Herpetology Notes, volume 11: 929-941 (2018) (published online on 11 November 2018)

Herpetofauna of protected areas in the Caatinga VII: Aiuaba

Ecological Station (Ceará, Brazil)

Taís Borges Costa1, Daniel Orsi Laranjeiras1, Francis Luiz Santos Caldas 2, Daniel Oliveira Santana1,
Cristiana Ferreira da Silva4, Edna Paulino de Alcântara4, Samuel Vieira Brito1, Jéssica Yara Galdino1, Daniel
Oliveira Mesquita1, Renato Gomes Faria2, Frederico Gustavo Rodrigues França3, Robson Waldemar Ávila4,
and Adrian Antonio Garda5,*

Abstract. We provide a list of amphibians and reptiles recorded in a strict protection area in the Brazilian semiarid Caatinga,
the Aiuaba Ecological Station, Ceará State, Brazil. We conducted herpetofauna surveys totaling 110 days of fieldwork using
pitfall trap arrays, glue traps, and active searches. We recorded 57 species distributed in 23 families: 16 lizards, 1 amphisbaena,
17 snakes, 20 amphibians, and 3 chelonians. Rarefaction curves for snakes and all herpetofauna combined suggest that species
richness may still be underestimated for the area, while amphibian and lizard species numbers are similar to values suggested
by diversity estimators. Despite its relatively small area, the Aiuaba Ecological Station serves as a herpetofaunal refuge,
protecting a representative set of Caatinga species.

Keywords. Anura, Squamate reptiles, chelonians, distribution, conservation

Introduction The Caatinga originally covered 10% of the Brazilian

territory, but today over 60% of its area is covered by
The Caatinga biome is the largest patch of Seasonally
anthropogenic ecosystems (Silva et al., 2017). The
Dry Tropical Forests (STDF), a highly fragmented
biome comprises the states of Ceará, most of Paraíba,
biome widespread from Mexico to Argentina (Prado,
Rio Grande do Norte and Pernambuco, southeastern
2000). Protected areas in the Caatinga encompass 7.4%
Piauí, western Alagoas and Sergipe, northern and
of the biome (Silva et al., 2017), but most are among the
least restrictive category in Brazil, known as APA (Área
de Proteção Ambiental, or Environmental Protection
Area), and all are critically underfunded (Oliveira and
Bernard, 2017). Moreover, the Caatinga is also the least
protected of Brazilian biomes, with under 2% of its 1
Universidade Federal da Paraíba, Centro de Ciências Exatas
territory inserted in strict protection areas (Fonseca et e da Natureza, Departamento de Sistemática e Ecologia,
al., 2017). Cidade Universitária, 58051-000, João Pessoa, PB, Brasil.
For a long time, the Caatinga’s fauna and flora were
2
Universidade Federal de Sergipe, Departamento de Biologia,
Laboratório de Cordados (Herpetologia/Ictiologia), Cidade
considered poor and with few endemic species and,
Universitária, Av. Marechal Rondon, s/n, Jardim Rosa Elze,
hence, given low priority for conservation (Vanzolini, 49100-000, São Cristóvão, SE, Brasil.
1976). This first impression resulted from insufficient 3
Universidade Federal da Paraíba, Centro de Ciências
inventories and was gradually abandoned as recent Aplicadas e Educação, Departamento de Engenharia e Meio
studies reported much higher species richness and Ambiente, 58297-000, Rio Tinto, PB, Brasil.
significant rates of endemism (Araujo and Silva, 2017; 4
Universidade Regional do Cariri, Departamento de Ciências
Carmignotto and Astúa, 2017; Garda et al., 2017; Lima Físicas e Biológicas. Rua Cel. Antonio Luiz Pimenta,
63105100, Crato, CE, Brasil.
et al., 2017; Mesquita et al., 2017). Nevertheless, the 5
Universidade Federal do Rio Grande do Norte, Laboratório
Caatinga is still neglected, receiving less financial
de Anfíbios e Répteis-LAR, Departamento de Botânica,
support, less scientific and conservation initiatives, Ecologia e Zoologia, Centro de Biociências, Campus
and having fewer research groups compared with other Universitário, Lagoa Nova,59078-900, Natal, RN, Brasil.
biomes in Brazil (Santos et al., 2011). *
Corresponding author. E-mail: pseudis@gmail.com
930
Figure 1. Aiuaba Ecological Station, Ceará, Brazil. The
numbers 1 to 12 refer to the sampling points. Legend: 1 = first
set of pitfall traps; 2, 8, 10, 11, 12 = active search points; 7 =
rock outcrop area near the dam; 3 = second set of pitfalls; 4
= third set of pitfalls; 5 = fourth set of pitfalls; 6 = fifth set of
pitfalls; 9 = protected area head office.
central Bahia, and northern Minas Gerais (Prado, 2003).
It is one of the most degraded biomes in Brazil (Leal
et al., 2005), and most of the local economy depends
on public services, with the lowest human development
indicators in the country (Silva et al., 2017)�.
For a long time, the scarcity of inventories with
adequate description of sampling efforts and comparable
methodologies in the Caatinga contributed to the lack
of knowledge about its herpetofauna (Rodrigues, 2003;
Albuquerque et al., 2012; Guedes et al., 2014)�������me
standardized inventories were recently conducted
in protected areas like Estação Ecológica Raso da
Catarina-BA (Garda et al., 2013), Parque Nacional da
Serra da Capivara-PI (Cavalcanti et al., 2014), Parque
Nacional do Catimbau-PE (Pedrosa et al., 2014),
Estação Ecológica do Seridó-RN (Caldas et al., 2016),
and Parque Nacional da Chapada da Diamantina-BA
(Magalhães et al., 2015).
These studies helped improve comparisons between
the herpetofaunas in different regions in the biome,
besides contributing directly to the knowledge of
Protected Area’s amphibian and reptile faunas. These
concerted efforts, along with other studies providing
lists and distribution data for many localities (Loebmann
and Haddad, 2010; Moura et al., 2010; Roberto et al.,
2013; Roberto and Loebmann, 2016), have allowed a
recent re-evaluation of the geographic distribution of
Taís Borges Costa et al.
amphibian and reptile species richness in the Caatinga
(Garda et al., 2017; Mesquita et al., 2017). Still, many
areas of the biome are understudied, and knowledge
gaps can hamper conservation strategies and resource
use prioritization.
The present study integrates the results of three
independent field surveys����������������������������
list of herpetofauna species for Aiuaba Ecological Station
in Ceará State, Brazil. Our results show a high diversity
of frogs and reptiles in a lowland area of Caatinga that
now likely represents one of the best sampled areas for
amphibian and reptiles in the biome.
Material and Methods
Study area.—The Aiuaba Ecological Station is a strict
use protection area located in Aiuaba Municipality,
Ceará State, Brazil (6° 35’ 58.1”S, 40° 07’ 20.5”W; Fig.
1). It encompasses 11,525 hectares between latitudes
06° 35’ and 06° 40’S and longitudes 40° 07’ and 40°
20’W. It is inserted in the “Sertão dos Inhamuns” and
“Depressão Sertaneja Setentrional”, presenting litholic
soils and flat reliefs where erosive formations are
more frequent and phyllites are the predominant rocks
(Velloso et al., 2002; Medeiros, 2004). The area is also
predominantly covered by arboreal Caatinga, although
presenting other phytophysiognomies like carrasco,
dense arboreal Caatinga, and arboreal-shrubby Caatinga
complex (Oliveira et al., 1983; Medeiros, 2004). The
rainy period usually occurs from October to April, with
mean annual precipitation ranging from 590 to 684 mm
and mean annual temperatures from 25.3°C to 25.9°C
(Velloso et al., 2002; Medeiros, 2004).
Sampling methods.—We conducted surveys during
different periods of the dry and rainy seasons (Table 1),
totaling 110 days of fieldwork. We used a combination
of different sampling methods, including active
searches, occasional encounters, pitfall traps with drift
fences, funnel traps, and glue traps. Funnel and glue
traps were implemented only in the September/October
2013 survey, while the remaining methods were used in
all surveys (Table 1).
Survey I— One of us (SVB) conducted two 10-day
surveys in the dry (October-November) and rainy (April-
May) seasons (Table 1), using 120 pitfall traps arranged
in Y shape connected with 5m plastic drift fences and
equally distributed in 3 areas (40 traps pear area). Day
and night actives searches were also conducted by two
researchers to complement trap captures.
Survey II— One of us (RWA) coordinated eight 5-day
surveys using a standardized protocol of the Biodiversity
Herpetofauna of protected areas in the Caatinga VII: Aiuaba Ecological Station 931
Table 1. Sampling
Table 1.procedures used in three
Sampling procedures usedseparate
in threefield surveys
separate fieldfor herpetofauna
surveys species at
for herpetofauna the Aiuaba
species Ecological Station,
at the Aiuaba
Ceará State,Ecological Station,
Brazil. Details Ceará State,
on duration, Brazil.methods,
sampling Details on duration,
season, andsampling
traps usedmethods,
are givenseason, andsurvey.
for each traps used are given
for each survey.

Survey Year Month Days in Sampling Methods Number and Season

field volume of traps
I 2010 October 10 pitfall traps with drift 30 arrays (four 30L- Dry season
fences buckets/array) Y
shape
I 2011 April 10 pitfall traps with drift 30 arrays (four 30L- Rainy
fences buckets/array) Y season
shape
II 2012 June and 5+5 pitfall traps with drift 10 arrays (eight 60L Rainy/dry
September fences, visual searches buckets/array) season
straight line
III 2013 September 30 pitfall traps with drift 37 arrays, (4 buckets, Dry season
and fences, visual searches, 60L, per array), Y
October hoop traps and glue shape
traps,
II 2013 January 5+5 pitfall traps with drift 10 arrays (eight 60L Rainy
and April fences, visual searches buckets/array) season
straight line
II 2013 June 5 pitfall traps with drift 10 arrays (eight 60L Rainy/dry
fences, visual searches buckets/array) season
straight line
II 2014 February 5 pitfall traps with drift 10 arrays (eight 60L Rainy
fences, visual searches buckets/array) season
straight line
II 2014 May and 5+5 pitfall traps with drift 10 arrays (eight 60L Rainy/dry
June fences, visual searches buckets/array) season
straight line

Table 2. Species richness estimates for amphibians, lizards, snakes and all herpetofauna combined for Aiuaba
Ecological Station using different estimators and the individual-based rarefaction methods.
Research Program from the Brazilian government arrays), in the driest area above the dam (10 arrays) and
(PPBio, see Magnusson et al., 2005). Ten plots (250 Mean near ±a Standard
hilly areaDeviation
(20 arrays). Our sampling points were
x 1 m, along the same elevational
Estimators Lizards similar to those
isocline) were
Amphibians of surveys I andHerpetofauna
Snakes III because of logistics
sampled using
ACEvisual searches during 16 ± 0 (road accessibility)
20 ± 0 both the diurnal 21.34 ± 0 and because all ±physiognomies
61.28 0 of
and nocturnal period. At the end of each sampling plot, the protected area are represented in these points (Fig.
CHAO 1 20 ± 0.14 16 ± 0.5 19.45 ± 2.85 58.62 ± 2.83
80 pitfall traps with drift fences consisting of eight 2). We conducted day and night active searches, starting
Jacknife1 22 ± 1 16 ± 0 22.88 ± 2.27 63 ± 2.64
buried buckets (60 L), 5 m apart in a straight line were 8:00 am and ending about 10:00 pm, avoiding warmest
installed. Jacknife2 20 ± 0 14 ± 0 hours of the23.94 ±0
day (11:00 63 ±pm).
am to 2:00 0 We surveyed as
CHAO1P 20 16 21.87 59.54
Survey III—A total of 37 pitfall trap arrays and 148 many physiognomies and habitats as possible, according
glue traps were
% deinstalled in the protected
Singletons 0 area in an effort
0 to the availability
35.3 of roads and12.5 logistic limitations. We
coordinated by AAG for 30 consecutive days. Each inspected microhabitats for herpetofauna (fallen logs,
trap array consisted of four buried buckets disposed bushes, rocks and litter) as well as the areas near pitfall
in a Y shape with 120º angles among 6-meter drift arrays, trails near hills and some areas in the buffer
fences connecting the central bucket to buckets at the zone of the protected area. As an additional method to
extremities. Within the 6-meter radius around the central collect chelonians we placed three hoop-traps (60 cm in
bucket we placed four glue traps: two on the ground, diameter and 2 m long) in the dam during five days with
above rocks and/or fallen tree trunks, and two on tree sardines as bait.
branches and trunks (Table 1, Fig. 2). We distributed Amphibians were euthanized with lidocaine (topic
trap arrays along three main tracks: near a dam (seven 5% cream) and reptiles with a lethal dose of Tiopental®,
932 Taís Borges Costa et al.

Figure 2. Sampled areas and traps used in the Aiuaba Ecological Station, Ceará, Brazil. A and B = shrubby caatinga associated
with rocky soils; C = dam; D = pitfall trap array; E = glue trap on fallen log; F = glue trap on tree trunk.

preserved in 10% formalin and stored in 70% ethanol.
Specimen collection was authorized through federal
permits issued to AAG (SISBIO #32575-1), FLSC
(SISBIO #36095-8), RWA (SISBIO #32758-2) and
TBC (SISBIO #29550-4). Voucher specimens were
housed in the herpetological collections of the Federal
University of Paraíba (CHUFPB) and the herpetological
collection of the Regional University of Cariri (URCA-
H).
Data analysis.—We constructed rarefaction curves
to evaluate species richness sampling efficacy.
These curves were estimated based on the number of
individual frogs, lizards, snakes and for the whole
Herpetofauna of protected areas in the Caatinga VII: Aiuaba Ecological Station
herpetofauna combined (Gotelli and Colwell, 2001).
Using the software ESTIMATES 9.1.0 (Colwell,
2013), we obtained species richness estimators based
on abundance data (CHAO 1, CHAO1P and ACE)
and based on nonparametric incidence (CHAO 2 and
JACKKNIFE 1 and 2) to determine the expected
richness of amphibians, reptiles and herpetofauna as a
whole. We conducted 1,000 randomizations without
replacement with the original data.
Results
We documented 2,732 individuals from 57 species
distributed in 23 families: 16 lizards, 1 amphisbaenian,
17 snakes, 3 chelonians and 20 amphibians (Appendix
1; Fig. 3, 4, 5). The rarefaction curves did not reach
the asymptote for snakes or for all the herpetofauna
combined, evidencing that more species still can
be added to the list as sampling effort increases.
Considering lizards and amphibians, rarefaction curves
reached the asymptote. The richness estimators ranged
between 20 and 22 species of amphibians and around
16 species of lizards close to the total recorded by us
(Table 2, Fig. 6).
Discussion
Amphibian and lizard species richness recorded in
Aiuaba Ecological Station are similar to other Caatinga
areas recorded in most lowland caatinga areas (e.g.
Moura et al., 2010; Garda et al., 2013; Roberto et al.,
2013; Roberto and Loebmann, 2016). In contrast, we
recovered more species of snakes at Aiuaba Ecological
Station than most other areas reported in the literature,
approaching richness values from areas with long term
surveys (Mesquita et al., 2013). This likely results from
the significant field effort we employed, compared to
short term inventories, as snakes are more secretive and
demand more intensive survey efforts to be appropriately
recorded. Nevertheless, simulations suggest that the
number of species we recorded is still underestimated
(Fig. 7), showing that snakes remain elusive even after
large efforts dedicated to survey a single area.
By combining more than one sampling method
species with different ecological requirements were
captured. Five amphibians, four snakes, and one turtle
were found only in occasional encounters. Three lizards
and four snakes were recorded only in active searches,
and one frog species was recorded solely in pitfall traps.
This reinforces the use of a combination of different
sample methodologies, because some species were
933
recorded by a single method. Indeed, active searches
and occasional encounters are efficient in collecting
fossorial or less abundant species like snakes, turtles,
and amphisbaenians, or even some amphibians, what is
corroborated by other studies conducted in the Caatinga
(Magalhães et al., 2013; Cavalcanti et al., 2014; Pedrosa
et al., 2014). Some species considered common for the
Caatinga, like Leptodactylus fuscus and Mabuya heathi,
were only recorded by us through active searches and
occasional encounters, what is in some ways counter
intuitive, but may be explained by one of two hypotheses.
First, during the time our inventories were conducted
the Caatinga underwent one of the most drastic droughts
ever recorded, with reduced rainfalls even during the
rainy seasons (Marengo et al., 2016). Second, regional
variation in species abundances and detectability
are only starting to be unveiled in the Caatinga. It is
possible that this severe drought and/or such unknown
regional patterns of abundances and detectability were
responsible for such otherwise common species in the
Caatinga being found only through active searches and
occasional encounters.
The Caatinga biome is characterized by seasonal,
temporally spaced and unpredictable periods of rainfall
(Ab’sáber, 2003; Prado, 2003). Because amphibians are
significantly associated with humidity, they are more
active during the rainy season and are frequently found
buried or inactive during dry seasons (Warburg, 1997;
Navas et al., 2004). Therefore, it is paramount that field
surveys targeting frogs in the Caatinga be conducted
during rainy seasons. Indeed, field work during rainy
periods has recovered much higher richness for this
group in the Caatinga (Caldas et al., 2016), even in
rapid inventories of a few days after the first heavy rains
(Santana et al., 2015).
Accordingly, most species of amphibians in the present
survey were registered during the rainy season, even with
scant rains because of the severe draught the region was
facing. These include common species for the Caatinga,
like Pleurodema diplolister and Physalaemus spp., but
also some less-common taxa. Proceratophrys aridus,
for example, whose distribution is known only from the
type locality (Milagres Municipality, Ceará State; Cruz
et al., 2012), was recorded, extending its distribution
about 200 km west. Elachistocleis piauienses, previously
known only from the states of Ceará, Maranhão, and
Piauí (Frost, 2018), and Pseudopaludicola pocoto,
recently described for the Caatinga (Magalhães et al.,
2014) are amongst the most abundant amphibians in this
part of the Caatinga and were also frequently recorded
934 Taís Borges Costa et al.

Figure 3. Amphibians collected in Aiuaba Ecological Station, Ceará, Brazil. A - Rhinella granulosa; B - Rhinella jimi; C –
Dendropsophus nanus; D - Dendropsophus soaresi; E - Boana raniceps; F - Phyllomedusa nordestina; G - Scinax x-signatus;
H - Proceratophrys aridus; I – Dermatonotus muelleri (juvenile); J - Elachistocleis piauiensis; K - Leptodactylus chaquensis; L
- Leptodactylus troglodytes; M - Leptodactylus vastus; N - Physalaemus albifrons; O - Pseudopaludicola pocoto.

in the protected area during our inventories.
The most abundant lizard species in the area are widely
distributed in the Caatinga, such as Tropidurus hispidus,
Gymnodactylus geckoides, Ameivula ocellifera,
Vanzosaura multiscutata, and Phyllopezus pollicaris
(Werneck et al., 2012; Oliveira et al., 2015; Mesquita
et al., 2017). Coleodactylus meridionalis, however, has
a relictual distribution in the Caatinga (Mesquita et al.,
2017), being typically found in forested habitats and
mesic enclaves (but see Ribeiro et al., 2013). This first
Herpetofauna of protected areas in the Caatinga VII: Aiuaba Ecological Station 935

Figure 4. Species of lizards, and amphisbaenians recorded in Aiuaba Ecological Station, Ceará, Brazil. A - Phyllopezus pollicaris;
B - Hemidactylus agrius; C - Lygodactylus klugei; D – Gymnodactylus geckoides; E - Coleodactylus meridionalis; F - Mabuya
heathi; G - Tropidurus hispidus; H – Tropidurus jaguaribanus; I – Micrablepharus maximiliani; J – Vanzosaura multiscutata; K
– Ameiva ameiva; L – Ameivula ocellifera; M – Iguana iguana; N – Polychrus acutirostris; O – Amphisbaena vermicularis.

report of C. meridionalis outside mesic enclaves or areas
under their influence in Ceará state shows that, despite
having a relictual distribution and a very small body
size, the species seems to tolerate climates otherwise
too harsh for mesic forest species. The second most
abundant lizard, Tropidurus jaguaribanus, is distributed
along the Jaguaribe river valley (Piauí and Ceará states)
and is endemic to the Caatinga biome (Passos et al.,
2011).
The Aiuaba Ecological Station presents one of the
936 Taís Borges Costa et al.

Figure 5. Snakes and chelonians recorded in Aiuaba Ecological Station, Ceará, Brazil. A - Epictia borapeliotes; B - Boa
constrictor; C - Leptodeira annulata; D - Erythrolamprus viridis; E - Erythrolamprus poecilogyrus; F - Leptophis ahaetulla;
G - Oxyrhopus trigeminus; H - Philodryas nattereri; I - Pseudoboa nigra; J - Micrurus ibiboboca; K - Crotalus durissus; L
- Phrynops geoffroanus.

richest herpetofauna reported for a Caatinga strictu
sensu site. Some areas, including sites in Ceará state,
have higher overall richness, but include many species
typical of mesic enclaves. For the Ibiapaba-Araripe
complex, for example, 38 amphibian and 84 reptile
species have been reported. However, this region has at
least five different phytophysiognomies, including areas
of Cerrado (savannas), Caatinga, and rainforest enclaves
(Loebmann and Haddad, 2010). Fifty-seven amphibians
and 126 reptiles are recognized throughout the state of
Ceará, occurring in many distinct phytophysiognomies
and under the influence of neighboring biomes, such
as the Amazon Forest and Cerrado (Roberto and
Loebmann, 2016).
The Aiuaba Ecological Station was recognized a
priority area for conservation in the Caatinga biome
but, until now, no survey reporting amphibian and
reptiles existed for the area. We hope the species list
 fences, visual searches buckets/array) season
straight line
II 2014 May and 5+5 pitfall traps with drift
10 arrays (eight 60L Rainy/dry
June fences, visual searches
buckets/array) season
Herpetofauna of protected areas in the Caatinga VII: Aiuaba Ecological
straightStation
line 937
Table 2. Species richness estimates for amphibians, lizards, snakes and all herpetofauna combined for Aiuaba Ecological Station
using different estimators
Table andrichness
2. Species the individual-based rarefactionlizards,
estimates for amphibians, methods.
snakes and all herpetofauna combined for Aiuaba
Ecological Station using different estimators and the individual-based rarefaction methods.

Mean ± Standard Deviation

Estimators Amphibians Lizards Snakes Herpetofauna
ACE 20 ± 0 16 ± 0 21.34 ± 0 61.28 ± 0
CHAO 1 20 ± 0.14 16 ± 0.5 19.45 ± 2.85 58.62 ± 2.83
Jacknife1 22 ± 1 16 ± 0 22.88 ± 2.27 63 ± 2.64
Jacknife2 20 ± 0 14 ± 0 23.94 ± 0 63 ± 0
CHAO1P 20 16 21.87 59.54
% de Singletons 0 0 35.3 12.5

we provide will assist the planning and implementation describe the areas local richness and effectively protect
of conservation actions in this area. An eventual its biodiversity. Also, areas outside the sites we sampled
management plan should aim to increase sampling for demand investigation, as our surveys were concentrated
some groups, especially snakes, in order to adequately in areas near roads and protected area’s facilities.

Figure 6. Species rarefaction curves based on abundance data for Amphibians, Lizards, and Snakes collected at Aiuaba Ecological
Station, Ceará, Brazil.
938
Acknowledgements. Many students, protected area’s staff
members, and researchers helped during field work, and we are in
debt with their kind support and effort. We specifically thank the
crew of ESEC AIUABA for logistic and personnel support during
our fieldwork. TBC thanks to CAPES for her PhD fellowship.
This work was supported by grants from CNPq/ICMBio to AAG
(Grant #552031/2011-9) and RWA (Grant #551993/2011-1).
References
Ab’sáber, A.N. (2003): Os Domínios de Natureza no Brasil:
Potencialidades Paisagísticas, São Paulo, Ateliê Editorial.
Albuquerque, U.P., Araújo, E.L., El-Deir, A.C., De Lima, A.L.,
Souto, A., Bezerra, B.M., et al. (2012): Caatinga revisited:
ecology and conservation of an important seasonal dry forest.
The Scientific World Journal 2012: 205182.
Araujo, H.F.P., Silva, J.M.C. (2017): The Avifauna of the Caatinga:
Biogeography, Ecology, and Conservation. In: Biodiversity,
Ecosystems Services and Sustainable Development in Caatinga:
The Largest Tropical Dry Forest Region in South America, p.
181–210. Silva, J.M.C., Leal, I. , Tabarelli, M., Ed., Berlin,
Springer-Verlag.
Caldas, F.L.S., Costa, T.B., Laranjeiras, D.O., Mesquita, D.O.,
Garda, A.A. (2016): Herpetofauna of protected areas in the
Caatinga V: Seridó Ecological Station (Rio Grande do Norte,
Brazil). Check List 12: 1929.
Carmignotto, A.P., Astúa, D. (2017): Mammals of the Caatinga:
Diversity, Ecology, Biogeography, and Conservation. In:
Biodiversity, Ecosystems Services and Sustainable Development
in Caatinga: The Largest Tropical Dry Forest Region in South
America, p. 211–254. Silva, J.M.C.D., Leal, I. , Tabarelli, M.,
Ed., Berlin, Springer-Verlag.
Cavalcanti, L.B.Q., Costa, T.B., Colli, G.R., Costa, G.C., França,
F.G.R., Mesquita, D.O., et al. (2014): Herpetofauna of protected
areas in the Caatinga II: Serra da Capivara National Park. Check
List 10: 18–27.
Colwell, R.K. (2013): EstimateS: Statistical Estimation of Species
Richness and Shared Species from Samples. University of
Connecticut, USA.
Cruz, C.A.G., Nunes, I., Juncá, F.A. (2012): Redescription of
Proceratophrys cristiceps (Müller, 1883)(Amphibia, Anura,
Odontophrynidae), with description of two new species without
eyelid appendages from Northeastern Brazil. South American
Journal of Herpetology 7: 110–122.
Fonseca, C.R., Antongiovanni, M., Matsumoto, M., Bernard, E.,
Venticinque, E.M. (2017): Conservation Opportunities in the
Caatinga. In: Biodiversity, Ecosystems Services and Sustainable
Development in Caatinga: The Largest Tropical Dry Forest
Region in South America, p. 429–443. Silva, J.M.C., Leal, I. ,
Tabarelli, M., Ed., Berlin, Springer-Verlag.
Garda, A.A., Costa, T.B., Santos-Silva, C.R., Mesquita, D.O.,
Faria, R.G., Conceicão, B.M et al. (2013): Herpetofauna of
protected areas in the Caatinga I: Raso da Catarina Ecological
Station. Check List 9: 405–414.
Garda, A.A., Stein, M.G., Machado, R.B., Lion, M.B., Juncá, F.A.,
Napoli, M.F. (2017): Ecology, Biogeography and Conservation
of Amphibians in the Semiarid Brazilian Caatinga. In:
Biodiversity, Ecosystems Services and Sustainable Development
Taís Borges Costa et al.
in Caatinga: The Largest Tropical Dry Forest Region in South
America, p. 133–149. Silva, J.M.C., Leal, I. , Tabarelli, M., Ed.,
Berlin, Springer-Verlag.
Gotelli, N.J., Colwell, R.K. (2001): Quantifying biodiversity:
procedures and pitfalls in the measurement and comparison of
species richness. Ecology Letters 4: 379–391.
Guedes, T.B., Nogueira, C., Marques, O.A. (2014): Diversity,
natural history, and geographic distribution of snakes in the
Caatinga, Northeastern Brazil. Zootaxa 3863: 1–93.
Leal, I.R., Silva, J.M.C., Tabarelli, M., Lacher, T.E. (2005):
Changing the course of biodiversity conservation in the Caatinga
of northeastern Brazil. Conservation Biology 19: 701–706.
Lima, S.M.Q., Ramos, T.P.A., Da Silva, M.J., Rosa, R.S. (2017):
Diversity, Distribution, and Conservation of the Caatinga
Fishes: Advances and Challenges. In: Biodiversity, Ecosystems
Services and Sustainable Development in Caatinga: The Largest
Tropical Dry Forest Region in South America, p. 97–131. Silva,
J.M.C., Leal, I. , Tabarelli, M., Ed., Berlin, Springer-Verlag.
Loebmann, D., Haddad, C.F.B. (2010): Amphibians and reptiles
from a highly diverse area of the Caatinga domain: composition
and conservation implications. Biota Neotropica 10: 227–256.
Magalhães, F.M., Dantas, A.K.B.P., Brito, M.R.M., Medeiros,
P.H.S., Oliveira, A.F., Pereira, et al. (2013): Anurans from an
Atlantic Forest-Caatinga ecotone in Rio Grande do Norte State,
Brazil. Herpetology Notes 6: 1–10.
Magalhães, F.M., Laranjeiras, D.O., Costa, T.B., Juncá, F.A.,
Mesquita, D.O., Röhr, D.L., et al. (2015): Herpetofauna of
protected areas in the Caatinga IV: Chapada Diamantina
National Park, Bahia, Brazil. Herpetology Notes 8: 243–261.
Magalhães, F.M., Loebmann, D., Kokubum, M.N.D.C., Haddad,
C.F.B., Garda, A.A. (2014): A new species of Pseudopaludicola
(Anura: Leptodactylidae: Leiuperinae) from northeastern Brazil.
Herpetologica 70: 77–88.
Magnusson, W.E., Lima, A.P., Luizão, R., Luizão, F., Costa,
F.R.C., Castilho, C.V., Kinupp, V.F. (2005): RAPELD: a
modification of the gentry method for biodiversity surveys in
long-term ecological research sites. Biota Neotropica 5: 19–24.
Marengo, J.A., Torres, R.R., Alves, L.M. (2016): Drought in
Northeast Brazil—past, present, and future. Theoretical and
Applied Climatology 129: 1189–1200.
Medeiros, J.B.L.P. (2004): Zoneamento fito-ecológico da Estação
Ecológica de Aiuaba – uma contribuição à educação ambiental
e à pesquisa científica. Unpublished Universidade Federal do
Ceará, Fortaleza, p. 141.
Mesquita, D.O., Costa, G.C., Garda, A.A., Delfim, F.R. (2017):
Species Composition, Biogeography and Conservation of
Caatinga Lizards. In: Biodiversity, Ecosystems Services and
Sustainable Development in Caatinga: The Largest Tropical
Dry Forest Region in South America, p. Silva, J.M.C., Leal, I. ,
Tabarelli, M., Ed., Springer-Verlag.
Mesquita, P.C.M.D., Passos, D.C., Borges-Nojosa, D.M., Cechin,
S.Z. (2013): Ecologia e história natural das serpentes de uma
área de Caatinga no Nordeste Brasileiro. Papéis Avulsos de
Zoologia (São Paulo) 53: 99–113.
Moura, G.J.B., Santos, E.M., Oliveira, M.A.B., Cabral, M.C.C.
(2010): Herpetofauna no Estado de Pernambuco, Brasília,
Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais
Renováveis.
Herpetofauna of protected areas in the Caatinga VII: Aiuaba Ecological Station
Navas, C.A., Antoniazzi, M.M., Jared, C. (2004): A preliminary
assessment of anuran physiological and morphological
adaptation to the Caatinga, a Brazilian semi-arid environment.
International Congress Series 1275: 298–305.
Oliveira, A.P.C., Bernard, E. (2017): The financial needs vs. the
realities of in situ conservation: an analysis of federal funding for
protected areas in Brazil’s Caatinga. Biotropica 49: 745–752.
Oliveira, E.F., Gehara, M., Sao Pedro, V.A., Chen, X., Myers,
E.A., Burbrink, F.T., et al. (2015): Speciation with gene flow
in whiptail lizards from a Neotropical xeric biome. Molecular
Ecology 24: 5957–5975.
Oliveira, J.G.B., Cesar, H.L., Nunes, E.P. (1983): Vegetação.
In: Projeto Aiuaba – Relatório técnico, p. 117–130. Oliveira,
J.G.B., Ed., Fortaleza,
Passos, D.C., Lima, D.C., Borges-Nojosa, D.M. (2011): A
new species of Tropidurus (Squamata, Tropiduridae) of the
semitaeniatus group from a semiarid area in Northeastern Brazil.
Zootaxa 2930: 60–68.
Pedrosa, I.M.M.C., Costa, T.B., Faria, R.G., França, F.G.R.,
Laranjeiras, D.O., Oliveira, et al. (2014): Herpetofauna of
protected areas in the Caatinga III: The Catimbau National Park,
Pernambuco, Brazil. Biota Neotropica 14: 1–12.
Prado, D.E. (2000): Seasonally dry forests of tropical South
America: from forgotten ecosystems to a new phytogeographic
unit. Edinburgh Journal of Botany 57: 437–461.
Prado, D.E. (2003): As Caatingas da América do Sul. In: Ecologia
e Conservação da Caatinga, p. 3–73. Leal, I.R., Tabarelli, M. ,
Silva, J.M.C., Ed., Recife, Ed. Universitária da UFPE.
Ribeiro, L.B., Goliath, M., Rodrigues, R.G., Barreto, R.M.F.,
Freire, E.M.X. (2013): Two new records of Coleodactylus
meridionalis (Boulenger, 1888) (Squamata, Sphaerodactylidae)
in north-eastern Brazil, including a map and comments regarding
its geographical distribution. Herpetology Notes 6: 23–27.
Roberto, I.J., Loebmann, D. (2016): Composition, distribution
patterns, and conservation priority areas for the herpetofauna
of the state of Ceará, northeastern Brazil. Salamandra 52: 134–
152.
939
Roberto, I.J., Ribeiro, S.C., Loebmann, D. (2013): Amphibians of
the state of Piauí, Northeastern Brazil: a preliminary assessment.
Biota Neotropica 13: 322–330.
Rodrigues, M.T. (2003): Herpetofauna da Caatinga. In: Ecologia e
Conservação da Caatinga, p. 181–236. Leal, I.R., Tabarelli, M. ,
Silva, J.M.C., Ed., Recife, Editora Universitária da UFPE.
Santana, D.J., Mângia, S., Silveira-Filho, R.R., Barros, L.C.S.,
Andrade, I., Napoli, M.F., Juncá, F., Garda, A.A. (2015):
Anurans from the middle Jaguaribe River region, Ceará State,
Northeastern Brazil. Biota Neotropica 15: 1–8.
Santos, J.C., Leal, I.R., Almeida-Cortez, J.S.D., Fernandes,
G.W., Tabarelli, M. (2011): Caatinga: the scientific negligence
experienced by a dry tropical forest. Tropical Conservation
Science 4: 276–286.
Silva, J.M.C., Barbosa, L.C.F., Leal, I.R., Tabarelli, M. (2017):
The Caatinga: Understanding the Challenges. In: Biodiversity,
Ecosystems Services and Sustainable Development in Caatinga:
The Largest Tropical Dry Forest Region in South America, p. 3–
19. Silva, J.M.C., Leal, I. , Tabarelli, M., Ed., Berlin, Springer-
Verlag.
Vanzolini, P.E. (1976): On the lizards of a Cerrado-Caatinga
contact: evolutionary and zoogeographical implications (Sauria).
Papéis Avulsos de Zoologia (São Paulo) 29: 111–119.
Velloso, A.L., Sampaio, E.V.S.B., Pareyn, F.G.S. (2002):
Ecorregiões: Propostas Para o Bioma Caatinga, Recife, The
Nature Conservancy do Brasil.
Warburg, M.R. (1997): Ecophysiology of Amphibians Inhabiting
Xeric Environments, Berlin, Springer.
Werneck, F.P., Gamble, T., Colli, G.R., Rodrigues, M.T., Sites Jr,
J.W. (2012): Deep diversification and long-term persistence in
South American ‘Dry Diagonal’: integrating continent-wide
phylogeography and distribution modeling of geckos. Evolution
66: 3014–3034.
940 Taís Borges Costa et al.

Appendix 1. Herpetofauna of Aiuaba Ecological Station, Ceará, Brazil. Abbreviations: AS= Active search; 65GT= glue trap; PT=
pitfall trap; OE= occasional encounter;
Appendix 1. Hoop-trap=HT.
Herpetofauna of Aiuaba Ecological Station, Ceará, Brazil. Abbreviations: AS= Active search;
GT= glue trap; PT= pitfall trap; OE= occasional encounter; Hoop-trap=HT.

Sampling
Taxon Voucher Abundance
Method
Amphibia (n= 20)
Bufonidae
Rhinella granulosa (Spix, 1824) URCA-H 7443 AS, PT, OE 40
Rhinella jimi (Stevaux, 2002) FSCHUFPB 6527 AS, OE 29
Hylidae
Corythomantis greeningi Boulenger, 1986 URCA-H 5550 AS, OE 12
Dendropsophus nanus (Boulenger, 1989) FSCHUFPB 3075 OE, AS 167
Dendropsophus soaresi (Caramaschi & Jim, 1983) URCA-H 7391 AS, OE 7
Boana raniceps (Cope, 1862) FSCHUFPB 3071 AS, OE, PT 34
Phyllomedusa nordestina Caramaschi, 2006 FSCHUFPB 6564 AS, OE 24
Scinax x-signatus (Spix, 1824) FSCHUFPB 3278 AS, OE, PT 36
Leptodactilydae
Leptodactylus fuscus (Schneider, 1799) URCA-H 7545 AS, OE 19
Leptodactylus chaquensis Cei, 1950 FSCHUFPB 3017 AS, OE 143
Leptodactylus troglodytes Lutz, 1926 FSCHUFPB 6361 AS, OE, PT 33
Leptodactylus vastus Lutz, 1930 FSCHUFPB 3028 AS, OE 45
Physalaemus albifrons (Spix, 1824) URCA-H 7373 AS, OE, PT 41
Physalaemus cicada Bokermann, 1966 URCA-H 7443 AS, OE, PT 50
Physalaemus cuvieri Fitzinger, 1826 URCA-H 7454 AS, OE, PT 29
Pleurodema diplolister (Peters, 1870) URCA-H 7388 AS, OE 21
Pseudopaludicola pocoto Magalhães et al. 2014 FSCHUFPB 2000 AS, OE, PT 263
Microhylidae
Dermatonotus muelleri (Boettger, 1885) URCA-H 7401 AS, OE, PT 8
Elachistocleis piauiensis Caramaschi and Jim, 1983 URCA-H 7461 AS, OE, PT 4
Odontophrynidae
Proceratophrys aridus Cruz, Nunes, and Juncá, 2012 URCA-H 7523 AS, OE 19
Lacertoidea (n=17)
Amphisbaenidae
Amphisbaena vermicularis Wagler, 1824 FSCHUFPB 3101 PT, AS 9
Gekkonidae
Hemidactylus agrius Vanzolini, 1978 FSCHUFPB 6368 PT, AS, GT, OE 35
Hemidactylus brasilianus (Amaral, 1935) FSCHUFPB 3161 PT, AS, GT, OE 10
Lygodactylus klugei (Smith, Martin & Swain, 1977) FSCHUFPB 3015 PT, AS, GT, OE 48
Gymnophthalmidae
Micrablepharus maximiliani (Reinhardt & Lütken, 1862) FSCHUFPB 3332 PT, AS, GT 28
Vanzosaura multiscutata (Amaral, 1933) FSCHUFPB 3016 PT, AS, GT 180
Iguanidae
Iguana iguana (Linnaeus, 1758) URCA-H 4765 AS 4

Phyllodactylidae
Gymnodactylus geckoides Spix, 1825 FSCHUFPB 3021 PT, AS, GT, OE 233
Phyllopezus pollicaris (Spix, 1825) FSCHUFPB 3061 PT, AS, GT, OE 162
Mabuyidae
Mabuya heathi (Schmidt & Inger, 1951) FSCHUFPB 6314 AS 13
Sampling
Taxon Voucher Abundance
Method
Polychrotidae
Polychrus acutirostris Spix, 1825 URCA-H 6536 OE 2
 Phyllodactylidae
HerpetofaunaGymnodactylus
of protectedgeckoides
areas inSpix,
the1825
Caatinga VII: AiuabaFSCHUFPB
Ecological Station
3021 PT, AS, GT, OE 233 941
Phyllopezus pollicaris (Spix, 1825) FSCHUFPB 3061 PT, AS, GT, OE 162
Appendix 1. Continued.
Mabuyidae
Mabuya heathi (Schmidt & Inger, 1951) FSCHUFPB 6314 AS 13
Sampling
Taxon Voucher Abundance
Method
Polychrotidae
Polychrus acutirostris Spix, 1825 URCA-H 6536 OE 2
Sphaerodactylidae
Coleodactylus meridionalis (Boulenger, 1888) FSCHUFPB 3301 PT, AS 9
Teiidae
Ameiva ameiva (Linnaeus, 1758) FSCHUFPB 3067 PT, AS 40
Ameivula ocellifera (Spix, 1825) FSCHUFPB 3030 PT, AS 197
Salvator merianae (Duméril & Bibron, 1839) URCA-H 5140 AS 2
Tropiduridae
Tropidurus hispidus (Spix, 1825) FSCHUFPB 3029 PT, AS, GT, OE 421
Tropidurus jaguaribanus Passos, Lima & Borges- FSCHUFPB 3268 PT, AS, OE 259
Nojosa, 2011
Snakes (n= 17)
Boidae
Boa constrictor Linnaeus, 1758 URCA-H 6735 OE, AS 2
Epicrates assisi Machado, 1945 OE 1
Colubridae
Oxybelis aeneus (Wagler, 1824) Road kill AS 1
Leptophis ahaetulla (Linnaeus, 1758) URCA-H 5539 AS, OE 1
Dipsadidae
Apostolepis cearensis Gomes, 1915 FSCHUFPB 3201 PT 1
Leptodeira annulata (Linnaeus, 1758) FSCHUFPB 3200 AS, OE 6
Oxyrhopus trigeminus Duméril, Bibron & Duméril, 1854 FSCHUFPB 3025 AS, OE 12
Philodryas nattereri Steindachner, 1870 FSCHUFPB 6618 AS 2
Philodryas olfersii (Lichtenstein, 1823) URCA-H 1774 AS 1
Pseudoboa nigra (Duméril, Bibron & Duméril, 1854) URCA-H 9536 AS 2
Erythrolamprus poecilogyrus (Wied-Neuwied, 1825) URCA-H 3739 AS 3
Erythrolamprus viridis (Günther, 1862) URCA-H 3740 AS, OE 1
Elapidae
Micrurus ibiboboca (Merrem, 1820) FSCHUFPB 3108 OE, AS, PT 4
Leptotyphlopidae
Epictia borapeliotes (Vanzolini, 1996) FSCHUFPB 3270 AS, PT 4
Viperidae
Bothrops erythromelas Amaral, 1923 URCA-H 3148 AS, OE 3
Bothrops leucurus Wagler, 1824 FSCHUFPB 3027 AS, OE 2
Crotalus durissus Linnaeus, 1758 FSCHUFPB 6401 AS 2
Testudines (n= 3)
Chelidae
Mesoclemmys tuberculata (Luederwaldt, 1926) URCA-H 1225 AS 1
Phrynops geoffroanus (SCHWEIGGER, 1812) FSCHUFPB 6637 HT, OE 6
Kinosternidae
Kinosternon scorpioides (Linnaeus, 1766) vestige/shell OE 1
FSCHUFPB: Field Series, Coleção Herpetológica da Universidade Federal da Paraíba; URCA-H: Coleção
Herpetológica da Universidade Regional do Cariri
Accepted by Anamarija Zagar