKS Improvement of Higher Education Personnel (CAPES). The literature on helminth parasites of lizards and amphisbaenians from South America COPYRIGHT is extensive, dating back to the 1920’s with © CEVAP 2010 the studies by Lauro Travassos. However, the information is scattered, and reviews of helminth SUBMISSION STATUS parasites are necessary because they provide a Received: June 1, 2010. basis for a better understanding on in Accepted: September 16, 2010. these groups for future research (180). The need Abstract published online: September 17, 2010. of updated knowledge becomes clear considering Full paper published online: November 30, 2010. the valuable information that parasites provide because it is synergistic with all the information CONFLICTS OF INTEREST about the natural history of the hosts (181). There is no conflict. Helminths are high diverse in vertebrates, but the species richness are poorly known (182, 183). FINANCIAL SOURCE For example, less than a half of species FAPESP (processes n. 04/03628-1 and 06/59692- are believed to be currently described (148). In 5) and CAPES provided the financial grants. this paper, a total of 644 records of helminths collected from lizards are reported. Since many CORRESPONDENCE TO species descriptions were made in the last decade, ROBSON WALDEMAR ÁVILA, Departamento the diversity of helminth parasites of lizards from de Parasitologia, Instituto de Biociências, UNESP, South America should be increased with further Distrito de Rubião Jr., s/n, CEP 18618-000, studies. Botucatu, SP, Brazil. Email: robsonavila@gmail. Of the 644 records in the present study, at least com. nine species of acantocephalans, 15 cestodes, 20 trematodes and 111 nematodes were reported, REFERENCES including records in which family or genus were 1. Bursey CR, Goldberg SR, Parmelee JR. not identified. From the 155 helminth species Gastrointestinal helminths from 13 species of reported, one was recorded from Chile and lizards from Reserva Cuzco Amazónico, Peru. French Guiana, three were from Colombia, three Comp Parasitol. 2005;72(1):50-68. were from Uruguay, eight were from Bolivia, nine 2. Goldberg SR, Bursey CR, Morando M. Metazoan were from Surinam, 13 were from Paraguay, 12 endoparasites of 12 species of lizards from were from Venezuela, 27 were from Ecuador, 17 Argentina. Comp Parasitol. 2004;71(2):208-14. were from Argentina, 39 were from Peru and 103 3. Vrcibradic D, Anjos LA, Vicente JJ, Bursey CR. were from Brazil. Five species considered species Helminth parasites of two sympatric lizards, Enyalius iheringii and E. perditus (Leiosauridae), inquirenda were also reported. from an Atlantic Rainforest area of southeastern Lizards and amphisbaenians comprises Brazil. Acta Parasitol. 2008;53(2):222-5. approximately 420 genera and more than 3,300 4. Travassos LP. Contribuições para o conhecimento species, most of which can be found in South da fauna helmintológica brasileira. IX. Sobre America (184, 185). Many South American as espécies do gênero Spinicauda. Mem Inst countries exhibit great lizard diversity, for Oswaldo Cruz. 1920;12:41-50. example Argentina presents 167 species while 5. Travassos LP. Pesquizas helminthologicas Brazil has 240 species, but data presented herein realizadas em Hamburgo. IX. Ensaio includes only 114 lizard species, which indicates monographico da familia Cosmocercidae that much work has to be done (186, 187). Travassos, 1925 (Nematoda). Mem Inst Oswaldo Cruz. 1931;25:237-98. ACKNOWLEDGEMENTS 6. Travassos LP, Freitas JFT, Kohn A. Trematodeos Both authors are thankful for the financial support do Brasil. Mem Inst Oswaldo Cruz 1969;67:1- 886. of The State of São Paulo Research Foundation 7. Baker MR. Synopsis of the Nematoda parasitic (FAPESP) and Robson W. Ávila is grateful for in amphibians and reptiles. Mem Univ the grant provided by Coordination for the Newfoundland Occas Pap Biol. 1987;11:1-325.

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de Janeiro state, southeastern Brazil. Amphib- gênero Cosmocerca Diesing, 1861, parasito de Reptil. 2000; 21(3):307-16. anfisbaenideo. Atas Soc Biol. Rio de Janeiro. 36. Barus V. Coy Otero A. Systematic survey of 1966;10:109-11. nematodes parasitizing lizards (Sauria) in Cuba. 50. Bursey CR, Goldberg SR, Vitt LJ. New species Helminthol. 1969;10(3):329-46. of Allopharynx (Digenea: Plagiorchiidae) and 37. Vicente JJ. Redescrição de Atractis cruciata other helminths in Uranoscodon superciliosus Linstow, 1902, em novo hospedeiro (Nematoda, (Squamata: Tropiduridae) from Amazonian Atractoidea). Atas Soc Biol Rio de Janeiro. Brazil. J Parasitol. 2005;91(5):1395-8. 1966;10(2):41-3. 51. Goldberg SR, Bursey CR, Vitt LJ. Helminths 38. Bursey CR, Flanagan JP. Atractis marquezi n. sp. of the brown-eared anole, Norops fuscoauratus (Nematoda: Atractidae) and a revision of Atractis (Squamata, Polychrotidae) from Brazil and Dujardin, 1845, sensu Baker, 1987. J Parasitol. Ecuador, South America. Phyllomedusa. 2002;88:320-4. 2006;5(1):83-6. 39. McAllister CT, Bursey CR, Freed PS. Helminth 52. Goldberg SR, Bursey CR, Vitt LJ. Parasite parasites of selected amphibians and reptiles communities of two lizard species, Alopoglossus from the Republic of Ecuador. Comp Parasitol. angulatus and Alopoglossus atriventris, from 2010;77(1):52-66. Brazil and Ecuador. Herpetol J. 2007;17(4):269- 40. Goldberg SR, Bursey CR, Vitt LJ. Parasites 72. of two lizard species, Anolis punctatus and 53. Baker MR. Dollfusnema amphisbaenia n. gen., n. Anolis transversalis (Squamata: Polychrotidae) sp. (Nematoda: Cosmocercinae) from the lizard from Brazil and Ecuador. Amphib-Reptil. Leposternon phocaena (Amphisbaenia) of Brazil. 2006;27(4):575-9. Can J Zool. 1981;59(1):138-40. 41. Ávila RW, Souza FL, Silva RJ. Helminths from 54. Baker MR. Africana chabaudi n. sp. (Nematoda, seven species of lizards (Reptilia: Squamata) at Heterakidae) in a Brazilian Iguana. Bull Mus Nat the Cerrado of Mato Grosso do Sul State, Brazil. His Nat Paris. 1981;3(3):783-7. Comp Parasitol. 2010;77(1):67-71. 55. Ávila RW, Silva RJ. A new species of Africana 42. Adamson ML, Baccam D. Systematic position of (Nematoda: Heterakidae) from lizards of southern the Atractidae sensu Chabaud (1978) (Nematoda: Amazon, Brazil. J Parasitol. 2009;95(5):1156-8. Cosmocercoidea): Maracaya belemensis n. sp. 56. Freitas JFT. Novo parasito de réptil da Ilha and Aplectana albae n. sp. from Amphisbaena Fernando de Noronha: Moaciria alvarengai g. n., alba in Brazil. Can J Zool. 1988;66:1857-64. sp. n. (Nematoda, Subuluroidea). Rev Bras Biol. 43. Boaumer S, Morand S. A new nematode 1956;16:335-9. (Nematoda: Cosmocercidae) from the 57. Días-Ungría C. Notas sobre nematodes de reptiles lizard, Chamaeleo inturensis (Squamata: en Venezuela. Rev Vet Venez. 1964;17(101):402. Chamaeleonidae) from the Democratic Republic 58. Rodrigues HO, Feijó LMF. Redescrição de of Congo. J Parasitol. 2006;92(2):346-9. Spinicauda spinicauda (Olfers, 1919) Travassos, 44. Ramallo GR, Bursey CR, Goldberg SR. A new 1920 (Nematoda, Oxyuroidea). Atas Soc Biol Rio species of Cosmocercidae (Ascaridida) in the de Janeiro.1976; 18:59-63. worm lizard, Amphisbaena bolivica (Squamata: 59. Travassos LP. Informações sobre a fauna Amphisbaenidae), from Argentina. J Parasitol. helminthologica de Mato Grosso. Folha Med. 2008;94(6):1361-3. 1923;4:58-60. 45. Goldberg SR, Bursey CR. Cercosaura argulus 60. Pereira C. Os Oxyurata parasitos de Lacertilia do endoparasites. Herpetol Rev. 2007;38(4):451. nordeste Brasileiro. Arch Inst Biol. 1935;6:5-27. 46. Ramalho ACO, Silva RJ, Schwartz H, Péres 61. Bursey CR, Goldberg SR, Telford Jr SR. AK. Helminths from an introduced species Strongyluris panamaensis n. sp. (Nematoda: (Tupinambis merianae), and two endemic species Hheterakidae) and other helminths from the (Trachylepis atlantica and Amphisbaena ridleyi) lizard, Anolis biporcatus (Sauria: Polychrotidae), from Fernando de Noronha archipelago, Brazil. J from Panama. J Parasitol. 2003;89(1):118-23. Parasitol. 2009;95(4):1026-8. 62. Vicente JJ. Helmintos de Tropidurus (Lacertilia, 47. Travassos LP. Aplecana raillieti, n. sp. C R Seanc Iguanidae) da coleção helmintólogica do Instituto Soc Biol. 1925;93:973-4. Oswaldo Cruz. II. Nematoda. Atas Soc Biol Rio 48. Sousa BM, Lima SS, Oliveira A. Gastrointestinal de Janeiro. 1981;22:7-18. helminth fauna of Enyalius perditus (Reptilia: 63. Harvey MB, Gutberlet Jr RL. Lizards of the Leiosauridae): Relation to host age and sex. J genus Tropidurus (Iguania: Tropiduridae) Parasitol. 2007; 93(1):211-3. from the serrania de Huanchaca, Bolivia: New 49. Freitas JFT, Vicente JJ. Nôvo nematódeo do species, natural history, and a key to genus.

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Biología. 2005;12(3):463-72. Reptil.1995;16(4):412-5. 91. Rodrigues HO, Pinto RM. Ocorrência de dois 104. Alho CJR, Moura VP. Oxyurata de faringodonídeos em Ameiva ameiva (L.) no lagartos do Planalto Central. Ocorrência estado da Guanabara (Nematoda, Oxyuroidea). de Pharyngodon cesarpintoi Pereira, 1935 Atas Soc Biol Rio de Janeiro.1967;11(3):123-5. (Nematoda, Subuluroidea). Atas Soc Biol Rio de 92. Vásquez PTS. Helmintos parásitos de Dicrodon Janeiro.1970;13(1-2):67-9. guttulatum, Dúmeril y Bibron, 1893 (SAURIA: 105. Freitas JFT, Ibáñez N. Fauna helmintológica del TEIIDAE) de la costa del Perú [dissertation]. Perú – Notas sobre “Pharyngodonidae” Travassos, Lima, Peru: Facultad Ciencias Biológicas, 1920 y descripción de Pharyngodon micrurus sp. Universidad Nacional Mayor de San Marcos; n. Rev. Universitaria 1963; 23 y 24. 2008. 63p. 106. Bursey CR, Goldberg SR. New species of 93. Alho CJR, Rodrigues HO. Sobre novo hospedeiro Skrjabinodon (Nematoda: Pharyngodonidae) in de Parapharyngodon sceleratus (Travassos, Uracentron flaviceps (Squamata: Iguanidae) from 1923) Freitas, 1957, com redescrição da espécie Ecuador and Peru. J Parasitol. 2007;93(4):866-9. (Nematoda, Oxyuroidea). Atas Soc Biol Rio de 107. Vicente JJ, Vrcibradic D, Muniz-Pereira LC, Pinto Janeiro.1963; 7:4-6. PM. Skrjabinodon heliocostai sp. n. (Nematoda, 94. Ribas SC, Rocha CFD, Teixeira-Filho PF, Pharyngodonidae) parasitizing Mabauya frenata Vicente JJ. Helminths (Nematoda) of the lizard (Cope) (Lacertilia, Scincidae) in Brazil and the Cnemidophorus ocellifer (Sauria: Teiidae): reallocation of Skrjabinodon capacyupanquii assessing the effect of rainfall, body size and (Freitas, Vicente & Ibanez) in the genus Thelandros sex in the nematode infection rates. Ciênc Cult. Wedl. Rev Bras Zool. 2000; 17:361-367. 1995;47(1-2):88-91. 108. Rocha CFD, Vrcibradic D, Vicente JJ, Cunha- 95. Van Sluys M, Rocha CFD, Ribas SC. Barros M. Helminths infecting Mabuya Nematodes infecting the lizard Tropidurus dorsivittata (Lacertilia, Scincidae) from a high- itambere in southeastern Brazil. Amphib- altitude habitat in Itatiaia National Park, Rio de Reptilia.1994;15(4):405-8. Janeiro State, southeastern Brazil. Braz J Biol. 96. Van Sluys M, Rocha CFD, Bergallo HG, 2003;63(1):129-32. Vrcibradic D, Ribas SC. Nematode infection in 109. Vicente JJ, Vrcibradic D, Rocha CFD, Pinto RM. three sympatric lizards in an isolated fragment of Description of Skrjabinodon spinosulus sp. n. restinga habitat in southeastern Brazil. Amphib- (Nematoda, Oxyuroidea, Pharyngodonidae) from Reptil.1997;18(4):442-6. the Brazilian lizard Mabuya dorsivittata Cope, 97. Goldberg SR, Bursey CR. Helminths from four 1862 (Scincidae). Rev Bras Zool. 2002;19(1):157- species of Phyllodactylus (Squamata: Gekkonidae) 62. from Peru. Comp Parasitol. 2010;77(1):91-3. 110. Ramallo GR, Bursey CR, Goldberg SR. Spauligodon 98. Freitas JFT. Sobre um novo nematódeo de reptil loboi n. sp. (Nematoda: Pharyngodonidae) da Bolivia: Parapharyngodon senisfaciecauduts parasite of Liolaemus spp. (Iguania: Liolaemidae) sp. n. (Nematoda, Oxyuroidea). Rev Bras Biol. from northwestern Argentina. J Parasitol. 1957;17:451-4. 2002;88(4):370-4. 99. Freitas JFT, Dobbin Jr, JE. Nova espécie do gênero 111. Vicente JJ, Ibañez HN. Nova espécie Parapharyngodon Chatterji, 1933 (Nematoda, do gênero Parathelandros Baylis, 1930 Oxyuroidea). An Soc Biol Pernambuco. (Nematoda, Oxyuroidea). Atas Soc Biol Rio de 1959;16:23-33. Janeiro.1968;11:185-7. 100. Rodrigues HO, Rodrigues SS, Faria Z. 112. Goldberg SR, Muñoz G, Garcias F, Bursey CR. Contribution to the knowledge of the Liolaemus lemniscatus (wreath tree iguana), helminthological fauna of vertebrates of Maricá, Liolaemus tenuis (thin tree iguana). Endoparasites. Rio de Janeiro State, Brazil. Mem Inst Oswaldo Herpetol Rev. 2001;32:41. Cruz.1990;85(1):115-6. 113. Bergmann PJ, Russell AP. Systematics and 101. Zaracho VH, Lamas MF. Tropidurus etheridgei. biogeography of the widespread Neotropical Endoparasites. Natural History. Herpetol Rev. gekkonid genus Thecadactylus (Squamata), with 2006; 37:473. the description of a new cryptic species. Zool J 102. Cruz FB, Silva S, Scrocchi GJ. Ecology of the lizard Linnean Soc. 2007;149:339-70. Tropidurus etheridgei (Squamata: Tropiduridae) 114. Goldberg SR, Bursey CR. Phyllodactylus reissi from the dry Chaco of Salta, Argentina. Herpetol Endoparasites. Herpetol Rev. 2004; 35:395. Nat Hist. 1998;6(1):23-31. 115. Ávila RW, Anjos LA, Gonçalves U, Freire EMX, 103. Rocha CFD. Nematode parasites of the Brazilian Almeida WO, Silva RJ. Nematode infection sand lizard, Liolaemus lutzae. Amphib- in the lizard Bogertia lutzae (Loveridge, 1941)

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