Behavioral description of the endangered lizard Liolaemus arambarensis in its natural Restinga environment through continuous filming

Abstract

Behavioral description is of key importance for the development of research and conservation projects for animals. Lizards are commonly classified by their foraging behavior, which can occur in a spectrum from “strictly ambush” to “wide-foraging”. Such characteristics are of taxonomic importance and play a crucial role in ecological function. Classic studies investigating foraging in lizards usually observe individuals in the field, but observer bias and other impacts may be poorly accounted for. We use surveillance cameras to describe the behavior of an endangered sand lizard. Continuous video was captured from 3 points within the study area on the dunes coasting the Patos Lagoon (Rio Grande do Sul, Brazil) for 48 days mostly during the hot season, which was screened for the appearance of the species, resulting in 7.35 hours of lizard footage. The behaviors performed were then identified and denoted. From this we observed that the individuals spent 90% of the time stationary, and in the remaining 10% they were found mostly dislocating, changing body posture or performing a sexual/territorial display. It was then possible to classify the species as a typical ambush predator. In addition, the predation footage obtained fit perfectly with that model. The recordings also showed other native vertebrate species as well as invasive wild-boars, which are a threat to the ecosystem. These findings inform the management and conservation needs of the Restinga habitat, that already faces many preservation challenges, and expand the prospects for future work both for the ethology and ecology of lizards.

Full text

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DOI: 10.55905/rdelosv17.n62-082
ISSN: 1988-5245
Originals received: 11/08/2024
Acceptance for publication: 12/02/2024
Revista DELOS, Curitiba, v.17, n.62, p. 01-19, 2024
Behavioral description of the endangered lizard Liolaemus arambarensis in
its natural Restinga environment through continuous filming
Descrição do comportamento do lagarto Liolaemus arambarensis, ameaçado
de extinção, em seu ambiente natural de restinga por meio de filmagem
contínua
Descripción del comportamiento de la lagartija en peligro de extinción
Liolaemus arambarensis en su entorno natural de Restinga mediante
filmación continua
Rafaela Kleinubing Gregol
Undergraduate Student in Biological Sciences
Institution: Universidade Federal do Rio Grande do Sul
Address: Porto Alegre - Rio Grande do Sul, Brasil
E-mail: rkgregol@gmail.com
Débora Newlands de Souza
Graduated in Biological Sciences
Institution: Universidade Federal do Rio Grande do Sul
Address: Porto Alegre - Rio Grande do Sul, Brasil
E-mail: debora.newlands@gmail.com
Laura Verrastro
Doctor in Ecology and Natural Resources
Institution: Universidade Federal de São Carlos
Address: Porto Alegre - Rio Grande do Sul, Brasil
E-mail: lauraver@ufrgs.br
ABSTRACT
Behavioral description is of key importance for the development of research and conservation
projects for animals. Lizards are commonly classified by their foraging behavior, which can
occur in a spectrum from “strictly ambush” to “wide-foraging”. Such characteristics are of
taxonomic importance and play a crucial role in ecological function. Classic studies investigating
foraging in lizards usually observe individuals in the field, but observer bias and other impacts
may be poorly accounted for. We use surveillance cameras to describe the behavior of an
endangered sand lizard. Continuous video was captured from 3 points within the study area on
the dunes coasting the Patos Lagoon (Rio Grande do Sul, Brazil) for 48 days mostly during the
hot season, which was screened for the appearance of the species, resulting in 7.35 hours of lizard
footage. The behaviors performed were then identified and denoted. From this we observed that
the individuals spent 90% of the time stationary, and in the remaining 10% they were found
mostly dislocating, changing body posture or performing a sexual/territorial display. It was then
possible to classify the species as a typical ambush predator. In addition, the predation footage
obtained fit perfectly with that model. The recordings also showed other native vertebrate species
as well as invasive wild-boars, which are a threat to the ecosystem. These findings inform the

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management and conservation needs of the Restinga habitat, that already faces many preservation
challenges, and expand the prospects for future work both for the ethology and ecology of lizards.
Keywords: behavioral description, endangered species, observer bias, sand lizard, novel
methodology, sit-and-wait.
RESUMO
A descrição do comportamento é de importância fundamental para o desenvolvimento de
projetos de pesquisa e conservação de animais. Os lagartos são comumente classificados por seu
comportamento de forrageamento, que pode ocorrer em um espectro que vai de “emboscada
estrita” a “forrageamento amplo”. Essas características são de importância taxonômica e
desempenham um papel crucial na função ecológica. Os estudos clássicos que investigam o
forrageamento em lagartos geralmente observam os indivíduos no campo, mas o viés do
observador e outros impactos podem não ser levados em conta. Usamos câmeras de vigilância
para descrever o comportamento de um lagarto de areia ameaçado de extinção. Foram capturados
vídeos contínuos de 3 pontos dentro da área de estudo nas dunas que margeiam a Lagoa dos Patos
(Rio Grande do Sul, Brasil) durante 48 dias, principalmente durante a estação quente, que foi
examinada quanto à aparência da espécie, resultando em 7,35 horas de filmagem do lagarto. Os
comportamentos executados foram então identificados e anotados. A partir disso, observamos
que os indivíduos passavam 90% do tempo parados e, nos 10% restantes, eram encontrados
principalmente se deslocando, mudando a postura corporal ou realizando uma exibição
sexual/territorial. Assim, foi possível classificar a espécie como um típico predador de
emboscada. Além disso, as imagens de predação obtidas se encaixam perfeitamente nesse
modelo. As gravações também mostraram outras espécies de vertebrados nativos, bem como
javalis invasores, que são uma ameaça ao ecossistema. Essas descobertas informam as
necessidades de gerenciamento e conservação do habitat da Restinga, que já enfrenta muitos
desafios de preservação, e ampliam as perspectivas de trabalhos futuros tanto para a etologia
quanto para a ecologia dos lagartos.
Palavras-chave: descrição comportamental, espécies ameaçadas de extinção, viés do
observador, lagarto da areia, nova metodologia, sit-and-wait.
RESUMEN
La descripción del comportamiento es de vital importancia para el desarrollo de proyectos de
investigación y conservación de animales. Las lagartijas se clasifican habitualmente por su
comportamiento de búsqueda de alimento, que puede darse en un espectro que va desde
«estrictamente emboscada» a «amplia búsqueda de alimento». Estas características tienen
importancia taxonómica y desempeñan un papel crucial en la función ecológica. Los estudios
clásicos que investigan la búsqueda de alimento en los lagartos suelen observar a los individuos
sobre el terreno, pero el sesgo del observador y otros impactos pueden estar mal contabilizados.
Utilizamos cámaras de vigilancia para describir el comportamiento de un lagarto de arena en
peligro de extinción. Se capturó vídeo continuo desde 3 puntos dentro del área de estudio en las
dunas que bordean la laguna de los Patos (Rio Grande do Sul, Brasil) durante 48 días, la mayoría
durante la estación cálida, en la que se examinó la aparición de la especie, lo que dio como
resultado 7,35 horas de metraje de lagartos. A continuación se identificaron y denotaron los
comportamientos realizados. A partir de esto observamos que los individuos pasaban el 90% del
tiempo inmóviles, y en el 10% restante se les encontraba principalmente dislocándose,

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cambiando de postura corporal o realizando un despliegue sexual/territorial. Así pues, fue posible
clasificar a la especie como un típico depredador de emboscada. Además, las grabaciones de
depredación obtenidas encajaban perfectamente con ese modelo. Las grabaciones también
mostraron otras especies de vertebrados autóctonos, así como jabalíes invasores, que constituyen
una amenaza para el ecosistema. Estos hallazgos informan sobre las necesidades de gestión y
conservación del hábitat de la Restinga, que ya se enfrenta a muchos retos de preservación, y
amplían las perspectivas de trabajo futuro tanto para la etología como para la ecología de los
lagartos.
Palabras clave: descripción del comportamiento, especies amenazadas, sesgo del observador,
lagarto de arena, metodología novedosa, sit-and-wait.
1 INTRODUCTION
Behavioral description is a useful tool for informing conservation initiatives for
endangered species, although small and inconspicuous species are disfavored in the literature
(Torr, 1994; Mesa-Avila & Molina-Borja, 2007; Festa-Bianchet & Apollonio, 2013; Breed &
Moore, 2021). For ethological and ecological studies to be well designed and analyzed,
understanding the basic behaviors of a species is crucial (Greenberg, 1978; Breed & Moore,
2021). Variations in the way lizards forage are indicative of the different ecological strategies
employed by species in different environments (Pough et al., 2006).
Foraging mode in lizards can be a determining factor of other behavioral patterns of a
species, and this characteristic is conserved within the different clades (Pianka & Vitt, 2003).
The two main types of foraging found in lizards are Active and Ambush (or sit-and-wait) (Perry,
1999). However, the feeding behavior of lizards can be nuanced, and some groups employ mixed
strategies (Pianka & Vitt, 2003; Cooper & Whiting, 1999). In addition, some species show
interpopulation variations in response to environmental factors (Wasiolka et al., 2009). Despite
this, statistical parameters have historically been used to assign a particular foraging mode to a
species (Perry, 2007; Bernstein et al., 1991). Perry (2007), following Pianka et al. (1979),
establishes two measures for quantifying foraging behavior in lizards: movements per minute
(MPM) and percentage of time in motion (PTM), and stresses the importance of observing
individuals for a sufficient amount of time (recommending at least ten minutes per individual).
However, statistical procedures can mask behavioral specificities of organisms, so a good prior
knowledge of the life history of the species studied is of great importance when evaluating

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foraging behavior, especially its period of activity and possible disturbances occurring during the
study period (Perry, 2007).
Observer effect is a variable that is difficult to quantify when it is not a research objective,
and is often considered null or ignored in the study of lizard behavior (Kerr, 2004; Fox et al.,
2003). In measuring foraging behavior, understanding the allocation of time by an animal is
essential, where, in addition to the fact that an ectothermic animal is rarely performing only one
activity (foraging, thermoregulation, territory defense, mate searching, etc.), the escape response
possibly triggered by the presence of researchers - in the field and in the laboratory - can be
significant in altering behavior, as demonstrated by Samia et al. (2015), Lion et al. (2012), Kerr
(2004) and Sugerman & Hacker (1980). The escape response and time spent in refuge by a lizard
in the wild may be related to the thermal condition of the environment and the individual, as well
as the presence of food/loss of the opportunity to feed (Cooper et al., 2004; Martín & López,
1999).
Liolaemus arambarensis (Verrastro, Veronese, Bujes & Martins Dias Filho, 2003) is a
sand lizard characterized by its extremely cryptic coloration and scale pattern in its environment,
as well as its association with thickets of grasses and shrubs, which it uses to perform various
vital functions (Verrastro et al., 2003). It is a small lizard, reaching a maximum rostro-cloacal
length of 60mm (Verrastro et al., 2003). The species distribution is restricted to areas of sand
dunes on the west coast of the Patos Lagoon, in the state of Rio Grande do Sul (Brazil), ensuring
its micro-endemic designation. Its small area of occurrence and the growing threats to its habitat
(anthropization and climate vulnerability) have classified the species as Endangered (EN)
(ICMBio, 2018; IUCN, 2014). In the year 2024, the entire area of distribution of the species was
hit by historical flooding, giving this and other works being developed about L. arambarensis, its
preservation and that of its coastal environment renewed importance. This work aims to describe
the behaviors performed by the species without observer effects, verify the foraging behavior of
L. arambarensis in the wild and report on the use of continuous recording cameras for the study
and behavioral description of sand lizards.

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2 THEORETICAL FRAMEWORK
Lion et al. (2012) showed that different observers caused different escape distances in
Tropidurus semitaeniatus (Tropiduridae). Putman et al. (2017) found that the color of the
researchers' clothing impacted the escape response and capture success in Sceloporus
occidentalis (Phrynosomatidae). Kerr (2004) studied the impacts of different levels of observer
presence and handling on the activity of Tiliqua rugosa (Scincidae), and reported that when the
lizard detected the researcher, after the initial response of remaining motionless, there was a 14
minute period of lower activity, followed by a period of higher activity, and that at all levels of
disturbance, the lizard's average activity levels remained higher for up to an hour, possibly
altering the allocation of time to vital functions, which may be significant for a species with few
hours of daily activity. The implications of these impacts on the study and quantification of
foraging behavior and mode are very little explored.
Perry (2007), in describing the methodology for quantifying foraging behavior, cites
several studies carried out with a variety of species whose data was obtained by observing
individuals in the field. Kerr (2004), when studying the observer effect, indicated that the use of
remote observation methods can prevent the impact of the researcher on behavior. More recent
studies have used video and photographic methods to observe and describe the habits of lizards
(Bertoia et al., 2023; Wester, 2019; Ebrahimi et al., 2015; Bennett, 2014). Regardless of the use
of Perry's (2007) statistical parameters, which were established for field observations, remote
monitoring of fauna enables a broad understanding that surpasses impacts caused by human
presence. In addition, the amount of time observed and the number of individuals can be much
greater, and further correlations can be drawn if other environmental data is available. Long-term
studies carried out using remote observation can provide important information to guide the
conservation of populations, making it possible to verify changes in the environment and human
presence over time (Ebrahimi et al., 2015).

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3 METHODOLOGY
3.1 STUDY AREA
The study was carried out in the dune area of the RPPN (Private Natural Patrimony
Reserve) Barba Negra (30°24'43“S and 51°13'03”W) (Figure 1), that belongs to Celulose
Riograndense LLC, in the municipality of Barra do Ribeiro, Rio Grande do Sul, one of the four
municipalities where the species occurs. The reserve is located between the Patos Lagoon and
the Guaíba River, with a sandy strip of land from the Quaternary period and restinga vegetation
(Celulose Riograndense, 2012).
Figure 1 - Satellite image showing the approximation of the Barba Negra RPPN area (30°24'43"S 51°13'03"W).
Adapted from Celulose Riograndense, 2018.

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3.2 DATA COLLECTION
Three security cameras were installed at different points in the dunes, near thickets of
herbaceous vegetation where the lizard was likely to be seen. The cameras were digital IP Onvif
cameras connected to a digital DVL recorder with a storage capacity of 1 terabyte. Two recorders
were used, which remained in the study area and were alternated periodically for image retrieval.
To obtain power, the equipment was connected to a 140-watt solar panel with a 150 ampere
stationary battery connected to a timer. Recordings were made in January, February, March and
June 2015, November and December 2016 and January, February and March 2017. The warm
season was chosen because it is the time when the species is most active. Image recording began
daily at 10h and ended at 18h. This time slot was chosen because L. arambarensis is diurnal,
with its peak activity around 12-14h (Verrastro et al., 2003). A total of 1304 hours were recorded
over 48 days within the study period.
3.3 DATA SORTING
The recordings were analyzed using Media Player Classic video viewing software. When
the lizard was observed, the section in which it appeared was cut out for analysis and the
behaviors performed were classified. We adapted the ethogram created by McElroy et al. (2011)
and Williams & McBrayer (2011) for classification (Table 1). In addition to the behaviors
performed by the individual, the date and time of the recording were also denoted. We used 160
unique recordings of L. arambarensis captured between 2016 and 2017 to represent the time slots
with the highest capture frequency.
Table 1 - Definitions of analyzed behaviors.
Behavior
Description
Move
A movement in which the lizard leaves its position
and moves to another point, covering a distance
greater than half its body length.
Pause
The lizard is motionless.
Head
The lizard moves its head horizontally while the rest
of its body remains still.

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Behavior
Description
Display
Territorial/sexual behavior in which the lizard is in a
fixed position, with forelegs extended and neck raised,
and/or when the head moves vertically quickly and
repeatedly.
Tail movement
Oscillatory or lifting movement of the lizard's tail,
with the rest of the body motionless.
Posture
Change in body posture without displacement.
Climb
Lizard climbs a vertical surface.
Jump
Jumping motion.
Attack
Attempt (successful or not) to capture and eat food.
Contact
Moment of contact between one individual and
another.
Substrate
Lizard rests the side of its head and/or tongue on the
substrate.
Other
Very rarely observed behaviors (≤0.1 minutes).
Source: The authors. Adapted from McElroy et al. (2011) and Williams & McBrayer (2011).
4 RESULTS AND DISCUSSIONS
A total of 313 clips were recorded in which the lizard was present. Individuals of
Liolaemus arambarensis were observed for approximately 7.35 hours (26457.76 seconds),
representing 0.563% of the total time screened. In addition to the lizard, other species of
vertebrate animals were also observed, such as Lycalopex gymnocercus (Pampa’s Fox/Graxaim-
do-Campo), Rhea americana (Greater rhea/Ema), Coendou spinosus (Hedgehog/Ouriço),
Salvator merianae (Argentine Tegu Lizard/Teiú) (Figure 2), Aspronema dorsivittatum (Paraguay
Mabuya/Scinco-dourado) and invasive species such as Sus scrofa (Boar/Javali), but never
simultaneously with L. arambarensis. Invertebrates from the orders Araneae, Coleoptera,
Lepidoptera and Odonata as well as from the Formicidae family, which make up the lizard’s diet,
were frequently seen alongside individuals (Schossler et al., 2000).
L. arambarensis was observed stationary (Pause behavior) most of the time, for a total of
23773.32 seconds (Figure 3). The frequencies of all the other behaviors described are shown in
Figure 4. The second most frequent behavior was displacement (6.31%). No individuals were

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observed while it rained. In the recordings made in the cold season (June 2015), the lizard was
only recorded for 5 seconds. The time slots with the highest number of recordings of the lizard
during the period analyzed are shown in Figure 5. Most of the recordings occurred between 12h
and 15h (50.6%), followed by those between 15h and 16h (16.8%). We provide a compilation of
clips of L. arambarensis and the identified behaviors obtained through this study through the
YouTube link https://youtu.be/lvR6IA4USdU .
Figure 2 - Recordings of A - Salvator merianae (Argentine tegu lizard), B - Coendou spinosus (Hedgehog), C -
Rhea americana (Greater rhea) and D - Sus scrofa (Boar), Barra do Ribeiro/RS, Brazil (30°24’43”S 51°13’03”W).
Source: The authors.

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Figure 3 - Total recording time of Liolaemus arambarensis individuals and time the lizard was stationary.
Source: The authors.
Figure 4 - Frequency of further analyzed behaviors of Liolameus arambarensis.
Source: The authors.

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Figure 5 - Number of unique recordings of L. arambarensis individuals in 2016 and 2017 by time slot, where each
column represents a 60-minute period, starting at 10:00h.
Source: The authors.
We recorded multiple attacks on prey, where the lizard performed this behavior in a
typical ambush fashion, waiting still for the most opportune moment when its prey is close
enough to capture. We were able to observe jump attacks (usually with flying prey) (Figure 6)
and attacks that consisted only of head movements and/or minimal movement to reach the
arthropod (Figure 7). The species is known to seldom feed on plant matter (Schossler et al.,
2000), so climbing the vegetation (Climb behavior) may be associated with the obtention of food
items.
Figure 6 - Jumping attack, Liolameus arambarensis, 19/02/2015, 30°24’43”S 51°13’03”W.
Source: The authors.

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Figure 7 - Attack with small displacement, Liolameus arambarensis, 23/02/2015, 30°24’43”S 51°13’03”W.
Source: The authors.
The results obtained, both numerically quantifiable and the individual observations, are
in line with what Verrastro et al. (2003) stipulated about the species in their description:
Liolaemus arambarensis is a lizard with strictly Ambush predation. This characteristic is in line
with what has been described for the Liolaemidae family and the Iguania clade (Verrastro & Ely,
2015; Huey & Pianka, 1981). All the predation events recorded fit perfectly into the “sit-and-
wait” model previously described, also corroborated by the fact that the individuals observed
spend most of their time standing still (Perry, 1995).
One function that is constant in lizards (and ectothermic animals in general) is
thermoregulation. We can say that the decision to remain in one spot is in line with the thermal
resource being offered at that moment, especially in the case of a heliothermic species such as L.
arambarensis (Vicente Liz et al., 2019). It was possible to observe lizards exposed in the sun,
refuged in the shade of vegetation and moving between these states. McConnachie et al. (2009)
states that, in a “sit-and-wait” lizard, the posture performed by the stationary lizard is indicative
of function: postures with the head elevated would be associated with the search for prey, and
postures with the head lowered would be related to thermoregulatory needs. Although the time
spent changing posture and moving the head were statistically small (since they are short
movements), they were very frequent occurrences, pointing to the multiplicity of functions
performed while standing still.

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We describe the locomotion of L. arambarensis as sprint or short-bursts, common in
ambush lizards, where they move in short bursts of running, rather than, for example, a long,
steady stride. Bertoia et al. (2023), also using monitoring cameras, demonstrated that this rapid
form of locomotion may be crucial for maintaining adequate body temperature. In addition, a
morphological aspect linked to the behavioral frequencies found is the cryptic coloration of L.
arambarensis. Verwaijen & Van Damme (2008) point out that cryptic morphology allows lizards
to depend on their non-detection by predators and prey in order to succeed in their feeding
strategy - the condition of cryptic morphology in conjunction with an ambush strategy being very
frequent among lizards. In order to cover relatively large distances, the lizard will make several
short runs interspersed with some time standing still. The escape response of L. arambarensis is
also short sprints to some vegetation that serves as a refuge, where it can often bury itself or just
remain motionless, relying on its almost identical coloration to the substrate. Notably said
response has been observed in field surveys, not in the present work. Garland (1999) associates
this condition with the “sit-and-wait mode” of foraging, since it is also necessary to make quick,
explosive movements when capturing prey.
The high number of recordings up to two hours beyond the peak activity time previously
described for the species by Verrastro et al. (2003) may be related to seasonality, since the
recordings were made during the hot season, justifying high levels of activity.
The exotic and invasive wild boar, Sus scrofa, represents a strong threat to biodiversity
in Rio Grande do Sul (Sordi & Lewgoy, 2017). The record of its occurrence, especially within a
protected area, alongside a species considered Endangered, serves as a basis for adapting
management so that it can be suppressed. The wild boar is a risk because it moves in large flocks
that trample the ground and destroy the herbaceous vegetation, which is closely related to the life
history of Liolaemus arambarensis.
When considering the STRANGE framework for behavioral studies (Webster & Rutz,
2020), we believe that the present methodology avoided most of the issues presented as
generating interference in animal behavior, since this study did not involve the handling of
individuals or any aspect that would influence microhabitat selection by the lizard. Santos (2019)
showed that Liolaemus arambarensis has a large area of use (in relation to its body size), with
males and females exceeding 560m2 in the breeding season, as well as having strongly
overlapping areas of occupation and being very abundant in its habitat. This information, in

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company with the low rate of lizard recordings over the study period, discredits the possibility
of pseudoreplication (i.e. recording the same individual repeatedly). However, it is likely that the
results were influenced by seasonality, since in colder periods, rates of activity, displacement and
sexual behavior may be intensely reduced.
5 CONCLUSION
Through the filming we were able to record various behaviors performed by the species
and observe the foraging and feeding of Liolaemus arambarensis, which we classify as strictly
Ambush, also corroborated by the frequency of the behaviors analyzed. The predominance of
Pause concurs with the designation of multiple functions performed by the lizard while immobile,
mainly thermoregulation and lurking for prey. The second most frequent behavior was
displacement, described as short-bursts, which is also associated with feeding behavior and
thermoregulation. The results are in line with those expected for the Liolaemidae family.
Ultimately, we hope to encourage the use of natural-habitat filming for behavioral studies
and expand the discussion on ethological methodology. In light of the recent climate catastrophe
that impacted the entire range of this already Endangered species, and the record of an invasive
threat, our work shines a spotlight on the fauna of the Restinga environment and mounts the
evidence for due importance to be given to maintaining it, both for drawing up conservation
guidelines and in environmental education endeavors geared towards local populations, which
are in direct contact with the species and need to be its ally if we want to see it survive and thrive.
ACKNOWLEDGEMENTS
We thank all of our laboratory peers who aided in the production of this study by providing field
and in-lab support to the authors.

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