
The nematode community in the Atlantic
rainforest lizard Enyalius perditus Jackson,

1978 from south-eastern Brazil

A.F. Barreto-Lima1*, G.M. Toledo2 and L.A. Anjos2

1UFRGS – Universidade Federal do Rio Grande do Sul – Instituto de
Biociências, Programa de Pós-Graduação em Ecologia, Avenida Bento

Gonçalves, n8 9500, Setor 4, Prédio 43422, Bairro Agronomia, Caixa Postal
15007, Porto Alegre, Rio Grande do Sul, Brazil: 2UNESP – Universidade

Estadual Paulista, Campus de Botucatu – Instituto de Biociências,
Departamento de Parasitologia, Botucatu, São Paulo, Brazil

(Received 10 March 2011; Accepted 4 September 2011; First Published Online 11 October 2011)

Abstract

Studies focusing on communities of helminths from Brazilian lizards are
increasing, but there are many blanks in the knowledge of parasitic fauna of wild
fauna. This lack of knowledge hampers understanding of ecological and
parasitological aspects of involved species. Moreover, the majority of research
has focused on parasitic fauna of lizards from families Tropiduridae and
Scincidae. Only a few studies have looked at lizards from the family
Leiosauridae, including some species of Enyalius. This study presents data on
the gastrointestinal parasite fauna of Enyalius perditus and their relationships
with ecological aspects of hosts in a disturbed Atlantic rainforest area in the state
of Minas Gerais, south-eastern Brazil. Two nematode species, Oswaldocruzia
burseyi [(Molineidae) and Strongyluris oscari (Heterakidae) were found. Nematode
species showed an aggregated distribution in this host population, with O. burseyi
being more aggregated than S. oscari. The present study extends the range of
occurrence of O. burseyi to the Brazilian continental area.

Introduction

The host–parasite relationship is an important par-
ameter in the study of animal communities, as such
relationships affect population dynamics and community
structure (Rocha et al., 2000; Anjos et al., 2005; Almeida
et al., 2009). The helminth fauna of lizards in South
America has received greater attention in recent years,
with many records of new hosts and localities as well as
the description of new species (Goldberg et al., 2004;
Bursey et al., 2005a, b; Durette-Desset et al., 2006; Bursey &
Goldberg, 2007; Vrcibradic et al., 2008; Ávila & Silva,
2010). Nevertheless, studies on helminth communities
and ecological aspects of host–parasite relationships

remain scarce in the literature (Rocha & Vrcibradic, 2003;
Anjos et al., 2005) and greater knowledge on the
parasitological and ecological features of different lizard
species is needed.

The genus Enyalius Wied, 1821 is comprised of nine
species (Sociedade Brasileira de Herpetologia, 2010) of
diurnal, insectivorous lizards (Sousa & Cruz, 2008;
Barreto-Lima, 2009) well distributed throughout different
biomes in Brazil, such as the Atlantic rainforest
(Etheridge, 1969; Vanzolini, 1972, 1974; Jackson, 1978),
Amazon (Ávila-Pires, 1995; Vitt et al., 1996), Caatinga
(semi-arid brush) and Cerrado (savanna-like vegetation)
(Bertolotto et al., 2002; Rodrigues et al., 2006). Despite
the wide distribution of Enyalius, few studies have
addressed its helminth fauna and parasite ecology
(Vicente et al., 1993; Durette-Desset et al., 2006; Sousa
et al., 2007; Vrcibradic et al., 2007, 2008). Enyalius perditus*E-mail: afblima@hotmail.com

Journal of Helminthology (2012) 86, 395–400 doi:10.1017/S0022149X11000599
q Cambridge University Press 2011


Jackson, 1978 is a small tropical lizard restricted to the
easternmost Atlantic rainforest in southern and south-
eastern Brazil (Jackson, 1978; Lima & Sousa, 2006) that has
been insufficiently studied with regard to many ecologi-
cal and behavioural aspects (Lima & Sousa, 2006).

As infection parameters, such as the prevalence and
number of parasites, may be influenced by the sex, age,
size (Ribas et al., 1995) and diet composition (Goldberg
et al., 1993, 1995) of the host, the aim of the present study
was to analyse helminth fauna from the gastrointestinal
tract of E. perditus and determine the relationship between
ecological aspects of the host, such as body mass, sex and
size, and the parasitic fauna.

Materials and methods

Host and parasite sampling

Lizards were collected with pitfall traps in a forest
fragment of the Santa Cândida Municipal Biological
Reserve (2184503500S, 4382005000W) in the city of Juiz de
Fora, state of Minas Gerais, Brazil, between 2002 and
2003. The lizard species were identified based on
Rodrigues et al. (2006). Animals were weighed on a
manual scale (accuracy: 0.10 g), euthanized with ether,
fixed in a 10% formalin solution and stored in 70%
ethanol. Snout–vent length (SVL), total body length (TBL)
and tail length (TL) were measured with a caliper to
the nearest 0.1 mm. Gonads were examined to determine
the sex. The gastrointestinal tract (stomach, small and
large intestines) was removed and examined for
helminths under a stereomicroscope. Nematodes were
removed, counted, treated with lactophenol for clarifica-
tion and examined under an optical microscope (Leicaw

DM5000B), using the LAS software program (Leica
Application Suite) for image analysis. The prevalence
and mean intensity of infection of each nematode species
were determined based on Bush et al. (1997).

Data analysis

Differences in body size and mean intensity of infection
between sexes were determined using the Mann–
Whitney test and Student’s t-test, respectively. A test for
proportions (Z-test) was used to determine whether
there was a significant difference in overall prevalence
between males and females (Zar, 1999). A Spearman
rank correlation (Zar, 1999) was performed to evaluate
the effect of host size (SVL) on the intensity of infection.
A Spearman rank correlation was performed to evaluate
the relationship between body mass and intensity
of infection, using the residual values obtained from
linear regression analysis between TL and body mass
(Zar, 1999). The discrepancy index (D) was calculated, as
suggested by Poulin (1993). This index has a minimum of
zero (D ¼ 0) when every host harbours the same number
of parasites. When all parasites are found in a single
host, aggregation is maximal (D ¼ 1). This index was
calculated using the Quantitative Parasitology 3.0 soft-
ware program (Rózsa et al., 2000). For all tests, the level
of significance was a # 0.05. Voucher specimens of the
lizard hosts were deposited in the Herpetological
Collection of the Zoology Laboratory of the Universidade

Federal de Juiz de Fora (1–5s/n.2002/UFJF, 59–61.2002/
UFJF, 233–240 CZ/UFJF, 243 CZ/UFJF, 245–255 CZ/
UFJF, 257–275 CZ/UFJF) and parasite specimens were
deposited in the Coleção Helmintológica do Instituto de
Biociências de Botucatu (CHIBB Lotes nos 5099 and 6000).

Results

Forty-nine specimens of E. perditus (31 males and 18
females) were examined: 27 adult males (SVL ¼ 72.2
^ 5.2 mm; mass ¼ 8.7 ^ 2.2 g), 15 adult females
(SVL ¼ 83.8 ^ 6.9 mm; mass ¼ 15.9 ^ 6.4 g) and seven
juveniles (SVL ¼ 37.4 ^ 5.7 mm; mass ¼ 1.5 ^ 0.9 g). Size
and body mass were different between adult males and

25

20

15

10

5

0

60 65 70 75

Body length (mm)

In
te

ns
ity

 o
f i

nf
ec

tio
n

80 85 90

Fig. 1. Regression between body length and intensity of infection
of Enyalius perditus from Reserva Biológica Municipal Fazenda
Santa Cândida, Juiz de Fora municipality, Minas Gerais state,

Brazil.

25

20

15

10

5

0

–4 –3 –2 –1 0

Residuals of mass

In
te

ns
ity

 o
f i

nf
ec

tio
n

1 2 3 4 5

Fig. 2. Regression between mass (independent of total body
length) and intensity of infection of Enyalius perditus from
Reserva Biológica Municipal Fazenda Santa Cândida, Juiz de

Fora municipality, Minas Gerais state, Brazil.

396 A.F. Barreto-Lima et al.


adult females (SVL: Z(U) ¼ 3.5, P , 0.001; body mass ¼
Z(U) ¼ 2.2, P , 0.05).

The overall prevalence of parasites was 55.1% (27/49):
70.4% in adult males, 40.0% in adult females and 28.6%
in juveniles. The difference between the prevalence of
parasites in adult males and adult females was non-
significant (Z ¼ 1.59, P ¼ 0.11). Mean intensity of infec-
tion was 7.1 ^ 6.8 (range: 1–26). The mean intensity of
infection was 11.3 ^ 8.9 in adult females and 6.3 ^ 6.0 in
adult males. However, this difference did not achieve
statistical significance (t ¼ 1.61, df ¼ 23, P ¼ 0.12). Two
juvenile lizards were infected by two and four nematodes.
The body size (SVL) of adult lizards was positively
associated with the intensity of infection (F ¼ 12.74,
P ¼ 0.003, r 2 ¼ 0.423) (fig. 1). After adjusting for the effect
of total body length (residual between TBL £ mass),
body mass of the adult hosts was positively associated
with the intensity of infection (F ¼ 4.68; P ¼ 0.045;
r 2 ¼ 0.187) (fig. 2).

Two nematode species were found in the stomach,
small intestine and large intestine of lizards: Oswaldocru-
zia burseyi Duretter-Desset, Anjos & Vrcibradic, 2006
(Molineidae) and Strongyluris oscari Travassos, 1923

(Heterakidae). The prevalence of O. burseyi was 62.7%
(n ¼ 17), with a mean intensity of infection of 3.1 ^ 2.9
(range 1–12) and discrepancy index of 0.621. The
prevalence of S. oscari was 74.1% (n ¼ 20), with a mean
intensity of infection of 6.75 ^ 6.4 (range 1–25) and
discrepancy index of 0.572. The prevalence of these two
nematode species was similar (Z ¼ 0.586; P ¼ 0.558).
However, S. oscari exhibited a significantly higher mean
intensity of infection (t ¼ 22.3; P ¼ 0.03). The discre-
pancy index (D) for both parasite species was 0.647
(O.burseyi, 0.621; S. oscari, 0.572).

Discussion

Lizards are hosts to a wide variety of gastrointestinal
nematodes (Goldberg & Bursey, 1992). The cosmopolitan
genus Oswaldocruzia Travassos, 1917 parasitizes amphi-
bians (Durette-Desset et al., 2006) and lizards from several
families, such as Gekkonidae, Gymnphthalmidae, Igua-
nidae, Leiosauridae, Polychrotidae, Teiidae and Tropi-
duridae (Ávila & Silva, 2010). Oswaldocruzia nematodes
are common parasites of lizards and new species have

Table 1. Helminth fauna of Enyalius spp. from Brazil; P(%), prevalence, MI, mean intensity of infection (^standard deviation).

Enyalius
bilineatus

Enyalius
iheringii

Enyalius
perditus

Sites of
infectionHelminth species P(%) MI P(%) MI P(%) MI

Acanthocephala
Acanthocephalus sp. 7.1 31 S, SI
Centrorhynchidae gen. sp. 3.7 2 S

Nematoda
Acuariidae gen. sp. – larvae 25.9 1.7 ^ 1.2 S, C

Cosmocercidade
Aplectana vellardi 6 4.8 ^ 0.52 SI, LI
Cosmocerca sp. 14.3 41 LI

Heterakidae
Strongyluris oscari 16.7 6 28.6 1.51 S, LI

58 7.6 ^ 7.32 SI, LI
Molineidade

Oswaldocruzia sp.a – –3 I
Oswaldocruzia fredi 33.3 5 S, SI
Oswaldocruzia burseyi 42.8 2.83 S, SI, LI
Oswaldocruzia benslimanei 7.4 1.5 SI
Oswaldocruzia subauricularis 29 3.2 ^ 3.12 S, SI, LI

Physalopteridae
Physaloptera lutzi 7.4 2 S, LI, LU, C
Physaloptera retusa 11.1 2.7 ^ 0.6 S, LI, C

Rhabdiasidae
Rhabdias sp. 33.3 2 ^ 1.1 33.3 1 14.3 21 LU

Locality (State) Marechal
Floriano (ES)

Ilha de São
Sebastião (SP)

1Ilha de São Sebastião (SP)
2Lima Duarte (MG)
3Rio de Janeiro (RJ)

References Durette-Desset et al.
(2006); Vrcibradic
et al. (2007)

Vrcibradic
et al. (2008)

Freitas (1955, 1956); Vicente et al. (1993);
Durette-Desset et al. (2006); Sousa et al.
(2007); Vrcibradic et al. (2008); Ávila & Silva
(2010)

Sites of infection: S, stomach, SI, small intestine; LI, large intestine; I, intestine; C, body cavity; LU, lungs.
Localities: ES, Espı́rito Santo State; MG, Minas Gerais; RJ, Rio de Janeiro State; SP, Sāo Paulo State.
Rhabdias sp. was found in E. catenatus in the Ibateguara locality, Alagoas state, by Freire (2008): P(%) ¼ 36.4, MI ¼ 3.7(^6.3).
a Oswaldocruzia subauricularis from the host Enyalius catenatus (without data), in Freitas (1955 and, probably, 1956) are both considered
as species inquirenda by Durette-Desset et al. (2006), and Enyalius catenatus was described as Enyalius perditus by Jackson (1978).

The nematode community in Enyalius perditus from south-eastern Brazil 397


been described infecting Enyalius, such as two new
species of Oswaldocruzia sampled from the stomach and
small intestine of E. iheringii and E. perditus on São
Sebastião Island off the state of São Paulo, Brazil (Durette-
Desset et al., 2006) (table 1). Vrcibradic et al. (2008)
reported that helminth assemblages in E. perditus and
E. iheringii on São Sebastião Island were depauperate and
dominated by generalist helminths with direct life cycles.
It is important to note that the host species E. catenatus
analysed by Freitas (1955) was later described as
E. perditus by Jackson (1978) and its nematode species
O. subauricularis was misidentified and is considered a
species inquirenda by Durette-Desset et al. (2006).

Oswaldocruzia burseyi was initially recorded for an
island habitat (São Sebastião Island) (Vrcibradic et al.,
2008) in an area of the Atlantic rainforest. The present
study reports the first occurrence of O. burseyi in the state
of Minas Gerais, extending its range of occurrence to the
Brazilian mainland. The other species recorded in the
present study, S. oscari, is a generalist parasite found in the
stomach and intestine of different species of South
American lizards: Ameiva ameiva, Anolis fuscoauratus,
A. punctatus, A. transversalis, E. iheringii, E. perditus,
Eurolophosaurus nanuzae, Mabuya agilis, Plica plica, P. umbra,
Stenocercus caducus, S. roseiventris, Tropidurus sp. T. guarani,
T. spinulosus, T. torquatus and T. melanopleurus (Sousa et al.,
2007; Vrcibradic et al., 2008, Ávila & Silva, 2010).

The present study reports the lowest richness of
helminth fauna associated with the gastrointestinal tract
of species of Enyalius. Parasites and predators are the first
groups to suffer from the environmental impact caused
by human activities (Gibb & Hochuli, 2002; Laurence et al.,
2002). The Santa Cândida Municipal Biological Reserve,
from which the lizards were sampled, is a small remnant
of secondary Atlantic rainforest surrounded by human
habitations. In the recent past, this area was covered by
coffee plantations. According to McKenzie (2007),
changes in anthropogenic land use affect the intermediate
host fauna and, consequently, the helminth fauna. Thus,
recent changes in the sampling area could have
influenced the availability and abundance of prey,
which may have reduced the richness of the helminth
fauna in comparison to fauna in a typical natural forest
(Vrcibradic et al., 2008).

Results of the present study confirm the hypothesis that
the intensity of infection increases with the body mass
and size of the lizard host, which corroborates findings
described in previous studies (Ribas et al., 1995; Sousa
et al., 2007). The prevalence of parasites was higher in
male lizards than in females. Similar findings are
described for Anolis sp. (Vogel & Bundy, 1987),
Cnemidophorus ocellifer (Ribas et al., 1995) and E. perditus
(Sousa et al., 2007). However, a greater mean intensity of
infection was found in females in the present study, which
differs from findings reported for E. perditus by Sousa et al.
(2007). This may be explained by local differences in the
specific characteristics of the host populations in each
study area and the small number of adult female hosts
sampled in the present study. Lizards acquire nematodes
through the ingestion of arthropods infected with the
larvae of these parasites (Goldberg & Bursey, 1992; Ribas
et al., 1995). Thus, gender differences in the prevalence
and intensity of helminths may be due to variations in

body size and differences in prey consumption in
qualitative and/or quantitative terms (Schoener, 1967;
Fitch, 1981; Ribas et al., 1995).

The habitat for parasites is not spatially continuous, but
rather consists of cells or discrete ‘islands’, i.e. the hosts
represent patches of favourable habitats in an uninhabi-
table environment. Parasites are also not evenly dis-
tributed among these islands. Some hosts contain more
parasites than average, others contain less, and many
individuals in the population may be free of parasites.
Whichever the index of aggregation chosen, the result is
usually the same; parasite populations are much more
aggregated among their hosts than if the parasites were
randomly distributed (Poulin, 1993, 1997). In the present
study on the parasite biology of E. perditus, the nematode
species exhibited a more aggregated distribution pattern
in this host population and O. burseyi was more
aggregated than individuals of S. oscari.

Acknowledgements

We thank Dr Reinaldo J. da Silva who facilitated the
helminthological analysis at the Laboratório de Para-
sitologia de Animais Silvestres (LAPAS), UNESP, and to
Dr Bernadete M. de Sousa who initially facilitated the
dissection of the lizards in one of the laboratories of the
Postgraduate Program (PPG), Comportamento e Biologia
Animal, UFJF. L.A.A. thanks Fundação de Amparo à
Pesquisa do Estado de São Paulo (FAPESP) for financial
support (postdoctoral scolarship, # 2008/50417-7). A.F.B.-
L. thanks Coordenação de Aperfeiçoamento de Pessoal de
Nı́vel Superior (CAPES), and PPG, Comportamento e
Biologia Animal, UFJF for support and facilities. Finally
we thank City Hall of Juiz de Fora for permission to access
the reserve, to Instituto Brasileiro dos Recursos Naturals
Renováveis (IBAMA) for permission to collect the animals
(227/2003, Fauna/MG) and to the Comissão de Ética na
Experimentação Animal da Pró-Reitoria de Pesquisa, UFJF
(47/2003). Finally we thank to Katherine Cameron and
the two anonymous referees for providing us with
constructive comments and suggestions.

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