65
Endoparasites of anurans in the seven cities national park
Neotropical Helminthology, Vol. 19, Nº1, jan - jun 2025
Neotropical Helminthology
Neotropical Helminthology, 2025, vol. 19 (1), 65-74
ORIGINAL ARTICLE / ARTÍCULO ORIGINAL
NEMATODES INFECTING
PHYLLOPEZUS POLLICARIS
(SPIX, 1825)
(REPTILIA: PHYLLODACTYLIDAE) IN THE PALMARES NATIONAL
FOREST, PIAUÍ STATE, BRAZIL
NEMATÓDEOS INFECTANDO
PHYLLOPEZUS POLLICARIS
(SPIX, 1825)
(REPTILIA: PHYLLODACTYLIDAE) NA FLORESTA NACIONAL
DE PALMARES, ESTADO DO PIAUÍ, BRASIL
NEMATODOS QUE INFECTAN A
PHYLLOPEZUS POLLICARIS
(SPIX, 1825)
(REPTILIA: PHYLLODACTYLIDAE) EN EL BOSQUE NACIONAL
DE PALMARES, ESTADO DE PIAUÍ, BRASIL
Brenda Milena Bezerra Costa
1
, Kássio de Castro Araújo
1
, Cristiana Ferreira-Silva
2
, Ronildo Alves
Benício
3
, Jussara da Conceição Eduardo
1
, Samuel Lopes Pereira
1
&Etielle Barroso de Andrade
1*
ISSN Versión Impresa 2218-6425 ISSN Versión Electrónica 1995-1403
DOI: https://dx.doi.org/10.62429/rnh20251911910
Universidad Nacional
Federico Villarreal
Volume 19, Number 1 (jan - jun) 2025
Este artículo es publicado por la revista Neotropical Helminthology de la Facultad de Ciencias Naturales y Matemática, Universidad Nacional Federico
Villarreal, Lima, Perú auspiciado por la Asociación Peruana de Helmintología e Invertebrados Af nes (APHIA). Este es un artículo de acceso abierto,
distribuido bajo los términos de la licencia Creative Commons Atribución 4.0 Internacional (CC BY 4.0) [https:// creativecommons.org/licenses/by/4.0/
deed.es] que permite el uso, distribución y reproducción en cualquier medio, siempre que la obra original sea debidamente citada de su fuente original.
ABSTRACT
Parasitological studies are important to understand the host-parasite interaction, providing information on the
coevolution, biogeography, morphology, and diet of the species. T e present study describes the endoparasite fauna of
the lizard
Phyllopezus pollicaris
(Spix, 1825) in the Palmares National Forest, Piauí State, Brazil, a tropical forest covered
by transitional vegetation between the Caatinga and Cerrado biomes. Furthermore, we investigated the inf uence of
1
Grupo de Pesquisa em Biodiversidade e Biotecnologia do Centro-Norte Piauiense -BIOTECPI, Instituto Federal de
Educação, Ciência e Tecnologia do Piauí - IFPI Campus Pedro II, 64255-000, Pedro II, Piauí, Brasil.
2
Núcleo Regional de Of ologia, Universidade Federal do Ceará - UFC, Bloco 905, Centro de Ciências, Campus PICI, 60455-
760,Fortaleza, Ceará, Brasil.
3
Laboratório de Herpetologia e Parasitologia de Animais Silvestres, Universidade Federal do Piauí - UFPI, Campus Picos,
64607-670, Picos, Piauí, Brasil.
* Corresponding author: brendamilena06@gmail.com
Brenda Milena Bezerra Costa:
https://orcid.org/0009-0004-3037-2769
Kássio de Castro Araújo:
https://orcid.org/0000-0003-4091-8521
Cristiana Ferreira-Silva:
https://orcid.org/0000-0002-3087-2614
Ronildo Alves Benício:
https://orcid.org/0000-0002-7928-2172
Jussara da Conceição Eduardo:
https://orcid.org/0009-0005-1233-0220
Samuel Lopes Pereira:
https://orcid.org/0009-0008-7618-4535
Etielle Barroso de Andrade:
https://orcid.org/0000-0002-5030-1675
66
Neotropical Helminthology, Vol. 19, Nº1, jan - jun 2025
Bezerra Costa
et al.
lizards’ size and sex on endoparasite infection patterns. We analyzed 23 individuals of
P. pollicaris
, of which 18 were
infected by at least one helminth, belonging to the Nematoda phylum:
Spauligodon oxkutzcabiensis
(Chitwood
, 1938),
Strongyloides
sp
.
, and
Parapharyngodon
sp.
Overall,
S. oxkutzcabiensis
showed the highest prevalence (73.9%), mean
infection intensity (8.64
± 12.7), and mean abundance (6.39 ± 11.5). We observed that the lizard size was positively
related to the abundance of endoparasites, while the sex of individuals had a poor efect on parasitological descriptors.
Tis study contributes information about the endoparasite fauna associated with
P. pollicaris
, composed exclusively of
nematodes. As a pioneering study, it can be considered the starting point for parasitological studies involving lizards in
Piauí State, Northeastern Brazil.
Keywords
: Geckos – helminths – lizards – parasites
RESUMO
Estudos parasitológicos são importantes para se compreender a relação parasito-hospedeiro, fornecendo informações
sobre a coevolução, biogeografa, morfologia e dieta das espécies. O presente estudo descreve a fauna endoparasitária
do lagarto
Phyllopezus pollicaris
(Spix, 1825) na Floresta Nacional (Flona) de Palmares, Piauí, Brasil, uma foresta
recoberta por vegetações de transição entre o Cerrado e a Caatinga. Além de investigar a infuência do sexo e do tamanho
dos lagartos nos padrões de infecções. Analisamos 23 exemplares de
P. pollicaris
dos quais 18 estavam infectados por
pelo menos um helminto, pertencente a três espécies de nematódeos:
Spauligodon oxkutzcabiensis
(Chitwood
, 1938),
Strongyloides
sp
.
e
Parapharyngodon
sp. No geral, a maior prevalência (73,9), intensidade média de infecção (8,64 ±
12,7) e abundância média (6,39 ± 11,5) foi de
S. oxkutzcabiensis
. Nós
observamos que o tamanho dos lagartos está
positivamente relacionado com a abundância de parasitos, enquanto que o sexo do indivíduo teve pouco efeito nos
descritores parasitológicos.
Este estudo contribui com informações sobre a fauna de endoparasitas associada a
P. pollicaris
,
sendo composta exclusivamente por nematoides. Por ser pioneiro, pode ser considerado o ponto de partida para estudos
parasitológicos envolvendo lagartos no estado do Piauí, Nordeste do Brasil.
Palavras-chave
: Gekkos – helmintos – lagartos – parasitos
RESUMEN
Los estudios parasitológicos son importantes para entender la relación parásito-huésped, proporcionando información
sobre la coevolución, biogeografía, morfología y dieta de la especie. El presente estudio describe la fauna endoparásita del
lagarto
Phyllopezus pollicaris
(Spix, 1825) en el Bosque Nacional de Palmares, Estado de Piauí, Brasil, un bosque cubierto
por vegetación de transición entre el Cerrado y la Caatinga. Además de investigar la infuencia del sexo y tamaño de las
lagartijas en los patrones de infección. Se analizaron 23 ejemplares de
P. pollicaris
, de los cuales 18 estaban infectados
por al menos un helminto, pertenecientes a tres especies de nematodos:
Spauligodon oxkutzcabiensis
(Chitwood
, 1938),
Strongyloides
sp. y
Parapharyngodon
sp. En general, la mayor prevalencia (73,9), intensidad media de la infección (8,64 ±
12,7) y abundancia media (6,39 ± 11,5) fue de
S. oxkutzcabiensis
. Observamos que el tamaño de la lagartija se relaciona
positivamente con la abundancia de parásitos, mientras que el sexo del individuo tuvo poco efecto en los descriptores
parasitológicos. Este estudio contribuye con información sobre la fauna endoparásita asociada a
P. pollicaris
, que está
compuesta exclusivamente por nematodos. Al ser pionero, puede considerarse el punto de partida para los estudios
parasitológicos que involucran lagartos en el estado de Piauí, en el noreste de Brasil.
Palabras clave
: Gekkos – helmintos – lagartos – parásito
67
Nematodes infecting
Phyllopezus pollicaris
Neotropical Helminthology, Vol. 19, Nº1, jan - jun 2025
INTRODUCTION
Parasites are organisms that need other living organisms to
survive, using the host as a food source and intermediate
or permanent home (Windsor, 1995). Tey are classifed
as ectoparasites when they live outside or in the host’s
pits (e.g., ticks) or endoparasites residing inside the host
(e.g., nematodes; Rey, 2018). Regarding endoparasites,
they are classifed as macro or microparasites, for
example, arthropods, plathelminths, nematodes, fungi,
and bacteria (Pantoja
et al
., 2015). Parasitological
studies provide information about the coevolution,
biogeography, morphology, and diet of species since
parasitic infections signifcantly afect their hosts (Poulin,
2021). Furthermore, endoparasites play an important role
in the ecosystems acting as environmental bioindicators
(Poulin, 1999; Zargar & Chishti, 2022).
Lizards are interesting ecological models for parasitological
studies because of their well-resolved taxonomy and for
acting as intermediate and defnitive hosts of diverse
endoparasite species (Wilson & Carpenter, 1996). Tis
interaction is usually associated with the host forage
mode and diet, in which they are infected directly by
larvae cutaneous penetration or by ingestion of infected
prey (González
et al.
, 2021; Dantas
et al
., 2024).
Lizards are one of the most diverse vertebrate groups,
with 7,724 species registered in the world (Uetz
et al
.,
2024). Morphophysiological adaptations and the water-
independence to reproduce contributed to the lizard
diversifcation worldwide (Hickman
et al
., 2013). Tere
are 292 lizard species in Brazil, of which 14 belong to
the Phyllodactylidae family (Guedes
et al
., 2023). Tese
lizards are characterized by having smooth skin with the
presence of granules, well-developed limbs, non-existent
eyelids, and adhesive lamellae that they use to attach
themselves to surfaces (Rocha, 1994; Gamble
et al
.,
2012;
Cacciali
et al
., 2018).
Among the Phyllodactylidae,
Phyllopezus pollicaris
(Spix,
1825) (Reptilia: Phyllodactylidae) is a mid-size Gecko
reaching up to 20 cm in total length, characterized by
having a depressed body, fattened head, snout with large
granules, well-developed limbs, and distinct coloring
between juveniles and adults (Rodrigues, 1985; Vanzolini
et al
.,1980). Tis species is widely distributed in Brazil,
being commonly recorded in the Caatinga and Cerrado
biomes (Gamble
et al
., 2012; Guedes
et al.
, 2023). It is a
nocturnal lizard that inhabits rocky outcrops, fallen tree
trunks, leaf litter, and also urban areas. In addition, it is a
“sit-and-wait” forager having an insectivore diet (Silva &
Araújo, 2008; Gamble
et al
., 2012; Carvalho
et al
., 2014).
Some studies already reported endoparasites infecting
P. pollicaris
, for exemple, nematodes and cestodes were
found parasitizing this lizard in the Estação Ecológica
de Aiuaba (Lima
et al
., 2017) and in an urban area of
Crato municipality (Sousa
et al
., 2014), both in Ceará
State, Northeastern Brazil. In these studies,
Spauligodon
oxkutzcabiensis
(Chitwood, 1938) had the highest
prevalence and mean infection intensity. Specifcally
in Piauí State, there is one only record of
Raillietiella
mottae
(Almeida, Freire & Lopes, 2008) parasitizing this
lizard (Sousa
et al.,
2010). Terefore, due to the lack
of parasitological studies dealing with lizard species in
Piauí State, the present study aimed to investigate the
endoparasites associated with
P. pollicaris
in the Palmares
National Forest, Piauí State, Northeastern Brazil, besides
testing how the size and sex of the individuals infuence
the infection patterns.
MATERIAL AND METHODS
Study area
Te present study was conducted in the Palmares
National Forest (Fig. 1), situated in the municipality of
Altos, Piauí State, Northeastern Brazil (05°03’25.90”S,
42°35’34.04”W, elev. 160 m). It is a small federal
conservation unit comprising 170 ha, created by
decree S/N of 21 February 2005, that aims to maintain
and protect wild animals and native forest species in
Piauí (ICMBio, 2022). It is a seasonal semideciduous
(ecotonal) forest inserted in a transition area between the
Cerrado and Caatinga biomes (IBGE, 2019). Te climate
is tropical with two well-defned seasons (rain and dry
seasons), annual rainfall ranging from 1200 to 1400 mm,
and annual mean temperature of 28º C (Brandão
et al.
,
2020; ICMBio, 2022; INMET, 2024).
Sampling
We searched for individuals of
P. pollicaris
using visual
active searches in microhabitats usually used by them,
such as trunks, trees, and sandstone outcrops (see
Recoder
et al.
, 2012; Gonçalves
et al.
, 2019). We carried
out two samplings of fve consecutive days in February
and March, totaling 10 collection days. Four researchers
conducted the samplings in the morning (9 to 12 h)
and night (18 to 23 h), totaling 320 hours of sampling
efort. We euthanatized the collected lizards following the
guidelines established by the National Council for the
Control of Animal Experimentation in accordance with
Law 11.794 of 2008, article 14 § 1º and with Normative
68
Neotropical Helminthology, Vol. 19, Nº1, jan - jun 2025
Bezerra Costa
et al.
Resolution 37, February 15, 2018 (Concea, 2018), and
the Brazilian Guide for the Production, Maintenance
or Use of Animals in Teaching or Scientifc Research
Activities (Jared
et al.
, 2023).
Figure 1.
Location map of the Palmares National Forest, Piauí State, Northeastern Brazil, highlighting
the main visiting trails.
Parasitological procedures
In the lab, we measured the snout-vent length of each
collected individual using a digital caliper (precision of
0.01 mm), and we necropsied the individuals with a
ventral incision and removed their organs with the help
of a stereomicroscope. Firstly, we analyzed the presence
of endoparasites in the oral cavity and, thereafter,
the following organs: digestive tract, lungs, heart,
liver, and kidneys (Amato
et al
., 1991). Te collected
endoparasites were fxed in 10% formalin, preserved
in 70% alcohol, and housed in the Helminthological
Collection of the Universidade Federal do Ceará,
Fortaleza, Brazil.
Data analyses
According to Bush
et al
. (1997), we used the following
parasitological descriptors: richness (total number of
endoparasite species), abundance (total number of
endoparasites), mean abundance (total number of a
given species of parasite in an individual/number of hosts
examined), prevalence (number of infected hosts/total
number of hosts analyzed), and mean infection intensity
(total number of a species of endoparasites/number of
hosts infected by the same parasite). After that, we used
the abundance data to investigate how the lizard sizes
infuence the endoparasite infection. For this, we used
the non-parametric regression of Kernel (Nadaraya,
1964) because our data did not follow the assumptions
for the linear regression test. We used the R packages
Vegan (Oksanen
et al.
, 2024) and ggplot2 (Wickham,
2016) to perform this analysis.
E
thic aspects
: All the procedures used in this work
comply with the ethical standards of the relevant
national and institutional guidelines on the care and
use of laboratory animals. Collection authorization
Chico Mendes Institute for Biodiversity Conservation
- ICMBio (#86665-2). Voucher specimens were housed
in the Biological Collection of Instituto Federal de
Educação, Ciência e Tecnologia do Piauí, Campus Pedro
II (CBPII).
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Nematodes infecting
Phyllopezus pollicaris
Neotropical Helminthology, Vol. 19, Nº1, jan - jun 2025
RESULTS
We collected 23
P. pollicaris
individuals (12 males and 11
females), of which 18 had at least one helminth parasite
(total prevalence = 78.2%). Overall, we found 149
nematodes with a mean abundance of 6.47 endoparasites
per individual and 8.27 mean infection intensity. Te
highest endoparasite abundance was found in the larger
intestine (139 nematodes), whereas 10 were recorded in
the stomach and fve in the small intestine (Table 1).
Table 1.
Endoparasites associated with
Phyllopezus pollicaris
in the Palmares National Forest, Piauí State, Northeastern
Brazil, with the respective number of helminths recorded (NH), prevalence (P), mean infection intensity (IM), mean
abundance (MA), minimum and maximum number of parasites per individual (Min-Max) and site of infection (IG).
EndoparasitosNHP%IM±SDMA±SDMin-MaxSI
Spauligodon oxkutzcabiensis
14773.98.64 ± 12
.
76.39 ± 11.51-41ST, LI, SI
Strongyloides
sp.14.3410.04 ± 0.201-1SI
Parapharyngodon
sp.14.3410.04 ± 0.201-1LI
TOTAL
14978.28.27 ± 12.46.47 ± 11.51-41
Legends:
standard deviation (SD), stomach (ST), large intestine (LI) and small intestine (SI)
.
Figure 2.
Nematode species parasitizing
Phyllopezus pollicaris
in the Palmares National Forest:
A)
Spauligodon
oxkutzcabiensis
,
B)
Strongyloides
sp., and
C)
Parapharyngodon
sp.
Phyllopezus pollicaris
was exclusively parasitized by
nematodes belonging to three species:
S. oxkutzcabiensis
,
Strongyloides
sp
.
, and
Parapharyngodon
sp. (Fig. 2).
Among these,
S. oxkutzcabiensis
had highest prevalence,
abundance, and mean infection intensity. Te other two
nematode species parasitized a single male individual.
Additionally, we observed that males had the highest
abundance (NH = 89), mean abundance (MA = 7.42),
prevalence (P = 83.3), and mean infection intensity (IM
= 8.90) than females (NH = 60, MA =5.45, P = 72.3, IM
= 7.50).
It is interesting to note that the abundance of
endoparasites was positively related to the lizard size (F
= 4.481, P = 0.025), thus, larger individuals tended to be
more parasitized (Fig. 3).
DISCUSSION
We found a high abundance of helminths parasitizing
P. pollicaris
belonging to three species, composed of
nematodes exclusively. Tese helminths are usually
found parasitizing reptiles (Anderson, 2000; Lacerda
et al
., 2023), and exclusively nematodes infection was
already reported for other lizards such as
Tropidurus
hispidus
(Spix, 1825) and
T. semitaeniatus
(Spix,
1825), and even Geckos such as
Hemidactylus
mabouia
(Moreau de Jonnés, 1818) (Ávila
et al.
, 2012; Anjos
et
al.,
2013).
70
Neotropical Helminthology, Vol. 19, Nº1, jan - jun 2025
Bezerra Costa
et al.
Among the
Phyllopezus
species, our results support the
fndings this lizard tend to be predominantly parasitized by
nematodes, mainly for
S. oxkutzcabiensis
in the Caatinga
biome (Ávila
et al.
, 2012; Sousa
et al
., 2014; Lima
et
al.
,
2017). Te high infections taxa of
S. oxkutzcabiensis
may be associated with its monoxenic life cycle, in which
parasitic infection can occur through the ingestion of
eggs (Anderson, 2000). Tis helminth had the highest
abundance and prevalence in the present study, and it is
commonly reported parasitizing other geckos in diferent
Brazilian biomes, as Cerrado, Caatinga, and Mata
Atlantica (Ávila & Silva, 2010; Ávila
et al
., 2012; Ávila
et al
., 2010; Araujo
et al
., 2020). It is noteworthy to note
that the host’s biological characteristics also infuence the
parasitic infection patterns (Aho, 1990).
We found only one
Strongyloides
sp. specimen parasitizing
the larger intestine of a single male individual of
P.
pollicaris
. Te genus
Strongyloides
can be usually found
in the gastrointestinal tract of reptiles, birds, amphibians,
and mammals (Little, 1966; Roca & Hornero, 1992).
Tese parasites have a complex life cycle with two phases:
in the free-living phase, male and female individuais are
found that develop through indirect development and
exchange gametes (sexual or heterogonic reproduction),
and in the parasitic phase, parthenogenetic females are
found only that produce eggs without the need for a male
(asexual or homogonic reproduction) (Viney & Lok,
2007). Tis is the frst report of this genus parasitizing
P.
pollicaris
, despite this nematode being frequently reported
to infect reptiles (Ávila & Silva, 2010; Mati
et al
.,
2013;
Mesquita
et al.
, 2020). Because we only found one female
with a broken posterior region, we were unable to count
the number of turns the posterior ovary makes around the
intestine, a characteristic crucial to species identifcation
(Little, 1966). As a result, we were unable to identify this
helminth to the species level.
Parapharyngodon
is a nematode genus widely distribut-
ed and commonly recorded infecting insectivore reptile
species (Adamson, 1981), including lizards such as
P.
pollicaris, Ameiva ameiva
(Linnaeus, 1758), and
T. semi-
taeniatus
(Lacerda
et al
., 2023). Once this helminth has
a monoxenic lifecycle (Anderson, 2000), host infection
probably occurs during foraging activity, where this lizard
accidentally ingested the eggs of infectious parasites that
were attached to its prey or in the environment. However,
further investigations are necessary to prove the validity
of this proposal.
Parapharyngodon
sp. was not identifed
to species level because only a single female was record-
ed, and the identifcation of
Parapharyngodon
species is
carried out based mainly on the reproductive characteris-
tics of males, such as morphology and size of the spicule,
number and arrangement of caudal papillae, presence or
absence of genital cone, and shape of the anterior lip of
the cloaca.
Figure 3
. Relationship between lizard size (SVL) and abundance of parasites associated with
Phyllopezus pollicaris
in the
Palmares National Forest, Piauí, Northeastern Brazil. We used logarithmized values for a better visualization.
71
Nematodes infecting
Phyllopezus pollicaris
Neotropical Helminthology, Vol. 19, Nº1, jan - jun 2025
We observed that larger lizards tended to be more infected
(highest endoparasite abundance). Based on the species-
area hypothesis (see Tjørve
et al.
, 2008), it is expected that
larger hosts will have more space and high niche diversity
to house endoparasites (Anjos
et al.
, 2012; Kamiya
et al.
,
2013; Campião
et al
., 2015). Tis pattern was already
reported in lizards, including
P. pollicaris
in an urban area
in Northeastern Brazil (Sousa
et al
., 2014). However,
further studies are still needed to understand the main
drivers of endoparasite infections in lizard species.
Overall, this study contributes to the knowledge of the
parasitic fauna of the lizard
P. pollicaris
, being the frst
study carried out in the Palmares National Forest, Piauí
State, Northeastern Brazil. Aspects related to seasonality,
foraging activity and feeding habits of hosts infuence
endoparasite infections, however, little is known about how
parasitological descriptors are afected by the environmental
characteristics of the Palmares National Forest. Terefore,
we reinforce the need of more parasitological studies in
this conservation unit aiming to understand the host-
endoparasite in lizard species of Piauí State.
ACKNOWLEDGMENTS
RAB and KCA thank Conselho Nacional de
Desenvolvimento Científco e Tecnológico (CNPq) and
Fundação de Amparo à Pesquisa do Estado do Piauí
(FAPEPI) for providing research fellowship (Processes:
301239/2022-3; 150013/2023-0). CFS thank to CNPq
(#150125/2023-2) and thank Fundação Cearense de
Apoio ao Desenvolvimento Científco e Tecnológico
(FUNCAP) (#FC3-0198-00006.01.00/22) for research
funding. We thank Andreia Andrade, Douglas Oliveira,
and João Lucas Ferreira for feldwork support. We also
thank the manager of the Palmares National Forest,
Gaspar da Silva Alencar, for logistical support.
Author contributions: CRediT (Contributor Roles
Taxonomy)
BMBC
= Brenda Milena Bezerra Costa
KCA
= Kássio de Castro Araújo
CFS
= Cristiana Ferreira-Silva
RAB
= Ronildo Alves Benício
JCE
= Jussara da Conceição Eduardo
SLP
= Samuel Lopes Pereira
EBA
= Etielle Barroso de Andrade
Conceptualization:
BMBC, KCA
Data curation:
BMBC, EBA
Formal Analysis:
BMBC, KCA
Funding acquisition:
RAB
Investigation:
BMBC, KCA, RAB, EBA
Methodology:
BMBC, KCA, RAB, EBA, CFS, JCE, SLP
Project administration:
KCA, EBA
Resources:
RAB, EBA
Software:
KCA, EBA
Supervision:
KCA, RAB, EBA
Validation:
KCA, RAB, EBA
Visualization:
BMBC, KCA, RAB, EBA
Writing – original draft:
BMBC, JCE, SLP, CFS
Writing – review & editing:
KCA, RAB, EBA, CFS
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