Introduction
The Neotropical region includes most of Latin America – central and southern Mexico, the Antilles and almost all of South America (Cabrera and Willink Reference Cabrera and Willink1973; Morrone Reference Morrone2015). It is composed of three subregions (Antilles, Brazilian, and Chaco), seven dominions (Mesoamerican dominion, Pacific dominion, Brazilian boreal dominion, Southern Brazilian dominion, Chaco dominion, Paraná dominion, and Southeastern Amazon dominion), two transition zones (Mexican and South American), and 57 provinces (Morrone Reference Morrone2015; Morrone et al. Reference Morrone, Escalante, Rodríguez-Tapia, Carmona, Arana and Mercado-Gómez2022). It also contains several genera and suprageneric groups of endemic taxa, with the largest number of flora and fauna in the world (Morrone Reference Morrone2017; Tundisi and Tundisi Reference Tundisi and Tundisi2008). It has the highest species richness of reptiles on the planet (Silva Reference Silva2014; Uetz et al. Reference Uetz, Freed and Hošek2021), which have a high potential for harboring large groups of parasites due to their significant diversity of prey (Taylor et al. Reference Taylor, Coop and Wall2017). There are eight families of terrestrial and freshwater chelonians in the Neotropical region: Chelidae, Chelydridae, Dermatemydidae, Emydidae, Geoemydidae, Kinosternidae, Podocnemididae, and Testudinidae, with the Chelidae family being the most diverse.
Parasites are of great importance to aquatic ecosystems, as they are considered hidden and essential elements of communities (Poulin Reference Poulin1999), participating in the energy flow (Pavanelli et al. Reference Pavanelli, Takemoto and Eiras2013; Thomas et al. Reference Thomas, Renaud, Rousset, Cezilly and De Meeûs1995) and shaping their host communities (Gregory et al. Reference Gregory, Keymer and Harvey1996; Poulin and Morand Reference Poulin and Morand1999). Their relationships with their hosts are crucial for understanding the dynamics of communities and the species that comprise them. (Gregory et al. Reference Gregory, Keymer and Harvey1996; Poulin and Morand Reference Poulin and Morand1999). Regarding the continental chelonians in South America, trematodes and nematodes are the most frequent, with 40 and 38 species recorded, respectively (Mascarenhas and Müller Reference Mascarenhas and Muller2021), as well as monogeneans and cestodes; the parasites are primarily found in the gastrointestinal tract, where they exhibit greater parasitism (Cubas et al. Reference Cubas, Silva and Dias2006; George Reference George, Music and Lutz1997; Mascarenhas et al. Reference Mascarenhas, Souza, Coimbra and Müller2013; Wilson and Carpenter Reference Wilson and Carpenter1996).
Surveying the parasite fauna is important for understanding biodiversity and its distribution (González et al. Reference González, Hamann and Duré2021), helping to illuminate the gaps in knowledge about parasites and their hosts. For chelonians, in the Neotropical region, only marine species have been reviewed for parasites by Werneck (Reference Werneck and Da Silva2016); for South America, in continental chelonians, more recently by Mascarenhas and Müller (Reference Mascarenhas and Muller2021) and by Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022) with new records for Peru; in Mexico, among reptiles, helminths parasitizing chelonians were mentioned by Paredes-León (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008), and in Costa Rica, helminths of chelonians were cited by Rodriguez-Ortiz et al. (Reference Rodriguez-Ortiz, Garcia-Prieto and Perez-Ponce de Leon2004). The aim of this study is to survey the literature on parasitic helminths (Nematoda, Trematoda, Monogenea, Acanthocephala, and Cestoda) associated with terrestrial and freshwater chelonians in the Neotropical region, seeking to compile the individuals reported both for their biogeographical distribution, as well as for the political units in which the Neotropical region is situated, verifying the main groups and regions that present helminths and adding a new record for the species Phrynops geoffroanus in the Paraíba hinterland, in northeastern Brazil.
Material and methods
List of helminth parasites of terrestrial and freshwater chelonians in the Neotropical region
To survey on helminths (Nematoda, Trematoda, Monogenea, Acanthocephala, and Cestoda) parasitizing terrestrial and freshwater chelonians in the Neotropical region, an extensive search was conducted of published records, including articles, books, monographs, and theses, using the search tools on the following websites: Scopus, SciELO, Web of Science, and Google Scholar. The key terms used were chelonians, parasites, nematoda, Neotropical region, ‘check list’, as well as terms for each Latin American country, host species, and family (e.g., ‘Phrynops geoffroanus and Brazil and nematoda’), considering the Neotropical region and transition areas, according to Morrone et al. (Reference Morrone, Escalante, Rodríguez-Tapia, Carmona, Arana and Mercado-Gómez2022).
A map of the Neotropical region was created in the Qgis 3.36.3 programme (QGIS Development Team 2021), incorporating the shapfile from the Morrone et al. (Reference Morrone, Escalante, Rodríguez-Tapia, Carmona, Arana and Mercado-Gómez2022) map, using the geographical coordinates available in each source. When these were unavaiable, the municipal coordinates were utilized. If this information was not provided, nor a precise location given, those entries were excluded from the occurrence map. Abbreviations were used for each biogeographic unit (dominion/region): Mexican transition zone (MTZ), Antillean subregion (AS), Mesoamerican dominion (MD), Pacific dominion (PCD), Boreal Brazilian dominion (BBD), South Brazilian dominion (SBD), Southeastern Amazonian dominion (SAD), Chacoan dominion (CD), Paraná dominion (PD), and South American transition zone (STZ). Furthermore, the taxonomic nomenclature used for the parasites in this study was in accordance with the WoRMS Editorial Board (2024); when it was not, classifications were made according to Moravec (Reference Moravec and Vargas-Vázquez1998), Anderson et al. (Reference Anderson2000) and Gibbons (Reference Gibbons2010), with modifications based on recent records.
The distribution of chelonian species was checked to identify those occurring in the Neotropical region, their taxonomic classification, and current status on the IUCN Red List of Threatened Species according to the Turtle Taxonomy Working Group (2021). The list of helminths and their hosts is organized in alphabetical order, detailing their classification and information on the site of infection, host, its biogeographical unit (province and dominion/region abbreviation), political unit (in order of country-state/province-city/location), and references. When more than one individual or reference is listed, hosts are marked with a superscript number for their location and reference.
Nematodes of Phrynops geoffroanus in the Brazilian semi-arid region
In order to add new records for the host, we utilized data collected in the field from a sampling of specimens conducted in November and December 2019 and December 2020 (SISBIO License: 70824-2), in urban stretches of the Piancó River in the city of Conceição (stretch 1: 7°33’44.3"S 38°30’27.4"W, and stretch 2: 7°34’00.9"S 38°30’08.9"W) and the Espinharas River in the municipality of Patos (stretch 1: 7°02’18.1"S 37°16’36. 8"W, and stretch 2: 7°01’59.2’S 37°16’25.2"W), in the hinterland of Paraíba. The specimens were captured using funnel traps, separated by sex according to morphological characteristics, insensitized using a container with water and ice for 20 minutes at a temperature of approximately 5ºC, then euthanized using propofol, fixed in 4% formalin, preserved in 70% alcohol, and transported in thermal boxes to the Hydrology, Microbiology and Parasitology Laboratory (LAHMP) at the Universidade Federal da Paraíba in João Pessoa. The research was approved by the Animal Use Ethics Committee of the Universidade Federal da Paraíba (CEUA/UFPB: no. 5117180719).
Parasitological analysis
The animals underwent a parasitological necropsy according to Work (Reference Work2000). Each organ was subsequently cut into parts and examined under a stereomicroscope. The endoparasites found were quantified, placed in containers with 70% ethanol, and labeled. Temporary slides were then mounted using lactic acid to lighten the nematodes and identify them using an optical microscope. The nematodes were identified according to Moravec (Reference Moravec and Vargas-Vázquez1998), Anderson (Reference Anderson2000) and Anderson et al. (Reference Anderson, Chabaud and Willmott2009), and added to the Invertebrate Collection Paulo Young (CIPY) of the Universidade Federal da Paraíba. The chelonians were preserved, cataloged, and added to the Herpetological Collection of the Universidade Federal da Paraíba (CHUFPB). Prevalence rates, mean intensity, mean infection abundance, and standard deviation were calculated using Quantitative Parasitology - Qpweb (Reiczigel et al. Reference Reiczigel, Marozzi, Fabian and Rózsa2019).
Results
Helminth parasites of terrestrial and freshwater chelonians in the Neotropical region
The bibliographical survey recovered 202 helminth taxa parasitizing terrestrial and freshwater chelonians in the Neotropical region: 98 nematode taxa, belonging to 18 families and 41 genera, with 72 taxa identified at the species level, 23 at the genus level, and three at the family level; 78 trematode taxa, belonging to 18 families and 33 genera, with 72 taxa identified at the species level, five at the genus level, and one at the family level; 18 monogenean taxa belonging to two families and five genera, with one taxa identified at the species level and one at the genus level; five taxa of acanthocephalans belong to two families and two genera, with four taxa identified at the species level and one at the genus level and three cestode taxa belonging to one family and one genus, with two taxa identified at the species level and one at the genus level. Nematoda had the highest number of taxa and occurred in all biogeographical units, with the second most diverse family of parasitic helminths, Atractidae (28 species), represented by the most commonly found species, Spiroxys contortus of the Gnathostomatidae family, in six hosts; Trematoda had the second highest number of taxa, featuring the most diverse family of parasitic helminths, Telorchiidae (29 species), represented by the most commonly found species, Nematophila grandis of the Cladorchiidae family, in 17 hosts; Acanthocephala was represented by the species Neoechinorhynchus emyditoides in two hosts; Cestoda by Ophiotaenia cohospes in one host, and Monogenea by Manotrema brasiliensis in three hosts.
A total of eight families, 18 genera, and 48 species of chelonians were recorded as being parasitized by helminths. Of these, 19 species are listed as Least Concern, nine as Vulnerable, nine as Lower Risk/near threatened, three Critically Endangered, three Endangered, and one Extinct in the IUCN Red List (Supplementary material 1). The family with the highest number of helminths was Chelidae (59 taxa), followed by Kinosternidae (52 taxa) and Podocnemididae (40 taxa) (Figure 1). In addition, Chelidae was the only family with all the helminth groups and the highest number of host species (14 hosts), followed by Kinosternidae (nine hosts), Emydidae and Geoemydidae (eight hosts). Nematodes and trematodes were present in all the chelonian families, with Testudinidae having the highest number of nematodes (32 taxa) and Chelidae the highest number of trematodes (27 taxa). Chelidae, Emydidae, and Kinosternidae were the only families to have acanthocephalans. Among the host genera, Kinosternon from the Kinosternidae family was the most parasitized (47 taxa), followed by Phrynops from the Chelidae family (39 taxa) and Podocnemis from the Podocnemididae family (37 taxa). The most parasitized species were the terrestrial chelonian Chelonoidis denticulatus and the freshwater chelonian Phrynops hilarii (25 taxa).
Number of records of helminth parasites per family of chelonians in the Neotropical region.

Helminth records in the biogeographical units of the Neotropical region
All ten biogeographic units recorded helminths associated with terrestrial and freshwater chelonians (Figures 2 and 3): Mesoamerican dominion showed the highest number of helminths (61 taxa), followed by Chacoan dominion (57 taxa), Pacific dominion (41 taxa), Paraná dominion (35 taxa), Boreal Brazilian dominion (30 taxa), South Brazilian dominion (27 taxa), Mexican transition zone (13 taxa), South American transition zone (six taxa), Antillean subregion (five taxa), and South Amazonian dominion (three taxa). Chacoan dominion recorded all the helminth groups and more nematodes, while the Mesoamerican dominion recorded more trematodes. Of the 57 provinces, 32 had records, with the Pampean province having the most helminth taxa (36 taxa) followed by Veracruzan province (35 taxa).
Number of records of helminth parasites of chelonians in the biogeographical units of the Neotropical region.

Map of records of helminth parasites of chelonians in the biogeographical units of the Neotropical region.

Helminth records in the political units encompassed by the Neotropical region
Considering the political units encompassed by the Neotropical region, 15 have studies on helminths of terrestrial and freshwater chelonians (Figure 4): Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, Grenada (island country), France-French Guiana, Mexico, Panama, Paraguay, Peru, Trinidad and Tobago (island country), Uruguay, and Venezuela. Brazil had the highest number of helminths (85 taxa) in all groups, followed by Mexico (56 taxa), Peru (32 taxa) and Argentina (27 taxa).
Helminth parasites of chelonians across political units in the Neotropical Region.

Publication history
This study found 199 publications related to helminths of chelonians in the Neotropical region between the years 1850 and 2024 (Figure 5). Of these, 107 were for Nematoda, 95 for Trematoda, 31 for Monogenea, 10 for Acanthocephala, and six for Cestoda. Publications for the Nematoda group showed irregular peaks after 1935 and a significant increase after the 1990s, making it the group with the most papers over the years. The Trematoda group presented more work between 1940 and 1980, with irregular peaks in publications over the years, increasing significantly in the last decade. The Monogenea group showed an increase in publications from the 2000s onward and, together with the Acanthocephala and Cestoda groups, exhibited low publication rates over the years, with a slight increase in the 2000s.
Number of publications by group of helminth parasites of chelonians over the years in the Neotropical region.

Field research: New record of Phrynops geoffroanus in the Brazilian semi-arid region
Two populations of P. geoffroanus were studied in November and December 2019 and December 2020, totaling 17 specimens captured (seven females, five males, and five juveniles). All the individuals were parasitized, resulting in a total of 432 helminths collected (Table 1). Regarding the diversity of endoparasites, they all belong to the Nematoda phylum. The nematodes recorded belong to the Camallanidae family - Camallanus sp., found in the stomach and intestine with the highest abundance (n=383) and prevalence (100%), and the Gnathostomatidae family - Spiroxys contortus, with the lowest abundance (n=49) and prevalence (64.7%) in the stomach. Camallanus sp. was identified at genus level because its characteristics differed from other species already recorded for reptiles, indicating the possibility of it being a new species. Spiroxys contortus is the first record of the species for the host P. geoffroanus.
Values of abundance (A), number of infected (NI), mean intensity of infection (MII), mean abundance (MA), standard deviation (SD) of average intensity, and site of infection (IS): stomach (St) and intestine (It) for nematodes of Phrynops geoffroanus in the semi-arid region of northeastern Brazil

Spiroxys contortus (Rudolphi, 1819) Schneider, 1866
Spiroptera contorta Rudolphi, 1819, Agamospirura ophidii Sharpilo, 1964
Taxonomic summary:
Host: Phrynops geoffroanus (Schweigger 1812).
Site of infection: Stomach
Locality: Conceição, Paraíba, Brazil (7°33’44.3’’S 38°30’27.4’’W) and Patos, Paraíba, Brazil (7°02’18.1’’ S 37°16’36.8’’W).
Specimens deposited: UFPB.NEMA 10 - 14
Description:
Males (eight specimens): Body length 14.79–29.98 mm and width 194.11–587.5 μm. Lips 36.75–71 mm long and 59–117.5 μm wide. Total oesophagus 2229–5044 μm wide, muscular section 770.6–811.8 and glandular 4015–4205 (two specimens measured) μm long. Nerve ring 370–776.5 and excretory pore 823.5–1100 (four specimens measured) μm from the anterior end. Tail curved ventrally, caudal wings poorly developed. 12 pairs of caudal papillae present: three pairs preanal, two adanal and six postanal. Spicules slender, slightly unequal in size. Larger spicule 2529–2941.25 μm long and smaller spicule 24.94–2926.5 μm long. Gubernaculum little sclerotized, 188.2–352.9 μm long (six specimens measured). Tail 200–352 μm long.
Females (five specimens): Body length 32.5–44.1 mm and width 529.5–661.75 μm. Lips length 47.05–59 by 53–59 (three specimens measured) μm wide. Total oesophagus is 3970.5–6677.5 μm long, muscular section 517.6–955.75 and glandular 4102.5–5882.5 μm. Nerve ring 517.6–955.75 and excretory pore 871–1367.75 (three specimens measured) μm from the anterior end. Vulva situated 19–28.35 mm from the anterior end. Vagina anteriorly directed, withventer opposite. Eggs 29.5–59 μm long by 41.2–55.85 μm wide (eight eggs measured). Tail 485.25–602.75 μm long, with one pair of dorsolateral phasmids.
Lists of Helminths of terrestrial and freshwater chelonians from the Neotropical region
PHYLUM NEMATODA Rudolphi, 1808
CLASS CHROMADOREA Inglis, 1983
Superfamily Ascaridoidea Baird, 1853
Family Anisakidae Railliet & Henry, 1912
Contracaecum Railliet & Henry, 1912
Contracaecum sp. larvae
Infection site: unknown
Host: Acanthochelys spixii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Henzel and Müller2017)
Contracaecum sp. larvae.
Infection site: body cavity
Host: Acanthochelys spixii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Family Ascarididae Baird, 1853
Angusticaecum Baylis, 1920
Angusticaecum brevispiculum Chapin, Reference Chapin1924
Infection site: intestine
Host: Chelonoidis denticulatus
Biogeographic unit: Rondônia province3, Ucayali province3 – SBD
Political unity: [BRAZIL] – (Pará): unspecified location1,2; [PERU] – (Loreto)3: Jenaro Herrera, Pullcapa and Yurimaguas; (San Martin): unspecified location4
References: Chapin (Reference Chapin1924)1; Vicente et al. (Reference Vicente, Rodrigues, Gomes and Pinto1993)2; Salizar and Sanchez (Reference Salizar and Sanchez2007)3; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)4
Angusticaecum holopterum (Rudolphi, 1819) Baylis, 1920
Infection site: small intestine and large intestine
Host: Chelonoidis denticulatus
Biogeographic unit: Ucayali province – SBD
Political unity: [PERU] – (Loreto): Iquitos
Reference: Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)
Angusticaecum sp.
Infection site: intestine
Host: Chelonoidis denticulatus
Political unity: [PERU] – (Loreto): unspecified location
References: Tantaleán et al. (Reference Tantaleán, Juarez and Cruz1983) cited by Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)
Angusticaecum sp.
Infection site: intestine
Host: Chelonoidis denticulatus
Biogeographic unit: Desert province –STZ
Political unity: [PERU] – Lima.
Reference: Chavez et al. (Reference Chavez, Serrano-Martínez, Tantaleán, Quispe and Casas2015)
Ophidascaris Baylis, 1921
Ophidascaris arndti Sprehn, 1929
Infection site: stomach, small intestine
Host: Chelonoidis denticulatus
Biogeographic unit: Ucayali province – SBD
Political unity: [PERU] – (Loreto): Iquitos.
Reference: Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)
Ophidascaris sp.
Infection site: intestine
Host: Chelonoidis denticulatus
Political unity: [PERU] – (Ucayali): unspecified location
Reference: Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)
Family Heterocheilidae Railliet & Henry, 1915
Brevimulticaecum Mozgovoy, 1951
Brevimulticaecum sp. Larvae
Infection site: stomach and large intestine
Host: Phrynops geoffroanus
Biogeographic unit: Parana Forest province – PD
Political unity: [BRAZIL] – (São Paulo): RPPN Foz do Rio Aguapeí
Reference: Silva (Reference Silva2014)
Superfamily Camallanoidea Travassos, 1920
Family Camallanidae Railliet & Henry, 1915
Camallanus Railliet & Henry, 1915
Camallanus emydidius Mascarenhas & Müller, 2017
Infection site: stomach, small intestine, large intestine, heart, lungs and liver
Host: Hydromedusa tectifera 1 and Trachemys dorbigni 2
Biogeographic unit: Pampean province1,2 – CD; Esteros del Iberá Province2 – PD
Political unity: [ARGENTINA] – (Entre Ríos)2: Concordia and Federal; [BRAZIL] – (Rio Grande do Sul)1: Capão do Leão2, Pelotas2, Rio Grande and Santa Vitória do Palmar
References: Mascarenhas and Müller (Reference Mascarenhas and Muller2017)2; Chaviel et al. (Reference Chaviel, Mascarenhas, Bernardon, Coimbra and Müller2020)1; Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)2; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2; Palumbo et al. (Reference Palumbo, Servián, Cassano and Diaz2024b)2
Camallanus pallidipectoris Palumbo, Servián, Cassano & Diaz, 2024
Infection site: stomach and small intestine
Host: Acanthochelys pallidipectoris and Phrynops hilarii
Biogeographic unit: Chaco province and Pampean province – CD
Political unity: [ARGENTINA] – (Entre Ríos province): Aldea Brasilera; (Santa Fe province): Vera
Reference: Palumbo et al. (Reference Palumbo, Servián, Cassano and Diaz2024b)
Camallanus sp.
Infection site: body cavity and small intestine
Host: Acanthochelys spixii and Hydromedusa tectifera
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande, and Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Souza, Coimbra and Müller2013)
Camallanus sp.
Infection site: stomach and small intestine
Host: Phrynops hilarii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Pelotas
Reference: Bernardon et al. (Reference Bernardon, Valente and Müller2013)
Camallanus sp.
Infection site: small intestine
Host: Trachemys dorbigni
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Pelotas
Reference: Bernardon et al. (Reference Bernardon, Valente and Müller2014)
Camallanus sp.
Infection site: small intestine
Host: Phrynops geoffroanus
Biogeographic unit: Parana Forest province – PD
Political unity: [BRAZIL] – (Minas Gerais): Juiz de Fora
Reference: Vieira et al. (Reference Vieira, Oliveira, Gonçalves, Sousa, Souza Lima and Novelli2016)
Camallanus sp.
Infection site: small intestine and large intestine
Host: Acanthochelys spixii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Camallanus sp.
Infection site: small intestine
Host: Kinosternon scorpioides
Biogeographic unit: Chaco province – CD
Political unity: [ARGENTINA] – (Chaco province): Formosa
Reference: Palumbo et al. (Reference Palumbo, Servián, Cassano and Diaz2024b)
Camallanus sp.
Infection site: stomach and intestine
Host: Phrynops geoffroanus
Biogeographic unit: Caatinga province – CD
Political unity: [BRAZIL] – (Paraiba): Conceição and Patos
Reference: Present study
Camallanus spiculatus Palumbo, Servián, Cassano & Diaz, 2024
Infection site: stomach and small intestine
Host: Acanthochelys spixii and Phrynops hilarii
Biogeographic unit: Esteros del Iberá province – PD
Political unity: [ARGENTINA] – (Corrientes province): Bella vista
Reference: Palumbo et al. (Reference Palumbo, Servián, Cassano and Diaz2024b)
Serpinema Yeh, 1960
Serpinema amazonicum (Ribiero, 1941)
Infection site: stomach and small intestine
Host: Mesoclemmys tuberculata 1, Phrynops geoffroanus 2, Podocnemis expansa 3, Podocnemis sp.4, and Podocnemis unifilis 5
Biogeographic unit: Atlantic province1,2 – PD; Caatinga province1,2 – CD; Pará province3 – BBD; Ucayali province – SBD3,5
Political unity: [BRAZIL] – (Sergipe)1,2: Areia Branca, Capela, Pirambu, Poço Redondo, and Tobias Barreto; (Pará): Belém3; [PERU] – (Loreto): Iquitos,3 Pacaya-Samiria National Reserve,5 and unspecified location4; (Ucayali): unspecified location4
References: Ribeiro (Reference Ribeiro1940)3; Tantaleán (Reference Tantaleán1998)3; Sanchez et al. (Reference Sánchez, Tantaleán, Vela and Méndez2006)5; Santana et al. (Reference Santana, Teixeira, Araújo-Filho, Teles, Pereira, Brito, Almeida and Mesquita2023)1,2; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)4
Serpinema cayennense Harnoster, Svitin & Preez, Reference Harnoster, Svitin and Preez2019
Infection site: intestine
Host: Rhinoclemmys punctularia
Biogeographic unit: Guianan Lowlands province – BBD
Political unity: [FRENCH GUIANA] – Cayenne
Reference: Harnoster et al. (Reference Harnoster, Svitin and Preez2019)
Serpinema kachugae (Baylis & Daubney, 1922)
Infection site: intestine
Host: Kinosternon scorpioides
Biogeographic unit: Pacific dominion
Political unity: [VENEZUELA] – Zulia
Reference: Díaz-Ungría (Reference Díaz-Ungría1978)
Serpinema magathi (Sprehn, Reference Sprehn1932 )
Infection site: stomach, small intestine, and large intestine
Host: Kinosternon integrum 1, Kinosternon leucostomum 2, and Kinosternon scorpioides 3
Biogeographic unit: Atlantic province3 – PD; Pacific Lowlands province2 – MD; Pará province3 – BBD; Roraima province3 – BBD
Political unity: [BOLIVIA] - unspecified location1; [BRAZIL] – (Pará): Ilha de Marajó3; (Pernambuco): Recife3; (Maranhão): São Bento3; [COSTA RICA] – Santa Rosa National Park2
References: Sprehn (Reference Sprehn1932)1; Alho (Reference Alho1965)3; Freitas and Dobbin Jr. (Reference Freitas and Dobbin1971)3; Bursey and Brooks (Reference Bursey and Brooks2011)2; Viana et al. (Reference Viana, Rodrigues, Madelaire, Santos and Sousa2016)3
Serpinema microcephalus (Dujardin, 1845)
Infection site: intestine
Host: Peltocephalus dumerilianus, Podocnemis expansa, and Podocnemis unifilis
Political unity: [BRAZIL] – unspecified location
Reference: Diesing (Reference Diesing1851)
Serpinema monospiculatus Freitas & Dobbin Jr., Reference Freitas and Dobbin1962
Infection site: stomach, small intestine, large intestine, lungs, and body cavity
Host: Kinosternon scorpioides 1, Mesoclemmys nasuta 2, Mesoclemmys tuberculata 3, and Phrynops geoffroanus 4
Biogeographic unit: Atlantic province2,3,4, Parana Forest province4 – PD; Caatinga province1,3,4 – CD
Political unity: [BRAZIL] – (Sergipe)3,4: Areia Branca, Capela, Pirambu, Poço Redondo, and Tobias Barreto; (Pernambuco): Cortês,4 Jaboatão dos Guararapes,3 Limoeiro,4 Paudalho,4 Recife,2,4 and Santa Cruz do Capibaribe4; (Ceará): Nova Olinda1,3,4; (São Paulo): RPPN Foz do Rio Aguapeí4
References: Freitas and Dobbin Jr. (Reference Freitas and Dobbin1971)2,3,4; Silva (Reference Silva2014)4; Pereira A et al. (Reference Pereira, Junior, Santana, Almeida, Pereira and Oliveira2018)1,3,4; Fonseca et al. (Reference Fonseca, Rojas, Vieira, Ferreira, Rodrigues, Barros, Silva Júnior, Moura and Oliveira2021)4; Santana et al. (Reference Santana, Teixeira, Araújo-Filho, Teles, Pereira, Brito, Almeida and Mesquita2023)3,4
Serpinema pelliculatus Silva, Jesus, Santos, Nandyara, Willkens, Santos & Melo, Reference Silva, Jesus, Santos, Nandyara, Willkens, Santos and Melo2023
Infection site: stomach and intestine
Host: Kinosternon scorpioides
Biogeographic unit: Para province1 and Roraima province2 – BBD
Political unity: [BRAZIL] – (Pará): Abaetetuba1 and Ilha do Marajó2
Reference: Silva et al. (Reference Silva, Jesus, Santos, Nandyara, Willkens, Santos and Melo2023)
Serpinema sp.
Infection site: stomach, small intestine, and large intestine
Host: Trachemys callirostris
Biogeographic unit: Magdalena province – PCD
Political unity: [COLOMBIA] – (Córdoba): Montería
Reference: Martínez et al. (Reference Martínez, Cordero, Velásquez, Bolaño, Otero, Monsalve, Mestra and Chacón-Pacheco2021)
Serpinema sp.
Infection site: small intestine
Host: Kinosternon scorpioides
Biogeographic unit: Chaco province – CD
Political unity: [ARGENTINA] – (Formosa province): Formosa
Reference: Palumbo et al. (Reference Palumbo, Servián, Cassano and Diaz2024b)
Serpinema trispinosum (Leidy, 1852)
Note: Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008) mention that this species was originally described as Camallanus magnorugosus by Caballero (Reference Caballero1939b) and Camallanus scabrae by (Caballero Reference Caballero1943b).
Infection site: stomach, intestine, and small intestine
Host: Dermatemys mawii 1, Kinosternon hirtipes 2, Rhinoclemmys areolata 3, Terrapene ornata 4, Trachemys scripta 5, and Trachemys venusta 6
Biogeographic unit: Transmexican Volcanic Belt province2 – MTZ; Veracruzan province1,3,4,6 – MD; Yucatán Peninsula province5 – MD
Political unity: [MEXICO]: (Veracruz): Alvarado – Laguna de Alvarado4; (Tabasco): Emiliano Zapata3, Laguna el espiño,6 and Río de Chilapa1; (Michoacán)2: Manantiales de Cointzio; Pátzcuaro – Laguna de Pátzcuaro, Zacapu – Lago de Zacapu and Zirahuén - Laguna de Zirahuén; (Yucatán): Aguada Xpoc5
References: Caballero (Reference Caballero1939b)4; Caballero (Reference Caballero1943b)1; Dyer and Carr (Reference Dyer and Carr1990a)3; Moravec and Vargas-Vázquez (Reference Moravec and Vargas-Vázquez1998)5; Pérez-Ponce de León et al. (Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)2; Aguilar-Aguilar et al. (Reference Aguilar-Aguilar, Báez-Valé, Mendoza-Palmero, Salgado-Maldonado and Barrios-Quiroz2003)6
Procamallanus (Spirocamallanus) Olsen, 1952
Spirocamallanus sp.
Infection site: small intestine
Host: Hydromedusa tectifera
Biogeographic unit: Parana florest province – PD
Political unity: [BRAZIL] – (Minas Gerais): Ingaí
Reference: Novelli et al. (Reference Novelli, Morton, Trindade, Diego, Silva and Vieira2014)
Superfamily Cosmocercoidea Railliet, 1916
Family Atractidae Railliet, 1917
Atractidae g. sp. larvae
Infection site: intestine
Host: Chelonoidis denticulatus (captivity)
Biogeographic unit: Desert province – STZ
Political unity: [PERU] – Lima
Reference: Chavez et al. (Reference Chavez, Serrano-Martínez, Tantaleán, Quispe and Casas2015)
Acanthoatractis Jesus and Melo, 2024
Acanthoatractis xinguensis Jesus & Melo in Jesus, Nandyara & Melo, 2024
Infection site: stomach
Host: Podocnemis unifilis
Biogeographic unit: Xingu-Tapajós province – SAD
Political unity: [BRAZIL] – (Pará): Vitória do Xingu – Xingu River
Reference: Jesus et al. (Reference Jesus, Santos, Nandyara, Santos and Melo2024b)
Atractis Dujardin, 1845
Atractis costaricaensis Gibbons & Platt, Reference Gibbons and Platt2006
Infection site: small intestine and large intestine
Host: Rhinoclemmys pulcherrima
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [COSTA RICA] – Santa Rosa National Park
References: Gibbons and Platt (Reference Gibbons and Platt2006); Bursey and Brooks (Reference Bursey and Brooks2011)
Atractis cruciata Linstow, 1902
Infection site: stomach
Host: Podocnemis expansa
Biogeographic unit: Cerrado province – CD
Political unity: [BRAZIL] – (Tocantins): Javaés River
Reference: Armond (Reference Armond2008)
Atractis dactyluris (Rudolphi, 1819) Dujardin, 1845
Infection site: intestine
Host: Mesoclemmys nasuta, Podocnemis expansa, and Podocnemis unifilis
Political unity: [BRAZIL] – unspecified location
Reference: Diesing (Reference Diesing1851)
Atractis impura Caballero, 1944
Infection site: digestive tract, intestine, large intestine
Host: Chelonoidis denticulatus 1, Gopherus sp.2, Kinosternon scorpioides cruentatum 3, Rhinoclemmys areolata 4, Rhinoclemmys diademata 5, and Rhinoclemmys pulcherrima 6
Biogeographic unit: Transmexican Volcanic Belt province2 – MTZ; Pacific Lowlands province6 – MD; Yucatán Peninsula province4 – MD; Ucayali province1 – SBD; Guajira province5, Puntarenas-Chiriquí province3 – PCD
Political unity: [COSTA RICA] – (San José): Santa Ana3; [MEXICO] – (Federal District): Mexico City2; (Guerrero): Acapulco6; (Quintana Roo): 1.1 km N of Coba road, on Hwy 3074; [PERU] – (Loreto): Iquitos1; (Ucayali): Pucallpa1; [VENEZUELA] – (Zulia): Lago de Maracaibo5
References: Caballero (Reference Caballero1944a)2; Brenes-Madrigal and Bravo-Hollis (Reference Brenes-Madrigal and Bravo-Hollis1960)3; Dyer and Carr (Reference Dyer and Carr1990a)4,5,6; Salizar and Sanchez (Reference Salizar and Sanchez2007)1
Atractis marquezi Bursey & Flanagan Reference Bursey and Flanagan2002
Infection site: colon, intestine, and large intestine
Host: Chelonoidis carbonarius 1, Chelonoidis denticulatus 2, Chelonoidis porteri 3, Mesoclemmys gibba 4, Phrynops geoffroanus 5, and Podocnemis unifilis 6
Biogeographic unit: Galápagos Islands province3 – PCD; Lesser Antilles province1 – AS; Ucayali province2 – SBD
Political unity: [ECUADOR] – (Galápagos): Isla Santa Cruz3; [GRANADA - West Indies]1; [PERU] – (Pasco); unspecified location6; (Ucayali): unspecified location4,5; (Loreto): Iquitos2
References: Bursey and Flanagan (Reference Bursey and Flanagan2002)3; Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)2; Springer et al. (Reference Springer, Kinsella, Vasuki and Sharma2020)1; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)4,5,6
Atractis sp.
Infection site: stomach
Host: Podocnemis expansa
Biogeographic unit: Cerrado province – CD
Political unity: [BRAZIL]: (Tocantins): Javaés River
Reference: Armond (Reference Armond2008)
Atractis sp.
Infection site: intestinal content
Host: Chelonoidis carbonarius
Biogeographic unit: Guianan Lowlands province – BBD
Political unity: [VENEZUELA] – (Bolívar): Upata
Reference: Pérez Mata et al. (Reference Pérez Mata, Anguirre, Piñango, Olivares, Camacho, Bautista and Núñez2014)
Atractis sp.
Infection site: intestine
Host: Chelonoidis sp.
Biogeographic unit: Galápagos Islands province – PCD
Political unity: [ECUADOR] – Galápagos
Reference: Fournié et al. (Reference Fournié, Goodman, Cruz, Cedeño, Vélez, Patiño, Millins, Gibbons, Fox and Cunningham2015)
Atractis sp.
Infection site: intestine (fecal samples)
Host: Chelonoidis carbonarius
Biogeographic unit: Lesser Antilles province – AS
Political unity: [GRANADA – West Indies]
Reference: Springer et al. (Reference Springer, Kinsella, Vasuki and Sharma2020)
Atractis thapari Petter, 1966
Infection site: small intestine and large intestine
Host: Chelonoidis carbonarius 1 and Chelonoidis denticulatus 2
Biogeographic unit: Pará province1,2 – BBD; Lesser Antilles province1 – AS
Political unity: [BRAZIL] – (Piauí): Teresina1,2; [GRANADA - West Indies]1
References: Leal et al. (Reference Leal, Freire, Knoff, Gomes, Santos and Mendonça2018)1,2; Freire et al. (Reference Freire, Leal, Batista, Rocha, Baêta and Mendonça2019)1,2; Springer et al. (Reference Springer, Kinsella, Vasuki and Sharma2020)1
Atractis trematophila Travassos, 1934
Infection site: stomach, small intestine, and large intestine
Host Mesoclemmys gibba 1, Peltocephalus dumerilianus 2, Podocnemis expansa 3, and Podocnemis unifilis 4
Biogeographic unit: Imerí province2 – BBD
Political unity: [BRAZIL] – unspecified location3; (Amazonas)2: Barcelos and Jaú National Park; [PERU] – (Pasco) – unspecified location1,4
References: Travassos (Reference Travassos1934a)3; Ferreira (Reference Ferreira2016)2; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)1,4; Menegaldo et al. (Reference Menegaldo, Arias-Pacheco, Perin, Tebaldi and Hoppe2024)2
Buckleyatractis Khalil & Gibbons Reference Khalil and Gibbons1988
Buckleyatractis marinkelli Khalil & Gibbons, Reference Khalil and Gibbons1988
Infection site: large intestine
Host: Podocnemis unifilis
Political unity: [COLOMBIA] – unspecified location
Reference: Khalil and Gibbons (Reference Khalil and Gibbons1988)
Klossinemella Goncalves Da Costa, Reference Costa1961
Klossinemella caballeroi (Brenes and Bravo Hollis, Reference Brenes-Madrigal and Bravo-Hollis1960) Bursey and Flanagan, Reference Bursey and Flanagan2002
Synonym: Atractis caballeroi Brenes & Bravo-Hollis, Reference Brenes-Madrigal and Bravo-Hollis1960
Infection site: digestive tract, intestine, and large intestine
Host: Chelydra serpentina 1, Kinosternon scorpioides cruentatum 2, Kinosternon leucostomum 3, Rhinoclemmys annulata 4, and Rhinoclemmys areolata 5
Biogeographic unit: Pacific Lowlands province1,4, Veracruzan province5 – MD; Puntarenas-Chiriquí province2 – PCD; Western Ecuador province3.4 – PCD
Political unity: [COSTA RICA] – Santa Rosa National Park1,4; (San José): Santa Ana2
[ECUADOR] – (Esmeraldas): San Jose de Cayapas4 and Concepcion3; [MEXICO] – (Chiapas): Catazajá5; (Tabasco): Emiliano Zapata5
References: Brenes-Madrigal and Bravo-Hollis (Reference Brenes-Madrigal and Bravo-Hollis1960)2; Dyer and Carr (Reference Dyer and Carr1990a)3,4,5; Bursey and Brooks (Reference Bursey and Brooks2011)1,4
Klossinemella conciliatus Alho, 1964
Infection site: stomach, small intestine, large intestine
Host: Peltocephalus dumerilianus 1, Podocnemis expansa,2 and Podocnemis unifilis 3
Biogeographic unit: Cerrado province2 – CD; Imerí province1 – BBD; Ucayali province3 – SBD
Political unity: [BRAZIL] – unspecified location2; (Amazonas)1: Barcelos and Jaú National Park; (Tocantins): Javaés River2; [PERU] – Samiria river3
References: Alho (Reference Alho1964a)2; Costa et al. (Reference Costa, Motta and Gomes1968)2; Salizar and Sanchez (Reference Salizar and Sanchez2007)3; Armond (Reference Armond2008)2; Ferreira (Reference Ferreira2016)1; Menegaldo et al. (Reference Menegaldo, Arias-Pacheco, Perin, Tebaldi and Hoppe2024)1
Klossinemella parvicapiticoronata - G. Caballero, 1971
Infection site: large intestine
Host: Chelonoidis carbonarius 1 and Staurotypus triporcatus 2
Biogeographic unit: Lesser Antilles province1 – AS
Political unity: [GRANADA – West Indies]1; [MEXICO] – (Veracuz): unspecified location2
References: Caballero-Rodríguez (Reference Caballero-Rodríguez1971)2; Springer et al. (Reference Springer, Kinsella, Vasuki and Sharma2020)1
Klossinemella sp.
Infection site: stomach, small intestine, and large intestine
Host: Peltocephalus dumerilianus
Biogeographic unit: Imerí province – BBD
Political unity: [BRAZIL] – (Amazonas): Barcelos
Reference: Ferreira (Reference Ferreira2016)
Klossinemella sp.
Infection site: stomach, small intestine, and large intestine
Host: Peltocephalus dumerilianus
Biogeographic unit: Imerí province – BBD
Political unity: [BRAZIL] – (Amazonas): Jaú National Park
Reference: Menegaldo et al. (Reference Menegaldo, Arias-Pacheco, Perin, Tebaldi and Hoppe2024)
Klossinemella travassosi Costa, Mota & Gomes, 1968
Infection site: intestine, large intestine
Host: Chelonoidis denticulatus 1 and Podocnemis unifilis 2
Biogeographic unit: Ucayali province1 – SBD
Political unity: [BRAZIL] – (Amazonas): unspecified location1; [PERU] – (Loreto)1: unspecified location and Iquitos; (Pasco): unspecified location2
References: Costa et al. (Reference Costa, Motta and Gomes1968)1; Muniz-Pereira et al. (Reference Muniz-Pereira, Vieira and Luque2009)1; Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)1; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)1,2.
Labiduris Schneider, 1866
Labiduris gulosa (Rudolphi, 1819)
Infection site: cecum, cardiac vessels, stomach, intestine, and large intestine
Host: Chelonoidis carbonarius 1 and Chelonoidis denticulatus 2
Biogeographic unit: Lesser Antilles province1 – AS; Ucayali province2 – SBD
Political unity: [BRAZIL] – unspecified location1; (Pará): unspecified location2; (Rio de Janeiro): unspecified location2; [GRANADA – West Indies]1; [PERU] – (Loreto): Iquitos2; (San Martin): unspecified location2; (Ucayali): unspecified location2
References: Diesing (Reference Diesing1851)1; Chapin (Reference Chapin1924)2; Muniz-Pereira et al. (Reference Muniz-Pereira, Vieira and Luque2009)2; Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)2; Springer et al. (Reference Springer, Kinsella, Vasuki and Sharma2020)1; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)2
Labiduris irineuta Costa, Reference Costa1961
Infection site: small intestine, large intestine
Host: Chelonoidis denticulatus
Biogeographic unit: Atlantic province – PD1; Ucayali province – SBD2
Political unity: [BRAZIL] – (Rio de Janeiro): Rio de Janeiro1; [PERU] – (Loreto): Iquitos2
References: Costa (Reference Costa1961)1 cited by Vicente et al. (Reference Vicente, Rodrigues, Gomes and Pinto1993); Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)2
Labiduris sp.
Infection site: intestine
Host: Chelonoidis sp.
Biogeographic unit: Galápagos Islands province – PCD
Political unity: [ECUADOR] – Galápagos islands
Reference: Fournié et al. (Reference Fournié, Goodman, Cruz, Cedeño, Vélez, Patiño, Millins, Gibbons, Fox and Cunningham2015)
Labiduris sp. larvae
Infection site: digestive system (fecal samples)
Host: Chelonoidis carbonarius and Chelonoidis denticulatus
Biogeographic unit: Atlantic province – PD
Political unity: [BRAZIL] – (Rio de Janeiro): Rio de Janeiro
Reference: Meireles et al. (Reference Meireles, Miguel, Mattos Junior, Barros and Magalhães2018)
Labiduris sp.
Infection site: small intestine
Host: Chelonoidis chilensis
Biogeographic unit: Monte province – STZ
Political unity: [ARGENTINA] – San Juan
Reference: Castillo et al. (Reference Castillo, Acosta, González-Rivas and Ramallo2020)
Labiduris zschokkei Linstow, 1899
Infection site: stomach, intestine, large intestine
Host: Chelonoidis denticulatus
Biogeographic unit: Ucayali province2 – SBD
Political unity: [BRAZIL] - unspecified location1; [PERU] – (Loreto): Iquitos2; (Ucayali): unspecified location3
References: Skrjabin (Reference Skrjabin, Shikhobalova and Lagodovskaya1964)1 cited by Vicente et al. (Reference Vicente, Rodrigues, Gomes and Pinto1993); Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)2; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)3
Orientatractis Petter, 1966
Orientatractis asymmetrica Gibbons & Platt, Reference Gibbons and Platt2006
Infection site: small intestine and large intestine
Host: Rhinoclemmys pulcherrima
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [COSTA RICA] - Santa Rosa National Park
Reference: Gibbons and Platt (Reference Gibbons and Platt2006)
Orientatractis bowseri Jesus & Melo in Jesus, Santos, De Almeida Pereira Oliveira, Félix‑Silva & Melo, 2023
Infection site: stomach
Host: Peltocephalus dumerilianus
Biogeographic unit: Imerí province – BBD
Political unity: [BRAZIL] – (Amazonas): Jaú National Park
Reference: Jesus et al. (Reference Jesus, Santos, Oliveira, Félix-Silva and Melo2024a)
Orientatractis leiperi Buckley, Reference Buckley1969
Infection site: stomach and intestine
Host: Podocnemis unifilis 1 and Podocnemis vogli 2
Biogeographic unit: Cerrado province1 – CD; Pará province1 – BBD
Political unity: [BRAZIL] – (Pará): Tucuruí – Tocantins River1; (Tocantins)1: Araguaia National Park/Cantão State Park-Javaés River and Tupiratins/Itapiratins – Tocantins River; [COLOMBIA] – unspecified location2
References: Buckley (Reference Buckley1969)2; Jesus et al. (Reference Jesus, Cardoso, Willkens, Furtado, Santos and Melo2020)1; Ribeiro et al. (Reference Ribeiro, Santana, Souza, Oliveira, Magalhães, Malvasio and Silva2023)1
Orientatractis matosi Jesus, Oliveira & Melo in Jesus, Oliveira, Ramos & Melo, 2023
Infection site: stomach, large intestine
Host: Podocnemis unifilis
Biogeographic unit: Pará province – BBD; Xingu-Tapajós province – SAD
Political unity: [BRAZIL] – (Pará): Senador José Porfírio – Xingu River and Tucuruí – Tocantins River
Reference: Jesus et al. (Reference Jesus, Oliveira, Ramos and Melo2023a)
Paraorientatractis Gibbons, Khalil & Marinkelle, Reference Gibbons, Khalil and Marinkelle1997
Paraorientatractis semiannulata Gibbons, Khalil & Marinkelle, Reference Gibbons, Khalil and Marinkelle1997
Infection site: stomach and intestine
Host: Mesoclemmys gibba 1, Phrynops geoffroanus,2 and Podocnemis unifilis 3
Biogeographic unit: Cerrado province3 – CD; Roraima province3 – BBD; Ucayali province3 – SBD
Political unity: [BRAZIL] – (Pará)3: Cuminá and Trombetas rivers; (Tocantins)3: Araguaia National Park/Cantão State Park-Javaés River and Tupiratins/Itapiratins – Tocantins River; [PERU] – (Pasco): unspecified location1,2; (Ucayali): Pucallpa3
References: Gibbons et al. (Reference Gibbons, Khalil and Marinkelle1997)3; Salizar and Sanchez (Reference Salizar and Sanchez2007)3; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)1,2; Ribeiro et al. (Reference Ribeiro, Santana, Souza, Oliveira, Magalhães, Malvasio and Silva2023)3
Paratractis Sarmiento, Reference Sarmiento1959
Paratractis hystrix (Diesing, Reference Diesing1851) Sarmiento, Reference Sarmiento1959
Infection site: stomach, small intestine, and large intestine
Host: Peltocephalus dumerilianus 1, Podocnemis erythrocephala 2, Podocnemis expansa 3, Podocnemis sp.4, Podocnemis unifilis 5, and Podocnemis vogli 6
Biogeographic unit: Cerrado province5 – CD; Imerí province1 – BBD; Ucayali province1 – SBD
Political unity: [BRAZIL] – (Amazonas): Barcelos1, Jaú National Park1 and unspecified location2; (Tocantins)5: Araguaia National Park/Cantão State Park-Javaés River and Tupiratins/Itapiratins – Tocantins River; [COLÔMBIA] – unspecified location3,5,6; [PERU] – (Loreto): unspecified location4,5; (Ucayali): Pucallpa1
References: Diesing (Reference Diesing1851)2; Sarmiento (Reference Sarmiento1959)1; Buckley (Reference Buckley1970)6; Khalil and Gibbons (Reference Khalil and Gibbons1988)3,5,6; Ferreira (Reference Ferreira2016)1; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)4,5; Ribeiro et al. (Reference Ribeiro, Santana, Souza, Oliveira, Magalhães, Malvasio and Silva2023)5; Menegaldo et al. (Reference Menegaldo, Arias-Pacheco, Perin, Tebaldi and Hoppe2024)1
Pneumoatractis Bursey, Reavill & Greiner, Reference Bursey, Reavill and Greiner2009
Pneumoatractis gibbonsae Jesus & Melo in Jesus, Willkens, dos Santos & Melo, 2023
Infection site: stomach and large intestine
Host: Podocnemis expansa and Podocnemis unifilis
Biogeographic unit: Xingu-Tapajós province1 – SAD; Pará province2 – BBD
Political unity: [BRAZIL] – (Pará): Senador José Porfírio – Xingu River1 and Tucuruí – Tocantins River2
Reference: Jesus et al. (Reference Jesus, Willkens, Santos and Melo2023b)
Pneumoatractis podocnemis Bursey, Reavill & Greiner, Reference Bursey, Reavill and Greiner2009
Infection site: lungs
Host: Podocnemis unifilis
Dristribution: [BRAZIL] – unspecified location
Reference: Bursey et al. (Reference Bursey, Reavill and Greiner2009)
Podocnematractis Gibbons, Khalil & Marinkelle, Reference Gibbons, Khalil and Marinkelle1995
Podocnematractis colombiaensis Gibbons, Khalil & Marinkelle, Reference Gibbons, Khalil and Marinkelle1995
Infection site: large intestine
Host: Podocnemis expansa and Podocnemis vogli
Political unity: [COLOMBIA] – unspecified location
Reference: Gibbons et al. (Reference Gibbons, Khalil and Marinkelle1995)
Podocnematractis ortleppi (Thapar, 1925) Gibbons, Khalil & Marinkelle, Reference Gibbons, Khalil and Marinkelle1995
Infection site: large intestine
Host: Podocnemis expansa, Podocnemis unifilis, and Podocnemis vogli
Political unity: [COLOMBIA] – unspecified location
Reference: Gibbons et al. (Reference Gibbons, Khalil and Marinkelle1995)
Proatractis Caballero, 1971
Proatractis sp. larvae
Infection site: intestine (fecal samples)
Host: Chelonoidis carbonarius
Biogeographic unit: Caatinga province – CD
Political unity: [BRAZIL] – (Rio Grande do Norte): Mossoró
Reference: Pereira AWS et al. (Reference Pereira, Junior, Santana, Almeida, Pereira and Oliveira2018)
Rhinoclemmysnema Gibbons & Platt, Reference Gibbons and Platt2006
Rhinoclemmysnema multilabiatum Gibbons & Platt, Reference Gibbons and Platt2006
Infection site: small intestine and large intestine
Host: Rhinoclemmys pulcherrima
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [COSTA RICA] – Santa Rosa National Park
Reference: Gibbons and Platt (Reference Gibbons and Platt2006)
Vogtnema Jesus, Santos, Nandyara & Melo in Jesus, Santos, Nandyara, Santos & Melo, Reference Jesus, Santos, Nandyara, Santos and Melo2022
Vogtnema asymmetricum Jesus, Santos, Nandyara & Melo in Jesus, Santos, Nandyara, Santos & Melo, Reference Jesus, Santos, Nandyara, Santos and Melo2022
Infection site: stomach and small intestine
Host: Rhinoclemmys punctularia
Biogeographic unit: Pará province – BBD
Political unity: [BRAZIL] – (Pará): Mosqueiro Island – Murubira river
Reference: Jesus et al. (Reference Jesus, Santos, Nandyara, Santos and Melo2022)
Family Cosmocercidae Railliet, 1916
Cosmocercoides Wilkie, 1930
Cosmocercoides variabilis Harwood, 1930
Infection site: large intestine
Host: Rhinoclemmys pulcherrima
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [COSTA RICA] – Santa Rosa National Park
Reference: Bursey and Brooks (Reference Bursey and Brooks2011)
Family Kathlaniidae Lane, 1914
Cruzia Travassos, 1917
Cruzia mexicana Khalil, 1926
Infection site: digestive tracts
Host: Rhinoclemmys areolata 1 and Rhinoclemmys pulcherrima 2
Biogeographic unit: Pacific Lowlands province2, Yucatán Peninsula province1 – MD
Political unity: [MEXICO] – (Guerrero): Acapulco2; (Quintana Roo): 1.1 km N of Coba road, on Hwy 3071;
Reference: Dyer and Carr (Reference Dyer and Carr1990a)
Falcaustra Lane, 1915
Falcaustra affinis (Leidy, 1856) Harwood, 1932
Infection site: heart, stomach, intestine, large intestine, and kidney
Host: Kinosternon hirtipes 1, Trachemys dorbigni,2 and Trachemys scripta 3
Biogeographic unit: Esteros del Iberá province2 – PD; Pampean province2 – CD; Transmexican Volcanic Belt province1 – MTZ; Yucatán Peninsula province3 – MD
Political unity: [ARGENTINA] – (Provincia de Buenos Aires): La Plata2; (Entre Rios)2: Concordia and Federal; [BRAZIL] – (Rio Grande do Sul)2: Capão do Leão and Pelotas; [MEXICO] – (Michoacán)1: Manantiales de Cointzio, Pátzcuaro – Lago de Pátzcuaro and Zirahuén – Lago de Zirahuén; (Yucatán): Aguada Xpoc3
References: Moravec and Vargas-Vázquez (Reference Moravec and Vargas-Vázquez1998)3; Pérez-Ponce de León et al. (Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)1; Mascarenhas and Müller (Reference Mascarenhas and Muller2015b)2; Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)2; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2
Falcaustra chelydrae Harwood, 1932
Infection site: large intestine
Host: Chelydra serpentina and Kinosternon leucostomum
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [COSTA RICA] – Santa Rosa National Park
Reference: Bursey and Brooks (Reference Bursey and Brooks2011)
Falcaustra guanacastensis Bursey & Brooks, Reference Bursey and Brooks2011
Infection site: rectum
Host: Rhinoclemmys annulata
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [COSTA RICA] - Santa Rosa National Park
Reference: Bursey and Brooks (Reference Bursey and Brooks2011)
Falcaustra intermedia (Caballero, 1939) Freitas & Lent, 1941
Synonym: Spironoura intermedia Caballero, 1939
Infection site: small intestine
Host: Kinosternon hirtipes
Biogeographic unit: Transmexican Volcanic Belt province – MTZ
Political unity: [MEXICO] – (Federal District): Xochimilco
Reference: Caballero (Reference Caballero1939a)
Falcaustra sp.
Infection site: digestive tracts
Host: Rhinoclemmys areolata
Biogeographic unit: Yucatán Peninsula province – MD
Political unity: [MEXICO] – (Quintana Roo): 1.1 km N of Coba road, on Hwy 307
Reference: Dyer and Carr (Reference Dyer and Carr1990a)
Falcaustra sp.
Infection site: intestine
Host: Rhinoclemmys nasuta
Biogeographic unit: Guajira province – PCD
Political unity: [COLÔMBIA] – (Sucre): Isla Palma
Reference: Pérez and Alegria (Reference Pérez and Alegria2011)
Falcaustra sp.
Infection site: small intestine
Host: Chelonoidis chilensis
Biogeographic unit: Monte province – STZ
Political unity: [ARGENTINA] – San Juan
Reference: Castillo et al. (Reference Castillo, Acosta, González-Rivas and Ramallo2020).
Falcaustra tikasinghi (Schroeder, Schmidt & Everard, 1977) Baker & Bain, Reference Baker and Bain1981
Infection site: digestive tract, cecum, ileum, and large intestine
Host: Rhinoclemmys annulata 1, Rhinoclemmys melanosterna 2, Rhinoclemmys nasuta,3 and Rhinoclemmys punctularia 4
Biogeographic unit: Pacific Lowlands province1 – MD; Pará province4 – BBD; Trinidad province4, Western Ecuador province1,2,3 – PCD
Political unity: [BRAZIL] – (Pará): Belém4; [COSTA RICA] – Santa Rosa National Park1; [ECUADOR] – (Esmeraldas): Carondelet – Tulubi River3, Concepcion2, San Jose de Cayapas1,3 and San José de Tagua1; (Guayas): Estero el Ceibo3; [TRINIDAD AND TOBAGO – West Indies] – (Trinidad): Aripo Savanas4
References: Schoenecker et al. (Reference Schoenecker, Schmidt and Everard1977)4; Baker and Bain (Reference Baker and Bain1981)4; Dyer and Carr (Reference Dyer and Carr1990a)1,2,3; Bursey and Brooks (Reference Bursey and Brooks2011)1
Falcaustra wardi (Mackin, 1936) Freitas & Lent, 1941
Synonym: Spironoura wardi Mackin, 1936
Infection site: stomach
Host: Kinosternon hirtipes
Biogeographic unit: Transmexican Volcanic Belt province – MTZ
Political unity: [MEXICO] – (Michoacán): Zacapu – Lago de Zacapu
Reference: Pérez-Ponce de León et al. (Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)
Superfamily Diplogasteroidea Micoletzky, 1922
Family Diplogastridae Micoletzky, 1922
Longibucca Chitwood, 1933
Longibucca sp.
Infection site: digestive tracts
Host: Rhinoclemmys pulcherrima
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [MEXICO] – (Guerrero): Acapulco
Reference: Dyer and Carr (Reference Dyer and Carr1990a)
Superfamily Dracunculoidea Stiles, 1907
Family Dracunculidae Stiles, 1907
Dracunculus Reichard, 1759
Dracunculus globocephalus Mackin, 1927
Infection site: mesentery
Host: Chelydra serpentina
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [COSTA RICA] - Santa Rosa National Park
Reference: Bursey and Brooks (Reference Bursey and Brooks2011)
Superfamily Gnathostomatoidea Railliet 1895
Family Gnathostomatidae Railliet 1895
Ancyracanthus Diesing, 1838
Ancyracanthus pinnatifidus Diesing, 1839
Infection site: stomach, small intestine, and large intestine
Host: Peltocephalus dumerilianus 1, Podocnemis expansa,2 and Podocnemis unifilis 3
Biogeographic unit: Cerrado province2 – CD; Imerí province1 – BBD; Ucayali province3 – SBD
Political unity: [BRAZIL] – (Amazonas)1: unspecified location2, Barcelos and Jaú National Park; (Goiás): Britânia2; [PERU] – (Loreto): Pacaya-Samiria National Reserve3
References: Diesing (Reference Diesing1851)1,2; Gomes and Kohn (Reference Gomes and Kohn1970)1 cited by Vicente et al. (Reference Vicente, Rodrigues, Gomes and Pinto1993); Vicente et al. (Reference Vicente, Rodrigues, Gomes and Pinto1993); Nomura (Reference Nomura1996)2; Sanchez et al. (Reference Sánchez, Tantaleán, Vela and Méndez2006)3; Ferreira (Reference Ferreira2016)1; Menegaldo et al. (Reference Menegaldo, Arias-Pacheco, Perin, Tebaldi and Hoppe2024)1
Gnathostoma Owen, 1836
Gnathostoma binucleatum Almeyda-Artigas, 1991
Infection site: muscle
Host: Claudius angustatus 1, Kinosternon acutum 2, Kinosternon integrum 3, Staurotypus triporcatus,4 and Trachemys scripta 5
Biogeographic unit: Pacific Lowlands province3,5, Veracruzan province1,2,3,4,5 – MD
Political unity: [MEXICO] – (Guerrero): Laguna de Tres Palos3; (Nayarit): Agua Brava3,5; (Oaxaca): Temascal4; (Sinaloa): Tecualilla3,5; (Veracruz): Tlacotalpan2; (Oaxaca/Veracruz): Temescal/Tlacotalpan – Cenca del Papaloapan1,3,4,5
References: Almeyda-Artigas et al. (Reference Almeyda-Artigas, Hernández-Viciconti and Saldaña-Martínez1992)4 cited by Miguel (Reference Miguel2015); Alvarez Guerrero and Alba-Hurtado (Reference Alvarez-Guerrero and Alba-Hurtado2007)3,5; Sánchez-Miranda (Reference Sánchez-Miranda2001)3; Mosqueda-Cabrera (Reference Mosqueda-Cabrera2003)1,3,4,5; Gaspar-Navarro (Reference Gaspar-Navarro2004)2,3 cited by Pérez-Álvarez et al. (Reference Pérez- Álvarez, García-Prieto, Osorio-Sarabia, Lamothe-Argumedo and León-Règagnon2008); Díaz–Camacho et al. (Reference Díaz-Camacho, Cruz-Otero, Torres-Montoya, Sánchez-Gonzales, Delgado-Vargas and Nawa2010)3,5
Gnathostoma sp. larvae
Infection site: small intestine
Host: Hydromedusa tectifera
Political unity: [BRAZIL] – (Rio Grande do Sul): unspecified location
Reference: Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021)
Gnathostoma turgidum Stossich, 1902
Infection site: muscle
Host: Kinosternon integrum
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [MEXICO] – (Guerrero): Laguna de Tres Palos
References: Sánchez-Miranda (Reference Sánchez-Miranda2001); Mosqueda-Cabrera (Reference Mosqueda-Cabrera2003); Mosqueda-Cabrera et al. (Reference Mosqueda-Cabrera, Miranda, Calderón and Ortiz Nájera2009)
Spiroxys Schneider, 1866
Spiroxys contortus (Rudolphi, 1819) Schneider, 1866
Infection site: body cavity, esophagus, stomach, small intestine, large intestine, and heart
Host: Acanthochelys spixii 1, Hydromedusa tectifera 2, Kinosternon hirtipes 3, Phrynops geoffroanus 4, Phrynops hilarii 5, Trachemys dorbigni,6 and Trachemys scripta 7
Biogeographic unit: Caatinga province4, Chaco province2,5, Pampean province1,2,5,6 – CD; Esteros del Iberá province5,6,7 – PD; Transmexican Volcanic Belt province3 – MTZ; Yucatán Peninsula province7 – MD
Political unity: [ARGENTINA] – (Buenos Aires Province): Arroyo Buñirigo2,5, La Plata6 and unspecified location2,5; (Córdoba)2: Flor Serrana and Tanti; (Corrientes province): Bella vista5; (Entre Ríos): Concordia6 and Diamante5; (Santa Fe): Vera5; [BRAZIL] – (Paraíba) 4: Conceição and Patos; (Rio Grande do Sul)1,2,5: Capão do Leão6, Pelotas6, Rio Grande and Santa Vitória do Palmar; [MEXICO] – (Michoacán)3: Manantiales de Cointzio, Pátzcuaro – Lago de Pátzcuaro, Zacapu – Lago de Zacapu and Zirahuén - Lago de Zirahuén; (Yucatán): Aguada Xpoc7
References: Moravec and Vargas-Vázquez (Reference Moravec and Vargas-Vázquez1998)7; Pérez-Ponce de León et al. (Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)3; Mascarenhas et al. (Reference Mascarenhas, Souza, Coimbra and Müller2013)1,2; Mascarenhas and Müller (Reference Mascarenhas and Muller2015)6; Palumbo et al. (Reference Palumbo, Capasso, Cassano, Alcalde and Diaz2016)2,5; Castillo et al. (Reference Castillo, Acosta, González-Rivas and Ramallo2020)2,5; Chaviel et al. (Reference Chaviel, Mascarenhas, Bernardon, Coimbra and Müller2020)5; Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)6; Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)1,2,5; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2,5,6; Present study4
Spiroxys corti Caballero, 1935
Infection site: stomach
Host: Kinosternon hirtipes and Kinosternon integrum
Biogeographic unit: Balsas Basin province – MD
Political unity: [MEXICO] – (Puebla): Izúcar de Matamoros
Reference: Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008)
Spiroxys figueiredoi Freitas & Dobbin Jr., 1962
Infection site: stomach, small intestine, large intestine, and coelomic cavity
Host: Kinosternon leucostomum 1, Kinosternon scorpioides 2, Mesoclemmys tuberculata,3 and Phrynops geoffroanus 4
Biogeographic unit: – Atlantic province2,3,4, Parana Forest province,4 – PD; Caatinga province3,4 – CD; Pacific Lowlands province1,2 – MD; Pará province2, Roraima province2 – BBD
Political unity: [BRAZIL] – (Ceará): Nova Olinda3,4; (Maranhão): São Bento2; (Pará): Ilha de Marajó2; (Pernambuco)4: Limoeiro, Paudalho, Santa Cruz do Capibaribe and Recife2; (São Paulo): RPPN Foz do Rio Aguapeí4; (Sergipe)3,4: Areia Branca, Capela, Pirambu, Poço Redondo and Tobias Barreto; [COSTA RICA] – Santa Rosa National Park1,2
References: Freitas and Dobbin Jr. (Reference Freitas and Dobbin1962)2; Alho (Reference Alho1965)2; Freitas and Dobbin Jr. (Reference Freitas and Dobbin1971); Bursey and Brooks (Reference Bursey and Brooks2011)1,2; Silva (Reference Silva2014)4; Viana et al. (Reference Viana, Rodrigues, Madelaire, Santos and Sousa2016)2; Pereira A et al. (Reference Pereira, Brito, Araujo Filho, Teixeira, Teles, Santana, Lima and Almeida2018)3,4; Fonseca et al. (Reference Fonseca, Rojas, Vieira, Ferreira, Rodrigues, Barros, Silva Júnior, Moura and Oliveira2021)4; Santana et al. (Reference Santana, Teixeira, Araújo-Filho, Teles, Pereira, Brito, Almeida and Mesquita2023)3,4
Spiroxys sp.
Infection site: unknown
Host: Rhinoclemmys pulcherrima
Political unity: [COSTA RICA] – unspecified location
Reference: Brenes-Madrigal (Reference Brenes-Madrigal1961)
Spiroxys sp.
Infection site: stomach
Host: Mesoclemmys vanderhaegei
Biogeographic unit: Cerrado province – CD
Political unity: [BRAZIL] – (Mato Grosso): Chapada dos Guimarães
Reference: Ávila et al. (Reference Ávila, Brito, Barrella, Strussmann and Silva2010)
Spiroxys sp.
Infection site: stomach and small intestine
Host: Phrynops hilarii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Pelotas
Reference: Bernardon et al. (Reference Bernardon, Valente and Müller2013)
Spiroxys sp.
Infection site: stomach and small intestine
Host: Trachemys dorbigni
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Pelotas
Reference: Bernardon et al. (Reference Bernardon, Valente and Müller2014)
Spiroxys sp.
Infection site: stomach, small intestine, and large intestine
Host: Trachemys callirostris
Biogeographic unit: Magdalena province – PCD
Political unity: [COLOMBIA] – (Córdoba): Montería
Reference: Martínez et al. (Reference Martínez, Cordero, Velásquez, Bolaño, Otero, Monsalve, Mestra and Chacón-Pacheco2021)
Spiroxys triretrodens Caballero & Zerecero, 1943
Infection site: stomach
Host: Kinosternon hirtipes
Biogeographic unit: Balsas Basin province – MD
Political unity: [MEXICO] – (Puebla): Izúcar de Matamoros
Reference: Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008)
Superfamily Habronematoidea Ivaschkin, 1961
Family Hedruridae Railliet, 1916
Hedruris Nitzch, 1821
Hedruris dratini Palumbo, Servián, Sánchez & Diaz, Reference Palumbo, Servián, Sánchez and Diaz2019
Infection site: stomach and intestine
Host: Hydromedusa tectifera 1, Phrynops hilarii,2 and Trachemys dorbigni 3
Biogeographic unit: Pampean province – CD
Political unity: [ARGENTINA] – (Buenos Aires Province)1: City Bell – stream Rodríguez2, La Plata3, Villa Elisa – stream Carnaval, Ringuelet – stream El Gato and unspecified location
References: Palumbo et al. (Reference Palumbo, Servián, Sánchez and Diaz2019)1,2; Castillo et al. (Reference Castillo, Acosta, González-Rivas and Ramallo2020)1; Palumbo et al. (Reference Palumbo, Capasso, Cassano, Alcalde and Diaz2021)1; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)1,3
Hedruris oriestae Moniez, 1889
Infection site: stomach
Host: Hydromedusa tectifera 1 and Phrynops hilarii 2
Biogeographic unit: Pampean province – CD
Political unity: [ARGENTINA] – (Buenos Aires Province)1: Arroyo Buñirigo2 and unspecified location
References: Palumbo et al. (Reference Palumbo, Capasso, Cassano, Alcalde and Diaz2016)1,2; Castillo et al. (Reference Castillo, Acosta, González-Rivas and Ramallo2020)1
Hedruris siredonis Baird, 1858
Infection site: stomach and intestine
Host: Kinosternon hirtipes
Biogeographic unit: Transmexican Volcanic Belt province – MTZ
Political unity: [MEXICO] – (Michoacán): Manantiales de Cointzio and Zacapu – Lago de Zacapu
Reference: Pérez-Ponce de León et al. (Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)
Hedruris sp.
Infection site: digestive tracts
Host: Rhinoclemmys nasuta
Biogeographic unit: Western Ecuador province -–PCD
Political unity: [ECUADOR] – (Guayas): Estero el Ceibo
Reference: Dyer and Carr (Reference Dyer and Carr1990a)
Superfamily Oxyuroidea Weinland, 1858
Family Pharyngodonidae Travassos, Reference Travassos1919
Pharyngodonidae gen. sp.
Infection site: large intestine
Host: Acanthochelys spixii
Political unity: [BRAZIL] – (Rio Grande do Sul): unspecified location
Reference: Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021)
Pharyngodonidae gen. sp.
Infection site: large intestine
Host: Acanthochelys spixii and Hydromedusa tectifera
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande, Santa Vitória do Palmar
References: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Pharyngodonidae gen. sp.
Infection site: stomach
Host: Kinosternon scorpioides
Biogeographic unit: Chaco province – CD
Political unity: [ARGENTINA] - (Chaco): Miraflores
Reference: Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)
Pharyngodonidae gen. sp.
Infection site: stomach
Host: Acanthochelys pallidipectoris
Biogeographic unit: Yungas province – SBD
Political unity: [ARGENTINA] – (Salta): Joaquín Victor González
Reference: Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)
Pharyngodonidae gen. sp.
Infection site: stomach
Host: Kinosternon scorpioides
Biogeographic unit: Yungas province – SBD
Political unity: [ARGENTINA] – (Salta): Las Lajitas
Reference: Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)
Alaeuris Thapar, 1925
Alaeuris auricularis (Walton, 1942)
Infection site: unknown
Host: Chelonoidis niger (extinct)
Biogeographic unit: Galápagos Islands province – PCD
Political unity: [ECUADOR] – Galápagos Islands
Reference: Bouamer and Morand (Reference Bouamer and Morand2006)
Alaeuris macroptera (Walton, 1942)
Infection site: unknown
Host: Chelonoidis niger (extinct)
Biogeographic unit: Galápagos Islands province – PCD
Political unity: [ECUADOR] – Galápagos islands
Reference: Bouamer and Morand (Reference Bouamer and Morand2006)
Alaeuris pharyngodentata (Walton, 1942)
Infection site: unknown
Host: Chelonoidis niger (extinct)
Biogeographic unit: Galápagos Islands province – PCD
Political unity: [ECUADOR] – Galápagos islands
Reference: Bouamer and Morand (Reference Bouamer and Morand2006)
Tachygonetria Wedl, 1862
Tachygonetria testudinis Walton, 1942
Infection site: unknown
Host: Chelonoidis niger (extinct)
Biogeographic unit: Galápagos Islands province – PCD
Political unity: [ECUADOR] – Galápagos islands
Reference: Bouamer and Morand (Reference Bouamer and Morand2006)
Tachygonetria tetrapapillata Caballero, 1941
Infection site: large intestine
Host: Gopherus sp.
Biogeographic unit: Transmexican Volcanic Belt province – MTZ
Political unity: [MEXICO] – (Federal District): Azcapotzalco
References: Caballero (Reference Caballero1944b)
Thaparia Ortlepp, 1933
Thaparia contortospicula Walton, 1942
Infection site: unknown
Host: Chelonoidis niger (extinct)
Biogeographic unit: Galápagos Islands province – PCD
Political unity: [ECUADOR] – Galápagos islands
Reference: Bouamer and Morand (Reference Bouamer and Morand2006)
Thelandros Wedl, 1861
Thelandros sp.
Infection site: intestinal content
Host: Chelonoidis carbonarius
Biogeographic unit: Guianan Lowlands province – BBD
Political unity: [VENEZUELA] – (Bolivar): Upata
Reference: Pérez Mata et al. (Reference Pérez Mata, Anguirre, Piñango, Olivares, Camacho, Bautista and Núñez2014)
Thelandros sp.
Infection site: stomach
Host: Acanthochelys pallidipectoris
Biogeographic unit: Chaco province – CD
Political unity: [ARGENTINA] – (Santa Fé): Vera
Reference: Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)
Thelandros sp.
Infection site: stomach
Host: Acanthochelys pallidipectoris
Biogeographic unit: Yungas province – SBD
Political unity: [ARGENTINA] – (Salta): Las Lajitas
Reference: Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)
Thelandros sp.
Infection site: stomach
Host: Phrynops hilarii
Biogeographic unit: Chaco province – CD
Political unity: [ARGENTINA] – (Chaco): General José de San Martin
Reference: Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)
Superfamily Physalopteroidea Railliet, 1893
Family Physalopteridae Railliet, 1893
Physaloptera Rudolphi, 1819
Physaloptera retusa Rudolphi, 1819
Infection site: stomach, small intestine, and large intestine
Host: Phrynops geoffroanus and Mesoclemmys tuberculata
Biogeographic unit: Caatinga province – CD
Political unity: [BRAZIL] – (Ceará): Nova Olinda
Reference: Pereira A et al. (Reference Pereira, Junior, Santana, Almeida, Pereira and Oliveira2018)
Physaloptera sp. larvae
Infection site: stomach
Host: Phrynops geoffroanus
Biogeographic unit: Parana Forest province – PD
Political unity: [BRAZIL] – (São Paulo): RPPN Foz do Rio Aguapeí
Reference: Silva (Reference Silva2014)
Physaloptera sp. larvae
Infection site: stomach.
Host: Acanthochelys spixii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
Reference: Chaviel et al. (Reference Chaviel, Mascarenhas, Bernardon, Coimbra and Müller2020)
Physaloptera sp. larvae
Infection site: stomach
Host: Acanthochelys spixii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Physaloptera sp. larvae
Infection site: intestine
Host: Chelonoidis denticulatus
Biogeographic unit: Ucayali province – SBD
Political unity: [PERU] – (San Martin)
Reference: Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)
Physaloptera sp. larvae
Infection site: stomach
Host: Mesoclemmys tuberculata and Phrynops geoffroanus
Biogeographic unit: Atlantic province – PD; Caatinga province – CD
Political unity: [BRAZIL] – (Sergipe): Areia Branca, Capela, Pirambu, Poço Redondo and Tobias Barreto
Reference: Santana et al. (Reference Santana, Teixeira, Araújo-Filho, Teles, Pereira, Brito, Almeida and Mesquita2023)
Superfamily Strongyloidea Baird, 1853
Family Ancylostomatidae Looss, 1905
Ancylostomatidae g. sp.
Infection site: intestine
Host: Rhinoclemmys annulata and Rhinoclemmys nasuta
Biogeographic unit: Cauca province – PCD
Political unity: [COLOMBIA] – (Valle del Cauca): Cali
Reference: Sierra et al. (Reference Sierra, Ramírez and Osorio2013)
Family Strongylidae Baird, 1853
Chapiniella Yamaguti, 1961
Chapiniella variabilis (Chapin, Reference Chapin1924) Yamaguti, 1961
Infection site: intestine and large intestine
Host: Chelonoidis carbonarius 1 and Chelonoidis denticulatus 2
Biogeographic unit: Pará province1,2 – BBD; Ucayali province2 – SBD
Political unity: [BRAZIL] – (Pará): unspecified location2; (Piauí): Teresina1,2; [PERU] – (Loreto): Iquitos2; (San Martín): unspecified location2; (Ucayali): unspecified location2
References: Chapin (Reference Chapin1924)2; Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)2; Freire et al. (Reference Freire, Leal, Knoff, Gomes, Santos, Giese, Silva and Mendonça2017)1,2; Freire et al. (Reference Freire, Leal, Batista, Rocha, Baêta and Mendonça2019)1,2; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)2
Sauricola Chapin, Reference Chapin1924
Sauricola sauriciola Chapin, Reference Chapin1924
Infection site: stomach, small intestine, and large intestine
Host: Chelonoidis denticulatus 1 and Rhinoclemmys annulata 2
Biogeographic unit: Pacific Lowlands province2 – MD; Ucayali province1 – SBD
Political unity: [BRAZIL] – (Pará): unspecified location1; [COSTA RICA] – Santa Rosa National Park2; [PERU] – (Loreto): Iquitos1
References: Chapin (Reference Chapin1924)1; Bursey and Brooks (Reference Bursey and Brooks2011)2; Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)1
Sauricola sp.
Infection site: intestine (intestinal content).
Host: Chelonoidis denticulatus
Biogeographic unit: Desert province – STZ
Political unity: [PERU] – Lima
Reference: Chavez et al. (Reference Chavez, Serrano-Martínez, Tantaleán, Quispe and Casas2015)
Sauricola sp.
Infection site: intestine and stomach
Host: Chelonoidis denticulatus
Political unity: [PERU] – (Ucayali) – unspecified location
Reference: Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)
Strongylus Müller, 1780
Strongylus sp.
Infection site: intestine (intestinal content)
Host: Chelonoidis denticulatus
Biogeographic unit: Desert province – STZ
Political unity: [PERU] – Lima
Reference: Chavez et al. (Reference Chavez, Serrano-Martínez, Tantaleán, Quispe and Casas2015)
Superfamily Strongyloidoidea Chitwood & McIntosh, 1934
Family Strongyloididae Chitwood & McIntosh, 1934
Strongyloides Grassi, 1879
Strongyloides sp.
Infection site: intestinal tract
Host: Chelonoidis carbonarius, Kinosternon leucostomum, Rhinoclemmys annulata, and Trachemys scripta (captivity)
Biogeographic unit: Cauca province – PCD
Political unity: [COLOMBIA] – (Valle del Cauca): Cali
Reference: Sierra et al. (Reference Sierra, Ramírez and Osorio2013)
Strongyloides sp. larvae
Infection site: digestive system
Host: Chelonoidis carbonarius and Chelonoidis denticulatus
Biogeographic unit: Atlantic province – PD
Political unity: [BRAZIL] – (Rio de Janeiro): Rio de Janeiro
Reference: Meireles et al. (Reference Meireles, Miguel, Mattos Junior, Barros and Magalhães2018)
CLASS ENOPLEA INGLIS, 1983
Superfamily Dioctophymoidea Railliet, 1915
Family Dioctophymidae Railliet, 1915
Dioctophyme Collet-Meygret, 1802
Dioctophyme renales (Goeze, 1782) Collet-Meygret, 1802
Infection site: body cavity, muscles, mesentery, stomach, stomach serosa, small intestine, and surface of various organs: esophagus, heart, lungs, liver, pancreas, spleen, and intestines
Host: Acanthochelys spixii 1, Hydromedusa tectifera 2, Phrynops hilarii,3 and Trachemys dorbigni 4
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul)1,2,3: Capão do Leão4, Pelotas4, Rio Grande and Santa Vitória do Palmar
References: Mascarenhas and Müller (Reference Mascarenhas and Muller2015a)4; Mascarenhas et al. (Reference Mascarenhas, Henzel and Müller2017)3; Chaviel et al. (Reference Chaviel, Mascarenhas, Bernardon, Coimbra and Müller2020)1; Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)4; Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)1,2,3
Eustrongylides Jägerskiöld, 1909
Eustrongylides sp. larvae
Infection site: stomach
Host: Trachemys dorbigni
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão and Pelotas
Reference: Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)
Superfamily Trichinelloidea Ward, 1907
Family Capillariidae Railliet, 1915
Capillaria Zeder, 1800
Capillaria sp.
Infection site: intestine
Host: Kinosternon cruentatum
Biogeographic unit: Puntarenas-Chiriquí province – PCD
Political unity: [COSTA RICA] – (San Jose): Santa Ana
Reference: Brenes-Madrigal and Bravo-Hollis (Reference Brenes-Madrigal and Bravo-Hollis1960)
PHYLUM PLATYHELMINTHES Gegenbaur, 1859
SUBPHYLUM RHABDITOPHORA
SUPERCLASS: NEODERMATA
CLASS CESTODA Van Beneden, 1849
ORDER ONCHOPROTEOCEPHALIDEA
Family Proteocephalidae La Rue, 1911
Ophiotaenia La Rue, 1911
Ophiotaenia cohospes Cordero, Reference Cordero1946
Infection site: intestine
Host: Hydromedusa tectifera
Biogeographic unit: Chaco province2, Pampean province2 – CD; Esteros del Iberá province1 – PD
Political unity: [ARGENTINA] – (Buenos Aires): Magdalena2; (Córdoba)2: Tanti and Flor Serrana; [URUGUAY] – (Montevideo): Montevideo city1
References: Cordero (Reference Cordero1946)1; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2
Ophiotaenia lopesi Rego, 1967
Infection site: small intestine
Host: Chelonoidis denticulatus
Political unity: [BRAZIL] – (Pará): unspecified location
References: Schmidt (Reference Schmidt1986) cited by Mascarenhas and Müller (Reference Mascarenhas and Muller2021); Muniz-Pereira et al. (Reference Muniz-Pereira, Vieira and Luque2009)
Ophiotaenia sp.
Infection site: small intestine
Host: Hydromedusa tectifera
Political unity: [BRAZIL] – (Rio Grande do Sul): unspecified location
Reference: Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021)
Ophiotaenia sp.
Infection site: small intestine
Host: Hydromedusa tectifera
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
CLASS TREMATODA Rudolphi, 1808
SUBCLASS DIGENEA
ORDER DIPLOSTOMIDA
Superfamily Diplostomoidea Poirier, 1886
Family Diplostomidae Poirier, 1886
Posthodiplostomum Dubois, Reference Dubois1936
Posthodiplostomum minimum (MacCallum, 1921) Dubois, Reference Dubois1936
Infection site: intestine
Host: Kinosternon hirtipes
Biogeographic unit: Transmexican Volcanic Belt province – MTZ
Political unity: [MEXICO] – (Michoacán): Lago de Pátzcuaro
References: Pérez-Ponce de León (Reference Pérez Ponce de León1992); Pérez-Ponce de León et al. (Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)
Family Liolopidae Odhner, Reference Odhner1912
Helicotrema Odhner, Reference Odhner1912
Helicotrema asymmetrica Travassos, 1922
Infection site: small intestine
Host: Rhinoclemmys melanosterna
Biogeographic unit: Guatuso-Talamanca province – PCD
Political unity: PANAMA – (Panama province): Panamá city
References: Caballero et al. (Reference Caballero, Zerecero and Grocott1958)
Helicotrema spirale (Diesing, Reference Diesing1850 ) Odhner, Reference Odhner1912
Infection site: stomach, intestine, small intestine
Host: Chelonoidis denticulatus 1 and Peltocephalus dumerilianus 2
Biogeographic unit: Imerí province2 – BBD; Ucayali province1 – SBD
Political unity: [BRAZIL] – unspecified location1,2; (Amazonas)2: Barcelos and Jaú National Park; [PERU] – (Loreto): Iquitos1
References: Odhner (Reference Odhner1912)1,2 cited by Fernandes and Kohn (Reference Fernandes and Kohn2014); Tantaleán and Forlong (Reference Tantaleán and Forlong2013)1; Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)1; Ferreira (Reference Ferreira2016)1,2; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)1; Menegaldo et al. (Reference Menegaldo, Arias-Pacheco, Perin, Tebaldi and Hoppe2024)2
Family Proterodiplostomidae Dubois, Reference Dubois1936
Herpetodiplostomum Dubois, Reference Dubois1936
Herpetodiplostomum argentinense (Palumbo & Diaz, Reference Palumbo and Diaz2018) Achatz, Brito, Fecchio & Tkach, Reference Achatz, Brito, Fecchio and Tkach2021
Synonym: Cheloniodiplostomum argentinensis Palumbo & Diaz, Reference Palumbo and Diaz2018
Infection site: intestine
Host: Phrynops hilarii
Biogeographic unit: Esteros del Ibera province1,2 – PD; Pampean province2 – CD
Political unity: [ARGENTINA] – (Buenos Aires Province): Arroyo Buñirigo1; La Plata2; (Entre Rios)2: Aldea Brasilera and Diamante; (Santa Fe): Helvecia2
References: Palumbo and Diaz (Reference Palumbo and Diaz2018)1; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2
Herpetodiplostomum breve (MacCallum, 1921) Zerecero, Reference Zerecero1947
Synonym: Cheloniodiplostomum breve (MacCallum, 1921) Sudarikov, 1960
Infection site: intestine
Host: Kinosternon scorpioides 1 and Phrynops geoffroanus 2
Biogeographic unit: Trinidad province1 – PCD
Political unity: [COLOMBIA] - unspecified location2; [TRINIDAD AND TOBAGO – West Indies] – Trinidad island1
References: Dubois (Reference Dubois1979)
Herpetodiplostomum delillei Zerecero, Reference Zerecero1947
Synonym: Cheloniodiplostomum delillei (Zerecero, Reference Zerecero1947) Sudarikov, 1960
Infection site: intestine, small intestine
Host: Chelydra serpentina 1 and Claudius angustatus 2
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Tabasco): Río Teapa1,2; (Veracruz)1: Laguna de Alvarado and unspecified location
References: Zerecero (Reference Zerecero1947)1; Thatcher (Reference Thatcher1963a)1,2; Dubois (Reference Dubois1979)1
Herpetodiplostomum duboisi Mañé-Garzón & Holcman-Spector, Reference Mañé-Garzón and Holcman-Spector1969
Infection site: intestine, small intestine
Host: Phrynops hilarii 1 and Phrynops williamsi 2
Biogeographic unit: Esteros del Iberá province1, Parana Forest province2 – PD
Political unity: [ARGENTINA] – (Misiones): Comandante Andresito2; [URUGUAY] – (Tacuarembó): Paso de los Toros1
References: Mañé-Garzón & Holcman (Reference Mañé-Garzón and Holcman-Spector1969)1; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2
Herpetodiplostomun ovalis Mañé-Garzón & Holcman-Spector, Reference Mañé-Garzón and Holcman-Spector1969
Infection site: small intestine
Host: Hydromedusa tectifera
Biogeographic unit: Pampean province – CD
Political unity: [URUGUAY] – (Canelones Departament): Arroyo de as Piedras and Banãdo Tropa Vieja
References: Mañé-Garzón and Holcman (Reference Mañé-Garzón and Holcman-Spector1969)
Herpetodiplostomum sp.
Infection site: small intestine and large intestine
Host: Acanthochelys spixii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
References: Mascarenhas et al. (Reference Mascarenhas, Bernardon and Muller2016); Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Herpetodiplostomum sp.
Infection site: small intestine
Host: Hydromedusa tectifera
Political unity: [BRAZIL] – (Rio Grande do Sul): unspecified location
Reference: Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021)
Herpetodiplostomum sp.
Infection site: intestine
Host: Hydromedusa tectifera and Phrynops hilarii
Biogeographic unit: Pampean province – CD
Political unity: [ARGENTINA] – (Buenos Aires): La Plata
Reference: Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)
Herpetodiplostomum sp.
Infection site: small intestine
Host: Hydromedusa tectifera
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande, Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Herpetodiplostomum sp.
Infection site: intestine
Host: Kinosternon sp.
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Oaxaca): Presa de Temascal
References: Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008)
Herpetodiplostomum sp.
Infection site: small intestine
Host: Phrynops geoffroanus
Biogeographic unit: Parana Forest province – PD
Political unity: [BRAZIL] – (Minas Gerais): Juiz de Fora
Reference: Novelli et al. (Reference Novelli, Sousa, Lima and Vieira2013)
Herpetodiplostomum sp.
Infection site: small intestine, large intestine
Host: Phrynops hilarii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Pelotas
Reference: Bernardon et al. (Reference Bernardon, Valente and Müller2013)
Herpetodiplostomum sp.
Infection site: small intestine
Host: Trachemys dorbigni
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – Rio Grande do Sul: Pelotas
Reference: Bernardon et al. (Reference Bernardon, Valente and Müller2014)
Herpetodiplostomum testudinis Dubois, Reference Dubois1936
Synonym: Cheloniodiplostomum testudinis (Dubois, Reference Dubois1936) Sudarikov, 1960
Infection site: small intestine and large intestine
Host: Hydromedusa tectifera 1, Phrynops geoffroanus 2, Phrynops hilarii,3 and Testudo sp.4
Biogeographic unit: Atlantic province2, Esteros del Ibera province1,3, Parana Forest province2 – PD; Caatinga province2, Pampean province3 – CD
Political unity: [ARGENTINA] – (Buenos Aires Province): Arroyo Buñirigo1,3; (Corrientes Province): unspecified location3; (Santa Fe)3: Cañada de Gomez and Santa Fe City; [BRAZIL] – unspecified location4; (Pernambuco)2: Cordeiro, Limoeiro, Paudalho and Santa Cruz do Capibaribe; (Rio Grande do Sul)3: Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar; (São Paulo): RPPN Foz do Rio Aguapeí2
References: Dubois (Reference Dubois1936)4; Lombardero & Moriena (Reference Lombardero and Moriena1977)3; Dubois (Reference Dubois1979)4; Silva (Reference Silva2014)2; Mascarenhas et al. (Reference Mascarenhas, Bernardon and Muller2016)3; Palumbo and Diaz (Reference Palumbo and Diaz2018)1,3; Fonseca et al. (Reference Fonseca, Rojas, Vieira, Ferreira, Rodrigues, Barros, Silva Júnior, Moura and Oliveira2021)2; Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)3; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)3
Herpetodiplostomum vogti Achatz, Brito, Fecchio & Tkach, Reference Achatz, Brito, Fecchio and Tkach2021
Infection site: small intestine
Host: Phrynops geoffroanus
Biogeographic unit: Cerrado province – CD
Political unity: [BRAZIL] – (Mato Grosso): Cuiabá
Reference: Achatz et al. (Reference Achatz, Brito, Fecchio and Tkach2021)
Herpetodiplostomum wolffhugeli Mañé-Garzón & Holcman-Spector, Reference Mañé-Garzón and Holcman-Spector1969
Infection site: small intestine
Host: Acanthochelys spixii
Biogeographic unit: Pampean province1 – CD
Political unity: [URUGUAY] – (Rocha Departament): La Coronilla1; (Tacuarembó): Rio Negro
Reference: Mañé-Garzón and Holcman (Reference Mañé-Garzón and Holcman-Spector1969)
Superfamily Schistosomatoidea Stiles & Hassall, 1898
Family Atamatamidae Bullard & Dutton, 2022
Atamatam Bullard & Roberts in Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant & Halanych, Reference Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant and Halanych2019
Atamatam amazoniensis Bullard & Roberts in Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant & Halanych, Reference Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant and Halanych2019
Infection site: blood vessels of kidney and mesentery
Host: Chelus fimbriata
Biogeographic unit: Ucayali province – SBD
Political unity: [PERU] – (Loreto): Iquitos
Reference: Bullard et al. (Reference Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant and Halanych2019)
Paratamatam Bullard & Roberts in Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant & Halanych, Reference Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant and Halanych2019
Paratamatam iquitosiensis Bullard & Roberts in Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant & Halanych, Reference Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant and Halanych2019
Infection site: blood vessels of kidney and mesentery
Host: Chelus fimbriata
Biogeographic unit: Ucayali province – SBD
Political unity: [PERU] – (Loreto): Iquitos
Reference: Bullard et al. (Reference Bullard, Roberts, Warren, Dutton, Whelan, Ruiz, Platt, Tkach, Brant and Halanych2019)
Pitiutrema Dutton & Bullard in Dutton et al. Reference Dutton, Warren and Bullard2019
Pitiutrema revelae Dutton & Bullard in Dutton et al. Reference Dutton, Warren and Bullard2019
Infection site: heart
Host: Podocnemis sextuberculata
Political unity: [PERU] – Amazon River Basin
Reference: Dutton et al. (Reference Dutton, Warren and Bullard2019)
Family Plattidae Bullard & Dutton, 2022
Hapalorhynchus Stunkard, 1922
Hapalorhynchus albertoi Lamothe-Argumedo, Reference Lamothe-Argumedo1978
Infection site: blood vessels
Host: Kinosternon leucostomum
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Tabasco): Villahermosa
References: Lamothe-Argumedo (Reference Lamothe-Argumedo1978)
ORDER PLAGIORCHIIDA
Superfamily Echinostomatoidea Looss, 1902
Family Echinostomatidae Looss, 1899
Echinostomatidae gen. sp.
Infection site: esophagus, lungs, stomach, and intestine
Host: Mesoclemmys tuberculata and Phrynops geoffroanus
Biogeographic unit: Atlantic province – PD; Caatinga province – CD
Political unity: [BRAZIL] – (Sergipe): Areia Branca, Capela, Pirambu, Poço Redondo and Tobias Barreto
Reference: Santana et al. (Reference Santana, Teixeira, Araújo-Filho, Teles, Pereira, Brito, Almeida and Mesquita2023)
Prionosomoides Teixeira de Freitas & Dobbin, Reference Freitas and Dobbin1967
Prionosomoides phrynopsis (Mané-Garzon & Gil, 1961) Teixeira de Freitas & Dobbin, Reference Freitas and Dobbin1967
Infection site: intestine, small intestine
Host: Phrynops hilarii
Biogeographic unit: Esteros del Ibera province1 – PD; Pampean province 2 – CD
Political unity: [ARGENTINA] – (Buenos Aires): Magdalena2; (Entre Rios)2: Diamante and Gualeguaychú; (Santa Fe)2: Rafaela and Santa Fe City; (Tucumán): San Miguel de Tucumán2; [URUGUAY] - (Tacuarembó): Paso de los Toros1
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1961a)1; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2
Prionosomoides scalaris Teixeira de Freitas & Dobbin, 1967
Infection site: small intestine and esophagus
Host: Phrynops geoffroanus1 and Phrynops hilarii 2
Biogeographic unit: Atlantic province1 – PD; Caatinga province1 – CD
Political unity: [ARGENTINA] – (Corrientes Province): unspecified location2; [BRAZIL] – (Pernambuco)1: Cordeiro, Limoeiro, Paudalho, Santa Cruz do Capibaribe and Recife
References: Freitas and Dobbin Jr. (Reference Freitas and Dobbin1967)1; Lombardero and Moriena (Reference Lombardero and Moriena1977)2 cited by Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021); Fonseca et al. (Reference Fonseca, Rojas, Vieira, Ferreira, Rodrigues, Barros, Silva Júnior, Moura and Oliveira2021)1
Prionosomoides sp.
Infection site: small intestine
Host: Phrynops hilarii
Political unity: [BRAZIL] – (Rio Grande do Sul): unspecified location
Reference: Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021)
Prionosomoides sp.
Infection site: small intestine
Host: Phrynops hilarii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Family Rhytidodidae Odhner, 1926
Rhytidodes Looss, 1901
Rhytidodes gelatinosus (Rudolphi, 1819) Looss, 1901
Infection site: intestine
Host Podocnemis expansa
Political unity: [BRAZIL] – unspecified location
Reference: Travassos et al. (Reference Travassos, Freitas and Kohn1969)
Superfamily Gorgoderoidea Looss, 1901
Family Braunotrematidae Yamaguti, 1958
Braunotrema Price, 1930
Braunotrema pulvinatum (Braun, 1899) Price, 1930
Infection site: small intestine
Host: Podocnemis expansa
Political unity: [BRAZIL] – (Pará): Amazon river
Reference: Lent and Freitas (Reference Lent and Freitas1938)
Family Dicrocoeliidae Looss, 1899
Infidum Travassos, 1916
Infidum similis Travassos, 1916
Infection site: bile duct and gallbladder
Host: Staurotypus triporcatus
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Tabasco): Rio Teapa
Reference: Thatcher (Reference Thatcher1963b)
Family Gorgoderidae Looss, 1899
Gorgoderina Looss, 1899
Gorgoderina sp.
Infection site: small intestine
Host: Phrynops geoffroanus
Biogeographic unit: Caatinga province – CD
Political unity: [BRAZIL] – (Ceará): Nova Olinda
Reference: Pereira A et al. (Reference Pereira, Brito, Araujo Filho, Teixeira, Teles, Santana, Lima and Almeida2018)
Superfamily Heronimoidea Ward, 1918
Family Heronimidae Ward, 1918
Heronimus MacCallum, 1902
Heronimus mollis (Leidy, 1856) Stunkard, 1954
Note: Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008) mention that this species was recorded as Heronimus chelydrae, which is considered a synonym of Heronimus mollis (Yamaguti, 1971)
Infection site: lungs and small intestine
Host: Claudius angustatus 1, Kinosternon cruentatum 2, Kinosternon hirtipes 3, Kinosternon leucostomum 4, Kinosternon scorpioides,5 and Trachemys callirostris 6
Biogeographic unit: Guatuso-Talamanca province5; Magdalena province6, Puntarenas-Chiriquí province2 – PCD; Transmexican Volcanic Belt province3 – MTZ; Veracruzan province1,4 – MD
Political unity: [COLOMBIA] - (Bolívar)6: Coyongal and San Nicolás; (Magdalena): Ricaurte6; [COSTA RICA] – (San José): Santa Ana2; [MEXICO] – (Michoacán): Uruapan3; (Tabasco)4: Río Teapa1 and Villahermosa; (Veracruz): Laguna de Alvarado4; [PANAMA] – (Panama province) – Panamá City5
References: Caballero (Reference Caballero1940a)1,3; Herrera-Rosales (Reference Herrera-Rosales1951)4; Caballero and Brenes-Madrigal (Reference Caballero and Brenes-Madrigal1957)2; Caballero et al. (Reference Caballero, Zerecero and Grocott1958)5; Thatcher (Reference Thatcher1963a)1,4; Parra-Rojas (Reference Parra-Rojas1983)4; Lenis and Vélez (Reference Lenis and Vélez2011)6
Superfamily Opisthorchioidea Looss, 1899
Family Cryptogonimidae Ward, 1917
Caimanicola Freitas & Lent, 1938
Caimanicola brauni (Mañé-Garzón & Gil, 1961) Brooks, 1980
Infection site: intestine, small intestine
Host: Acanthochelys spixii 1 and Phrynops hilarii 2
Biogeographic unit: Esteros del Ibera province2 – PD; Pampean province – CD1,2
Political unity: [ARGENTINA] – (Buenos Aires): Magdalena2; (Santa fé): Rafaela2; [BRAZIL] – (Rio Grande do Sul)1.2: Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar; [URUGUAY] - (Tacuarembó): Paso de los Toros2
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1961a)2; Chaviel et al. (Reference Chaviel, Mascarenhas, Bernardon, Coimbra and Müller2020)1,2; Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)1,2; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2
Timoniella Rebecq, 1960
Timoniella ostrowskiae Brooks & Holcman, Reference Brooks and Holcman1993
Infection site: small intestine
Host: Phrynops hilarii
Biogeographic unit: Esteros del Iberá province2 – PD; Pampean province3,4 – CD
Political unity: [BRAZIL] – (Rio Grande do Sul)3,4: Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar; [URUGUAY] – (Tacuarembó): Paso de los Toros2 and unspecified location1
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1961a)1; Brooks and Holcman (Reference Brooks and Holcman1993)2; Chaviel et al. (Reference Chaviel, Mascarenhas, Bernardon, Coimbra and Müller2020)3; Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)4
Timoniella scyphocephala (Braun, 1899) Brooks, 1980
Synonym : Acanthochasmus scyphocephalus (Braun, 1899) Braun, Reference Braun1901; Proctocaecum scyphocephalum (Braun, 1899)
Infection site: intestine
Host: Chelus fimbriata
Political unity: [BRAZIL] – unspecified location
References: Ostrowski de Núñez (Reference Ostrowski de Núñez1986)
Superfamily Paramphistomoidea Fischoeder, 1901
Family Cladorchiidae Fischoeder, 1901
Caballerodiscus Sey, 1988
Caballerodiscus resupinatus (Caballero y C., 1940)
Synonym: Schizamphistomoides resupinatus Caballero y C., 1940
Infection site: large intestine
Host: Dermatemys mawii
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado1 and Tlacotalpan2; (Tabasco): Río Teapa4 and Villahermosa3
References: Caballero (Reference Caballero1940b)1; Caballero (Reference Caballero1942)2; Caballero-Rodríguez (Reference Caballero-Rodríguez1960)3; Thatcher (Reference Thatcher1963a)4 cited by Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008)
Caballerodiscus tabascensis (Caballero y C. & Sokoloff, 1934) Sey, 1988
Synonym : Schizamphistomoides tabascensis Caballero y. C. & Sokoloff, 1934
Infection site: stomach, small intestine, large intestine, and cloaca
Host: Dermatemys mawii 1, Kinosternon leucostomum 2, Terrapene ornata 3, Trachemys ornata,4 and Trachemys scripta 5
Biogeographic unit: Veracruzan province1,2,3,5 – MD; Guatuso-Talamanca province4 – PCD
Political unity: [MEXICO] – (Tabasco)1: Comalcalco, Río Teapa2,5 and Villahermosa; (Veracruz)1: Laguna de Alvarado3, Río Blanco and Tlacotalpan; [PANAMA] – (Panama province): Panamá city4
References: Caballero (Reference Caballero1938)3; Caballero (Reference Caballero1943a)1; Herrera-Rosales (Reference Herrera-Rosales1951)1; Caballero et al. (Reference Caballero, Zerecero and Grocott1958)4; Caballero-Rodríguez (Reference Caballero-Rodríguez1960)1; Thatcher (Reference Thatcher1963a)2,5; Thatcher (Reference Thatcher1964)1 cited by Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008); Parra-Rojas (Reference Parra-Rojas1983)1; Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008)1
Halltrema Lent & Teixeira de Freitas, 1939
Halltrema avitellinum Lent & Teixeira de Freitas, 1939
Infection site: digestive tract, stomach, intestine
Host: Chelonoidis denticulatus 1, Podocnemis expansa 2, Podocnemis sp.,3 and Podocnemis unifilis 4
Biogeographic unit: Cerrado province2 – CD; Roraima province1 – BBD; Sabana province – PCD3; Ucayali province1,2,4 – SBD
Political unity: [BRAZIL] – unspecified location2; (Pará): Ilha do Marajó1 and unspecified location2; (Tocantins): Javaés River2; [PERU]: (Ucayali): unspecified location1; (Loreto): Iquitos1,2,4; [VENEZUELA] – (Guarico): Orinoco river3
References: Lent and Freitas (Reference Lent and Freitas1939)2; Freitas and Lent (Reference Freitas and Lent1942)2; Caballero and Diaz-Ungria (Reference Caballero and Diaz-Ungria1958)3; Alho (Reference Alho1965)1; Armond (Reference Armond2008)2; Tantaleán et al. (Reference Tantaleán, Sánchez and Pineda Catalan2011)2,4; Julca et al. (Reference Julca, Casas, Chavera, Sánchez, Sánchez and Batalla2014)1; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)1
Halltrema heteroxenum (Cordero & Vogelsang, Reference Cordero and Vogelsang1940) Jones, 2005
Infection site: digestive tract, stomach
Host: Podocnemis sp1., Podocnemis unifilis2, and Rhinoclemmys nasuta 3
Biogeographic unit: Ucayali province2 – SBD; Western Ecuador province3 – PCD
Political unity: [ECUADOR] – (Provincia de Esmeraldas)3; [PERU] – (Loreto): Iquitos2; [VENEZUELA] – (Guarico): unspecified location1
References: Cordero and Vogelsang (Reference Cordero and Vogelsang1940)1 cited by Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021); Dyer and Carr (Reference Dyer and Carr1990b)3; Tantaleán and Forlong (Reference Tantaleán and Forlong2013)2
Nematophila Travassos, 1934
Nematophila argentinum (Cordero & Vogelsang, Reference Cordero and Vogelsang1940) Lenis & Vélez, Reference Lenis and Vélez2011
Infection site: mouth, esophagus, stomach, small intestine, large intestine, and cloaca
Host: Phrynops sp.1, Podocnemis lewyana,2 and Trachemys callirostris 3
Biogeographic unit: Magdalena province2,3 – PCD
Political unity: [ARGENTINA] – (Misiones Province): unspecified location1; (Córdoba): Montería3; [COLOMBIA] – (Bolívar)2,3: Coyongal and San Nicolás
References: Cordero and Vogelsang (Reference Cordero and Vogelsang1940)1 cited by Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021); Lenis and Vélez (Reference Lenis and Vélez2011)2,3; Martínez et al. (Reference Martínez, Cordero, Velásquez, Bolaño, Otero, Monsalve, Mestra and Chacón-Pacheco2021)3
Nematophila grandis (Diesing, 1839) Travassos, 1934
Infection site: stomach, small intestine, large intestine
Host: Chelonoidis denticulatus 1, Chelus fimbriata 2, Hydromedusa tectifera 3, Kinosternon scorpioides 4, Mesoclemmys gibba 5, Mesoclemmys nasuta 6, Peltocephalus dumerilianus 7, Phrynops geoffroanus 8, Phrynops hilarii 9, Podocnemis erythrocephala 10, Podocnemis expansa 11, Podocnemis sp.12, Podocnemis unifilis 13, Podocnemis vogli 14, Rhinoclemmys areolata 15, Rhinoclemmys melanosterna 16, Rhinoclemmys nasuta,17 and Rhinoclemmys punctularia 18
Biogeographic unit: Atlantic province8, Parana Forest province8 – PD; Caatinga province8, Cerrado province13; Pampean province9 – CD; Sabana province12, Guatuso-Talamanca province16, Western Ecuador province17, – PCD; Ucayali province11,13 – SBD; Guianan Lowlands province18, Imerí province7,13, Pará province4,18, Roraima province4,13,18 – BBD; Veracruzan province15 – MD
Political unity: [ARGENTINA] – (Corrientes Province): unspecified location9; (Buenos Aires province): Ciudad Autónoma de Buenos Aires9; [BRAZIL] - unspecified location2,5,6,7,8,10,11; (Amazonas)7: Barcelos and Jaú National Park; (Pará)4,18: Bajuru river and Ilha do Marajó; (Pernambuco)8: Cordeiro, Limoeiro, Paudalho and Santa Cruz do Capibaribe; (São Paulo): RPPN Foz do Rio Aguapeí8; (Tocantins)13: Araguaia National Park/Cantão State Park - Javaés river and Tupiratins/Itapiratins - Tocantins river; [ECUADOR] – (Esmeraldas Province): unspecified location17; [FRENCH GUIANA]: Iracoubo18; [MEXICO] - (Chiapas): Catazajá15; (Tabasco): Emiliano Zapata15; [PANAMA] – (Panama province):Panamá city16; [PARAGUAY]- unspecified location3; [PERU] – (San Martin): unspecified location1; (Loreto)13: Iquitos11, Pacaya-Samiria National Reserver, Putumayo River and Samiria River; (Manu province): Madre de Dios river13; (Ucayali): unspecified location7; [VENEZUELA] - unspecified localities11,13,14; (Guarico)12: Apurito river, Bolivar city and Orinoco river; (Zulia): unspecified location4
References: Diesing (Reference Diesing1850)10; Travassos (Reference Travassos1934b)2,5,6,7.8; Caballero et al. (Reference Caballero, Zerecero and Grocott1958)16; Caballero and Diaz-Ungria (Reference Caballero and Diaz-Ungria1958)12, Heyneman et al. (Reference Heyneman, Brenes and Diaz-Ungria1960)13,14; Alho (Reference Alho1964b)4,18; Travassos et al. (Reference Travassos, Freitas and Kohn1969)2,8,11; Masi-Pallarés et al. (Reference Masi-Pallarés, Benitez-Usher and Maciel1976)3 cited by Fernandes and Kohn (Reference Fernandes and Kohn2014); Lombardero and Moriena (Reference Lombardero and Moriena1977)9; Díaz Ungría (Reference Díaz-Ungría1978)4,11; Dyer and Carr (Reference Dyer and Carr1990b)15,17,18; Salizar and Sanchez (Reference Salizar and Sanchez2004)13; Sánchez et al. (Reference Sánchez, Tantaleán, Vela and Méndez2006)13; Tantaleán et al. (Reference Tantaleán, Sánchez and Pineda Catalan2011)11,13; Silva (Reference Silva2014)8; Ferreira (Reference Ferreira2016)7; Fonseca et al. (Reference Fonseca, Rojas, Vieira, Ferreira, Rodrigues, Barros, Silva Júnior, Moura and Oliveira2021)8; Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022)1,7,13; Ribeiro et al. (Reference Ribeiro, Santana, Souza, Oliveira, Magalhães, Malvasio and Silva2023)13; Menegaldo et al. (Reference Menegaldo, Arias-Pacheco, Perin, Tebaldi and Hoppe2024)7; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)9
Nematophila venezuelensis (Cordero & Vogelsang, Reference Cordero and Vogelsang1940) Yamaguti, 1958
Infection site: stomach
Host: Podocnemis lewyana 1 and Podocnemis sp.2
Biogeographic unit: Magdalena province1 – PCD
Political unity: [COLOMBIA] – (Bolívar): San Nicolás1; (Magdalena): Ricaurte1; [VENEZUELA] – (Bolívar): unspecified location2; (Guarico): unspecified location2
References: Cordero and Vogelsang (Reference Cordero and Vogelsang1940)2 cited by Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021); Lenis and Vélez (Reference Lenis and Vélez2011)1
Oriximinatrema Knoff, Brooks, Mullins & Gomez, 2012
Oriximinatrema noronhae Knoff, Brooks, Mullins & Gomez, 2012
Infection site: stomach, intestine
Host: Podocnemis expansa
Biogeographic unit: Roraima province – BBD
Political unity: [BRAZIL] - (Pará): Oriximiná
Reference: Knoff et al. (Reference Knoff, Brooks, Mullins and Gomes2012)
Parachiorchis Caballero, 1943
Parachiorchis parviacetabulatus Caballero, 1943
Infection site: large intestine
Host: Dermatemys mawii
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado
References: Caballero (Reference Caballero1943a)
Pseudocleptodiscus Caballero, Reference Caballero1961
Pseudocleptodiscus margaritae Caballero, Reference Caballero1961
Note: Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008) mention that this species was originally described as Pseudocleptodiscus bravoae (Caballero-Rodríguez, Reference Caballero-Rodríguez1960) and Dadaytrema sphaerorchidum (Thatcher, 1963)
Infection site: small intestine, large intestine
Host: Dermatemys mawii 1 and Rhinoclemmys nasuta 2
Biogeographic unit: Veracruzan province1 – MD; Western Ecuador province2 – PCD
Political unity: [ECUADOR] - (Esmeraldas Province)2; [MEXICO] - (Tabasco)1: Río Teapa and Villahermosa
References: Caballero-Rodríguez (Reference Caballero-Rodríguez1960)1; Caballero (Reference Caballero1961)1 cited by Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008); Thatcher (Reference Thatcher1963a)1; Dyer and Carr (Reference Dyer and Carr1990b)2
Pseudonematophila Lenis & Vélez, Reference Lenis and Vélez2011
Pseudonematophila ovalis (Cordero & Vogelsang, Reference Cordero and Vogelsang1940) Lenis & Vélez, Reference Lenis and Vélez2011
Infection site: stomach
Host: Podocnemis lewyana1 and Podocnemis sp.2
Biogeographic unit: Magdalena province1, Sabana province2 – PCD
Political unity: [COLOMBIA] – (Bolívar): San Nicolás1; (Magdalena): Ricaurte1; [VENEZUELA] – (Guárico)2: Apurito river and unspecified location
References: Cordero and Vogelsang (Reference Cordero and Vogelsang1940)2 cited by Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021); Lenis and Vélez (Reference Lenis and Vélez2011)1,2
Schizamphistomoides Stunkard, 1925
Schizamphistomoides sp.
Infection site: large intestine
Host: Podocnemis expansa
Biogeographic unit: Cerrado province – CD
Political unity: [BRAZIL] – (Goiás): Britânia
Reference: Nomura (Reference Nomura1996)
Family Microscaphidiidae Looss, 1900
Dictyangium Stunkard, 1943
Dictyangium chelydrae Stunkard, 1943
Infection site: small intestine
Host: Terrapene ornata
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Estero Tamacuil
References: Caballero (Reference Caballero1946)
Neodeuterobaris Brooks, Reference Brooks1976
Neodeuterobaris pritchardae Brooks, Reference Brooks1976
Infection site: stomach
Host: Podocnemis lewyana
Biogeographic unit: Magdalena province – PCD
Political unity: [COLOMBIA] – (Bolívar)2: Coyongala and San Nicolás; (Caldas): La dorada1; (Magdalena): Ricaurte2
References: Brooks (Reference Brooks1976)1; Lenis and Vélez (Reference Lenis and Vélez2011)2
Octangioides Price, Reference Price1937
Octangioides skrjabini Price, Reference Price1937
Infection site: intestine, large intestine
Host: Dermatemys mawii
Biogeographic unit: Pacifc Lowlands province, Veracruzan province – MD
Political unity: [MEXICO] – (Oaxaca): Tehuantepec1; (Tabasco): Villahermosa2
References: Price (Reference Price1937)1; Caballero-Rodríguez (Reference Caballero-Rodríguez1960)2
Octangioides tlacotalpensis Caballero y C., 1942
Infection site: intestine, large intestine
Host: Dermatemys mawii 1 and Rhinoclemmys nasuta 2
Biogeographic unit: Veracruzan province1 – MD; Western Ecuador province2 – PCD
Political unity: [MEXICO] – (Veracruz)1: Laguna de Alvarado and Tlacotalpan; (Tabasco): Río Teapa1; [ECUADOR] – (Esmeraldas Province)2
References: Caballero (Reference Caballero1942)1; Caballero (Reference Caballero1943a)1; Thatcher (Reference Thatcher1963a)1; Dyer and Carr (Reference Dyer and Carr1990b)2
Podocnemitrema Alho & Vicente, Reference Alho and Vicente1964
Podocnemitrema papillosum Alho & Vicente, Reference Alho and Vicente1964
Infection site: stomach
Host: Podocnemis expansa 1 and Podocnemis unifilis 2
Biogeographic unit: Ucayali province – SBD
Political unity: [PERU] – (Loreto): Iquitos1,2; [BRAZIL] – (Amazonas): unspecified location1
References: Alho and Vicente (Reference Alho and Vicente1964)1; Tantaleán et al. (Reference Tantaleán, Sánchez and Pineda Catalan2011)1,2
Family Paramphistomidae Fischoeder, 1901
Dermatemytrema Price Reference Price1937
Dermatemytrema trifoliata Price Reference Price1937
Infection site: stomach, intestine, large intestine
Host: Dermatemys mawii
Biogeographic unit: Veracruzan province2,3,4,5, Pacifc Lowlands province1 – MD
Political unity: [MEXICO] – (Oaxaca): Tehuantepec1; (Tabasco): Río Teapa5 and Villahermosa4; (Veracruz): Laguna de Alvarado2,3
References: Price (Reference Price1937)1; Caballero (Reference Caballero1943a)2; Herrera-Rosales (Reference Herrera-Rosales1951)3, Caballero-Rodríguez (Reference Caballero-Rodríguez1960)4; Thatcher (Reference Thatcher1963a)5
Superfamily Plagiorchioidea Lühe, 1901
Family Plagiorchiidae Lühe, 1901
Cheloniotrema Caballero, Zerecero & Grocott, Reference Caballero, Zerecero and Grocott1956
Cheloniotrema tropicum Caballero, Zerecero & Grocott, Reference Caballero, Zerecero and Grocott1956
Infection site: esophagus, duodenum, and common bile duct
Host: Kinosternon integrum 1 and Kinosternon scorpioides 2
Biogeographic unit: Veracruzan province1 – MD; Guatuso-Talamanca province/Puntarenas-Chiriquí province2 – PCD
Political unity: [MEXICO] – (Oaxaca): Tuxtepec1; [PANAMA] – Valle de Antón2
References: Caballero et al. (Reference Caballero, Zerecero and Grocott1956)
Family Telorchiidae Looss, 1899
Loefgrenia Travassos, 1920
Loefgrenia loefgreni Travassos, 1920
Infection site: intestine
Host: Podocnemis unifilis
Political unity: [BRAZIL] – unspecified location
References: Travassos (Reference Travassos1919) cited by Fernandes and Kohn (Reference Fernandes and Kohn2014)
Telorchis Lühe, 1899 nec Looss, 1899
Telorchis achavali Mañé-Garzón & Holcman- Spector, 1973
Infection site: small intestine
Host: Trachemys dorbigni
Biogeographic unit: Pampean province2,3 – CD
Political unity: [BRAZIL] – (Rio Grande do Sul)2,3: Capão do Leão and Pelotas; [URUGUAY] – Tacuarembó1
References: Mañé-Garzón and Holcman-Spector (Reference Mañé-Garzón and Holcman-Spector1973)1 cited by Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021); Mascarenhas and Müller (Reference Mascarenhas and Muller2013)2; Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)3
Telorchis aculeatus (von Linstow, 1879) Braun, Reference Braun1901
Infection site: intestine
Host: Podocnemis unifilis
Political unity: [VENEZUELA] – (Sucre): unspecified location
References: Nasir (Reference Nasir1974)
Telorchis attenuatus Goldberger, 1911
Infection site: small intestine
Host: Trachemys scripta
Biogeographic unit: Yucatán Peninsula province – MD
Political unity: [MEXICO] – (Yucatán): Aguada Xpoc
References: Moravec & Vargas-Vázquez (Reference Moravec and Vargas-Vázquez1998)
Telorchis bifurcus (Braun, 1899) Braun, Reference Braun1901
Infection site: small intestine
Host: Podocnemis expansa
Political unity: [BRAZIL] – unspecified location
References: Braun (Reference Braun1901)
Telorchis birabeni Mañé-Garzón & Gil, 1961
Infection site: intestine, small intestine
Host: Phrynops geoffroanus 1, Phrynops hilarii,2 and Phrynops williamsi 3
Biogeographic unit: Atlantic province1, Esteros del Iberá province2, Parana Forest province1,3 – PD; Caatinga province1, Pampean province2 - CD
Political unity: [ARGENTINA] – (Buenos Aires Province): La Plata2; (Corrientes Province): unspecified location2; (Entre Rios): Gualeguaychú2; (Misiones): Comandante Andresito3; (Santa Fe)2: Cañada de Gomez, Helvecia, Rafaela and Santa Fe City; [BRAZIL] – (Pernambuco)1: Cordeiro, Limoeiro, Paudalho and Santa Cruz do Capibaribe; (Rio Grande do Sul)2: Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar; (São Paulo): RPPN Foz do Rio Aguapeí1; [URUGUAY] – (Tacuarembó): unspecified location2
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1961c)2, Lombardero and Moriena (Reference Lombardero and Moriena1977)2 cited by Mascarenhas and Müller (Reference Mascarenhas and Muller2021); Silva (Reference Silva2014)1; Mascarenhas et al. (Reference Mascarenhas, Bernardon and Muller2016)2; Fonseca et al. (Reference Fonseca, Rojas, Vieira, Ferreira, Rodrigues, Barros, Silva Júnior, Moura and Oliveira2021)1; Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)2; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2,3
Telorchis bravoae Caballero y C. & Zerecero y D., 1961
Infection site: small intestine
Host: Staurotypus triporcatus
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Tabasco): Río Mezcalapa
References: Caballero and Zerecero (Reference Caballero and Zerecero1960)
Telorchis caballeroi Rosales, 1951
Infection site: small intestine
Host: Rhinoclemmys areolata
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado
References: Herrera-Rosales (Reference Herrera-Rosales1951)
Telorchis corti Stunkard, 1915
Infection site: small intestine and large intestine
Host: Chelydra serpentina 1, Claudius angustatus 2, Kinosternon hirtipes 3, Kinosternon integrum 4, Kinosternon leucostomum 5, Kinosternon sp.6, Rhinoclemmys areolata 7, Trachemys callirostris 8, Trachemys dorbigni 9, Trachemys ornata,10 and Trachemys scripta 11
Biogeographic unit: Pampean province9 – CD; Balsas Basin province4, Veracruzan province1,2,5,6,7,11 – MD, Transmexican Volcanic Belt province3 – MTZ; Guatuso-Talamanca province10, Magdalena province8 – PCD
Political unity: [BRAZIL] – (Rio Grande do Sul)9: Capão do Leão and Pelotas; [COLOMBIA] – (Magdalena): Ricaurte8; [MEXICO] – (Puebla): Izúcar de Matamoros4; (Oaxaca): Presa de Temascal6; (Veracruz): Laguna de Alvarado7; (Tabasco): Río Teapa1,2,5,11; (Michoacán)3: Lago de Pátzcuaro, Lago de Zacapu, Lago de Zirahuén and Manantiales de Cointzio; [PANAMA] – (Panama province): Panamá city10
References: Bravo-Hollis (Reference Bravo-Hollis1944)4; Herrera-Rosales (Reference Herrera-Rosales1951)7; Caballero et al. (Reference Caballero, Zerecero and Grocott1958)10; Thatcher (Reference Thatcher1963a)1,2,5,11; Pérez-Ponce de León et al. (Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)3; Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008)6; Lenis and Vélez (Reference Lenis and Vélez2011)8; Mascarenhas and Müller (Reference Mascarenhas and Muller2013)9; Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)9
Telorchis devincenzii Mañé-Garzón & Gil, 1961
Synonym: Pseudotelorchis devincenzii (Mañé-Garzón & Gil, 1961)
Infection site: intestine, small intestine
Host: Hydromedusa tectifera
Biogeographic unit: Pampean province – CD
Political unity: [ARGENTINA] – (Buenos Aires): La Plata2; [URUGUAY] – (Canelones): Arroyo Las Piedras 1
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1961b)1; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2
Telorchis dhongokii (Mehra & Bokhari, 1932) Wharton, 1940
Infection site: large intestine
Host: Rhinoclemmys areolata
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado
References: Herrera-Rosales (Reference Herrera-Rosales1951)
Telorchis diaphanus Teixeira de Freitas & Dobbin, Reference Freitas and Dobbin1959
Infection site: intestine, small intestine
Host: Kinosternon scorpioides 1, Phrynops hilarii 2 and Trachemys dorbigni 3
Biogeographic unit: Esteros del Iberá province2,3 – PD
Political unity: [ARGENTINA] – (Corrientes): Bella Vista2; (Entre Rios) Federal3; [BRAZIL] – (Pernambuco): unspecified location1
References: Freitas and Dobbin Jr. (Reference Freitas and Dobbin1959)1; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2,3
Telorchis dissentaneus Caballero y C. & Herrera Rosales, 1947
Infection site: small intestine
Host: Kinosternon integrum
Biogeographic unit: Balsas Basin province – MD
Political unity: [MEXICO] – (Morelos): Casasano
References: Caballero and Herrera-Rosales (Reference Caballero and Herrera-Rosales1947)
Telorchis dissimilis (Caballero y C, 1938) Hughes, Higginbotham & Clary, 1941
Synonym: Cercorchis dissimilis Caballero y C., 1938
Infection site: intestine, small intestine
Host: Terrapene ornata 1 and Rhinoclemmys areolata 2
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado
References: Caballero (Reference Caballero1938)1; Herrera-Rosales (Reference Herrera-Rosales1951)2
Telorchis dubius Mañé-Garzón & Holcman-Spector, 1968
Infection site: intestine, small intestine
Host: Trachemys dorbigni
Biogeographic unit: Esteros del Iberá province2 – PD; Pampean province1 – CD
Political unity: [ARGENTINA] – (Entre Rios)2: Concordia and Federal; [URUGUAY] – (Flores): Arroyo del Perdido1
References: Mañé-Garzón and Holcman-Spector (Reference Mañé-Garzón and Holcman-Spector1968a)1; Palumbo et al. (Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a)2
Telorchis grocotti Caballero y C. & Zerecero, Reference Caballero and Zerecero1954
Infection site: intestine
Host: Kinosternon scorpioides
Biogeographic unit: Guatuso-Talamanca province – PCD
Political unity: [PANAMA] – (Panama Oeste Province): Arraijón
References: Caballero and Zerecero (Reference Caballero and Zerecero1954)
Telorchis hagmanni Lent & Teixeira de Freitas, 1937
Infection site: digestive tract, stomach, small intestine
Host: Peltocephalus dumerilianus 1, Podocnemis expansa 2, Podocnemis lewyana,3 and Podocnemis unifilis 4
Biogeographic unit: Imerí province1, Pará province2 – BBD; Magdalena province3 – PCD; Ucayali province4 – SBD
Political unity: [BRAZIL] – Amazon River2; (Amazonas)1: Barcelos and Jaú National Park; (Pará): Belém2; [COLOMBIA] – (Bolívar): San Nicolás3; (Magdalena): Ricaurte3; (Loreto): Iquitos4
References: Lent and Freitas (Reference Lent and Freitas1937)2; Alho (Reference Alho1965)2; Lenis and Vélez (Reference Lenis and Vélez2011)3; Tantaleán et al. (Reference Tantaleán, Sánchez and Pineda Catalan2011)4; Ferreira (Reference Ferreira2016)1; Menegaldo et al. (Reference Menegaldo, Arias-Pacheco, Perin, Tebaldi and Hoppe2024)1
Telorchis medius Stunkard, 1915
Infection site: intestine
Host: Rhinoclemmys melanosterna
Biogeographic unit: Guatuso-Talamanca province – PCD
Political unity: [PANAMA] – (Panama province): Panamá city
References: Caballero et al. (Reference Caballero, Zerecero and Grocott1958)
Telorchis membranaceus (Caballero y C., 1940) Yamaguti, 1958
Synonym: Cercorchis membranaceus Caballero y C., 1940
Infection site: large intestine
Host: Terrapene ornata
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado
References: Caballero (Reference Caballero1940b)
Telorchis panamensis Caballero y C., Zerecero & Grocott, Reference Caballero and Brenes-Madrigal1957
Infection site: intestine
Host: Kinosternon scorpioides
Biogeographic unit: Pacific Dominion
Political unity: [PANAMA] – El Valle
Reference: Caballero et al. (Reference Caballero, Zerecero and Grocott1956)
Telorchis patonianus (Caballero y C., 1935) Wharton, 1940
Synonym: Cercorchis patonianus Caballero y C., 1935
Infection site: intestine
Host: Claudius angustatus and Staurotypus triporcatus
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Tabasco): Río Teapa
Reference: Thatcher (Reference Thatcher1963a)
Telorchis platensis Mañé-Garzón & Gil, 1961
Infection site: large intestine, small intestine
Host: Acanthochelys spixii 1 and Hydromedusa tectifera 2
Biogeographic unit: Pampean province1 – CD
Political unity: [BRAZIL] – (Rio Grande do Sul)1: Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar; [URUGUAY] – unspecified location2.
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1961b)2; Mascarenhas et al. (Reference Mascarenhas, Bernardon and Muller2016)1; Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)1
Telorchis pleroticus (Braun, 1899) Braun, Reference Braun1901
Infection site: intestine
Host: Freshwater turtle
Political unity: [BRAZIL] – unspecified location
References: Braun (Reference Braun1901)
Telorchis productus Mañé-Garzón & Gil, 1961
Infection site: small intestine
Host: Phrynops hilarii
Biogeographic unit: Esteros del Iberá province – PD
Political unity: [URUGUAY] – (Tacuarembó): Paso de los Toros
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1961b)
Telorchis pseudoaculeatus Dollfus, 1929
Infection site: small intestine
Host: Kinosternon cruentatum
Biogeographic unit: Puntarenas-Chiriquí province – PCD
Political unity: [COSTA RICA] – (San José): Santa Ana
References: Caballero and Brenes-Madrigal (1957)
Telorchis rapidulus Dobbin, Reference Dobbin1957
Infection site: small intestine
Host: Kinosternon scorpioides
Biogeographic unit: Atlantic province – PD
Political unity: [BRAZIL] – (Pernambuco): Cortês
References: Dobbin Jr. (Reference Dobbin1957)
Telorchis reelfooti (Parker, 1941) Rosales, 1951
Synonym: Cercorchis reelfooti Parker, 1941
Infection site: small intestine
Host: Kinosternon integrum
Biogeographic unit: Balsas Basin province – MD
Political unity: [MEXICO] – (Morelos): Cuautla
References: Herrera-Rosales (Reference Herrera-Rosales1951)
Telorchis robustus Goldberger, 1911
Infection site: large intestine
Host: Rhinoclemmys areolata
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado
References: Herrera-Rosales (Reference Herrera-Rosales1951)
Telorchis singularis (Bennett, 1935) Wharton, 1940
Note: This record was considered dubious (Nasir, Reference Nasir1974; Paredes-León et al. Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008)
Infection site: small intestine
Host: Rhinoclemmys areolata
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado
References: Herrera-Rosales (Reference Herrera-Rosales1951)
Telorchis sp.
Infection site: small intestine
Host: Hydromedusa tectifera
Political unity: [BRAZIL] – (Rio Grande do Sul): unspecified location
Reference: Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021)
Telorchis sp.
Infection site: intestine
Host: Hydromedusa tectifera
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
Reference: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Telorchis sp.
Infection site: small intestine
Host: Trachemys dorbigni
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão and Pelotas
Reference: Mascarenhas and Müller (Reference Mascarenhas and Muller2013)
Telorchis sp.
Infection site: small intestine
Host: Trachemys dorbigni
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão and Pelotas
Reference: Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)
Telorchis sp.
Infection site: intestine
Host: Kinosternon integrum
Biogeographic unit: Pacific Lowlands province - MD
Political unity: [MEXICO] – (Jalisco): Tequila
Reference: Paredes-León et al. (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008)
Superfamily Pronocephaloidea Looss, 1899
Family Pronocephalidae Looss, 1899
Choanophorus Caballero, Reference Caballero1942
Choanophorus rovirosai Caballero, Reference Caballero1942
Infection site: intestine, small intestine, duodenum
Host: Dermatemys mawii
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Tabasco): Río Teapa3 and Villahermosa2; (Veracruz): Tlacotalpan1
References: Caballero (Reference Caballero1942)1; Caballero-Rodríguez (Reference Caballero-Rodríguez1960)2; Thatcher (Reference Thatcher1963a)3
CLASS MONOGENOIDEA Bychowsky, 1937
ORDER DACTYLOGYRIDEA
Family Iagotrematidae Mañe-Garzon & Gil, 1962
Iagotrema Mañe-Garzon & Gil, 1962
Iagotrema uruguayense Mañe-Garzón & Gil, 1962
Infection site: urinary bladder
Host: Hydromedusa tectifera
Political unity: [URUGUAY] – (Tacuarembó): unspecified location
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1962b)
ORDER POLYSTOMATIDEA
Family Polystomatidae Gamble, 1896
Fornixtrema Du Preez & Verneau, 2020
Fornixtrema fentoni (Platt, Reference Platt2000) Du Preez & Verneau, 2020
Synonym: Neopolystoma fentoni Platt, Reference Platt2000
Infection site: conjunctival sacs
Host: Kinosternon leucostomum 1 and Rhinoclemmys pulcherrima 2
Biogeographic unit: Pacific Lowlands province – MD
Political unity: [COSTA RICA] – unspecified locations1; (Guanacaste): Guanacaste Conservation Area1,2
References: Platt (Reference Platt2000)1,2; Verneau et al. (Reference Verneau, Palacios, Platt, Alday, Billard, Allienne, Basso and Preez2011)1
Fornixtrema guianense (Du Preez, Badets, Héritier & Verneau, Reference Du Preez, Badets, Héritier and Verneau2017) Du Preez & Verneau, 1920
Synonym: Neopolystoma guianensis Du Preez, Badets, Héritier & Verneau, Reference Du Preez, Badets, Héritier and Verneau2017
Infection site: conjunctival sacs, conjunctival sacs of the eye
Host: Rhinoclemmys punctularia
Biogeographic unit: Guianan Lowlands province – BBD
Political unity: [FRENCH GUIANA] – Cayenne
References: Du Preez et al. (Reference Du Preez, Badets, Héritier and Verneau2017); Chaabane et al. (Reference Chaabane, Du Preez, Johnston and Verneau2022)
Fornixtrema scorpioides (Du Preez, Badets, Héritier & Verneau, Reference Du Preez, Badets, Héritier and Verneau2017) Du Preez & Verneau, 2020
Synonym: Neopolystoma scorpioides Du Preez, Badets, Héritier & Verneau, Reference Du Preez, Badets, Héritier and Verneau2017
Infection site: conjunctival sacs, conjunctival sacs of the eye
Host: Kinosternon scorpioides
Biogeographic unit: Guianan Lowlands province – BBD
Political unity: [FRENCH GUIANA]- Cayenne1; Roura2
References: Du Preez et al. (Reference Du Preez, Badets, Héritier and Verneau2017)1; Chaabane et al. (Reference Chaabane, Du Preez, Johnston and Verneau2022)2
Manotrema du Preez, Domingues & Verneau, 2022
Manotrema brasiliensis (Viera, Novelli, Sousa & SouzaLima, 2008) du Preez, Domingues & Verneau, 2022
Synonym: Polystomoides brasiliensis Vieira, Novelli, Sousa & SouzaLima, Reference Vieira, Novelli, Sousa and SouzaLima2008
Infection site: buccal and pharyngeal cavities, esophagus
Host: Hydromedusa maximiliani 1, Mesoclemmys tuberculata 2 and Phrynops geoffroanus 3
Biogeographic unit: Atlantic province2,3, Parana Forest province1,3 – PD; Caatinga province2,3 - CD
Political unity: [BRAZIL] – (Pernambuco)3: limoeiro, Paudalho, Recife and Santa Cruz do Capibaribe; (Minas Gerais): Juiz de Fora1,3; (São Paulo): RPPN Foz do Rio Aguapeí3; (Rio de Janeiro): unspecified location1; (Sergipe)2,3: Areia Branca, Pirambu, Poço Redondo and Tobias Barreto
References: Vieira et al. (Reference Vieira, Novelli, Sousa and SouzaLima2008)1,3; Muniz-Pereira et al. (Reference Muniz-Pereira, Vieira and Luque2009)1; Silva (Reference Silva2014)3; Santana et al. (Reference Santana, Iwama, Teixeira, Moura, Faria and Mesquita2019)2,3; Fonseca et al. (Reference Fonseca, Rojas, Vieira, Ferreira, Rodrigues, Barros, Silva Júnior, Moura and Oliveira2021)3; Santana et al. (Reference Santana, Teixeira, Araújo-Filho, Teles, Pereira, Brito, Almeida and Mesquita2023)2
Manotrema fuquesi (Mañe-Garzón & Gil, 1962) du Preez, Domingues & Verneau, 2022
Synonym: Polystomoides fuquesi Mañe-Garzón & Gil, 1962
Infection site: oral cavity
Host: Phrynops hilarii
Biogeographic unit: Esteros del Iberá province – PD
Political unity: [URUGUAY] – (Artigas Departament) – Artigas
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1962a)
Manotrema uruguayensis (Mañe-Garzón & Gil, 1961) du Preez, Domingues & Verneau, 2022
Synonym: Polystomoides uruguayensis Mañe-Garzón & Gil, 1961
Infection site: oral cavity
Host: Phrynops hilarii
Biogeographic unit: Pampean province – CD
Political unity: [URUGUAY] – (Artigas Departament): Arroyo Cuaró
References: Mañé-Garzón and Gil (Reference Mañé-Garzón and Gil1961d)
Polystomoidella Price, 1939
Polystomoidella oblonga (Wright, 1879) Price, 1939
Infection site: urinary bladder
Host: Kinosternon hirtipes 1, Kinosternon integrum,2 and Kinosternon leucostomum 3
Biogeographic unit: Transmexican Volcanic Belt province1 – MTZ; Balsas Basin province2, Veracruzan province1,3 – MD
Political unity: [MEXICO] – unspecified location1; (Federal District): Lago de Xochimilco1; (Guerrero): Cacahuamilpa2; (Michoacán)1: Lago de Pátzcuaro, Lago de Zacapu, Lago de Zirahuén and Manantiales de Cointzio; (Nayarit): Villa Hidalgo - Río Grande1; (Veracruz): Laguna de Alvarado3.
References: Caballero (Reference Caballero1940b)2; Herrera-Rosales (Reference Herrera-Rosales1951)3; Lamothe-Argumedo (Reference Lamothe-Argumedo1972)1; Parra-Rojas (Reference Parra-Rojas1983)1; Pérez-Ponce de León et al. (Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)1
Polystomoidella whartoni Price, 1939
Infection site: urinary bladder
Host: Kinosternon hirtipes 1, Kinosternon integrum,2 and Kinosternon leucostomum 3
Biogeographic unit: Balsas Basin province2,3 – MD
Political unity: [MEXICO] – (Guanajuato): Río Lerdo1; (Morelos): Casasano2,3
References: Caballero (Reference Caballero1940b)1; Herrera-Rosales (Reference Herrera-Rosales1951)2,3
Polystomoides Ward, 1917
Polystomoides cayensis (Du Preez, Badets, Héritier & Verneau, Reference Du Preez, Badets, Héritier and Verneau2017) Chaabane, Du Preez, Johnston & Verneau, Reference Chaabane, Du Preez, Johnston and Verneau2022
Synonym: Neopolystoma cayensis Du Preez, Badets, Héritier & Verneau, Reference Du Preez, Badets, Héritier and Verneau2017
Infection site: urinary bladder
Host: Rhinoclemmys punctularia
Biogeographic unit: Guianan Lowlands province – BBD
Political unity: [FRENCH GUIANA] – Cayenne
References: Du Preez et al. (Reference Du Preez, Badets, Héritier and Verneau2017); Chaabane et al. (Reference Chaabane, Du Preez, Johnston and Verneau2022)
Polystomoides coronatus (Leidy, 1888) Ozaki, 1935
Infection site: oral cavity
Host: Trachemys scripta
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Tabasco): Río Teapa
Reference: Thatcher (Reference Thatcher1963a)
Polystomoides cyclovitellum (Caballero, Zerecero & Grocott, 1957) Chaabane, Du Preez, Johnston & Verneau, Reference Chaabane, Du Preez, Johnston and Verneau2022
Infection site: urinary bladder
Host: Rhinoclemmys melanosterna
Biogeographic unit: Guatuso-Talamanca province – PCD
Political unity: [PANAMA] – (Panama province): Panamá city
Reference: Caballero et al. (Reference Caballero, Zerecero and Grocott1956)
Polystomoides domitilae (Caballero, Reference Caballero1938) Chaabane, Du Preez, Johnston & Verneau, Reference Chaabane, Du Preez, Johnston and Verneau2022
Synonym: Neopolystoma domitilae (Caballero, Reference Caballero1938) Price, 1939
Infection site: urinary bladder and cloaca
Host: Chelydra serpentina 1, Terrapene ornata,2 and Trachemys scripta 3
Biogeographic unit: Veracruzan province – MD;
Political unity: [MEXICO] – (Tabasco): Río Teapa1.3; (Veracruz): Laguna de Alvarado2
References: Caballero (Reference Caballero1938)2; Thatcher (Reference Thatcher1963a)1,3
Polystomoides magdalenensis Lenis & Garca-Prieto, Reference Lenis and Garcia-Prieto2009
Infection site: oral cavity
Host: Trachemys callirostris
Biogeographic unit: Magdalena province – PCD
Political unity: [COLOMBIA] – (Bolívar): Coyongal and Santa Cruz de Mompox; (Magdalena): Ricaurte
References: Lenis and García-Prieto (Reference Lenis and Garcia-Prieto2009)
Polystomoides orbiculare (Stunkard, 1916) Chaabane, Du Preez, Johnston & Verneau, Reference Chaabane, Du Preez, Johnston and Verneau2022
Synonym: Neopolystoma orbiculare (Stunkard, 1916) Price, 1939
Infection site: urinary bladder
Host: Kinosternon leucostomum 1 and Trachemys scripta 2
Biogeographic unit: Veracruzan province- MD
Political unity: [MEXICO] – (Oaxaca): Tuxtepec1; (Tamaulipas): Río Tamesi2
References: Zerecero (Reference Zerecero1948)1; Lamothe-Argumedo (Reference Lamothe-Argumedo1972)2
Polystomoides rohdei Mañe-Garzón & Holcman-Spector, 1968
Infection site: oral cavity
Host: Trachemys dorbigni
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul)2: Capão do Leão and Pelotas; [URUGUAY] – unspecified location3; (Florida): unspecified location1; (Rivera): unspecified location1; (Soriano): Arroyo del Perdido1
References: Mañé-Garzón and HolcmanSpector (Reference Mañé-Garzón and Holcman-Spector1968b)1; Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)2; Chaabane et al. (Reference Chaabane, Du Preez, Johnston and Verneau2022)3
Polystomoides santamariensis Lignon, Cohen, Justo, Du Preez & Monteiro in Lignon et al. Reference Lignon, Cohen, Justo, Du Preez, Comarella, Nishimaru, Souza, Ataíde, Müller, Brun and Silvia Gonzalez Monteiro2023
Infection site: urinary bladder
Host: Trachemys dorbigni
Biogeographic unit: Parana Forest province – PD
Political unity: [BRAZIL] – (Rio Grande do Sul): Santa Maria
Reference: Lignon et al. (Reference Lignon, Cohen, Justo, Du Preez, Comarella, Nishimaru, Souza, Ataíde, Müller, Brun and Silvia Gonzalez Monteiro2023)
Polystomoides sp.
Infection site: oral cavity
Host: Acanthochelys spixii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar
References: Chaviel et al. (Reference Chaviel, Mascarenhas, Bernardon, Coimbra and Müller2020); Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Polystomoides sp.
Infection site: conjunctival sacs
Host: Kinosternon leucostomum
Political unity: [COSTA RICA] – unspecified location
Reference: Héritier et al. (Reference Héritier, Badets, Du Preez, Aisien, Lixian, Combes and Verneau2015)
Polystomoides sp.
(original record: Neopolystoma sp.)
Infection site: unspecified site of infection
Host: Mesoclemmys vanderhaegei
Biogeographic unit: Cerrado province – CD
Political unity: [BRAZIL] – (Mato Grosso): Chapada dos Guimarães
Reference: Ávila et al. (Reference Ávila, Brito, Barrella, Strussmann and Silva2010)
Polystomoides sp.
(original record: Neopolystoma sp.)
Infection site: urinary bladder
Host: Phrynops geoffroanus
Biogeographic unit: Parana Forest province – PD
Political unity: [BRAZIL] – (São Paulo): RPPN Foz do Rio Aguapeí
Reference: Silva (Reference Silva2014)
Polystomoides sp.
Infection site: oral cavity
Host: Phrynops hilarii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão, Pelotas, Rio Grande and Santa Vitória do Palmar; unspecified location
References: Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)
Polystomoides sp.
Infection site: oral cavity
Host: Phrynops hilarii
Political unity: [BRAZIL] – (Rio Grande do Sul): unspecified location
References: Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021)
Polystomoides sp.
Infection site: urinary bladder
Host: Trachemys dorbigni
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): Capão do Leão and Pelotas
References: Mascarenhas et al. (Reference Mascarenhas, Silva and Müller2021)
PHYLUM ACANTHOCEPHALA RUDOLPHI, 1808
ORDER NEOECHINORHYNCHIDA
Family Neoechinorhynchidae Ward, 1917
Neoechinorhynchus Hamann in Stiles & Hassall, 1905
Neoechinorhynchus (Neoechinorhynchus) emydis (Leidy, 1851) Van Cleave, 1919
Infection site: intestine
Host: Terrapene ornata
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado
References: Bravo-Hollis (Reference Bravo-Hollis1946)
Neoechinorhynchus (Neoechinorhynchus) emyditoides Fisher, 1960
Infection site: intestine, small intestine
Host: Trachemys scripta 1 and Trachemys scripta venusta 2
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Veracruz): Laguna de Alvarado1 and Lago de Catemaco1; (Tamaulipas): Purificación river1,2; (Tabasco): Centla Swamps1
References: Barger (Reference Barger2004)1; García-Varela et al. (Reference García-Varela, García-Prieto and Rodríguez2011)1; Pinacho-Pinacho et al. (Reference Pinacho-Pinacho, Hernández-Orts, Sereno-Uribe, Pérez-Ponce de León and García-Varela2017)1; García-Varela and Pinacho-Pinacho (Reference García-Varela and Pinacho-Pinacho2018)2
Neoechinorhynchus (Neoechinorhynchus) schmidti Barger, Thatcher & Nickol, Reference Barger, Thatcher and Nickol2004
Infection site: intestine
Host: Trachemys scripta 1, Trachemys venusta 2
Biogeographic unit: Veracruzan province – MD
Political unity: [MEXICO] – (Tabasco)1: Centla Swamps and 24 km S from Villahermosa2
References: Barger (Reference Barger2004)2; Barger et al. (Reference Barger2004)1; Martínez-Aquino et al. (Reference Martínez-Aquino, Reyna-Fabián, Rosas-Valdez, Razo-Mendivil, Pérez-Ponce de León and García-Varela2009)1
ORDER POLYMORPHIDA
Family Polymorphidae Meyer, 1931
Polymorphidae gen. sp. immature
Infection site: small intestine
Host: Acanthochelys spixii
Biogeographic unit: Pampean province – CD
Political unity: [BRAZIL] – (Rio Grande do Sul): unspecified location1, Capão do Leão2, Pelotas2, Rio Grande2 and Santa Vitória do Palmar2
References: Mascarenhas and Müller (Reference Mascarenhas, Silva and Müller2021)1; Mascarenhas et al. (Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022)2
Arhythmorhynchus Lühe, 1911
Arhythmorhynchus brevis Van Cleave, 1916
Synonym: Polymorphus brevis (Van Cleave, 1916) Travassos, 1926
Infection site: body cavity, mesentery
Host: Kinosternon hirtipes
Biogeographic unit: Transmexican Volcanic Belt province – MTZ
Political unity: [MEXICO] – (Michoacán): Lago de Zacapu
Reference: Pérez-Ponce de León et al. ( Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001)
Discussion
In the literature, only reviews of marine species have been published for chelonians in the Neotropical region by Werneck (Reference Werneck and Da Silva2016), with other reviews focusing on political regions, such as in Mexico, for reptiles by Paredes-León (Reference Paredes-León, García-Prieto, Guzmán-Cornejo, León-Règagnon and Pérez2008), Costa Rica, by Rodriguez-Ortiz (Reference Rodriguez-Ortiz, Garcia-Prieto and Perez-Ponce de Leon2004), and in South America more recently, for continental chelonians by Mascarenhas and Müller (Reference Mascarenhas and Muller2021). Finally, Cuellar et al. (Reference Cuellar, Sáez, Cantú, Sánchez, Mendoza, Conga, Cruces, Luque and Chero2022) added new records to their review of helminths associated with reptiles in Peru. This study presents the first checklist of parasitic helminth species (Nematoda, Trematoda, Monogenea, Acanthocephala, and Cestoda) of freshwater and terrestrial chelonians for the Neotropical region, addressing their biogeographical and political distribution, including a new occurrence in a chelonian host in the hinterland of Paraíba, Brazil.
Main groups of helminths parasitizing terrestrial and freshwater chelonians in the Neotropical region
In this study, Nematoda was the group that occurred in all biogeographical units, had the highest number of taxa (98 taxa), with the second most diverse family of parasitic helminths, Atractidae (28 species), represented by the most recorded species in the group: Spiroxys contortus, in a total of six hosts. The Nematoda phylum exhibits a great taxonomic diversity, being found in marine, terrestrial, and freshwater environments, with parasitic and free-living representatives, possessing enormous economic, ecological, and zoonotic importance (Gómez-Valdez et al. Reference Gómez-Valdez, Carvalho-Saucedo, Ocampo and Cruz-Villacorta2019; Hodda Reference Hodda2011; Onaka Reference Onaka and Moraes2004). They are common endoparasites of freshwater fish (Luque Reference Luque2004), but they also frequently parasitize reptiles and other vertebrates. Species of the genus Spiroxys Schneider, 1866 are often found parasitizing reptiles and amphibians, mainly in the stomachs of chelonians (Roca and García Reference Roca and García2008). In the Neotropical region, S. contortus infects the gastrointestinal tract of seven host species and is present in four biogeographic units: Chacoan dominion, Mesoamerican dominion, Mexican transition zone, and Paraná dominion.
Trematoda was the second group to present the most taxa (78 taxa), with the most diverse family of parasitic helminths, Telorchiidae (29 species), represented by the most commonly found species, Nematophila grandis. Trematodes are among the most diverse groups in the Platyhelminth phylum that parasitize vertebrates (Pérez-Ponce De León et al. Reference Pérez Ponce de León, García-Prieto and Mendoza-Garfias2007). Nematophila grandis infects the gastrointestinal tract of various aquatic chelonians in South and Central America (Lombardero and Moriena, Reference Lombardero and Moriena1977; Salízar and Sanchez Reference Salizar and Sanchez2004; Sánchez et al. Reference Sánchez, Tantaleán, Vela and Méndez2006). It was the species of parasitic helminths that had the highest number of hosts, found in 17 species and five families of chelonians in the Neotropical region, as well as being present in nine countries and six biogeographic units: Boreal Brazilian dominion, Chacoan dominion, Mesoamerican dominion, Pacific dominion, Paraná Dominion, and South Brazilian dominion.
Terrestrial and freshwater chelonian hosts of parasitic helminths in the Neotropical region
Among the eight families of chelonians recorded, the one with the greatest diversity of helminths was Chelidae, which is also the only family to include all groups of parasitic helminths, with the highest occurrence of trematodes, and also the highest number of host species recorded, followed by Kinosternidae, which had the highest rate of nematodes.
The Chelidae family is represented by the freshwater terrapins of the Pleurodira suborder and is made up of 14 genera and 64 species worldwide (Uetz et al. Reference Uetz, Freed and Hošek2021). In the Americas, this family is geographically restricted to South America and includes of eight genera (Acanthochelys, Chelus, Hydromedusa, Mesoclemmys, Phrynops, Platemys, Ranacephala, and Rhinemys) and 24 species. It also includes the second genus, Phrynops, which features one of the most parasitized species, Phrynops hilarii. Many species, particularly P. geoffroanus, are capable to surviving in anthropogenic environments, are widely distributed, share habitats with other species, and possess a diverse range of prey, in addition to exhibiting scavenging habits (Martins et al. Reference Martins, Souza and Costa2010; Medem Reference Medem1960; Souza and Abe Reference Souza and Abe2000; Souza Reference Souza2004;). These factors may contribute to the occurrence of all helminth groups in the family.
Chelonians of the Kinosternidae family include the smallest turtle species exclusive to the New World (Castro Reference Castro2006). They are semi-aquatic freshwater chelonians with a distribution that ranges from Canada to South America (Berry and Iverson Reference Berry and Iverson2011). This family comprises a total of four genera, 30 species and five subspecies (Uetz et al. Reference Uetz, Freed and Hošek2021), of which 19 species and three genera – Claudius, Kinosternon, and Staurotypus – are found in the Neotropical region. Its extensive distribution, along with the great diversity of species, may contribute to the number of records found, with the most parasitized host genus, Kinosternon, in the Neotropical region, primarily in Mexico.
Along with the freshwater chelonian Phrynops hilarii, the terrestrial chelonian Chelonoidis denticulatus from the Testudinidae family was the most parasitized species, with a total of 25 helminth taxa. Jabutis are terrestrial chelonians that, in addition to inhabiting diverse ecosystems and having omnivorous eating habits, are widely kept as pets (Oliveira et al. Reference Oliveira, Nascimento, Sousa, Carvalho, Silva, Oliveira and Oliveira2023). In the literature, most records are of animals in captivity or used as pets. When maintained in captivity, they are susceptible to outbreaks of parasitic diseases (Fowler Reference Fowler1986), acquiring more parasites than free-living animals (Menezes Reference Menezes2000).
Biogeographical context
We found helminths associated with terrestrial and freshwater chelonians in all ten biogeographical units recorded, particularly the Mesoamerican dominion and its province, the Veracruzan province, as well as the Chaco dominion with its province, the Pampean province. Both domains and provinces show the highest concentrations of research related to helminth parasites in chelonians within the Neotropical region.
The Mesoamerican dominion recorded the highest number of helminth taxa (61 taxa), and its province, the Veracruzan province, recorded the second highest number of taxa (35 taxa). This dominion covers the lowlands of southern and central Mexico and most of Central America, including Guatemala, Belize, Honduras, El Salvador, and northern Nicaragua. It encompasses the Pacific lowlands, the Balsas basin, Veracruzan, the Yucatán Peninsula, and the Mosquito provinces (Morrone Reference Morrone2014a; Morrone Reference Morrone2014b). The dominion has endemic species from the Kinosternidae family: Dermatemys mawii, Claudius angustatus, Kinosternon abaxillare, Kinosternon oaxacae, Staurotypus salvinii, and Staurotypus triporcatus; from the Chelydridae family: Chelydra rossignonii; and from the Geoemydidae family: Rhinoclemmys funerea and Rhinoclemmys rubida.
The Chaco dominion recorded the second highest rate of helminths (57 taxa), while its province, the Pampean province, recorded the highest rate of helminths (36 taxa). The dominion covers a large area in South America, encompassing northern and central Argentina, southeastern Bolivia, western and central Uruguay, Paraguay, and southern, central, and northeastern Brazil (Morrone Reference Morrone2006, Reference Morrone2014a; Morrone and Coscarón, Reference Morrone and Coscarón M1996). It is divided into four provinces: Caatinga, Cerrado, Chaco, and Pampa (Morrone Reference Morrone2015). Examples of species found in the domain include the chelonians Acanthochelys pallidipectoris, Acanthochelys spixii, Chelonoidis carbonarius, Hydromedusa tectifera, Kinosternon scorpioides, Mesoclemmys tuberculata, Phrynops geoffroanus, Phrynops hilarii, and Trachemys dorbigni.
Literature review
The literature search found studies on parasitic helminths between 1850 and 2024. Nematoda and Trematoda were the groups with the most reports. Most of the records were irregular after the 1940s until they peaked and grew from 2000 onwards. Many old records lack information on the host species and their origins or have not been published in articles, remaining as dissertations and theses, making it difficult to be more precise in the knowledge and distribution of the species and their hosts in the Neotropical region.There has been a significant increase in work on all groups in recent years, with a notable rise in the nematode group and the description of new genera and species: Vogtnema asymmetrica (Jesus et al. Reference Jesus, Santos, Nandyara, Santos and Melo2022), Orientatractis matosi (Jesus et al. Reference Jesus, Oliveira, Ramos and Melo2023a); Polystomoides santamariensis (Lignon et al. Reference Lignon, Cohen, Justo, Du Preez, Comarella, Nishimaru, Souza, Ataíde, Müller, Brun and Silvia Gonzalez Monteiro2023); Pneumoatractis gibbonsae (Jesus et al. Reference Jesus, Willkens, Santos and Melo2023b); Serpinema pelliculatus (Silva et al. (Reference Silva, Jesus, Santos, Nandyara, Willkens, Santos and Melo2023); Camallanus pallidipectoris (Palumbo et al. Reference Palumbo, Servián, Cassano and Diaz2024b); Orientatractis bowseri (Jesus et al. Reference Jesus, Santos, Oliveira, Félix-Silva and Melo2024a); Acanthoatractis xinguensis (Jesus et al. Reference Jesus, Santos, Nandyara, Santos and Melo2024b). This increase in new species shows that there is still a lot to discover about the helminth fauna in the Neotropical region, especially in areas with gaps or a lack of records.
Field research: First record of Spiroxys contortus for Phrynops geoffroanus
For the field research, this study recorded a low helminth diversity for the P. geoffroanus species: Camallanus sp., from the Camallanidae family, predominant in the small intestine with the highest prevalence (100%) and abundance and Spiroxys contortus, from the Gnathostomatidae family, in the stomach, marking the first record of the species for the host. The nematode has previously been reported in Argentina in Hydromedusa tectifera in the provinces of Buenos Aires (Castillo et al. Reference Castillo, Acosta, González-Rivas and Ramallo2020; Palumbo et al. Reference Palumbo, Capasso, Cassano, Alcalde and Diaz2016) and Córdoba (Palumbo et al. Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a); Phrynops hilarii in the provinces of Buenos Aires (Castillo et al. Reference Castillo, Acosta, González-Rivas and Ramallo2020; Palumbo et al. Reference Palumbo, Capasso, Cassano, Alcalde and Diaz2016), Corrientes, Entre Ríos and Santa Fe (Palumbo et al. Reference Palumbo, Alcalde, Bonino, Lescano, Montes, Solari and Inés Diaz2024a); in Brazil in Peltocephalus dumerilianus (Gomes and Kohn Reference Gomes and Kohn1970); in the state of Rio Grande do Sul in Phrynops hilarii (Chaviel et al. Reference Chaviel, Mascarenhas, Bernardon, Coimbra and Müller2020; Mascarenhas et al. Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022), Acanthochelys spixii and Hydromedusa tectifera (Mascarenhas et al. Reference Mascarenhas, Souza, Coimbra and Müller2013; Mascarenhas et al. Reference Mascarenhas, Chaviel, Bernardon, Wolter, Coimbra and Müller2022) and Trachemys dorbigni (Mascarenhas et al. Reference Mascarenhas and Muller2015b; Mascarenhas et al. Reference Mascarenhas, Silva and Müller2021); in Mexico in Kinosternon hirtipes, in the state of Michoacán (Pérez-Ponce de León et al. Reference Pérez-Ponce De León, Jiménez-Ruiz, Mendoza-Garfias and García-Prieto2001) and Trachemys scripta in the state of Yucatán (Moravec and Vargas-Vázquez, Reference Moravec and Vargas-Vázquez1998).
Finally, despite the recent increase in research on helminths of chelonians in the Neotropical region, with the description of new genera and species, the literature review showed that there are still gaps in the knowledge of helminths parasitic to chelonians, particularly in groups with few studies over the years and in localities with no records in the Neotropical region. The helminth fauna of terrestrial and freshwater chelonians in the Neotropical region is mainly composed of nematodes and trematodes, with the Chelidae family presenting all groups of helminths and the Chelonoidis denticulatus and Phrynops hilarii species being the most parasitized. In the biogeographical context, the biogeographic units with the most taxa and the highest concentrations of research were the Mesoamerican dominion and the Chacoan dominion. This study contributes to the knowledge of helminths of terrestrial and freshwater chelonians in the Neotropical region and adds to their biogeographical distribution. It also provides the first record of the nematode Spiroxys contortus in the host P. geoffroanus.
Supplementary material
The supplementary material for this article can be found at http://doi.org/10.1017/S0022149X25000392.
Acknowledgements
We would like to thank the Universidade Federal da Paraíba (UFPB), the biologist Gilson Melo and our colleagues at the Hydrobiology, Microbiology and Parasitology Laboratory (LAHMP), Invertebrate Collection Paulo Young (CIPY), and the Herpetological Collection of the Universidade Federal da Paraíba (CHUFPB) for their support during the research and the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for granting the research scholarship.
Financial support
This work was supported by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – CAPES (P.H.I.B., master’s research fellow scholarship DS/CAPES 88882.440357/2019-01). A.C.F.L. is supported by PELD-RIPA (Rio Paraíba Integrado, Chamada FAPESQ/PELD Nº 21/2020, nº termo de outorga: 403/2021).
Competing interest
Authors declare they have no financial interests.
Ethical standards
All authors gave their consent to participation in the study. The study was approved by the Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) with permission to collect the animals and in accordance with the ethical standards of relevant national and institutional guidelines on the care and use of laboratory animals.