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ISSN Versión impresa 2218-6425
ISSN Versión Electrónica 1995-1043
REVIEW ARTICLE / ARTICULO DE REVISIÓN
CHECKLIST OF HELMINTHS ASSOCIATED WITH CONTINENTAL TESTUDINES FROM SOUTH
AMERICA
LISTA DE HELMINTOS ASOCIADOS A LOS TESTUDINES CONTINENTALES DE AMÉRICA DEL
SUR
1
Laboratório de Parasitologia de Animais Silvestres (LAPASIL), Departamento de Microbiologia e Parasitologia, Instituto de
Biologia, Universidade Federal de Pelotas (UFPel), Caixa Postal: 354, CEP 96010-900, Pelotas, RS, Brazil.
*Corresponding author: phrybio@hotmail.com
Carolina S. Mascarenhas: https://orcid.org/0000-0002-7045-0926
Gertrud Müller: https://orcid.org/0000-0002-2213-6721
11
Carolina S. Mascarenhas* & Gertrud Müller
Neotropical Helminthology
97
doi:10.24039/rnh20201511047
Neotropical Helminthology, 2021, 15(1), ene-jun:97-126.
D
ABSTRACT
This study collected records of 135
taxa
of parasitic helminths (Nematoda, Trematoda, Cestoda,
Monogenoidea and Acanthocephala) and ecto-symbionts (Temnocephalida) associated with continental
Testudines from South America. Eighty-nine helminths were identified at the species level while others
were identified up to genus or family levels. The greatest diversity of helminths associated with
Testudines was reported in Brazil. Chelidae was the family with the largest number of helminth species.
Regarding the conservation status, 17 Testudines species with helminth records are cited in the IUCN Red
List of Threatened Species. Knowledge of helminth biodiversity and of relations between these organisms
and Testudines can provide important data on host biology. Therefore, information generated by studies of
helminths can contribute to research which aims at the conservation of organisms and their habitats.
D
Keywords:
Acanthocephala – biodiversity – Cestoda – Chelidae – conservation – Digenea –ecto-symbiont – Emydidae – freshwater turtle –
Geoemydidae – IUCN – Kinosternidae – Monogenoidea – Nematoda – parasite – Podocnemididae – Temnocephala – Testudinidae – tortoise
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Among helminths that can be associated with
Testudines, there are parasites (e.g., Nematoda,
Trematoda) (Vicente
et al
., 1993; Fernandes &
Kohn, 2014) and ectosimbionts (e.g.,
Temnocephalida) (Martínez-Aquino
et al.
, 2014).
Parasitic organisms represent a significant part of
biodiversity; according to Windsor (1998), they
represent the largest number of species on Earth.
There is at least a kind of parasite associated with
every free-living species (Poulin, 1999).
According to Dobson
et al.
(2008), there are
between 75,000 and 300,000 species of helminths
parasitizing vertebrates; this estimate suggests that
there may be more parasitic species than free-living
ones, reinforcing Windsor's findings (1998).
Temnocephalida are composed of freshwater
ectosymbionts with specificity to the host group or
species (Martínez-Aquino
et al.
, 2014).
Testudines comprise 478
taxa
(356 species and 122
subspecies) worldwide; seven species and three
subspecies, i. e., 10
taxa
(2.1%), have been extinct
(Rhodin
et al.
, 2017). In South America,
continental Testudines are represented by 71
taxa
;
37 (35 threatened ones and two extinct ones) out of
71
taxa
are cited in the International Union for
Conservation of Nature (IUCN) Red List
of
Threatened Species
(Rhodin
et al.
, 2017; IUCN,
98
RESUMEN
En este estudio se recogieron registros de 135
taxones
de helmintos parásitos (Nematoda, Trematoda,
Cestoda, Monogenoidea y Acanthocephala) y ectosimbiontes (Temnocephalida) asociados a Testudines
continentales de Sudamérica. Se identificaron 89 helmintos a nivel de especie, mientras que otros se
identificaron hasta el nivel de género o familia. La mayor diversidad de helmintos asociados a los
Testudines se registró en Brasil. Chelidae fue la familia con el mayor número de especies de helmintos. En
cuanto al estado de conservación, 17 especies de Testudines con registros de helmintos están citadas en la
Lista Roja de Especies Amenazadas de la IUCN. El conocimiento de la biodiversidad de los helmintos y
de las relaciones entre estos organismos y los Testudines puede aportar datos importantes sobre la biología
de los hospedadores. Por lo tanto, la información generada por los estudios de los helmintos puede
contribuir a la investigación que tiene como objetivo la conservación de los organismos y sus hábitats.
INTRODUCTION
2020). Continental Testudines undergo several
negative impacts that affect their populations. The
main threats are: fragmentation of terrestrial or
aquatic habitats, degradation of water quality,
disorderly occupation of spawning grounds,
consumption of meat and eggs, illegal trade for pets
and death by running over (Gibbons
et al.
, 2000;
Gibbs & Shriver, 2002; Steen & Gibbs, 2004;
Bujes & Verrastro, 2008). Anthropic actions, such
as habitat change, loss of biodiversity, pollution,
climate change and introduction of species, can
also impact helminth fauna (Lafferty & Kuris,
2005). Dobson
et al
. (2008) estimated that from 3%
to 5% of parasitic helminths will be threatened with
extinction in the next 50 to 100 years.
In the literature, there is no checklist of parasitic
and ectosymbiont helminths associated with
continental Testudines from South America. Cohen
et al.
(2013) carried out a checklist of
Monogenoidea, while Fernandes & Kohn (2014)
introduced one of Trematoda from South America
and Martínez-Aquino
et al.
(2014) listed the
records of Temnocephalida in the Neotropical
region. Therefore, this checklist, which gathered
the records of parasitic and ectosymbiont
helminths associated with continental Testudines
from South America, aiming at contributing to the
knowledge of the diversity of helminths associated
with the group and at encouraging
helminthological studies with group of vertebrates.
Neotropical Helminthology, 2021, 15(1), ene-jun
Palabras clave:
Acanthocephala – biodiversidad – Cestoda – Chelidae – conservación – Digenea – ectosimbionte – Emydidae – Geoemydidae –
IUCN – Kinosternidae – Monogenoidea – Nematoda parasito – Podocnemididae – Temnocephala – Testudinidae – tortuga de agua dulce
Mascarenhas & Müller
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99
The checklist resulted from information published
in articles, books, theses and dissertations. Some
specimens deposited in the Helminth Collection
that belongs to the Parasitology Laboratory of Wild
Animals (CHLAPASIL) at the Microbiology and
Parasitology Department in the Institute of Biology
at the Universidade Federal de Pelotas (UFPel),
located in Pelotas, RS, Brazil, were also examined.
Classification and systematization of helminths
were carried out in agreement with Anderson
et al.
(2009) for Nematoda; Gibson
et al.
(2002), Jones
et
al.
(2005), and Bray
et al.
(2008) for Digenea;
Khalil
et al.
(1994) for Cestoda; Boeger & Kritsky
(1993) for Monogenoidea; Petrochenko (1971) for
Acanthocephala; and Tyler
et al.
(2006-2019) for
Temnocephalida. Classification and
systematization of hosts followed Rhodin
et al.
(2017).
Helminth
taxa
are shown in alphabetical order in
their respective superfamilies, followed by host
taxa
, site of infection (parasitic helminths) or
infestation (ectosymbionts), geographical location
and respective bibliographical references (between
parentheses and chronological order). Citation of
helminth and host species in this list does not mean
that the authors agree with their validity and
taxonomy. The list of host-helminth includes the
conservation status of Testudines species in
agreemen with the Red List of Threatened Species
issued by the IUCN (2020).
One hundred and thirty-five helminths were
recorded in association with 39 continental
Testudines (34 species, three identified at the genus
level and two unidentified) from nine South
American countries. Eighty-nine helminths (40
Trematoda, 38 Nematoda, six Monogenoidea,
three Temnocephalida and two Cestoda) were
identified at the species level, while the others were
identified up to genus (44
taxa
) and family levels (2
taxa
).
The greatest diversity of helminths associated with
Testudines was reported in Brazil (94
taxa
),
MATERIAL AND METHODS
followed by Uruguay (17
taxa
), Peru (15
taxa
),
Colombia (14
taxa
), Argentina (13
taxa
),
Venezuela (10
taxa
), Ecuador (10
taxa
), French
Guiana (2
taxa
), Paraguay (1
taxa
) and Bolivia (1
taxa
). Figure 1 shows the number of helminths
(according to the group) registered in each country.
No records of helminthological studies were found
in Chile, Suriname and Guyana. Chelidae was not
only the family with the greatest diversity of
recorded species (32), but also the group with the
largest number of hosts under study (12 species)
(Fig. 2). Podocnemidae and Emydidae had records
of 30 and 14 species of helminths, respectively
(Fig. 2). The largest numbers of
taxa
were
associated with
Phrynops hilarii
(Duméril &
Bibron, 1835) (21
taxa
) and
Hydromedusa
tectifera
Cope, 1870 (20
taxa
) (Chelidae) in
records from Brazil, Uruguay and Argentina.
Nematoda and Trematoda occurred in all groups of
Testudines (Fig. 2); their main site of infection was
the digestive tract of the hosts. Monogenoidea was
mainly represented by
Polystomoides
Ward, 1917
species (Polystomatidae) parasites of the oral
cavity of Chelidae species. Two Cestoda species, in
Geoemydidae and in Chelidae, were recorded.
Acanthocephala was only recorded in a Testudines
species,
Acanthochelys spixii
(Duméril & Bibron,
1835) (Chelidae). Ectosymbiont helminths,
Temnocephalidae, were recorded in six Chelidae
species and in an Emydidae species. The digenetic
trematode
Nematophila grandis
(Diesing, 1839)
was the taxon recorded in the largest number of
host species (17) belonging to Kinosternidae,
Geoemydidae, Testudinidae, Chelidae and
Podocnemidae. The species, which has wide
geographical distribution, was found in Brazil,
French Guyana, Venezuela, Ecuador, Peru,
Paraguay and Argentina. Atractidae was the group
of Nematoda with the largest number of
representatives (27
taxa
), which were mainly
registered in Podocnemidae species in Brazil.
Among Testudines reported by helminthological
studies in South America, 17 were cited in the
IUCN Red List of Threatened Species. However,
three species,
Kinosternon integrum
Le Conte,
1854 (Kinosternidae),
Emys orbicularis
(Linnaeus,
1758) (Emydidae) and
Kinixys erosa
(Schweigger,
1812) (Testudinidae), are not native to the region.
Figure 3 shows the number of helminths associated
with species native to South America that were
cited in the Red List. Most of these Testudines have
RESULTS
Neotropical Helminthology, 2021, 15(1), ene-jun
Helminths associated with continental Testudines
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Figure 2.
Number of helminth species associated with continental Testudines families from South America.
Numbers of species studied in each family are between parentheses.
100
Figure 1.
Number of helminths associated with continental Testudines from South America.
Neotropical Helminthology, 2021, 15(1), ene-jun
Mascarenhas & Müller
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101
few records of helminths. However, Nematoda and
Trematoda are the most frequently reported ones
(Fig. 3).
HELMINTH-HOST LIST
Phylum Nematoda (Rudolphi, 1808)
Superfamily Dioctophymatoidea Railliet, 1916
Family Dioctophymatidae (Railliet, 1915)
Dioctophyme
Collet-Meygret, 1802
Dioctophyme renale
(Goeze, 1782) (larvae)
Acanthochelys spixii
(Duméril & Bibron, 1835),
large intestine, BRAZIL (Rio Grande do Sul State)
(Chaviel
et al.,
2020).
Hydromedusa tectifera
Cope, 1870, body cavity
(surface of stomach, and muscles) and serous of
stomach BRAZIL (Rio Grande do Sul State)
(present study, CHLAPASIL 882).
Phrynops hilarii
(Duméril & Bibron, 1835),
muscles, body cavity (surface of esophagus,
stomach, lung, liver) and serous of stomach,
BRAZIL (Rio Grande do Sul State) (Mascarenhas
et al.
, 2017).
Trachemys dorbigni
(Duméril & Bibron, 1835),
body cavity, muscles, mesentery, stomach serous
lining and on surfaces of the lung, heart, liver,
pancreas, spleen and intestine, BRAZIL (Rio
Grande do Sul State) (Mascarenhas & Müller,
2015a).
Eustrongylides
Jägerskiöld, 1909
Eustrongylides
sp. (larvae)
Trachemys dorbigni
(Duméril & Bibron, 1835),
stomach (serous), BRAZIL (Rio Grande do Sul
State) (Mascarenhas, 2014).
Superfamily Strongyloidea Weinland, 1858
Family Strongylidae Baird, 1853
Chapiniella
Yamaguti, 1961
Chapiniella variabilis
(Chapin, 1924)
Chelonoidis carbonarius
(Spix, 1824), large
intestine, BRAZIL (Piauí State) (Freire
et al.,
2017;
Freire
et al
., 2019).
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, BRAZIL (Pará State) (Chapin, 1924).
Figure 3.
Number of registered helminths in association with native continental Testudines from South
America cited in the IUCN Red List of Threatened Species (2020). LR – Lower Risk; NT – Near threatened; VU
– Vulnerable; CR – Critically Endangered.
Neotropical Helminthology, 2021, 15(1), ene-jun
Helminths associated with continental Testudines
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102
Chelonoidis denticulatus
(Linnaeus, 1766),
large
intestine, BRAZIL (Piauí State) (Freire
et al.,
2017;
Freire
et al
., 2019).
Sauricola
Chapin, 1924
Sauricola sauricola
Chapin, 1924
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, BRAZIL (Pará State) (Chapin, 1924).
Superfamily Oxyuroidea
Railliet, 1916
Family Pharyngodonidae Travassos, 1919
Thelandros
Wedl, 1862
Thelandros
sp.
Chelonoidis carbonarius
(Spix, 1824), intestine,
VENEZUELA (Bolivar State) (Pérez Mata
et al
.,
2014).
Pharyngodonidae gen. sp.
Acanthochelys spixii
(Duméril & Bibron, 1835),
large intestine, BRAZIL (Rio Grande do Sul State)
(present estudy, CHLAPASIL, 889).
Superfamily Cosmocercoidea Skrjabin &
Shikhobalova, 1951
Family Kathlaniidae (Lane, 1914)
Falcaustra
Lane, 1915
Falcaustra affinis
(Leidy, 1856)
Trachemys dorbigni
(Duméril & Bibron, 1835),
large intestine, heart, kidney, stomach, BRAZIL
(Rio Grande do Sul State) (Mascarenhas & Müller,
2015b).
Falcaustra tikasinghi
(Schroeder, Schmidt &
Everard, 1977)
Rhinoclemmys annulata
(Gray, 1860), digestive
tract, ECUADOR (São Jose de Tagua, and Playa
Grande) (Dyer & Carr, 1990a).
Rhinoclemmys melanosterna
(Gray, 1861),
digestive tract, ECUADOR (Rio Bogotá) (Dyer &
Carr, 1990a).
Rhinoclemmys nasuta
(Boulenger, 1902),
digestive tract, ECUADOR (Sarria, Estero el
Ceibo, and Playa Grande) (Dyer & Carr, 1990a).
Rhinoclemmys punctularia
(Daudin, 1801),
unspecified site of infection, BRAZIL (Pará State)
(Baker & Bain, 1981).
Falcaustra
sp.
Chelonoidis chilensis
(Gray, 1870), small
intestine, ARGENTINA (San Juan Province)
(Castillo
et al
., 2020).
Family Atractidae Travassos, 1919
Atractis
Dujardin, 1845
Atractis caballeroi
Brenes & Bravo-Hollis, 1960
Kinosternon leucostomum
Duméril & Bibron,
1851, digestive tract, ECUADOR (Rio Bogotá)
(Dyer & Carr, 1990a).
Rhinoclemmys annulata
(Gray, 1860), digestive
tract, ECUADOR (Playa Grande) (Dyer & Carr,
1990a).
Atractis cruciata
Linstow, 1902
Podocnemis expansa
(Schweigger, 1812),
stomach, BRAZIL (Tocantins State) (Armond,
2008).
Atractis dactyluris
Rudolphi
1819
Mesoclemmys nasuta
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Diesing,
1851).
Peltocephalus dumerilianus
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Diesing,
1851).
Podocnemis expansa
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Diesing,
1851).
Atractis impura
(Caballero, 1944)
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, PERU (Loreto and Ucayali
Departament) (Salizar & Sanchez, 2007).
Rhinoclemmys diademata
(Mertens, 1954),
digestive tract, VENEZUELA (Lake Maracaib
basin) (Dyer & Carr, 1990a).
Atractis marquezi
Bursey & Flanagan 2002
Chelonoidis porteri
(Rothschild, 1903), colon,
ECUADOR (Isla Santa Cruz, Galapagos) (Bursey
& Flanagan, 2002).
Atractis thapari
Petter, 1966
Chelonoidis carbonarius
(Spix, 1824), small and
large intestines, BRAZIL (Piauí State) (Leal
et al.,
2018; Freire
et al
., 2019).
Chelonoidis denticulatus
(Linnaeus, 1766), small
and large intestines, BRAZIL (Piauí State) (Leal
et
al.,
2018; Freire
et al
., 2019).
Atractis trematophila
Travassos, 1934
Amazon river turtle, digestive tract of
Nematophila
grande
(Digenea), BRAZIL (unspecified locality)
(Travassos, 1934a).
Peltocephalus dumerilianus
(Schweigger, 1812),
Neotropical Helminthology, 2021, 15(1), ene-jun
Mascarenhas & Müller
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103
stomach, small and large intestine, BRAZIL
() (Ferreira, 2016).
Amazonas State
Atractis
sp.
Podocnemis expansa
(Schweigger, 1812),
stomach, BRAZIL (Tocantins State) (Armond,
2008).
Atractis
sp.
Chelonoidis carbonarius
(Spix, 1824), intestine,
VENEZUELA (Bolivar State) (Pérez Mata
et al
.,
2014).
Atractis
sp.
Chelonoidis porteri
(Rothschild, 1903), intestine,
ECUADOR (Isla Santa Cruz, Galapagos) (Fournié
et al
., 2015).
Buckleyatractis
Khalil & Gibbons 1988
Buckleyatractis marinkelli
Khalil & Gibbons
1988
Podocnemis unifilis
Troschel, 1848, large intestine,
COLOMBIA (unspecified locality) (Khalil &
Gibbons, 1988).
Labiduris
Schneider, 1866
Labiduris gulosa
(Rudolphi, 1819)
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, BRAZIL (Pará State) (Chapin, 1924).
Mesoclemmys nasuta
(Schweigger, 1812),
intestine, BRASIL (unspecified locality) (Diesing,
1851).
Labiduris irineuta
Costa, 1961
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, BRAZIL (Rio de Janeiro State) (Costa,
1961 cited by Vicente
et al.
, 1993).
Labiduris zschokkei
Linstow, 1899
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, BRAZIL (unspecified locality) (Skrjabin
et al.,
1964 cited by Vicente
et al.,
1993).
Labiduris
sp.
Chelonoidis porteri
(Rothschild, 1903), intestine,
ECUADOR (Isla Santa Cruz, Galapagos) (Fournié
et al
., 2015).
Labiduris
sp.
Chelonoidis chilensis
(Gray, 1870), small
intestine, ARGENTINA (San Juan Province)
(Castillo
et al
., 2020).
Klossinemella
Costa, 1961
Klossinemella conciliatus
Alho, 1964
Peltocephalus dumerilianus
(Schweigger 1812),
stomach, small and large intestine, BRAZIL
() (Ferreira, 2016).
Amazonas State
Podocnemis expansa
(Schweigger, 1812),
stomach, BRAZIL (Amazonas State) (Alho 1964a;
Costa
et al.,
1968).
Podocnemis expansa
(Schweigger, 1812),
stomach, BRAZIL (Tocantins State) (Armond,
2008).
Podocnemis unifilis
Troschel, 1848, stomach,
PERU (Samiria river) (Salizar & Sanchez, 2007).
Klossinemella travassosi
Costa, Mota & Gomes,
1968
Chelonoidis denticulatus
(Linnaeus, 1766), large
intestine, BRAZIL (Amazonas State) (Costa
et al.,
1968; Muniz-Pereira
et al.,
2009).
Klossinemella
sp.
Peltocephalus dumerilianus
(Schweigger, 1812),
stomach, small and large intestine, BRAZIL
() (Ferreira, 2016).
Amazonas State
Klossinemella
sp.
Peltocephalus dumerilianus
(Schweigger, 1812),
stomach, small and large intestine, BRAZIL
() (Ferreira, 2016).
Amazonas State
Klossinemella
sp.
Peltocephalus dumerilianus
(Schweigger, 1812),
stomach, small and large intestine, BRAZIL
() (Ferreira, 2016).
Amazonas State
Orientatractis
Petter, 1966
Orientatractis leiperi
Buckley, 1969
Podocnemis vogli
Müller, 1935, "stomach"
(presumably colon), COLOMBIA (unspecified
locality) (Buckley, 1969).
Podocnemis unifilis
Troschel, 1848, stomach,
BRAZIL (Tocantins River, Pará State) (Jesus
et al.
,
2020).
Paraorientatractis
Gibbons, Khalil &
Marinkelle, 1997
Paraorientatractis semiannulata
Gibbons,
Khalil & Marinkelle, 1997
Podocnemis unifilis
Troschel, 1848, large intestine,
BRAZIL (Cuminá and Trombetas rivers near Pará
State) (Gibbons
et al.,
1997).
Podocnemis unifilis
Troschel, 1848, intestine,
Neotropical Helminthology, 2021, 15(1), ene-jun
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104
PERU (Ucayali Departament) (Salizar & Sanchez,
2007).
Paratractis
Sarmiento, 1959
Paratractis hystrix
(Diesing, 1851)
Peltocephalus dumerilianus
(Schweigger, 1812),
intestine, PERU (Ucayali Departament)
Pucallpa,
(Sarmiento, 1959).
Peltocephalus dumerilianus
(Schweigger, 1812),
stomach, small and large intestine, BRAZIL
() (Ferreira, 2016).
Amazonas State
Podocnemis erythrocephala
(Spix, 1824),
intestine, BRAZIL (Amazonas State) (Diesing,
1851).
Podocnemis expansa
(Schweigger, 1812), large
intestine COLOMBIA (unspecified locality)
(Khalil & Gibbons, 1988).
Podocnemis unifilis
Troschel, 1848, large intestine,
COLOMBIA (unspecified locality) (Khalil &
Gibbons, 1988).
Podocnemis vogli
Müller, 1935, unspecified site of
infection, COLOMBIA (unspecified locality)
(Buckley, 1969; Buckley, 1970).
Podocnemis vogli
Müller, 1935, large intestine
COLOMBIA (unspecified locality) (Khalil &
Gibbons, 1988).
Pneumoatractis
Bursey, Reavill & Greiner, 2009
Pneumoatractis podocnemis
Bursey, Reavill &
Greiner, 2009
Podocnemis unifilis
Troschel, 1848, lungs, South
America (Unknown, wild-caught, confiscated
South America turtle) (Bursey
et al.,
2009).
Podocnematractis
Gibbons, Khalil &
Marinkelle, 1995
Podocnematractis colombiaensis
Gibbons,
Khalil & Marinkelle, 1995
Podocnemis expansa
(Schweigger, 1812), large
intestine, COLOMBIA (unspecified locality)
(Gibbons
et al.,
1995).
Podocnemis vogli
Müller, 1935, large intestine,
COLOMBIA (unspecified locality) (Gibbons
et
al.,
1995).
Podocnematractis
ortleppi
(Thapar, 1925)
Podocnemis expansa
(Schweigger, 1812), large
intestine, COLOMBIA (unspecified locality)
(Gibbons
et al.,
1995).
Podocnemis unifilis
Troschel, 1848, large intestine,
COLOMBIA (unspecified locality) (Gibbons
et
al.,
1995).
Podocnemis vogli
Müller, 1935, large intestine,
COLOMBIA (unspecified locality) (Gibbons
et
al.,
1995).
Superfamily Ascaridoidea Railliet & Henry,
1915
Family Anisakidae Skrjabin & Karokhin, 1945
Contracaecum
Railliet & Henry, 1912
Contracaecum
sp. (larvae)
Acanthochelys spixii
(Duméril & Bibron, 1835),
unspecified site of infection, BRAZIL (Rio Grande
do Sul State) (Mascarenhas
et al.,
2017).
Family Ascarididae Baird, 1853
Angusticaecum
Baylis, 1920
Angusticaecum brevispiculum
Chapin, 1924
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, BRAZIL (Pará State) (Chapin, 1924).
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, PERU (Loreto and Ucayali
Departament) (Salizar & Sanchez, 2007).
Brevimulticaecum
Mozgovoy, 1951 in Skrjabin,
Shikhobalova & Mozgovoi, 1952
Brevimulticaecum
sp. (larvae)
Phrynops geoffroanus
(Schweigger, 1812), small
intestine, BRAZIL (São Paulo State) (Silva, 2014).
Superfamily Camallanoidea Travassos, 1920
Family Camallanidae Railliet & Henry, 1915
Camallanus
Railliet & Henry, 1915
Camallanus
emydidius
Mascarenhas & Müller,
2017
Hydromedusa tectifera
Cope, 1870, small
intestine, BRAZIL (Rio Grande do Sul State)
(Mascarenhas
et al
., 2013; Chaviel
et al.,
2020).
Trachemys dorbigni
(Duméril & Bibron, 1835),
small intestine, BRAZIL (Rio Grande do Sul
State), (Mascarenhas & Müller, 2017).
Camallanus kachugae
Baylis & Daubney, 1922
Kinosternon scorpioides
(Linnaeus, 1766),
intestine, VENEZUELA (Zulia State) (Díaz-
Ungría, 1978).
Camallanus
sp.
Phrynops hilarii
(Duméril & Bibron, 1835),
stomach and small intestine, BRAZIL (Rio Grande
do Sul State) (Bernardon
et al.,
2013).
Camallanus
sp.
Trachemys dorbigni
(Duméril & Bibron, 1835),
Neotropical Helminthology, 2021, 15(1), ene-jun
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105
small intestine, BRAZIL (Rio Grande do Sul State)
(Bernardon
et al.,
2014).
Camallanus
sp.
Acanthochelys spixii
(Duméril & Bibron, 1835),
small intestine, BRAZIL (Rio Grande do Sul State)
(Mascarenhas
et al.,
2013).
Camallanus
sp.
Phrynops geoffroanus
(Schweigger, 1812), small
intestine, BRAZIL (Minas Gerais State) (Vieira
et
al.,
2016).
Serpinema
Yeh, 1960
Serpinema amazonicus
(Ribeiro, 1940)
Podocnemis expansa
(Schweigger, 1812), small
intestine, BRAZIL (Pará State) (Ribeiro, 1940).
Podocnemis expansa
(Schweigger, 1812),
intestine, PERU (Loreto Departament) (Tantaleán,
1998).
Podocnemis unifilis
Troschel, 1848, small
intestine, PERU (Loreto Departament) (Sánchez
et
al.,
2006).
Serpinema cayennensis
Harnoster, Svitin &
Preez, 2019
Rhinoclemmys punctularia
(Daudin, 1801),
intestine, FRENCH GUIANA (Cayenne)
(Harnoster
et al.,
2019).
Serpinema maghati
(Sprehn, 1932)
Kinosternon integrum
Le Conte, 1854, intestine,
BOLIVIA (unspecified locality) (Sprehn, 1932).
Kinosternon scorpioides
(Linnaeus, 1766), small
intestine, BRAZIL (Pernambuco, and Pará States)
(Alho, 1965; Freitas & Dobbin Jr., 1971).
Kinosternon scorpioides
(Linnaeus, 1766),
stomach, small and large intestine, BRAZIL
(Maranhão State) (Viana
et al.,
2016).
Serpinema microcephalus
(Dujardin, 1845)
Peltocephalus dumerilianus
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Diesing,
1851).
Podocnemis expansa
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Diesing,
1851).
Serpinema monospiculatus
Freitas & Dobbin
Jr., 1962
Kinosternon scorpioides
(Linnaeus, 1766), large
intestine, BRAZIL (Ceará State) (Pereira
et al.,
2018).
Mesoclemmys nasuta
(Schweigger, 1812), small
intestine, BRAZIL (Pernambuco State) (Freitas &
Dobbin Jr., 1971).
Mesoclemmys turbeculata
(Lüderwaldt, 1926),
small intestine, BRAZIL (Pernambuco State)
(Freitas & Dobbin Jr., 1971).
Mesoclemmys turbeculata
(Lüderwaldt, 1926),
small and large intestines, BRAZIL (Ceará State)
(Pereira
et al.
, 2018).
Phrynops geoffroanus
(Schweigger, 1812), small
intestine, BRAZIL (Pernambuco State) (Freitas &
Dobbin Jr., 1971).
Phrynops geoffroanus
(Schweigger, 1812), small
intestine, BRAZIL (São Paulo State) (Silva, 2014).
Phrynops geoffroanus
(Schweigger, 1812),
stomach, small and large intestines, lungs, body
cavity BRAZIL (Ceará State) (Pereira
et al.,
2018).
Spirocamallanus
Olsen, 1952
Spirocamallanus
sp.
Hydromedusa tectifera
Cope, 1870, small
intestine, BRAZIL (Minas Gerais State) (Novelli
et
al.,
2014).
Superfamily Gnathostomatoidea
Railliet, 1895
Family Gnathostomatidae
Railliet, 1895
Ancyracanthus
Diesing, 1858
Ancyracanthus pinnatifidus
Diesing, 1839
Peltocephalus dumerilianus
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Diesing,
1851).
Peltocephalus dumerilianus
(Schweigger, 1812),
stomach and intestine, BRAZIL (unspecified
locality) (Gomes & Kohn, 1970 cited by Vicente
et
al.,
1993).
Peltocephalus dumerilianus
(Schweigger, 1812),
stomach, BRAZIL () (Ferreira,
Amazonas State
2016).
Podocnemis expansa
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Diesing,
1851).
Podocnemis unifilis
Troschel, 1848, small
intestine, PERU (Loreto Departament) (Sánchez
et
al.,
2006).
Gnathostoma
Owen, 1836
Gnathostoma
sp. (larvae)
Hydromedusa tectifera
Cope, 1870, small
intestine, BRAZIL (Rio Grande do Sul State)
(present study, CHLAPASIL 881).
Spiroxys
Schneider, 1866
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Spiroxys contortus
(Rudolphi, 1819)
Acanthochelys spixii
(Duméril & Bibron, 1835),
stomach
, BRAZIL (Rio Grande do Sul State)
(Mascarenhas
et al.,
2013).
Hydromedusa tectifera
Cope, 1870,
esophagus,
stomach, small and large intestines, and cavity
,
BRAZIL (Rio Grande do Sul State) (Mascarenhas
et al.,
2013).
Phrynops hilarii
(Duméril & Bibron, 1835),
stomach, ARGENTINA (Buenos Aires Province)
(Palumbo
et al.,
2016).
Phrynops hilarii
(Duméril & Bibron, 1835),
stomach, small and large intestines, BRAZIL (Rio
Grande do Sul State) (Chaviel
et al.,
2020).
Trachemys dorbigni
(Duméril & Bibron, 1835),
esophagus, stomach, small and large intestines,
heart, BRAZIL (Rio Grande do Sul State)
(Mascarenhas & Müller, 2015b).
Spiroxys figueiredoi
Freitas & Dobbin Jr., 1962
Kinosternon scorpioides
(Linnaeus, 1766),
stomach, BRAZIL (Pará State) (Alho, 1965).
Kinosternon scorpioides
(Linnaeus, 1766),
stomach, BRAZIL (Pernambuco State) (Freitas &
Dobbin Jr., 1962; Freitas & Dobbin Jr., 1971).
Kinosternon scorpioides
(Linnaeus, 1766),
stomach, small and large intestine, BRAZIL
(Maranhão State) (Viana
et al.,
2016).
Mesoclemmys turbeculata
(Lüderwaldt, 1926),
stomach, small intestine, BRAZIL (Ceará State)
(Pereira
et al.,
2018).
Phrynops geoffroanus
(Schweigger, 1812),
stomach, BRAZIL (Ceará State) (Pereira
et al.,
2018).
Phrynops geoffroanus
(Schweigger, 1812),
stomach, BRAZIL (São Paulo State) (Silva, 2014).
Spiroxys
sp.
Mesoclemmys vanderhaegei
(Bour, 1973),
stomach, BRAZIL (Mato Grosso State) (Ávila
et
al.,
2010).
Spiroxys
sp.
Phrynops hilarii
(Duméril & Bibron, 1835),
stomach and small intestine, BRAZIL (Rio Grande
do Sul State) (Bernardon
et al.,
2013).
Spiroxys
sp.
Trachemys dorbigni
(Duméril & Bibron, 1835),
stomach and small intestine, BRAZIL (Rio Grande
do Sul State) (Bernardon
et al.,
2014).
Superfamily Physalopteroidea Sobolev 1949
Family Physalopteridae Leiper, 1908
Physaloptera
Rudolphi, 1819
Physaloptera retusa
Rudolphi, 1819
Mesoclemmys turbeculata
(Lüderwaldt, 1926),
large intestine, BRAZIL (Ceará State) (Pereira
et
al.,
2018).
Phrynops geoffroanus
(Schweigger, 1812),
stomach, small intestine, BRAZIL (Ceará State)
(Pereira
et al.,
2018).
Physaloptera
sp. (larvae)
Acanthochelys spixii
(Duméril & Bibron, 1835),
stomach, BRAZIL (Rio Grande do Sul State)
(Chaviel
et al.,
2020).
Physaloptera
sp. (larvae)
Phrynops geoffroanus
(Schweigger, 1812),
stomach, BRAZIL (São Paulo State) (Silva, 2014).
Superfamily Habronematoidea
Railliet &
Henry, 1915
Family Hedruridae Railliet, 1916
Hedruris
Nitzch, 1821
Hedruris dratini
Palumbo, Servián, Sánchez &
Diaz, 2019
Hydromedusa tectifera
Cope, 1870, stomach,
ARGENTINA (Buenos Aires Province) (Palumbo
et al.,
2019).
Phrynops hilarii
(Duméril & Bibron, 1835),
stomach, ARGENTINA (Buenos Aires Province)
(Palumbo
et al.,
2019).
Hedruris orestiae
(Moniez, 1889)
Hydromedusa tectifera
Cope, 1870, stomach,
ARGENTINA (Buenos Aires Province) (Palumbo
et al.
, 2016).
Phrynops hilarii
(Duméril & Bibron, 1835),
stomach, ARGENTINA (Buenos Aires Province)
(Palumbo
et al.
, 2016).
Hedruris
sp.
Rhinoclemmys nasuta
(Boulenger, 1902), stomach,
ECUADOR (Estero el Ceibo) (Dyer & Carr,
1990a).
Phylum Platyhelminthes Gegenbaur, 1859
Class Trematoda Rudolphi, 1808
Subclass Digenea Carus, 1863
Superfamily Diplostomoidea Poirier, 1886
Family Proterodiplostomidae Dubois, 1936
Cheloniodiplostomum
Sudaricov, 1960
Cheloniodiplostomum argentinensis
Palumbo &
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107
Diaz, 2018
Phrynops hilarii
(Duméril & Bibron, 1835),
anterior portion of intestine,
ARGENTINA
(Buenos Aires Province)).
(Palumbo & Diaz, 2018
Cheloniodiplostomum brevis
(MacCallum, 1921)
Phrynops geoffroanus
(Schweigger, 1812),
unspecified site of infection, COLOMBIA
(unspecified locality) (Dubois, 1979).
Cheloniodiplostomum testudinis
(Dubois, 1936)
Hydromedusa tectifera
Cope, 1870,
anterior
portion of intestine,
ARGENTINA (Buenos Aires
Province) (Palumbo
et al.
, 2018).
Phrynops geoffroanus
(Schweigger, 1812), small
and large intestines, BRAZIL (São Paulo State)
(Silva, 2014).
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, BRAZIL (Rio Grande do Sul State)
(Mascarenhas
et al.
, 2016).
Phrynops hilarii
(Duméril & Bibron, 1835),
intestine, ARGENTINA (Corrientes Province)
(Lombardero & Moriena, 1977).
Phrynops hilarii
(Duméril & Bibron, 1835),
anterior portion of intestine,
ARGENTINA
(Buenos Aires Province) (Palumbo
et al.
, 2018).
Testudo
sp.,
unspecified site of infection, BRAZIL
(unspecified locality) (Dubois 1936 cited by
Fernandes & Kohn, 2014).
Cheloniodiplostomum
sp.
Phrynops hilarii
(Duméril & Bibron, 1835),
stomach and small intestine, BRAZIL (Rio Grande
do Sul State) (Bernardon
et al.
, 2013).
Cheloniodiplostomum
sp.
Phrynops geoffroanus
(Schweigger, 1812), small
intestine, BRAZIL (Minas Gerais State) (Novelli
et
al.,
2013).
Cheloniodiplostomum
sp.
Acanthochelys spixii
(Duméril & Bibron, 1835),
small and large intestine, BRAZIL (Rio Grande do
Sul State) (Mascarenhas
et al.
, 2016).
Cheloniodiplostomum
sp.
Trachemys dorbigni
(Duméril & Bibron, 1835),
small intestine, BRAZIL (Rio Grande do Sul State)
(Bernardon
et al.
, 2014).
Cheloniodiplostomum
sp.
Hydromedusa tectifera
Cope, 1870, small
intestine, BRAZIL (Rio Grande do Sul State)
(present study, CHLAPASIL 885).
Superfamily Schistosomatoidea Stiles &
Hassall, 1898
Family Spirorchiidae Stunkard, 1921
Atamatam
Bullard & Roberts, 2019
Atamatam amazoniensis
Bullard & Roberts,
2019
Chelus fimbriata
(Schneider, 1783), blood vessels
of kidney and mesentery, PERU (Iquitos
Departament) (Bullard
et al.
, 2019).
Paratamatam
Bullard & Roberts, 2019
Paratamatam iquitosiensis
Bullard & Roberts,
2019
Chelus fimbriata
(Schneider, 1783), blood vessels
of kidney and mesentery, PERU (Iquitos
Departament) (Bullard
et al
., 2019).
Superfamily Echinostomatoidea Looss, 1899
Family Echinostomatidae Looss, 1899
Prionosomoides
Freitas & Dobbin Jr., 1967
Prionosomoides
phrynopsis
(Mañé-Garzón &
Gil, 1961)
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, URUGUAY (Tacuarembó Departament)
(Mañé-Garzón & Gil, 1961a).
Prionosomoides scalaris
Freitas & Dobbin Jr.,
1967
Phrynops geoffroanus
(Schweigger, 1812), small
intestine, BRAZIL (Pernambuco State) (Freitas &
Dobbin Jr., 1967).
Phrynops hilarii
(Duméril & Bibron, 1835),
esophagus, ARGENTINA (Corrientes Province)
(Lombardero & Moriena, 1977).
Prionosomoides
sp.
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, BRAZIL (Rio Grande do Sul State)
(present study, CHLAPASIL, 886-887).
Family Rhytidodidae Odhner, 1926
Rhytidodes
Looss, 1901
Rhytidodes gelatinosus
(Rudolphi, 1819)
Podocnemis expansa
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality)
(Travassos
et al.,
1969).
Superfamily Heronimoidea Ward, 1917
Family Heronimidae Ward, 1917
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108
Heronimus
MacCallum, 1902
Heronimus mollis
(Leidy, 1856)
Trachemys callirostris
(Gray, 1856), lungs,
COLOMBIA (unspecified locality) (Lenis &
Vélez, 2011).
Superfamily Microscaphidioidea Looss, 1900
Family Microscaphidiidae Looss, 1900
Neodeuterobaris
Brooks, 1976
Neodeuterobaris pritchardae
Brooks, 1976
Podocnemis lewyana
Duméril, 1852, stomach,
COLOMBIA (Caldas Departament) (Brooks,
1976).
Podocnemis lewyana
Duméril, 1852, stomach,
COLOMBIA (Magdalena river) (Lenis & Vélez,
2011).
Octangioides
Price, 1937
Octangioides tlacotalpensis
Caballero, 1942
Rhinoclemmys nasuta
(Boulenger, 1902), large
intestine, ECUADOR (Esmeraldas Province)
(Dyer & Carr, 1990b).
Podocnemitrema
Alho & Vicente, 1964
Podocnemitrema papillosus
Alho & Vicente,
1964
Podocnemis expansa
(Schweigger, 1812),
stomach, BRAZIL (Amazonas State) (Alho &
Vicente, 1964).
Podocnemis expansa
(Schweigger, 1812),
digestive tract, PERU (Iquitos Departament)
(Tantaleán
et al.,
2011).
Podocnemis unifilis
Troschel, 1848, digestive
tract, PERU (Iquitos Departament) (Tantaleán
et
al.,
2011).
Superfamily Paramphistomatoidea Fischoeder,
1901
Family Cladorchiidae Fischoeder, 1901
Halltrema
Lent & Freitas, 1939
Halltrema avitellina
Lent & Freitas, 1939
Chelonoidis denticulatus
(Linnaeus, 1766),
stomach, BRAZIL (Pará State) (Alho, 1965).
Podocnemis expansa
(Schweigger, 1812),
stomach, BRAZIL (Tocantins State) (Armond,
2008).
Podocnemis expansa
(Schweigger, 1812),
unspecified site of infection, BRAZIL (Pará State)
(Lent & Freitas, 1939; Freitas & Lent, 1942 cited
by Fernandes & Kohn, 2014).
Podocnemis expansa
(Schweigger, 1812),
digestive tract, PERU (Iquitos Departament)
(Tantaleán
et al.
, 2011).
Podocnemis unifilis
Troschel, 1848, digestive
tract, PERU (Iquitos Departament) (Tantaleán
et
al.
, 2011).
Podocnemis
sp., unspecified site of infection,
VENEZUELA (unspecified locality) (Caballero &
Diaz-Ungria, 1958 cited by Fernandes & Kohn,
2014).
Halltrema heteroxenus
(Cordero & Vogelsang,
1940)
Podocnemis unifilis
Troschel, 1848, digestive
tract, PERU (Loreto Departament) (Tantaleán &
Forlong, 2013).
Podocnemis
sp., stomach, VENEZUELA (Guarico
State) (Cordero & Vogelsang, 1940).
Rhinoclemmys nasuta
(Boulenger, 1902),
digestive tract, ECUADOR (Esmeraldas Province)
(Dyer & Carr, 1990b).
Nematophila
Travassos, 1934
Nematophila argentinum
(Cordero &
Vogelsang, 1940)
Phrynops
sp., intestine, ARGENTINA (Misiones
Province) (Cordero & Vogelsang, 1940).
Podocnemis lewyana
Duméril, 1852, large
intestine, COLOMBIA (Bolívar Departament)
(Lenis & Vélez, 2011).
Trachemys callirostris
(Gray, 1856), large
intestine, COLOMBIA (Bolívar Departament)
(Lenis & Vélez, 2011).
Nematophila grandis
(Diesing, 1839)
Chelus fimbriata
(Schneider, 1783), intestine,
BRAZIL (unspecified locality) (Travassos, 1934b;
Travassos
et al.
,
1969).
Hydraspis schopfii
, intestine, BRAZIL
(unspecified locality) (Travassos, 1934b).
Hydromedusa tectifera
Cope, 1870, unspecified
site of infection, PARAGUAY (unspecified
locality) (Masi-Pallarés
et al.
, 1976 cited by
Fernandes & Kohn, 2014).
Kinixys erosa
(Schweigger, 1812), intestine,
BRAZIL (unspecified locality) (Travassos
et al.,
1969; Fernandes & Kohn, 2014).
Kinosternon scorpioides
(Linnaeus, 1766), small
and large intestine, BRAZIL (Pará State) (Alho,
1964b).
Kinosternon scorpioides
(Linnaeus, 1766),
stomach, VENEZUELA (Zulia State) (Díaz-
Ungría, 1978).
Mesoclemmys gibba
(Schweigger, 1812), intestine,
Neotropical Helminthology, 2021, 15(1), ene-jun
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109
2014).
Rhinoclemmys nasuta
(Boulenger, 1902), small
and large intestines, ECUADOR (Esmeraldas
Province) (Dyer & Carr, 1990b).
Rhinoclemmys punctularia
(Daudin, 1801),
large
intestine, BRAZIL (Pará State) (Alho, 1964b).
Rhinoclemmys punctularia
(Daudin, 1801),
digestive tract, GUYANA FRANCESA (Iracoubo)
(Dyer & Carr, 1990b).
Nematophila venezuelenisis
(Cordero &
Vogelsang, 1940)
Podocnemis lewyana
Duméril, 1852, stomach,
COLOMBIA (Bolívar and Magdalena
Departaments) (Lenis & Vélez, 2011).
Podocnemis
sp., stomach, VENEZUELA
(Guarico, and Bolívar State) (Cordero &
Vogelsang ,1940).
Oriximinatrema
Knoff, Brooks, Mullins &
Gomes, 2012
Oriximinatrema noronhae
Knoff, Brooks,
Mullins & Gomes, 2012
Podocnemis expansa
(Schweigger, 1812), stomach
and intestine, BRAZIL (Pará State) (Knoff
et al
.,
2012).
Pseudocleptodiscus
Caballero, 1961
Pseudocleptodiscus margaritae
Caballero, 1961
Rhinoclemmys nasuta
(Boulenger, 1902), small
and large intestines, ECUADOR (Esmeraldas
Province) (Dyer & Carr, 1990b).
Pseudonematophila
Lenis & Vélez, 2011
Pseudonematophila ovalis
(Cordero &
Vogelsang, 1940)
Podocnemis
sp., stomach, VENEZUELA (Guárico
State) (Cordero & Vogelsang, 1940).
Podocnemis lewyana
Duméril, 1852, stomach,
COLOMBIA (Bolívar and Magdalena
Departaments) (Lenis & Vélez, 2011).
Superfamily Allocreadioidea Looss, 1902
Family Opecoleidae Ozaki, 1925
Helicotrema
Odhner, 1902
Helicotrema spirale
(Diesing, 1850)
Chelonoidis denticulatus
(Linnaeus, 1766), small
intestine, PERU (Iquitos Departament) (Tantaleán
& Forlong, 2013).
Chelonoidis denticulatus
(Linnaeus, 1766),
intestine, BRAZIL (unspecified locality) (Odhner,
1912 cited by Fernandes & Kohn, 2014).
BRAZIL (unspecified locality) (Travassos, 1934b;
Travassos
et al.
,
1969).
Mesoclemmys nasuta
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality)
(Travassos, 1934b; Travassos
et al.
,
1969).
Peltocephalus dumerilianus
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality)
(Travassos, 1934b; Travassos
et al.
,
1969).
Peltocephalus dumerilianus
(Schweigger, 1812),
stomach, small and large intestine, BRAZIL
() (Ferreira, 2016).
Amazonas State
Phrynops geoffroanus
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality)
(Travassos, 1934b; Travassos
et al.
,
1969).
Phrynops geoffroanus
(Schweigger, 1812), large
intestine, BRAZIL (São Paulo State) (Silva, 2014).
Phrynops hilarii
(Duméril & Bibron, 1835),
intestine, ARGENTINA (Corrientes Province)
(Lombardero & Moriena, 1977).
Podocnemis expansa
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality)
(Travassos
et al.,
1969).
Podocnemis expansa
(Schweigger, 1812),
stomach, PERU (Loreto Departament) (Tantaleán
et al.,
2011).
Podocnemis expansa
(Schweigger, 1812),
stomach, VENEZUELA (Zulia State) (Díaz-
Ungría, 1978).
Podocnemis erythrocephala
(Spix, 1824),
unspecified site of infection, BRAZIL (unspecified
locality) (Diesing, 1850 cited by Fernandes &
Kohn, 2014).
Podocnemis unifilis
Troschel, 1848, stomach, and
intestine, PERU (Loreto, and Madre de Dios
Departaments) (Salizar & Sanchez, 2004).
Podocnemis unifilis
Troschel, 1848, stomach, and
large intestine, PERU (Loreto Departament)
(Sánchez
et al.
, 2006).
Podocnemis unifilis
Troschel, 1848, stomach,
PERU (Loreto Departament) (Tantaleán
et al.
,
2011).
Podocnemis unifilis
Troschel, 1848, unspecified
site of infection, VENEZUELA (unspecified
locality) (Heyneman
et al.
, 1960 cited by
Fernandes & Kohn, 2014).
Podocnemis vogli
Müller, 1935, unspecified site of
infection, VENEZUELA (unspecified locality)
(Heyneman
et al.
, 1960 cited by Fernandes &
Kohn, 2014).
Podocnemis
sp., unspecified site of infection,
VENEZUELA (unspecified locality) (Caballero &
Diaz-Ungria, 1958 cited by Fernandes & Kohn,
Neotropical Helminthology, 2021, 15(1), ene-jun
Helminths associated with continental Testudines
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110
Podocnemis expansa
(Schweigger, 1812), small
intestine, BRAZIL (Pará State) (Lent & Freitas,
1938).
Superfamily Plagiorchioidea Lühe, 1901
Family Telorchiidae Looss, 1899
Loefgrenia
Travassos, 1920
Loefgrenia loefgreni
Travassos, 1919
Podocnemis unifilis
Troschel, 1848, intestine,
BRAZIL (unspecified locality) (Travassos, 1919
cited by Fernandes & Kohn, 2014).
Pseudotelorchis
Yamaguti, 1971
Pseudotelorchis devincenzii
(Mañé-Garzón &
Gil, 1961)
Hydromedusa tectifera
Cope, 1870, small
intestine, URUGUAY (Canelones Departament)
(Mañé-Garzón & Gil, 1961b).
Telorchis
Lühe, 1899
Telorchis achavali
Mañé-Garzón & Holcman-
Spector, 1973
Trachemys dorbigni
(Duméril & Bibron, 1835),
small intestine, URUGUAY (Tacuarembó
Departament) (Mañé-Garzón & Holcman-Spector,
1973).
Trachemys dorbigni
(Duméril & Bibron, 1835),
small intestine, BRAZIL (Rio Grande do Sul State)
(Mascarenhas & Müller, 2013).
Telorchis aculeatus
(Linstow, 1879)
Podocnemis unifilis
Troschel, 1848, intestine,
VENEZUELA (Sucre State) (Nasir, 1974).
Telorchis bifurcus
(Braun, 1899)
Podocnemis expansa
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Braun,
1901 cited by Fernandes & Kohn, 2014).
Telorchis birabeni
Mañé-Garzón & Gil, 1961
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, URUGUAY (Tacuarembó Departament)
(Mañé-Garzón & Gil, 1961c).
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, ARGENTINA (Corrientes Province)
(Lombardero & Moriena, 1977).
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, BRAZIL (Rio Grande do Sul State)
(Mascarenhas
et al.
, 2016).
Phrynops geoffroanus
(Schweigger, 1812), small
intestine, BRAZIL (São Paulo State) (Silva, 2014).
Telorchis corti
Stunkard, 1915
Peltocephalus dumerilianus
(Schweigger, 1812),
intestine, BRAZIL (unspecified locality) (Odhner,
1912 cited by Fernandes & Kohn 2014).
Peltocephalus dumerilianus
(Schweigger, 1812),
small intestine, BRAZIL ()
Amazonas State
(Ferreira, 2016).
Superfamily Opisthorchioidea Looss, 1899
Family Cryptogonimidae Ward,1917
Acanthostomum
Looss, 1899
Acanthostomum scyphocephalus
(Braun, 1899)
Chelus fimbriata
(Schneider, 1783), intestine,
BRAZIL (unspecified locality) (Ostrowski de
Núñez, 1986).
Caimanicola
Freitas & Lent, 1938
Caimanicola
brauni
(Mañé-Garzón & Gil, 1961)
Acanthochelys spixii
(Duméril & Bibron, 1835),
small intestine, BRAZIL (Rio Grande do Sul State)
(Chaviel
et al.,
2020).
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, URUGUAY (Tacuarembó Departament)
(Mañé-Garzón & Gil, 1961a).
Phrynops hilarii
(Duméril & Bibron, 1835),
ARGENTINA (Ostrowski de Núñez 1987
[experimental study] cited by Fernandes & Kohn,
2014).
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, BRAZIL (Rio Grande do Sul State)
(Chaviel
et al.,
2020).
Timoniella
Rebecq, 1960
Timoniella ostrowski
Brooks & Holcman, 1993
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, URUGUAY (Tacuarembó Departament)
(Mañé-Garzón & Gil, 1961a; Brooks & Holcman,
1993).
Phrynops hilarii
(Duméril & Bibron, 1835), small
intestine, BRAZIL (Rio Grande do Sul State)
(Chaviel
et al.,
2020).
Superfamily Gorgoderoidea Looss, 1899
Family Gorgoderidae Looss, 1899
Gorgoderina
Looss, 1902
Gorgoderina
sp.
Phrynops geoffroanus
(Schweigger, 1812), small
intestine, BRAZIL (Ceará State) (Pereira
et al.,
2018).
Family Braunotrematidae Yamaguti, 1958
Braunotrema
Price, 1930
Braunotrema pulvinatum
(Braun, 1899)
Neotropical Helminthology, 2021, 15(1), ene-jun
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111
intestine, URUGUAY (Tacuarembó Departament)
(Mañé-Garzón & Gil, 1961b).
Telorchis rapidulus
Dobbin Jr., 1957
Kinosternon scorpioides
(Linnaeus, 1766), small
intestine, BRAZIL (Pernambuco State) (Dobbin
Jr., 1957).
Telorchis
sp.
Hydromedusa tectifera
Cope, 1870, small
intestine, BRAZIL (Rio Grande do Sul State)
(present study, CHLAPASIL 893-896).
Telorchis
sp.
Trachemys dorbigni
(Duméril & Bibron, 1835),
small intestine, BRAZIL (Rio Grande do Sul State)
(Mascarenhas, 2014).
Class Cestoda Van Beneden, 1849
Order Proteocephalidea Mola, 1928
Family Proteocephalidae La Rue, 1911
Ophiotaenia
La Rue, 1911
Ophiotaenia cohospes
Cordero, 1946
Hydromedusa tectifera
Cope, 1870, intestine,
URUGUAY (Montevideo) (Cordero 1946).
Ophiotaenia lopesi
Rego, 1967
Chelonoidis denticulatus
(Linnaeus, 1766),
small
intestine, BRAZIL (Pará State) (Schmidt, 1986;
Muniz-Pereira
et al.,
2009).
Ophiotaenia
sp.
Hydromedusa tectifera
Cope, 1870, small
intestine, BRAZIL (Rio Grande do Sul State)
(present study, CHLAPASIL, 890-892).
Class Monogenoidea Bychowsky, 1937
Order Polystomatidea Lebedev, 1988
Family Polystomatidae Gamble, 1896
Polystomoides
Ward, 1917
Polystomoides brasiliensis
Vieira, Noveli, Sousa
& Souza-Lima, 2008
Hydromedusa maximiliani
(Mikan, 1820), buccal
and pharyngeal cavities, BRAZIL (Minas Gerais
State) (Vieira
et al.
,
2008).
Mesoclemmys turbeculata
(Lüderwaldt, 1926),
buccal and pharyngeal cavities, BRAZIL (Sergipe
State) (Santana
et al.
,
2019).
Phrynops geoffroanus
(Schweigger, 1812), buccal
and pharyngeal cavities, BRAZIL (Minas Gerais
State) (Vieira
et al.
,
2008).
Phrynops geoffroanus
(Schweigger, 1812), mouth
Trachemys callirostris
(Gray, 1856), small
intestine, COLOMBIA (unspecified locality)
(Lenis & Vélez, 2011).
Trachemys dorbigni
(Duméril & Bibron, 1835),
small intestine, BRAZIL (Rio Grande do Sul State)
(Mascarenhas & Müller, 2013).
Telorchis diaphanus
Freitas & Dobbin Jr., 1959
Kinosternon scorpioides
(Linnaeus, 1766), small
intestine, BRAZIL (Pernambuco State) (Freitas &
Dobbin Jr., 1959).
Telorchis dubius
Mañé-Garzón & Holcman-
Spector, 1968
Trachemys dorbigni
(Duméril & Bibron, 1835),
small intestine, URUGUAY (Flores Departament)
(Mañé-Garzón & Holcman-Spector, 1968a).
Telorchis hagmanni
Lent & Freitas, 1937
Peltocephalus dumerilianus
(Schweigger, 1812),
small intestine, BRAZIL ()
Amazonas State
(Ferreira, 2016).
Podocnemis expansa
(Schweigger, 1812),
stomach, and small intestine, BRAZIL (Pará State)
(Lent & Freitas, 1937; Alho, 1965).
Podocnemis lewyana
Duméril, 1852, small
intestine, COLOMBIA (Bolívar and Magdalena
Departaments) (Lenis & Vélez, 2011).
Podocnemis unifilis
Troschel, 1848, digestive
tract, PERU (Iquitos Departament) (Tantaleán
et
al.
, 2011).
Telorchis parvus
Braun, 1901
Emys orbicularis
(Linnaeus, 1758), intestine,
BRAZIL (unspecified locality) (Braun, 1901 cited
by Fernandes & Kohn, 2014).
Telorchis platensis
Mañé-Garzón & Gil, 1961
Acanthochelys spixii
(Duméril & Bibron, 1835),
large intestine, BRAZIL (Rio Grande do Sul State)
(Mascarenhas
et al.
, 2016).
Hydromedusa tectifera
Cope, 1870, small
intestine, URUGUAY (Canelones Departament)
(Mañé-Garzón & Gil, 1961b).
Telorchis pleroticus
(Braun, 1899)
Freshwater turtle, intestine, BRAZIL (unspecified
locality) (Braun, 1901 cited by Fernandes & Kohn,
2014).
Telorchis productus
Mañé-Garzón & Gil, 1961
Phrynops hilarii
(Duméril & Bibron, 1835), small
Neotropical Helminthology, 2021, 15(1), ene-jun
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112
State) (Mascarenhas, 2014).
Neopolystoma
sp.
Mesoclemmys vanderhaegei
(Bour, 1973),
unspecified site of infection, BRAZIL (Mato
Grosso State) (Ávila
et al.
, 2010).
Superfamily Iagotrematoidea Mañé-Garzón &
Gil, 1962
Family Iagotrematidae Mañé-Garzón & Gil,
1962
Iagotrema
Mañé-Garzón & Gil, 1962
Iagotrema uruguayensis
Mañé-Garzón & Gil,
1962
Hydromedusa tectifera
Cope, 1870, urinary
bladder, URUGUAY (Tacuarembó Department)
(Mañé-Garzón & Gil, 1962b).
Infraclass Rhabdocoela Ehrenberg, 1831
Order Dalytyphloplanida Williems
et al.
, 2006
Infraorder Temnocephalida Blanchard, 1849
Superfamily Temnocephaloidea Baer, 1953
Family Temnocephalidae Monticelli, 1899
Temnocephala
Blanchard, 1849
Temnocephala brevicornis
Monticelli 1889
Acanthochelys radiolata
(Mikan, 1820), BRAZIL
(unspecified locality) (Monticelli 1889 cited by
Martínez-Aquino
et al.
, 2014).
Acanthochelys spixii
(Duméril & Bibron, 1835),
skin of the neck, axillary, inguinal region, BRAZIL
(Rio Grande do Sul State) (Yuki
et al.
, 1993).
Hydromedusa tectifera
Cope, 1870, plastron and
the skin of the axillary, inguinal and anal regions,
BRAZIL (São Paulo State) (Pereira & Cuocolo,
1940).
Hydromedusa tectifera
Cope, 1870, URUGUAY
(Tacuarembó Departament) (Dioni, 1967 cited by
Martínez-Aquino
et al
., 2014).
Hydromedusa tectifera
Cope, 1870, URUGUAY
(Montevideo) (Cordero, 1946).
Hydromedusa tectifera
Cope, 1870, surface,
ARGENTINA (Buenos Aires Province) (Brusa &
Damborenea, 2000; Martínez-Aquino
et al
.,
2014).
Hydromedusa maximiliani
(Mikan, 1820), plastron
and the skin of the axillary, inguinal and anal
regions, BRAZIL (Rio de Janeiro) (Pereira &
Cuocolo, 1940).
Hydromedusa maximiliani
(Mikan, 1820), plastron
and the skin of the axillary, inguinal and anal
regions, BRAZIL (Minas Gerais State) (Pereira &
Cuocolo, 1940).
Hydromedusa maximiliani
(Mikan, 1820),
and esophagus, BRAZIL (São Paulo State) (Silva,
2014).
Phrynops geoffroanus
(Schweigger, 1812), buccal
and pharyngeal cavities, BRAZIL (Sergipe State)
(Santana
et al.,
2019).
Polystomoides fuquesi
Mañé-Garzón & Gil,
1962
Phrynops hilarii
(Duméril & Bibron, 1835), oral
cavity, URUGUAY (Artigas Departament) (Mañé-
Garzón & Gil, 1962a).
Polystomoides magdalenensis
Lenis & García-
Prieto, 2009
Trachemys callirostris
(Gray, 1856), oral cavity,
COLOMBIA (Bolivar Departament) (Lenis &
García-Prieto, 2009).
Polystomoides rohdei
Mañé-Garzón &
Holcman-Spector, 1968
Trachemys dorbigni
(Duméril & Bibron, 1835),
oral cavity, URUGUAY (Florida, Soriano, and
Rivera Departaments) (Mañé-Garzón & Holcman-
Spector, 1968b).
Trachemys dorbigni
(Duméril & Bibron, 1835),
oral cavity, BRAZIL (Rio Grande do Sul State),
(Mascarenhas, 2014).
Polystomoides uruguayensis
Mañé-Garzón &
Gil, 1961
Phrynops hilarii
(Duméril & Bibron, 1835), oral
cavity, URUGUAY (Artigas Departament) (Mañé-
Garzón & Gil, 1961d).
Polystomoides
sp.
Acanthochelys spixii
(Duméril & Bibron, 1835),
oral cavity, BRAZIL (Rio Grande do Sul State)
(Chaviel
et al
., 2020).
Polystomoides
sp.
Phrynops geoffroanus
(Schweigger, 1812),
urinary
bladder
, BRAZIL (São Paulo State) (Silva, 2014).
Polystomoides
sp.
Phrynops hilarii
(Duméril & Bibron, 1835), oral
cavity, BRAZIL (Rio Grande do Sul State) (present
study, CHLAPASIL 902).
Neopolystoma
Price, 1939
Neopolystoma
sp.
Trachemys dorbigni
(Duméril & Bibron, 1835),
urinary bladder
, BRAZIL (Rio Grande do Sul
Neotropical Helminthology, 2021, 15(1), ene-jun
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image/svg+xml
113
of the carapace and epidermis of the bases of neck
and limbs
,
BRAZIL (Rio Grande do Sul State)
(present study, CHLAPASIL 897-901).
Phylum Acanthocephala Rudolphi, 1808
Class Palaeacanthocephala Meyer, 1931
Order Polymorphida Petrochenko, 1956
Family Polymorphidae Meyer, 1931
Polymorphidae gen. sp.
(immature)
Acanthochelys spixii
(Duméril & Bibron, 1835),
small intestine, BRAZIL (Rio Grande do Sul State)
(present study, CHLAPASIL 888).
HOST-HELMINTH LIST
Cryptodira
Kinosternoidea
Kinosternidae
Kinosternon integrum
(IUCN:
Least concern)
Serpinema maghati
Kinosternon leucostomum
Atractis caballeroi
Kinosternon scorpioides
Camallanus kachugae
Nematophila grandis
Serpinema maghati
Serpinema monospiculatus
Spiroxys figueiredoi
Telorchis diaphanus
Telorchis rapidulus
Testudinoidea
Emydidae
Emys orbicularis
(IUCN: near threatened)
Telorchis parvus
Trachemys callirostris
Heronimus mollis
Nematophila argentinum
Polystomoides magdalenensis
Telorchis corti
Trachemys dorbigni
Camallanus
emydidius
Camallanus
sp.
Cheloniodiplostomum
sp.
Dioctophyme renale
(larvae)
Eustrongylides
sp. (larvae)
Falcaustra affinis
Neopolystoma
sp.
Polystomoides rohdei
epidermis of the neck, axillary, inguinal and anal
region and plastron, BRAZIL (Minas Gerais State)
(Novelli
et al.
, 2009).
Mesoclemmys gibba
(Schweigger, 1812), BRAZIL
(Monticelli, 1899 cited by Pereira & Cuocolo,
1940).
Phrynops hilarii
(Duméril & Bibron, 1835),
ARGENTINA (Buenos Aires Province) (Martínez-
Aquino
et al.
, 2014).
Trachemys dorbigni
(Duméril & Bibron, 1835),
skin of the neck, axillary, inguinal and anal areas,
and plastron, BRAZIL (Rio Grande do Sul State)
(Yuki
et al.
, 1993).
Temnocephala cuocoloi
Volonterio, 2010
Hydromedusa tectifera
Cope, 1870, surface of the
plastron, URUGUAY (Canelones Departament)
(Volonterio, 2010).
Temnocephala pereirai
Volonterio, 2010
Hydromedusa tectifera
Cope, 1870, surface of the
plastron, URUGUAY (Canelones Departament)
(Volonterio, 2010).
Trachemys dorbigni
(Duméril & Bibron, 1835),
skin of the neck, axillary, inguinal and perianal
areas, and in the surface of plastron, BRAZIL (Rio
Grande do Sul State) (Seixas
et al.
, 2014;
Mascarenhas
et al.
, 2018).
Temnocephala
sp.
Trachemys dorbigni
(Duméril & Bibron, 1835),
skin of the neck, axillary, inguinal and perianal
areas, and plastron, BRAZIL (Rio Grande do Sul
State) (Mascarenhas
et al.
, 2018).
Temnocephala
sp.
Hydromedusa tectifera
Cope, 1870, body, limbs
and hull,
BRAZIL (Rio Grande do Sul State)
(Soares
et al.
, 2007).
Temnocephala
sp.
Hydromedusa tectifera
Cope, 1870, dorsal surface
of the plastron, ventral surface of the carapace and
bridges, epidermis adjacent to the ventral surface
of the carapace and epidermis of the bases of neck
and limbs,
BRAZIL (Rio Grande do Sul State)
(Huckembeck & Quintela, 2013).
Temnocephala
sp.
Hydromedusa tectifera
Cope, 1870, surface of the
plastron, ventral surface of the carapace and
bridges, epidermis adjacent to the ventral surface
Neotropical Helminthology, 2021, 15(1), ene-jun
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114
Klossinemella travassosi
Labiduris gulosa
Labiduris irineuta
Labiduris zschokkei
Ophiotaenia lopesi
Sauricola sauricola
Chelonoidis porteri
(IUCN: Critically
endangered)
Atractis marquezi
Atractis
sp.
Labiduris
sp.
Kinixys erosa
(IUCN: Data deficient)
Nematophila grandis
Testudo
sp.
Cheloniodiplostomum testudinis
Pleurodira
Chelidae
Acanthochelys radiolata
(IUCN: Near threatened)
Temnocephala brevicornis
Acanthochelys spixii
(IUCN: Near threatened)
Caimanicola
brauni
Camallanus
sp.
Cheloniodiplostomum
sp.
Contracaecum
sp. (larvae)
Dioctophyme renale
(larvae)
Pharyngodonidae gen. sp.
Physaloptera
sp. (larvae)
Polymorphidae gen. sp. (immature)
Polystomoides
sp.
Spiroxys contortus
Telorchis
platensis
Temnocephala brevicornis
Chelus fimbriata
Acanthostomum scyphocephalus
Atamatam amazoniensis
Nematophila grandis
Paratamatam iquitosiensis
Mesoclemmys gibba
Nematophila grandis
Temnocephala brevicornis
Mesoclemmys nasuta
Atractis dactyluris
Labiduris gulosa
Nematophila grandis
Spiroxys
contortus
Spiroxys
sp.
Telorchis achavali
Telorchis corti
Telorchis dubius
Telorchis
sp.
Temnocephala brevicornis
Temnocephala pereirai
Temnocephala
sp.
Geoemydidae
Rhinoclemmys annulata
(IUCN: near threatened)
Atractis caballeroi
Falcaustra tikasinghi
Rhinoclemmys diademata
Atractis impure
Rhinoclemmys melanosterna
Falcaustra tikasinghi
Rhinoclemmys nasuta
(IUCN: near threatened)
Falcaustra tikasinghi
Halltrema
heteroxenus
Hedruris
sp.
Nematophila grandis
Octangioides tlacotalpensis
Pseudocleptodiscus margaritae
Rhinoclemmys punctularia
Falcaustra tikasinghi
Nematophila grandis
Serpinema cayennensis
Testudinidae
Chelonoidis carbonarius
Atractis thapari
Atractis
sp.
Chapiniella variabilis
Thelandros
sp.
Chelonoidis chilensis
(IUCN: Vulnerable)
Falcaustra
sp.
Labiduris
sp.
Chelonoidis denticulatus
(IUCN: Vulnerable)
Angusticaecum brevispiculum
Atractis impure
Atractis thapari
Chapiniella variabilis
Halltrema avitellina
Helicotrema spirale
Neotropical Helminthology, 2021, 15(1), ene-jun
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image/svg+xml
115
Phrynops
sp.
Nematophila argentinum
Hydromedusa maximiliani
(IUCN: Vulnerable)
Polystomoides brasiliensis
Temnocephala brevicornis
Hydromedusa tectifera
Camallanus
emydidius
Cheloniodiplostomum testudinis
Cheloniodiplostomum
sp.
Dioctophyme renale
(larvae)
Gnathostoma
sp. (larvae)
Hedruris dratini
Hedruris orestiae
Iagotrema uruguayensis
Nematophila grandis
Ophiotaenia cohospes
Ophiotaenia
sp.
Pseudotelorchis devincenzii
Spirocamallanus
sp.
Spiroxys contortus
Telorchis platensis
Telorchis
sp.
Temnocephala brevicornis
Temnocephala cuocoloi
Temnocephala pereirai
Temnocephala
sp.
Hydraspis schopfii
Nematophila grandis
Podocnemididae
Peltocephalus dumerilianus
(IUCN: Vulnerable)
Ancyracanthus pinnatifidus
Atractis dactyluris
Atractis trematophila
Helicotrema spirale
Klossinemella conciliatus
Klossinemella
sp.
Nematophila grandis
Paratractis hystrix
Serpinema microcephalus
Telorchis hagmanni
Podocnemis erythrocephala
(IUCN: Vulnerable)
Nematophila grandis
Paratractis hystrix
Podocnemis expansa
(IUCN: Lower
Risk/conservation dependen)
Ancyracanthus pinnatifidus
Serpinema monospiculatus
Mesoclemmys turbeculata
Physaloptera retusa
Polystomoides brasiliensis
Serpinema monospiculatus
Spiroxys figueiredoi
Mesoclemmys vanderhaegei
(IUCN: Near
threatened)
Neopolystoma
sp.
Spiroxys
sp.
Phrynops geoffroanus
Brevimulticaecum
sp. (larvae)
Camallanus
sp.
Cheloniodiplostomum brevis
Cheloniodiplostomum testudinis
Cheloniodiplostomum
sp.
Gorgoderina
sp.
Nematophila grandis
Physaloptera
sp. (larvae)
Physaloptera retusa
Polystomoides brasiliensis
Polystomoides
sp.
Prionosomoides scalaris
Serpinema monospiculatus
Spiroxys figueiredoi
Telorchis birabeni
Phrynops hilarii
Caimanicola brauni
Camallanus
sp.
Cheloniodiplostomum argentinensis
Cheloniodiplostomum testudinis
Cheloniodiplostomum
sp.
Dioctophyme renale
(larvae)
Hedruris dratini
Hedruris orestiae
Nematophila grandis
Polystomoides fuquesi
Polystomoides uruguayensis
Polystomoides
sp.
Prionosomoides phrynopsis
Prionosomoides scalaris
Prionosomoides
sp.
Spiroxys
contortus
Spiroxys
sp.
Telorchis birabeni
Telorchis productus
Temnocephala brevicornis
Timoniella ostrowoski
Neotropical Helminthology, 2021, 15(1), ene-jun
Helminths associated with continental Testudines
image/svg+xml
116
Podocnemis
sp.
Halltrema avitellina
Halltrema
heteroxenus
Nematophila grandis
Nematophila venezuelenisis
Pseudonematophila ovalis
Freshwater turtle
Telorchis pleroticus
Amazon river turtle
Atractis trematophila
Atractis cruciate
Atractis dactyluris
Atractis
sp.
Braunotrema pulvinatum
Halltrema avitellina
Klossinemella conciliatus
Nematophila grandis
Oriximinatrema noronhae
Paratractis hystrix
Podocnematractis colombiaensis
Podocnematractis
ortleppi
Podocnemitrema papillosus
Rhytidodes gelatinosus
Serpinema amazonicus
Serpinema microcephalus
Telorchis bifurcus
Telorchis hagmanni
Podocnemis lewyana
(IUCN: Critically
Endangered)
Nematophila argentinum
Nematophila venezuelenisis
Neodeuterobaris pritchardae
Pseudonematophila ovalis
Telorchis hagmanni
Podocnemis unifilis
(IUCN: Vulnerable)
Ancyracanthus pinnatifidus
Buckleyatractis marinkelli
Halltrema avitellina
Halltrema heteroxenus
Klossinemella conciliatus
Loefgrenia loefgrenia
Nematophila grandis
Nematophila
sp.
Orientatractis leiperi
Paraorientatractis semiannulata
Paratractis hystrix
Pneumoatractis podocnemis
Podocnematractis
ortleppi
Podocnemitrema papillosus
Serpinema amazonicus
Telorchis aculeatus
Telorchis hagmanni
Podocnemis vogli
Nematophila grandis
Orientatractis leiperi
Paratractis hystrix
Podocnematractis colombiaensis
Podocnematractis
ortleppi
Cohen
et al.
(2013) carried out a list of
Monogenoidea associated with Testudines from
South America in which they cited eight records.
This checklist added nine records of
Polystomatidae. Fernandes & Kohn (2014)
introduced a checklist of Trematoda, in which they
cited 34
taxa
registered in the region. This checklist
added 15
taxa
to this group of parasites. Martínez-
Aquino
et al.
(2014) gathered the records of
Temnocephalida in the Neotropical region and
cited four
taxa
associated with freshwater turtles
from South America. This checklist added three
taxa
associated with these vertebrates and included
the record of
Temnocephala pereirai
in
Trachemys
dorbigni
in Brazil (Seixas
et al.
, 2014;
Mascarenhas
et al.
, 2018). In addition, this
checklist has gathered the records of Nematoda,
Cestoda and Acanthocephala parasites of
Testudines from South America. Thus, it should be
highlighted that these vertebrates host a little-
known rich fauna of helminths, that many
Testudines have been poorly studied and that
several studies have only been carried out with the
examination of their digestive tract (e.g. Sanchez
et
al.
, 2006; Tantaleán
et al.
, 2011; Tantaleán &
Forlong, 2013; Viana
et al.
, 2016). Besides the
stomach and intestines (main reported sites of
infection), parasitic helminths can be found in
blood vessels (Bullard
et al.
, 2019), lungs (Bursey
et al.,
2009; Lenis & Vélez, 2011), the urinary
bladder (Mañé-Garzón & Gil, 1962; Mascarenhas,
2014) and in other sites of infection that have not
been reported yet.
Concerning the conservation status of continental
Testudines from South America, many species are
DISCUSSION
Neotropical Helminthology, 2021, 15(1), ene-jun
Mascarenhas & Müller
image/svg+xml
117
inventories of wildlife associated helminths,
particularly in biomes and host species that have
not been sampled because the missing information
is even greater than the current state of knowledge,
as observed in the present review. In this context,
helminthological studies of Testudines generate
information that can be used in studies that aim at
the conservation of the host species and,
consequently, of their associated organisms, as
well as the sites that sustain these invaluable
relations.
The authors would like to thank Marco Antonio
Afonso Coimbra for his assistance. This study was
supported by CAPES (Coordenação de
Aperfeiçoamento do Pessoal de Nível Superior,
process number 32/2010).
threatened. According to Rhodin
et al.
(2017),
Testudines are one of the most threatened major
groups of vertebrates, in general more than birds,
mammals, cartilaginous or bony fishes and
amphibians. In this scenario, several helminths
associated with these hosts can either be potentially
impacted or even extinct without at least being
described or recorded. Less than half of native
South American species cited in the IUCN list have
had helminthological records.
Peltocephalus
dumerilianus
(VU),
Podocnemis unifilis
(VU) and
Chelonoides denticulatus
(VU) are the most
studied ones, while
Chelonoides chilensis
(VU),
Podocnemis erythrocephala
(VU),
Podocnemis
lewyana
(CR) and
Chelonoides porteri
(CR) are
the least studied ones; two records of helminths in
th
P. erythrocephala
date from the 19 century
(Diesing, 1850 cited by Fernandes & Kohn, 2014;
Diesing, 1851). Studies of
P. lewyana
,
C. chilensis
,
and
C. porteri
are more recent (Brooks, 1976;
Bursey & Flanagan, 2002; Lenis & Vélez, 2011;
Fournié
et al
., 2015; Castillo
et al
., 2020). As for
ectosymbionts, it should be highlighted that
Temnocephala
comprises freshwater species
endemic to the Neotropical region that have
specificity regarding the host group and/or species
(Martínez-Aquino
et al.
, 2014). Only seven out of
71 continental Testudines
taxa
from South America
were reported as hosts of
Temnocephala
; three
species were listed as near threatened (NT) by the
IUCN. Therefore, conservation actions related to
Testudines can contribute to the conservation of
several other organisms that depend on these
vertebrates.
Parasites are regulators of host populations and
powerful agents that maintain the stability of
ecosystems. Many species of helminths, for
example, have complex life cycles which involve
one or more intermediate hosts and must be
ingested by the final host to ensure transmission
and survival of the parasite organism. Thus,
occurrence of a heteroxenic helminth may reflect
prey-predator interactions and show the position of
hosts in the trophic chain (Brooks & Hoberg, 2001;
Marcogliese, 2004). Since parasites can provide a
lot of information about host organisms and their
habitats, they contribute to the knowledge of host
biology, environmental stress, trophic chains and
biodiversity (Poulin, 1999; Marcogliese, 2004;
Horwitz & Wilcox, 2005). Anjos (2011)
highlighted the importance of basic research and
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