ISSN Versión impresa 2218-6425 ISSN Versión Electrónica 1995-1043
Neotropical Helminthology, 2020, 14(2), jul-dic:129-141.
ORIGINAL ARTICLE / ARTÍCULO ORIGINAL
1Graduate Course of Ecology and Natural Resources, Department of Biology, Pici Campus,
Federal University of Ceará, Fortaleza - CE Zip Code 60440-900, Brazil.
2Graduate Course of Biology, Department of Biology, Regional University of Cariri, Crato - CE
Zip Code 63105-000, Brazil.
*Corresponding author: kassio.ufpi@gmail.com
1,* 2 1
Kássio de Castro Araújo ; Charles de Sousa Silva ; Heitor Tavares de Sousa Machado ,
1 1
Cícero Ricardo de Oliveira & Robson Waldemar Ávila
ABSTRACT
Keywords: Checklist – Delta do Parnaíba – Neotropical – Parasites – Snakes
Endoparasites checklists might contribute to biodiversity understanding. Endoparasites infection might
be associated with the host evolutionary history or vary according to host diet and habitat. It was important
to investigate these infections in distinct environments since environmental conditions and resource
availability might influence the helminth composition and structure in the snake assemblages. We aimed
to investigate the endoparasites associated with Philodryas olfersii (Lichtenstein, 1823), and we present a
bibliographic revision of the endoparasites associated with the genus Philodryas. We surveyed the
endoparasites of the snake P. olfersii in the Parnaíba River Delta, coastal zone of Piauí state, northeastern
Brazil. We analyzed the presence of ectoparasites under the epidermis and oral cavity and endoparasites of
the digestive tract, lungs, heart, liver, and kidneys, and we used the parasitological descriptors:
prevalence, abundance, mean abundance, and mean infection intensity to describe the endoparasite
community. We recorded 312 endoparasite specimens belonging to nine taxa: Kalicephalus costatus
Rudolphi, 1819, Strongyloides ophidiae Pereira 1929, Physaloptera sp., Physalopteroides venancioi
Lent, Freitas & Proença, 1946, Cosmocercidae larvae, Raillietiella furcocerca (Diesing, 1835),
Ophiotaenia sp., cystacanths, and a digenetic trematode not identified. Nematodes and Pentastomids were
the most abundant taxa infecting P. olfersii. Our results contribute to the increasing knowledge base
concerning the endoparasites of P. olfersii. This is the first research about parasitism in reptiles from the
Parnaíba River Delta. We encourage additional parasitic studies in order to improve the understanding of
the host-parasite relationships in Brazilian snakes, mainly in coastal environments.
Neotropical Helminthology
129
doi:10.24039/rnh2020142720
ÓrganooficialdelaAsociaciónPeruanadeHelmintologíaeInvertebradosAfines(APHIA)
Lima-Perú
VersiónImpresa:ISSN2218-6425VersiónElectrónica:ISSN1995-1043
Volume14,Number2(jul-dec)2020
ENDOPARASITES OF PHILODRYAS OLFERSII (LICHTENSTEIN, 1823) IN RESTINGA
ENVIRONMENTS OF THE PARNAÍBA RIVER DELTA, NORTHEASTERN BRAZIL
ENDOPARÁSITOS DE PHILODRYAS OLFERSII (LICHTENSTEIN, 1823) EN AMBIENTES DE
RESTINGA DEL DELTA DEL RÍO PARNAIBA, NORDESTE DE BRASIL
INTRODUCTION
130
RESUMEN
Palabras clave: Inventario – Delta del Parnaíba – Neotropical – Parásitos – Serpientes
Lista de endoparásitos son fundamentalmente importantes porqué contribuyen para la comprensión de la
biodiversidad. Las infecciones endoparásitas pueden estar asociadas a la historia evolutiva del huésped
del parásito o variar según la dieta y el hábitat; así, es importante investigar tales infecciones en distintos
ambientes, a la vez que las condiciones ambientales y disponibilidad de recursos pueden influir en la
composición y estructura de los helmintos en ensambles de serpientes. Aquí, investigamos los
endoparásitos asociados con Philodryas olfersii (Lichtenstein, 1823), y presentamos la revisión
bibliográfica de los endoparásitos asociados con el género Philodryas. Buscamos los endoparásitos
relaccionados con la serpiente P. olfersii en el Delta del Parnaíba, región costera del estado de Piauí,
noreste de Brasil. Hemos analizado la presencia de ectoparásitos en la epidermis y la cavidad oral, y
endoparásitos en los siguientes órganos: tracto digestivo, pulmones, corazón, hígado y riñones, utilizando
los descriptores parasitológicos: prevalencia, abundancia, abundancia promedio e intensidad promedio
de infección, para describir la comunidad de endoparásitos. Registramos 312 especímenes de
endoparásitos pertenecientes a nueve tasas: Kalicephalus costatus Rudolphi, 1819, Strongyloides
ophidiae Pereira 1929, Physaloptera sp., Physalopteroides venancioi Lent, Freitas & Proença, 1946,
larva de Cosmocercidae, Raillietiella furcocerca (Diesing, 1835), Ophiotaenia sp., cistacantos y un
trematodo digenea no identificado. Nematodos y pentastómidos fueron los taxones más abundantes
infectando P. olfersii. Nuestros resultados contribuyen a la expansión del conocimiento acerca de los
endoparásitos de P. olfersii, con destaque para la naturaleza pionera del presente estudio, el primero en
investigar el parasitismo en reptiles del Delta del Parnaíba; por consiguiente, alentamos más estudios
parasitológicos con vista a mejorar la comprensión de la relación parásito-huésped en serpientes
brasileñas, sobre todo en ambientes costeros.
Neotropical Helminthology, 2020, 14(2), jul-dic
Endoparasites associated with snakes are
commonly acquired through their diet or direct
infection (Aho, 1990) and these endoparasites
might be associated with the host evolutionary
history or vary according to the host diet and
habitat (Fontenot & Font, 1996; Jiménez-Ruiz et
al., 2002). Therefore, it is important to investigate
the endoparasites infection in distinct
environments, since environmental conditions and
resource availability might influence the helminth
composition and structure in snake assemblages.
The restinga environments are coastal sandy
ecosystems with floristical and physiognomical
distinct communities (Falkenberg, 1999);
however, the endoparasite loads associated with
reptiles in these environments is still undersampled
(Van Sluys et al., 2000; Menezes et al., 2004; Dias
et al., 2005; Almeida et al., 2009; Viana et al.,
2013).
Brazil harbors a rich snake fauna (405 species),
being the Family Dipsadidae the most diverse
Endoparasites checklists are fundamentally
important because they might contribute to
biodiversity understanding and also are considered
environmental quality bioindicators (Galli et al.,
2001; Poulin & Morand, 2004; Brandão, 2017).
Furthermore, due to the considerable number of
dead snakes by parasitic infection (Klingenberg,
1993; Sánchez et al., 2004; Chinnadurai et al.,
2008; Vimalaraj et al., 2015) and the role of
endoparasites on distinct levels of environmental
organizations (Wood & Johnson, 2015), it is
essential to study the endoparasites associated with
Brazilian snakes. Nematodes, digenetic and
cestodes are the main causes of these infections
(Benson, 1999; Wilson & Carpenter, 1996; Silva,
2000). These endoparasites produce negative
effects on the host as anemia, anorexia, survival
and fecundity reduction (Vitt & Caldwell, 2009;
Matias et al., 2018).
Castro Araujo et al.
131
MATERIAL AND METHODS
RESULTS
Endoparasites of Philodryas olfersii
Neotropical Helminthology, 2020, 14(2), jul-dic
(Costa & Bérnils, 2018). Despite this diversity,
endoparasites associated with Brazilian snakes
have been neglected, with an increase in number of
studies recently (e.g. Kuzmin et al., 2014; Mati et
al., 2015; Mendoza-Roldan & Fiorillo, 2016;
Borges-Nojosa et al., 2017; Carvalho et al., 2018;
Emmerich et al., 2018; Matias et al., 2018). Among
Dipsadidae, the genus Philodryas Wagler, 1830 is
widely distributed and diverse in Brazil, with 13
species recorded (Costa & Bérnils, 2018).
However, endoparasites have been reported to only
five species: Philodryas aestiva Rossellini, 2007,
P. nattereri Tavares et al. 2017, P. patagoniensis
Sprent, 1978, P. psammophidea Lunaschi &
Drago, 2007 and P. olfersii Travassos, 1917.
Furthermore, there are endoparasites records for P.
baroni Berg, 1895 in North America (Hartdegen &
Gamble, 2002) and P. chamissonis Wiegmann,
1835 in Chile (Travassos, 1917; Fredes & Raffo,
2005).
For Philodryas olfersii a semi-arboreal snake with
diurnal habits preying on small vertebrates
(Hartmann & Marques, 2005), there are eight
parasites recorded. Herein, we aimed to investigate
the endoparasites associated with P. olfersii from a
restinga environment in the Parnaíba River Delta,
Northeastern Brazil, and we present a
bibliographic revision of endoparasites associated
with the genus Philodryas.
Study area
Fieldwork took place in the Environmental
Protection Area (EPA) Delta do Parnaíba, Ilha
Grande municipality, Piauí State, Northeastern
Brazil (2°50'9.62"S and 41°48'46.20"W, 5 masl).
The study area presents quaternary habitats
characterized by sandy soils with high salt
concentrations covered predominantly by
herbaceous and shrubby xerophytic vegetation
(Santos-Filho et al., 2010).
Sampling
Five specimens of P. olfersii were collected
manually at the end of rain season (July/2017). The
specimens were euthanized following ethical
procedures of Federal Council of Veterinary
Medicine - CFMV (2013) after approval by the
Ethics Committee of Universidade Regional do
Cariri (CEUA/URCA, process #00260/2016.1).
We analyzed the presence of ectoparasites under
the epidermis and oral cavity and endoparasites in
the following organs: digestive tract, lungs, heart,
liver and kidneys, according to Amato et al. (1991).
The fixation techniques and parasite preservations
were conducted according to each taxonomic
group (Vidal-Martínez et al., 2001). Nematodes
were clarified with lactic acid (Andrade, 2000),
Trematoda and Cestoda were dehydrated in an
increasing series of alcohol, stained with the
hydrochloric carmine technique and diaphanized
in eugenol. Then, we mounted on temporary slides
and examined in light microscope. For species
identification, we used the following literature:
Yamaguti (1961), Sprent (1978), Vicente et al.
(1993), Smales (2007), Gibbons (2010), and Gamil
(2012). The P. olfersii specimens were deposited in
the Coleção Herpetológica of Universidade
Regional do Cariri (voucher specimens URCA-H
13474-13478) and the endoparasites specimens
were deposited in the Helminthological Collection
of the laboratory of Zoology of Universidade
Regional do Cariri, URCA, Ceará State, Brazil.
The parasitological descriptors prevalence,
abundance, mean abundance, and mean infection
intensity were measured according to Bush et al.
(1997), using the software Quantitative
Parasitology 3.0 (Rózsa et al., 2000).
All P. olfersii specimens analyzed (three females
and two males, both adults) were infected by at
least one parasite taxon. We recorded 312
endoparasite specimens belonging to nine taxa
from the following taxonomic groups:
Aca n thocep h ala, C e stoda, Nema t oda,
Pentastomida, and Trematoda (Table 1).
The highest endoparasite abundance of P olfersii
was recorded in stomach (n = 137), small intestine
(n = 72) and large intestine (n = 55). Adult
endoparasites were recorded at all these organs
132
analyzed, whereas cystacanths were recorded only
in stomach (n = 3). The most abundant taxa
infecting P. olfersii were Nematoda (n = 256),
followed by Pentastomida (Raillietiella
furcocerca, n = 48, Table 1).
The endoparasite species most abundant were
Physaloptera sp. (n = 170), Strongyloides ophidiae
(n = 54), R. furcocerca (n = 48), and
Cosmocercidae larvae (n = 28). These species also
had the highest values of prevalence, being S.
ophidiae recorded for all specimens studied. The
mean infection intensity in P. olfersii ranged from 1
to 42.5, in which Physaloptera sp. also was the
most infectious species, and the endoparasites
infection amplitude ranged 1 to 46 (Table 1).
Table 1. Endoparasites associated with Philodryas olfersii. Values of prevalence (P %), number of endoparasites
(NE), mean infection intensity and standard deviation (MII ± SD), mean abundance (MA), site of infection (SI) and
range (R). Sto= stomach; Lun=lung; Lgi= large intestine; Smi= small intestine; * new record for P. olfersii.
Endoparasites P% NE MII± DP
AM
SI R
Acanthocephala
Cystacanth 20
3 3 0.6
Sto 3
Cestoda
Ophiotaenia
sp.* 40
3 1.5 ± 0.7
0.6
Smi 1 – 2
Nematoda
Cosmocercidae larvae 80
28 7 ± 2.5 5.6
Smi/Lgi
3 – 8
Kalicephalus
costatus* 20
2 2 0.4
Smi 2
Strongyloides ophidiae
100
54 10.8
±
13
10.8
Sto/Smi/Lgi
1 – 21
Physaloptera sp.
80
170
42.5 ±
6
34
Sto/Smi/Lgi
2 – 46
Physalopteroides venancioi*
40
2
1±0
0.4
Smi
1
Pentastomida
Raillietiella furcocerca*
80
48
12
±
12.3
9.6
Lun
4 – 30
Trematoda
Unidentified digenetic 20 2 2 0.4 Lgi 2
genus is able to be a host for further endoparasite
diversity.
Regarding nematodes, we registered two
specimens of P. venancioi in the host small
intestine. The Physalopteroides genus is
distributed worldwide (Bursey & Goldberg, 1994;
Al-Moussawi, 2016); however, only P. venancioi is
registered in South America (Vrcibradic et al.,
2000; Ávila & Silva, 2010; Cabral et al., 2018).
Despite the P. venancioi life-cycle remains poorly
known (Bursey et al., 2005), this genus has
heteroxenic life-cycle, with an intermediated
invertebrate host and a vertebrate (amphibian,
lizard, bird or small rodent) as a final host
(Anderson, 2000; Al-Moussawi, 2016). Thus, as
DISCUSSION
Despite the low sample size of the present study, we
recorded a relative abundance and richness of
endoparasites, since at least 41 endoparasites
species are recorded for the genus Philodryas and
about 20 % of these were listed in the present study
(Table 2). We present the first record of the
following endoparasites associated with P. olfersii:
Ophiotaenia sp., Kalicephalus costatus,
Physalopteroides venancioi, and R. furcocerca. In
addition, we present the first record of P. venancioi
infecting the genus Philodryas. However,
considering its wide distribution in South
American (Uetz et al., 2020) we believe that this
Neotropical Helminthology, 2020, 14(2), jul-dic Castro Araujo et al.
133
Host Endoparasites References
Philodryas aestiva (Duméril, Bibron & Duméril, 1854)
Caryospora brasiliensis
Carini, 1932 Lainson &
Shaw
(1973)
Hastospiculum digiticaudum
Freitas, 1956 Rosselini (2007)
Infidum infidum
Faria, 1910 Noronha et al. (2009)
P. baroni
Berg, 1895
Cephalobaena tetrapoda
Heymons, 1922
Haffner &
Rack (1971)
Hexametra boddaertii
Baird, 1860
Hartdegen & Gamble (2002)
P. chamissonis
(Wiegmann, 1835)
Raillietiella sp.
Fredes &
Raffo (2005)
P. nattereri
Steindachner, 1870
Caryospora brasiliensis
Carini, 1932
Lainson &
Shaw (1973)
Cephalobaena tetrapoda
Almeida et al. (2008a)
Hepatozoon musa
Miller, 1908
Borges-Nojosa et al. (2017)
Oligacanthorrhynchus sp.
Araújo-Filho et al. (2018)
Physaloptera sp.
Oliveira et al. (2019)
Raillietiella furcocerca
Almeida et al. (2008a)
P. olfersii
(Linchtenstein, 1823)
Caryospora brasiliensis
Lainson & Shaw (1973)
Caryospora olfersii
Viana et al. 2013
Viana et al. (2013)
Catadiscus longicoecalis
Poumarau, 1965
Caubisens-Paumorau (1965)
Centrorhynchus
sp.
Silva &
Muller (2012)
Cosmocercidae
larvae
Present study
Cystacanth
Present study
Infidum similis
Travassos, 1916
Caubisens-Paumorau (1968)
Isospora decipiens
Lainson & Shaw 1973
Lainson & Shaw (1973)
Kalicephalus costatus
Present study
Opisthogonimus lecithonotus
Lühe, 1900
Silva & Muller (2012)
Ophiotaenia
sp.
Present study
Physaloptera
sp.
Oliveira et al. (2020)
Physalopteroides venancioi
Present study
Raillietiella furcocerca
Present study
Strongyloides ophidiae
Mati &
Melo (2014)
Westella serpentis
Artigas, Ruiz & Leão,
1943
Rosselini (2007)
Unidentified digenetic
Present study
P. patagoniensis
(Girard, 1858)
Caryospora brasiliensis
Lainson &
Shaw (1973)
Dioctophyma renale Goeze,1782 Mascarenhas et al. (2018)
Hepatozoon philodryasi Carini, 1910 Carini (1910)
Hexametra boddaerti Sprent (1978)
Table 2. Bibliographic revision of endoparasites associated with the genus Philodryas.
Endoparasites of Philodryas olfersii
Neotropical Helminthology, 2020, 14(2), jul-dic
Kalicephalus costatus
Vicente et al. (1993)
Kalicephalus inermis
Molin, 1861
Vicente et al. (1993)
Ophiotaenia sp.
Ammann &
Chambrier (2008)
Ophidascaris sicki
Freitas, 1951
Sprent (1988)
Opisthogonimus afranioi
Pereira, 1929
Rosselini (2007)
Opisthogonimus fonsecai
Ruiz & Leão, 1942
Rosselini (2007)
Opisthogonimus interrogativus
Nicoll, 1914
Rosselini (2007)
Opisthogonimus lecithonotus
Lunaschi &
Drago (2007)
Opisthogonimus pereirai
Ruiz & Leão, 1942
Rosselini (2007)
Ochetosoma heterocoelium
Travassos, 1921
Rosselini (2007)
Paradistomus parvissimum
Travassos,
1918
Noronha et al. (2009)
Renifer heterocoelium
Travassos, 1921
Pinto et al. (2012)
Rhabdias vellardi
Pereira, 1928
Rosselini (2007)
Styphlodora condita
Faria, 1911
Lunaschi &
Drago (2007)
Travtrema stenocotyle
Cohn, 1902
Lunaschi &
Drago (2007)
Westella philodryadum West, 1896 Rosselini (2007)
P. psammophidea Gunther, 1872
Infidum infidum Lunaschi & Drago (2007)
Opisthogonimus artigasi Ruiz & Leao, 1942 Lunaschi & Drago (2007)
Philodryas sp.
Catadiscus dolichocotyle Cohn, 1903 Lunaschi & Drago (2016)
Infidum similis Lunaschi & Drago (2016)
Travtrema stenocotyle Lunaschi & Drago (2016)
134
registered obstructing the snakes' intestinal tract
(Matos-Junior et al., 2004).
Strongyloides infection on snakes also occurs
through cutaneous penetration, ingestion of
infected preys and contaminated water (Mitchell,
2007; Mihalca et al., 2010; Taylor et al., 2010).
Despite not having a detailed biology (Santos et al.,
2010), it is known that this parasite presents direct
and indirect lifecycle, being the first one most
common (Mati & Melo, 2014). This helminth was
already recorded infecting birds, mammals,
amphibians, reptiles and occasionally humans
around the world (Little, 1966; Urquhart et al.,
1998). Nevertheless, only S. ophidiae has been
recorded for Neotropical snakes (e.g. Pereira,
1929; Santos et al., 2010; Mati & Melo, 2014).
Among the Brazilian Squamata, the helminth S.
ophidiae was registered infecting the snakes
Crotalus durissus, Erythrolamprus miliaris,
Oxyrhopus guibei Hoge & Romano, 1977, and also
supposed in previous studies with other snake hosts
(Al-Moussawi, 2016; Matias et al., 2018) and
considering that P. olfersii feed on small
vertebrates (Leite et al., 2009), it also may have
acquired this helminth through ingestion of
infected prey.
Likewise, we also registered only two K. costatus
specimens parasitizing one individual host. It is a
digestive tract helminth of snakes, being recorded
for different species such as Crotalus durissus
Linnaeus, 1758, Erythrolamprus miliaris
Linnaeus, 1758, Macrelaps microlepidotus
Gunther, 1860, and Xenodon sp. (Dias et al., 2004;
Ramallo, 2005; McAllister et al., 2010; Mati et al.,
2015; Tavares et al., 2017). Thus, this is the first
record of K. costatus infecting P. olfersii. This
nematode has direct life-cycle (Anderson, 2000),
adhering to the mucosa, feeding on blood
(Mitchell, 2007). This endoparasite might be
considered unhealthy because it was already
Neotropical Helminthology, 2020, 14(2), jul-dic Castro Araujo et al.
135
P. olfersii (see: Pereira, 1929; Santos et al., 2010;
Mati & Melo, 2014; Mati et al., 2015).
Nematodes belong to Physaloptera Rudolphi,
1819 genus are commonly registered infecting
snakes (Goldberg et al., 2004; Barbosa et al.,
2006), being recorded parasitizing distinct serpents
in Brazil, counting Philodryas species (e.g. Ávila
et al., 2013; Silva, 2014; Oliveira et al., 2019;
2020). Physaloptera spp. use invertebrates as
intermediate hosts, which are consumed by
amphibians and lizards (Duellman & Trueb, 1986;
Werneck et al., 2009), common preys of P. olfersii
(Hartmann & Marques, 2005). Infection by
Physaloptera spp. may cause hemorrhages and
excessive digestive secretions (Schell, 1952;
Widmer, 1970; Goldberg & Bursey, 1989; Araújo
et al., 1999).
About 80% of the hosts were infected by
Cosmocercidae larvae, however, these helminths
were not identified at species level due to their
immature stage, although are usually registered
parasitizing amphibian and reptile species (Avila et
al., 2010; Rizvi, 2009; Rizvi & Bursey, 2014;
Bursey et al., 2015). Likewise, the digenetics
recorded in the large intestine were not identified at
the species level due to their development stage.
Trematodes were frequently reported infecting
snakes (see. Travassos et al., 1969; Rossellini,
2007; Pinto et al., 2012; Fernandes & Kohn, 2014),
including Philodryas olfersii (Silva & Müller,
2012).
We also recorded cystacanths of Acanthocephala
on the snake stomach, which was extensively
reported for other snakes (Smales, 2007; Carvalho
et al., 2018; Matias et al., 2018). Acanthocephala
species belonging to the genus Centrorhynchus
Lühe, 1911 and Oligacanthorhynchus Travassos,
1915 were already registered parasitizing
individuals of P. olfersii and P. nattereri,
respectively (Silva & Muller, 2012; Araújo-Filho
et al., 2018). Silva & Muller (2012) considered P.
olfersii as a paratenic host for acanthocephalans,
reaching their adult stage mainly in birds (Baker,
2007).
The genus Ophiotaenia La Rue, 1911 has six
species which five were reported infecting snakes
(O. calmetti Barrois, 1898, O. elongate Fuhrmann,
1927, O. flava Rudin, 1917, O. hyaline Rudin, 1917
and O. macrobothria Rudin, 1917) and O. lopesi
Rego, 1967 was found infecting lizards (Schmidt,
1986; Silva et al., 2006). These endoparasites have
wide distribution, being reported for different
snake hosts (e.g. Chambrier et al., 2010;
McAllister et al., 2010), including records for
Brazilian snakes (e.g. Silva et al., 2006; Matias et
al., 2018). This species is a small intestine
endoparasite and it could difficult the nutrient
absorption by the host, obstructing their intestinal
tract (Tantaleán & Gozalo, 1985), and there are
important records of snake death due to possible
cestodes intensive infection (Murvanidze, 2013;
Sánchez et al., 2004; Vimalaraj et al., 2015).
Pentastomids are reported to different snakes
belong to genus Philodryas (e.g. Almeida et al.,
2008a; 2008b; Brito et al., 2012). The species R.
furcocerca infected about 80% of P. olfersii hosts in
present study. The higher infection levels reported
here might be considered unhealthy because
pentastomids are specific lung parasites, which
could cause serious lung injuries (Riley, 1986;
Almeida et al., 2007; 2008b).
The Environmental Protection Area Delta do
Parnaíba, northeastern Brazil harbors rich
herpetofauna (Loebmann & Mai, 2008; Andrade et
al., 2016, Araújo et al., 2018), however, there is a
lack of information about the reptile parasite load
in restinga environments, mainly in northeastern
region. Our results contribute to increase the
knowledge of endoparasites infecting P. olfersii,
and this is the first research about parasitism in
reptiles from the Parnaíba River Delta; thus, we
encourage further parasitological studies in order
to improve the understanding of the parasite-host
relationships in Brazilian snakes, mainly in coastal
environments.
We are grateful to the ICMBio for the collection
license (Permit: ICMBio 72384, process: 29613).
We are grateful to the Conselho Nacional de
Desenvolvimento Científico e Tecnológico-CNPq,
Coordenação de Aperfeiçoamento de Pessoal de
Nível Superior - Brasil (CAPES) and the FUNCAP
( F u n d a ç ã o C e a r e n s e d e A p o i o a o
Desenvolvimento Científico e Tecnológico) for
ACKNOWLEDGEMENTS
Endoparasites of Philodryas olfersii
Neotropical Helminthology, 2020, 14(2), jul-dic
partial financial support (PROTAX-Processes
4 4 0 5 1 1 / 2 0 1 5 - 1 ; 5 5 7 4 6 8 5 / 2 0 1 7 ;
88882.156872/2016-01). R.W.A thanks CNPq for
the research grant (PQ#303622/2015-6;
305988/2018-2).
136
Ammann, M & Chambrier, AD. 2008. Ophiotaenia
g i l b e r t i s p . n ( E u c e s t o d a :
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Received June 11, 2020.
Accepted July 27, 2020.
Endoparasites of Philodryas olfersii
Neotropical Helminthology, 2020, 14(2), jul-dic