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The lizard genus Enyalius (Leiosauridae) currently
comprises nine species distributed mainly in the
Atlantic Forest domain of South America, and in the
Brazilian Amazon rainforest, Cerrado, and Caatinga
(Frost et al., 2001; Rodrigues et al., 2006; Uetz, 2013).
Species of Enyalius are insectivorous, diurnal, with
arboreal or semi-arboreal habits (Jackson, 1978; Vitt
et al., 1996; Zamprogno et al., 2001; Van Sluys et al.,
2004; Rautenberg and Laps, 2010; Barreto-Lima and
Sousa, 2011). Enyalius iheringii Boulenger, 1885 and
E. perditus Jackson, 1978 are two species endemic
to the Brazilian Atlantic forest in southeastern Brazil
(Jackson, 1978), whose natural history, especially
their reproductive aspects, is poorly known (Marques
and Sazima, 2004; Barreto-Lima and Sousa, 2006;
Rautenberg and Laps, 2010; Sturaro and Silva, 2010).
Herein, we provide novel information on reproductive
aspects of E. iheringii and E. perditus from São Paulo,
southeastern Brazil.
On 3 November 2012 at 11:10 h (mid-spring), a
couple of E. perditus (both with snout-vent length
between 120-130 mm, approximately) were found in
a copulation attempt, during a herpetofaunal survey in
a fragment of Atlantic Forest with approximately 52
km2 at Juquiazinho district, in Juquitiba, São Paulo (-
23.97203, -47.02294; datum=WGS84; 730 m a.s.l.),
south-eastern Brazil. The couple was on a tree trunk
about 1 m above ground. The male was biting the
female’s neck when suddenly both fell from the trunk
on the leaf litter. After the fall, the male remained on
the back of the female, biting her neck in an attempt to
immobilize her (Fig. 1). Both remained in this position
for about 30 seconds. After this, the female disengaged
from the male and escaped, followed by the male a few
seconds later. Effective copulation was not observed.
Courtship was previously reported for at least three
species of Enyalius. Vitt et al. (1996) reported a possible
courtship in eld conditions for E. leechii (Boulenger,
1885), whereas Grantsau (1966) and Barreto-Lima and
Sousa (2006) reported courtship and mating in captive
conditions for E. catenatus and E. perditus. To our
knowledge, our observation was the rst courtship
conrmed taken in eld conditions for a species of
Enyalius. In both captive reports, male bit females
during attempted copulation. The pattern of courtship
behaviour observed in nature was similar to those taken
in captive conditions (Grantsau, 1966; Barreto-Lima
and Sousa, 2006).
The period of courtship (spring) was coincident with
the period of vitellogenesis and courtship/mating in
captivity previously reported for the species (Barreto-
Lima and Souza, 2006; Sturaro and Silva, 2010;
Table 1). Coincidently, the same female involved in
this courtship/mating episode was observed in the
previous night. The individual was recognized for
being in process of shedding skin (Fig. 2), in addition
to the presence of a callosity in the tail. The shedding
process is associated with the mating season of lizards
and snakes (Mason and Gutzke, 1990; Marques et al.,
2009), and several authors have suggested that shedding
is part of the oestrus signal (Aldridge and Duvall,
2002). The proposed mechanism is that reproductive
pheromones released at the time of shedding stimulate
male courtship behaviour (Mason and Gutzke, 1990;
Mason, 1992). Additionally, this female showed some
light orange spots on the neck and throat during the
copulation attempt, similarly to the observed by Sturaro
and Silva (2010) in reproductive females, suggesting
that this feature is an important courtship stimulator for
Herpetology Notes, volume 7: 273-276 (2014) (published online on 25 April 2014)
Reproductive aspects of two Enyalius
lizards from the Atlantic forest in Southeastern Brazil
Serena Najara Migliore1,2, Henrique Bartolomeu Braz1 and Selma Maria Almeida-Santos1
1Laboratório de Ecologia e Evolução, Instituto Butantan - Av.
Dr. Vital Brazil, 1500 – Butantã, São Paulo-SP, CEP 05503-
2Corresponding author:;
Serena Najara Migliore et al.
males (Baird, 2004). Taken together, shedding process
and colour signal suggest that the female of E. perditus
was probably signalling her condition to the male.
Reproductive observations on Enyalius iheringii were
taken from a female (voucher specimen: IBSPCR 0425;
Instituto Butantan, São Paulo, Coleção de Referência)
collected on 3 March 2012, in the Serra do Mar State
Park, Ubatuba, São Paulo, southeastern Brazil. The
female measured 103 mm of snout-vent length (SVL)
and 203 mm of tail length (TL). It was kept in a cage
containing branches and leaf litter. On 2 April 2012, it
laid 18 eggs within the cage. Five eggs were infertile.
The remaining 13 eggs were measured, using a digital
calliper (to the nearest 0.1 mm) and weighed, using a
digital balance (to the nearest 0.01 g). Eggs averaged
15.0 ± 0.5 mm in length (range = 14.1–15.7 mm), 9.7 ±
0.7 mm in width (range = 8.8–11.2 mm), and 0.81 ± 0.10
g in mass (range = 0.70–1.05 g). After oviposition, the
female weighed 24.0 g. The relative clutch mass (RCM;
total clutch mass/maternal body mass after oviposition
+ total clutch mass; following Vitt and Price, 1982) was
0.378. Thirteen eggs were incubated. Eggs were housed
in a plastic container, half-buried with moistened
vermiculite, and maintained at room temperature
Figure 1. Copulation attempt in Enyalius perditus (Juquiazinho, Juquitiba, São Paulo, Brazil). Notice the male (green individual)
biting female’s neck (brownish individual) in attempt to immobilize her.
Table 1.
Species Courtship Mating Vitellogenesis Gravidity or
Corpus luteum Hatching Clutch size
E. perditus Spring
(Dec) and
summer (Jan)
(Feb and Mar)
- 7-14
E. iheringii - - Spring
(Dec) and
Summer (Jan)
(Jan and Feb) and
Autumn (Apr)
- Winter
(1) Barreto-Lima and Sousa (2006); (2) Sturaro and Silva (20 10); (3) Marques and Sazima (2004); (4) Rautenberg and Laps (2010); (5) Present study.
Table 1. Timing of reproductive events recorded for Enyalius perditus and E. iheringii in the literature and present study.
averaging 2 C. Eight eggs spoiled over incubation
period due to fungal contamination. Hatchings (n =
4) occurred between 23 and 28 August 2012, after an
incubation period ranging from 143 to 148 days. One egg
failed to hatch. This egg was dissected, and it contained
one full-term dead embryo with a malformation in the
tail. Hatchlings averaged 27.5 ± 3.5 mm SVL (range =
25–30 mm), 49.0 ± 8.5 mm TL (range = 43–55 mm),
and 0.77 ± 0.01 g (range = 0.76–0.78 g).
There are no data on RCM for other Enyalius but the
value observed here in E. iheringii is high and close
to the upper limit observed in other lizards (Vitt and
Price, 1982). In lizards, RCM and foraging modes
are associated. RCM tends to be relatively low within
species that forage actively, and relatively high within
species using the “sit-and-wait” foraging mode (Vitt and
Price, 1982). Since females of Enyalius species appears
to be “sit-and-wait” foragers (Sousa and Cruz, 2008;
Borges et al., 2013) and move shorter distances than
males (Barreto-Lima et al., 2013), the high RCM value
observed here in E. iheringii appears to be congruent
with such association.
Estimates on gravidity period and clutch size available
in the literature for Enyalius were largely obtained from
dissection of preserved females (Rand, 1982; Vitt et
al., 1996; Marques and Sazima, 2004; Rautenberg and
Laps, 2010; Sturaro and Silva, 2010). This is the rst
report of egg size, incubation period, and hatchling size
of an Enyalius species. Gravid females of E. iheringii
were previously found in January and February
(Table 1) suggesting that the species presents some
reproductive seasonality (Marques and Sazima, 2004;
Rautenberg and Laps, 2010; R. Rautenberg, pers. com.).
Our observation of egg laying in April corroborates this
assumption but suggests that oviposition may be further
extended (at least to early autumn) than previously
thought. Considering the period of encounter of
gravid females of E. iheringii and our hatching report,
recruitment appears to occur from autumn to mid-winter
(Table 1).
The few information available for E. perditus and E.
iheringii impair broad comparisons between species and
denitive generalization on the timing of reproductive
events. However, taken together our data and literature
Figure 2. (A) The female Enyalius perditus observed one night before the copulation attempt. (B) Detail of the female’s head
highlighting the piece of skin in the head above the eye, indicating the process of shedding.
Reproductive aspects of two Enyalius lizards from the Atlantic forest 275
information indicate certain similarity between species,
at least on the timing of vitellogenesis and gravidity
(Table 1). However, gravidity period appears to extend
further (to early autumn) in E. iheringii. Detailed studies
are clearly needed to fully understand the reproductive
cycles of these species and test this assumption.
Acknowledgements. We thank M. T. Rodrigues, F. F. Curcio,
and V. J. Germano for taxonomic identication of the specimens,
and R. Recoder for the critical reading and comments on the
manuscript. We also thank R. Rautenberg, A. F. Barreto-Lima,
and B. Rocha for information on Enyalius specimens, and A.
Silveira for assistance with hatchlings. PAP (Programa de
Aprimoramento Prossional) provided a fellowship to S. N.
Migliore, and FAPESP (Fundação de Amparo à Pesquisa do
Estado de São Paulo) to H. B. Braz (grant no. 2009/54478-3).
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Serena Najara Migliore et al.
Accepted by Diogo Provete;
Managing Editor: Diogo Provete
... Enyalius perditus is commonly found in Atlantic forest areas in southeastern Brazil, where it may occur in sympatry with E. brasiliensis, E. iheringii, and E. bilineatus (Barreto-Lima, 2012). Many studies have addressed several aspects of the natural history of the species, including feeding ecology, activity patterns, microhabitat use, sexual dimorphism, and behavior (Barreto-Lima and Sousa, 2006, 2011, Sturaro and Silva, 2010Barreto-Lima et al., 2013;Migliore et al., 2014). Recently, Migliore et al. (2014) summarized the reproductive information available for E. perditus and E. iheringii and found that published data is limited to punctual observations on clutch size, courtship and mating behavior, and timing of mating and gravidity. ...
... Many studies have addressed several aspects of the natural history of the species, including feeding ecology, activity patterns, microhabitat use, sexual dimorphism, and behavior (Barreto-Lima and Sousa, 2006, 2011, Sturaro and Silva, 2010Barreto-Lima et al., 2013;Migliore et al., 2014). Recently, Migliore et al. (2014) summarized the reproductive information available for E. perditus and E. iheringii and found that published data is limited to punctual observations on clutch size, courtship and mating behavior, and timing of mating and gravidity. This limited information impairs both an overview on the reproduction of the species and broad comparisons across other species. ...
... These females were kept in terrarium containing branches and leaf litter and allowed to oviposit naturally. Procedures for egg measurements and incubation were used according to Migliore et al. (2014). ...
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Enyalius perditus is a semi-arboreal lizard species whose reproduction is poorly known. Here, we combine information obtained from preserved and live specimens to describe the reproductive timing (vitellogenesis, gravid-ity, and egg-laying) and fecundity (clutch size, egg size, and relative clutch mass) in females of E. perditus. Female reproduction is remarkably seasonal and occurs in the warmer and wetter periods of the year. Secondary vitellogenesis occurs from mid to late spring, whereas gravidity and egg-laying occur in early summer. Mating appears to be synchronized with secondary vitellogenesis, indicating an associated reproductive cycle. We suggest that E. perditus females produce only a single clutch per reproductive season. Clutch size ranged from three to 11 eggs and was positively correlated with female body size. Finally, the relative clutch mass was high, a recurrent feature to "sit-and-wait" foragers.
... Courtship and copulation behaviours of different species in the genus Enyalius may be similar, both in captivity (Grantsau, 1966;Barreto-Lima and Sousa, 2006) and under natural conditions (Vitt et al., 1996;Migliore et al., 2014;Novelli et al., 2015) (Table 1). In captivity (terraria), copulation in an E. catenatus pair (Grantsau, 1966), as well as copulations of E. perditus (Barreto-Lima and Sousa, 2006) were observed. ...
... In one instance, Vitt et al. (1996) documented a possible post-copulation of E. leechii, in which lizards were found sleeping in a posture similar to that observed in copulations of Polychrus acutirostris by Vitt and Lacher (1981). A second case involved courtship display without copulation success in E. perditus (Migliore et al., 2014). Also, courtship display followed by copulation in E. bilineatus on bare soil was reported, with the male biting and holding onto the female with its feet and toes (Novelli et al., 2015). ...
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Enyalius boulengeri Etheridge, 1969 occurs in the Atlantic Forest, in the states of Espírito Santo and Minas Gerais, in southeastern Brazil (Costa and Bérnils, 2018), and its conservation status has not yet been assessed, since basic information on its biology is still scarce or non-existent in the literature. We here describe for the first time a copulation of E. boulengeri under natural conditions, in a protected area in the state of Espírito Santo. In addition, we discuss the courtship and copulation behaviours of some other Enyalius species.
... ). Oviparous, no detailed data is known about the number of eggs laid by the species, but it could be similar to other Enyalius(Migliore et al. 2014).Type locality. State of Bahia, Brasil. ...
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The Caatinga is the largest seasonal dry tropical forest in South America and it has been historically neglected in terms of its bio- diversity. Regarding lizards, different studies led to the current knowledge of diversity and endemism in Caatinga, but detailed syntheses are scarce in the literature. We present the most detailed and up-to-date synthesis of knowledge about Caatinga lizards by providing a detailed (i) list of species; (ii) taxonomic richness patterns; (iii) knowledge gaps and spatial biases; and (iv) detailed distribution maps of all species that contain at least one occurrence record within Caatinga limits. We created a distribution database using occurrences of lizards in Caatinga based on scientific collections, field collection, and literature. We produce up-to-date dis- tribution maps, calculate the Extent of Occurrence and provide the environmental and bioclimatic profile for each species recorded. We draw taxonomic richness and sampling gap maps. Our database has 20,538 records of occurrence of lizards of the Caatinga. We recorded 93 lizard species (13 families), 52.7% of which are endemic. Forthy-four percent of the species present restricted distri- butions. We identified that 53% of the Caatinga area (or 70% of the municipalities) has no record of occurrence of lizards. The data presented are an important step towards synthesizing in detail the accumulated knowledge about Caatinga lizards and is crucial for accurate strategies for the conservation planning. It directs actions to advance our knowledge on Caatinga lizards: to concentrate inventories in sample void areas; continuous update of the species occurrence database, advance in the generation of autoecology data for species.
... Herpetological Bulletin 158 (2021) 21 The mating behaviour of T. atlantica is similar to that described for several skinks (e.g., Fitch, 1954;Evans, 1961) and unrelated lizards (Barreto-Lima & Sousa, 2006;Sánchez-Hernández et al., 2012;Migliore et al., 2014). Prolonged neck bites are likely to be advantageous and suggestive of mate guarding in other lizard species, including skinks (Bull, 2000). ...
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The skink Trachylepis atlantica is endemic to the Fernando de Noronha archipelago. Although this species is abundant in the archipelago, it is increasingly threatened by invasive species, predation, and anthropic interference. However, little is known about its natural history. Here we report on territory use, male-male combat, and mating behaviour of freeranging T. atlantica. During the dry season, we observed two bouts of male combat and two copulations, which suggests some mating seasonality. In male combat, T. atlantica displays ritualised agonistic behaviour with escalated aggressiveness that include - i) visualisation, approaching or following, ii) visual display, iii) bite and body confrontation, and iv) dominance. Adult males were always alert to invaders within their territories. After combat, resident males showed scars on their heads resulting from agonistic encounters. During the two copulations, we observed courtship, immobilisation, and mating.
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Studies on the natural history of the lizard Enyalius iheringii Boulenger, 1885, as well as other tropical lizards, are rare. In this study, some aspects of the natural history of this endemic species from the Atlantic forest are reported in areas of Vale do Itajai, state of Santa Catarina, Brazil. Twenty individuals were found, of which 18 were collected. Most of them were found over the vegetation (n= 17) and on the ground (n= 3). The main defensive strategy displayed was camouflage (n= 16). Jumping (n= 1), jumping and running (n= 1) and running (n= 2) were also observed in some individuals. When handled, lizards exhibited mouth wide open, hissing, and occasionally biting, as well as color change in males. Regarding its diet, the numerically most important prey was beetles (Coleoptera), followed by Lepidoptera larvae. Beetles, lepidopteran larvae and spiders were the most frequent food items. Males and females did not differ in size. Three sexually mature females (100-113 mm SVL) were found in December and January.
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ZOOCIÊNCIAS 8(2): 193-197 , dezembro 2006 Revista Brasileira de Scientific Note Court and copulation behaviors of Enyalius perditus Jackson, 1978 (Squamata, Leiosauridae) in captivity conditions 1 André Felipe Barreto Lima 2 & Bernadete Maria de Sousa 2 Enyalius Wied, 1821 is an arboreal lizard genus, diurnal, and insectivorous, which lives in old forests in the East Side of South America, Regions of Atlantic Forest, South of Amazon Forest, and Brazilian Cen-tral Region (ETHERIDGE 1969; VANZOLINI 1972; JACKSON 1978; ÁVILA-PIRES 1995; VITT et al. 1996). Almost nothing is known about the ecology of the lizards, the main information available for Enyalius species is restricted to account of feeding habits and microhabitat use (VANZOLINI 1972; JACKSON 1978; SAZIMA & HADDAD 1992; VITT et al. 1996; SOUSA 2000; ZAMPROGNO et al. 2001; VAN SLUYS et al. 2004). Enyalius perditus Jackson, 1978 is a small arboreal tropical lizard restricted to the forest areas, its history is unfamiliar (JACKSON 1978; SOUSA 2000). This lizard species, which is scarce and endemic in Brazilian East Coast, may be found in forests pertaining to the Atlantic Forest biome, in limited areas of Southeast (predominantly) and South regions of Brazil. This leiosaurid lizard occurs in the states of Rio de Janei-ro, São Paulo, Minas Gerais, and Paraná (JACKSON 1978; SOUSA 2000). However, little information is known and/or published on biological, ecological, and, mainly, behavioral aspects of this species, that can be considered insufficiently studied. Therefore this study aimed to elucidate the court and copulation 1
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We assessed the ecology, morphology and diet of Enyalius perditus. The data were compared among and within gender and collection periods. The specimens were collected in an anthropized biological reserve in the Atlantic forest biome, Brazil. The stomach contents were analyzed for number of prey, frequency, mass and volume. Sexual dimorphism occurred. No positive correlation between lizard size and volume of prey, nor between diet and collection period was found. Enyalius perditus diet was diverse comprising predominantly Formicidae, Orthoptera, Isoptera and other insect prey. In an urban forest, E. perditus was an opportunist predator of terrestrial arthropods. Differences in diet across gender were not found. Intersexual trophic similarities suggest there is no food resource partitioning in the population's diet studied herein.
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The present study was carried out to describe the diet, and its variations according to food availability, in Enyalius perditus Jackson, 1978 from Parque Estadual do Ibitipoca, State of Minas Gerais, Brazil. Three areas were chosen in the forest fragments; pit falls and glue traps were used to capture lizards and their preys. Lizards (n=55) were dissected in order to analyze the stomach content. The Electivity Index showed that insect larvae were volumetrically important in the diet of E. perditus, while ants and woodlices where numerically important. The quantity of diet items have differed significantly between males and females, perhaps, as a consequence of the higher quantity of ants ingested by female lizards and larvae ingested by males lizards. Based upon alimentary behavior, the male lizards resemble the active forragers and the females sit and wait forragers.
Thirteen species of iguanid lizards (excluding Anolis) from Brazil were examined. Species from the Amazonian region have fewer and larger eggs than do those from south of the Amazonian forests. These differences presumably have evolved because young lizards in tropical forests suffer less nonselective mortality than they do in other environments, attributed to the greater stability or predictability of the tropical forest environments. -from Author
Relative clutch mass (RCM) is an important life-history characteristic in lizards. Differential survivorship of females carrying clutches of eggs should partially determine RCM, and foraging mode and predator escape tactics play important roles in selection for a given RCM. RCM may vary within a population due to variation in resource availability or among populations of a species due to resource availability or geographic variation in escape tactics. Within species that forage widely, RCM is relatively low. RCM tends to be relatively high in species using the sit-and-wait foraging tactic. RCM variation in the latter group correlates with the manner in which crypsis is manifested. Where a streamlined morphology is advantageous (thin branch mimics or fossorial species), RCM may be low. Where crypsis does not involve a streamlined morphology, RCM tends to be high. Energetic costs associated with changes in RCM and variation in predation rates were estimated with a model of the daily net energy-intake of a lizard. Increasing RCM and/or predation intensity on gravid females reduces daily net energy-intake of wide foragers, but has little effect on species which are sit-and-wait foragers. -Authors