Introduction

Amphibians, especially Anurans, can inhabit many environments, from high altitudinal forests to sandbanks. They adopt different reproductive strategies resulting in a combination of morphological, physiological and behavioral traits (Duellman and Trueb, 1994). An aquatic and terrestrial life render amphibians among the first vertebrates to suffer the consequences of habitat loss and fragmentation (Becker et al., 2007), in which urbanization can play a major role (Teixeira et al., 2009).

Urbanization increases the density of the human population, which results in the destruction of natural environments. Farmland and natural areas are converted into commercial, industrial, and residential areas (e.g. Shochat et al., 2006). Habitat loss (e.g. Shochat et al., 2006) is the primary threat to amphibian populations in the Neotropics (Young et al., 2001; Stuart et al., 2004). Although many wild species have been extirpated as a result of urbanization, many species persist and may even prosper in urban settings (Adams, 1994; Rodda and Tyrrell, 2008).

The magnitude of population fluctuations may indicate the risk of local extinction from demographic or environmental stochasticity (Leigh, 1981; Doak et al., 1994; Engen and Saether, 1998). Fluctuations in population decreases the power of monitoring programs to detect real declines (Gerrodette, 1987) and may confuse the decisions of which monitoring strategy is best suited for a given population (Gibbs et al., 1998). Most worrying in this regard geographical distributions of known species are also under-dimensioned (Auricchio and Salomão, 2002), or virtually unknown, a deficiency similarly characterized as Wallacean deficit by Lomolino et al. (2005).

In this context, in recent years, many researchers have focused to produce lists of anuran species in many areas of Espirito Santo state, southeastern Brazil (e.g. Ramos and Gasparini, 2004; Gasparini et al., 2007; Rödder et al., 2007; Teixeira et al., 2007; Teixeira et al., 2008; Ferreira et al., 2010; Tonini et al., 2010; Almeida et al., 2011). However, there are many inventory gaps in this region, particularly in urban places and other neglected areas. In this study our goal was to inventory the anuran fauna of forest fragments in an urban landscape located in the coastal region of Espirito Santo.

Materials and methods

Study site

The study was conducted at Costa Bela, municipality of Serra, state of Espírito Santo, southeastern Brazil at 20°06’07’’ S and 40°10’56’’ W, 6.51 m (Figure 1), in adjacent forest fragments which together correspond to 69.94 ha. The climate is characterized as tropical hot and humid with a rainy season in summer and low precipitation in the winter, average of 900 to

Herpetology Notes, volume 4: 435-444 (2011) (published online on 10 December 2011)

Anurans from Costa Bela, state of Espírito Santo, southeastern Brazil: inventory at an urban area and the re-discovery

of Allobates in the state

João Filipe Riva Tonini1*, Israel Souza Mendonça2, Alyson Borges Coutinho2 and João Luiz Gasparini3

1 Departamento de Ciências Biológicas, Universidade Federal do Espírito Santo, Av. Marechal Campos 1468, Maruípe, CEP 29043-900, Vitória, ES, Brasil;

2 Departamento de Ciências Biológicas, Faculdades Integradas São Pedro, Rodovia Serafim Derenzi 3115, São Pedro, CEP 29030-026, Vitória, ES, Brasil;

3 Departamento de Oceanografia e Ecologia, Universidade Federal do Espírito Santo, Av. Fernando Ferrari 514, Goia-beiras, CEP 29075-910, Vitória, ES, Brasil.

* Corresponding author: e-mail: jfrtonini@gmail.com

Abstract. Our aim was to inventory the fauna of anurans from forest fragments in an urban landscape at Costa Bela, state of Espírito Santo, southeastern Brazil. Between December 2008 and November 2009 we conducted 26 field expeditions that resulted in 31 records of anuran species. Among these, 15 (48.5%) are restricted to the Atlantic forest, nine (29%) are widely distributed in Atlantic forest and seven (22.5%) range other biomes beyond the Atlantic forest. Allobates olfersioides was re-discovered in the state of Espirito Santo and is listed as Vulnerable in IUCN Red List of Threatened Species.

Keywords. Amphibians, Atlantic forest, biodiversity, South America, urban ecology, population flutuations.

João Filipe Riva Tonini et al. 346

1,200 mm³ and mean annual temperature 24 °C. The dominant vegetation is Lowland Rain Forest (Oliveira-Filho and Fontes, 2000). The forest fragments are located in an urban area surrounded by human constructions and suffer with the growth of forestry plantations.

Sampling procedure

We carried out 26 field expeditions each lasting five hours, one every 15 days between December 2008 and November 2009. Our sampling methods were active search and pitfall traps. Active search was conducted using flashlights to locate the specimens in environments where they usually take shelter, such as tree cavities, crevices between rocks, under rocks and logs, soil, litter, on bromeliads and marginal vegetation along the watercourses. The expeditions occurred between 17:30 and 22:30 h, with a total sampling effort of 130 h. Occasionally, additional field expeditions were carried out on rainy days. The pitfall traps were installed during October 2009 and consisted of 11 buckets (20 L), 5 m distant from each other, connected by 50 m of 1 m high black plastic. The buckets were opened three days per week over eight weeks with a total sampling effort of 1,728 hours. During the sampling intervals the buckets were closed.

Following Prado & Pombal (2005), we classify the monthly occurrence of species along the sampling period. The species with records in ten or more months were considered “frequent”, species occurring within three to nine months were considered “occasional” and species occurring in less than three months were considered as “sporadic”. We considered the species with records in one to three states covering the Atlantic forest as having a “restricted distribution”. Species that occur in more than three states of the Atlantic forest have a “wide distribution”, and species that occur in other biomes are designated “range also includes other biomes beyond Atlantic forest” (Dixo and Verdade, 2006).

Specimens were identified by comparing morphological characters with specimens deposited in research collections and original articles describing the species. The species distribution

and taxonomic arrangement follows Frost (2010). The individuals were sampled according to license No. 20044-3 from Instituto Brasileiro de Meio Ambiente e dos Recursos Hídricos (IBAMA). After analysis and identification, the specimens were deposited in the Museu de Biologia Mello Leitão (MBML), Santa Teresa, Espirito Santo and in the Amphibian Collection at Museu Nacional (MNRJ). Tissues collected during the setting process of specimens are deposited in the Coleção Fiel Depositária de Tecidos e de DNA (CTA-UFES) at Universidade Federal do Espírito Santo, Vitória, Espírito Santo.

Results

We recorded 149 specimens belonging to 31 species of anurans (Table 1, Figures 2-5) distributed in seven families: Aromobatidae (1), Bufonidae (2), Cycloramphidae (1), Hylidae (19), Leiuperidae (1), Leptodactylidae (4), Microhylidae (3) (vouchers listed in Appendix 1). Among these species, 15 (48.5%) have a restricted distribution within the Atlantic forest, nine (29%) have a wide distribution within the Altantic forest and seven (22.5%) inhabit other biomes beyond the Atlantic forest. According to the occurrence throughout the sampling period we consider five species as frequent, 18 species as occasional and seven species as sporadic (Table 1), and in the rainy season (mostly in October 2009) we recorded more than 80 % of the total species. The species Scinax aryreornatus and Chiasmocleis cf. capixaba were the most abundant with 21 individuals each, and Dendropsophus berthalutzae and Proceratophrys laticeps was less abundant with only two individuals each.

Discussion

In the regional list of threatened species of anurans from the state of Espirito Santo, Gasparini et al. (2007) commented that since the description of Allobates capixaba (Bokermann, 1967), later called Allobates olfersioides by Verdade and Rodrigues (2007), the species has not been observed in this state, even in recent surveys carried out near the type locality. After 43 years without new records of this species in the state, it was found in our study site at Costa Bela, and in Aracruz (J.L. Gasparini and A.P. Almeida, personal communication). Other authors reported population fluctuations for anurans in Espirito Santo(e.g. Weygoldt, 1989; Papp and Papp, 2000). However, populations of some species listed in Weygoldt (1989) were re-discovered in recent fieldwork by Tonini et al. (2010), and in many unpublished studies - Phasmahyla exilis, Crossodactylus gaudichaudii and Hylodes lateristrigatus, and Allobates olfersioides (this study)

Figure 1. Costa Bela, municipality of Serra, state of Espírito Santo, southeastern Brazil. BA, Bahia; ES, Espírito Santo; MG, Minas Gerais; RJ, Rio de Janeiro.

Anurans from Costa Bela 437

in the lowland. Eterovick et al. (2005) reports that cases of Brazilian

anuran population fluctuations in regions of medium and high altutides (≥ 500 m above sea level) have different causes, and a few observations relate these fluctations to habitat loss and landscape changes. However, these authors suggest that the rapid rate of destruction of Brazil’s natural ecosystems, through habitats loss and habitat modification, may present much larger problems than those documented thus far (see Becker et al., 2007; Becker et al., 2010). They also suggest that ecological processes, such as competition among species, can lead to population decline

(Eterovick et al., 2005). Therefore, the rediscovery of new populations of missing species must be followed by the concern to keep them viable. In the state of Espírito Santo the populations of Allobates are still in danger, mainly because of habitat splitting and habitat loss caused by real estate speculation and urban growth in the region.

Amphibian species have been re-discovered around the world (e.g. Rodrígues-Contreras et al., 2008; Abarca et al., 2010), and some of these population fluctuations can be attributed to the biodiversity crisis resulting mainly from the destruction, modification and fragmentation of habitats, species introductions,

Species Distribution OccurenceAromobatidae

Allobates olfersioides (Lutz, 1925) ++ OccasionalBufonidae

Rhinella crucifer (Wied-Neuwied, 1821) +++ OccasionalRhinella granulosa (Spix, 1824) +++ Occasional

CycloramphidaeProceratophrys laticeps Izecksohn & Peixoto, 1981 + Sporadic

HylidaeAparasphenodon brunoi Miranda-Ribeiro, 1920 + OccasionalDendropsophus berthalutzae (Bokermann, 1962) + SporadicDendropsophus bipunctatus (Spix, 1824) ++ SporadicDendropsophus branneri (Cochran, 1948) ++ FrequentDendropsophus cf. decipiens ++ SporadicDendropsophus elegans (Wied-Neuwied, 1824) ++ OccasionalDendropsophus haddadi (Bastos & Pombal, 1996) ++ OccasionalDendropsophus seniculus (Cope, 1868) + SporadicHypsiboas albomarginatus (Spix, 1824) + FrequentHypsiboas pombali (Caramaschi, Pimenta & Feio, 2004) +++ FrequentHypsiboas semilineatus (Spix, 1824) + OccasionalPhylomedusa burmeisteri Boulenger, 1882 ++ SporadicScinax alter (Lutz, 1973) ++ FrequentScinax argyreornatus (Miranda-Ribeiro, 1926) ++ OccasionalScinax cuspidatus (Lutz, 1925) + OccasionalScinax eurydice (Bokermann, 1968) + OccasionalScinax fuscovarius (Lutz, 1925) + SporadicSphaenorynchus planicola (Lutz & Lutz, 1938) +++ OccasionalTrachycephalus sp. + Occasional

LeiuperidaePhysalaeumus crombiei Heyer & Wolf, 1989 +++ Occasional

LeptodactylidaeLeptodactylus fuscus (Schneider, 1799) + OccasionalLeptodactylus latrans (Linnaeus, 1758) ++ OccasionalLeptodactylus natalensis Lutz, 1930 +++ OccasionalLeptodactylus thomei Almeida & Ângulo, 2006 +++ Sporadic

MicrohylidaeArcovomer cf. passarellii + OccasionalChiasmocleis cf. capixaba + FrequentStereocyclops incrassatus Cope, 1870 + Occasional

(+) restricted distribution within the Atlantic forest, (++) wide distribution within the Altantic forest and (+++) range also includes other biomes beyond Atlantic forest.

Table 1. Anurans from Costa Bela, municipalitiy of Serra, state of Espírito Santo, southeastern Brazil.

João Filipe Riva Tonini et al. 438

Figure 2. Anuran species recorded from Costa Bela photographed along the study period. A) Allobates olfersioides; B) Rhinella crucifer; C) Rhinella granulosa; D) Proceratophrys laticeps; E) Aparasphenodon brunoi; F) Dendroposophus berthalutzae; G) Dendropsophus bipunctatus; H) Dendropspophus branneri.

Anurans from Costa Bela 439

Figure 3. Anuran species recorded from Costa Bela photographed along the study period. A) Dendropsophus haddadi; B) Dendropsophus elegans; C) Dendropsophus cf. decipiens; D) Dendropsophus seniculus; E) Hypsiboas albomarginatus; F) Hypsiboas pombali; G) Hypsiboas semilineatus; H) Phyllomedusa burmeisteri.

João Filipe Riva Tonini et al. 440

Figure 4. Anuran species recorded from Costa Bela photographed along the study period. A) Scinax alter; B) Scinax argyreornatus; C) Scinax cuspidatus; D) Scinax eurydice; E) Scinax fuscovarius; F) Sphaenorhynchus planicola; G) Trachycephalus sp.; H) Physalaemus crombiei.

and overexploration. Other potential factors include climate change, increased UV-B radiation, chemical contaminants, emerging infectious diseases and congenital or malformations (Collins and Storfer, 2003). Weygoldt (1989) correlated the fluctuations of anurans in the state of Espirito Santo to factors including severe frost, epidemic diseases, pollution and acid rain, associated with a low pH detected in the water of some streams in the municipality of Santa Teresa in 1989. This author also discusses the effects of climatic changes and prolonged droughts on the larvae

of many stram-breeding species, which are known to take a long time to metamorphose. Parts of the streams in Santa Teresa dried completely in 1987. Moreover, the possibility of prolonged dry periods could have led to changes in population size, and ultimately local extintion (Weygoldt, 1989).

The value of 31 species of anurans correspond to 23% of the anurans listed to state of Espírito Santo (Almeida et al. 2011). This results compared to other localities in the state, like Reserva Biológica de Duas Bocas (52 species; 2,910 ha; Tonini et al.,

Anurans from Costa Bela 441

Figure 5. Anuran species recorded from Costa Bela photographed along the study period. A) Leptodactylus fuscus; B) Leptodactylus latrans; C) Leptodactylus natalensis; D) Leptodactylus thomei; E) Arcovomer cf. passarellii; F) Chiasmocleis cf. capixaba.

2010), Área de Proteção Ambiental Goiapaba-Açu (41 species; 3,740 ha; Ramos and Gasparini, 2004), Área de Proteção Ambiental de Setiba (38 species; 1,500 ha; Gasparini et al., 2007) and municipality of Santa Teresa (104 species; more than 500 ha; Rödder et al., 2007; Peres et al. 2010), can be considered low. However, these studies were conducted mostly within protected areas of great relevance to the state. When the richness of Costa Bela was compared with other studies in impacted areasin the state, such as the campus of Universidade Federal do Espírito Santo (33 species; 9.3 ha; Ferreira et al. 2010), Anchieta (21 species; 0.35 ha; Teixeira et al., 2007) and Vale do Suruaca (26 species; 0.075 ha; Teixeira et al., 2008), we realized that can be considered high. Ferreira et al. (2010) found anuran species richness values similar to ours near the present study in disturbed areas of the municipality of Vitória,

Variables such as the size of the study area, proximity to conservation areas, altitudinal differences, sampling procedures and conservation status of the study area, affect the comparisons between inventories, but they provide an important tool in assessing the ecological potential and local area management. An example is Costa Bela, where a fragment of 69.94 ha of forest houses a higher species richness than some conservation areas in the state of Espírito Santo, and still houses the genus Allobates that had not been captured since its description and is listed as “vulnerable” on the IUCN Red List of Threatened Species (Verdade, 2008).

Regarding the productivity of the environment, at a slightly less extreme level of urbanization, landscape management of green spaces (e.g. parks, gardens and golf courses) often elevates local productivity relative to surrounding wildland, with such green spaces lying close to the highest end of local productivity continua

(Shochat et al., 2006). On the other hand, this increase in primary productivity causes the attenuation of seasonal variations, thereby increasing the abundance of opportunistic species that compete with native species (Shochat et al., 2006). As a result of this process there is a range extension of good colonizers, have its distribution extended and can also occupy new habitats, either for reproduction or protection from predators (e.g. Ferreira and Tonini, 2010).

We suspect that the species Trachycephalus sp. is a hybrid between the species T. nigromaculatus and T. mesopaheus, this two species have its distribution overlapping in the state of Espírito Santo. It were already noticed in sympatry by others authors that previously notice the possibility of hybridization between T. nigromaculatus and T. mesophaeus (Ramos and Gasparini 2004). Faivovich et al. (2006) comparing the morphology and the phylogenetic relationship of the Phrynohyas mesophea (latter called T. mesophaeus) with T. nigromaculatus, reported that the paraphyly between this two taxa can be attributed to the possibility of hybrids and introgression. Preliminary data using the gene Cytochrome Oxidase 1 (gene used in the Barcode project; Hebert et al., 2003) grouped Trachycepahlus sp. with T. typhonius. However, future cariotipic data, new mitochondrial and nuclear genes can help to elucidated these uncertainties.

The study area consists of a small forest fragment, located in an urban landscape surrounded by human constructions. It suffers with the growth of forestry plantations and has clear-cut timber, causing considerable change in the original vegetation. However, the environment was still able to harbor several species considered as restricted to the Atlantic forest (Table 1). Souza et al. (2008) proposes that intermediate levels of disturbance have suggested

João Filipe Riva Tonini et al. 442

Figure 6. (A) Stereocyclops incrassatus performing the stiff-legged defensive behavior (B).

a lift of biodiversity in some environments because they generate more heterogeneity in the fragments, which can accommodate a larger number of species by increasing the amount of microhabitats and the diversity of resources. On the other hand, environments with low levels of disturbance have their niches occupied by taxa experts limiting the occupation of other species (Souza et al., 2008).

In spite of the advanced degree of urbanization, Costa Bela presents a high level of resilience, since there is a considerable number of anuran species with a restricted distribution and at least one endangered species that succeeded to remain in that environment. However, human impact is rapidly increasing, and with exclusive competition imposed by potentially newly colonizing species, endemic species of anurans might face elimination. Changes in ecological processes can alter the selective forces in cities, leading to genetic differentiation between native and urban populations (Shochat et al., 2006). Because there is no protected area in the surroundings of the forest fragments of Costa Bela, the area acts as an important repository of biodiversity, especially for anurans. Future studies should focus on the processes of establishment for new species and long-term population fluctuations.

Acknowledgements. We are grateful to C. Mattedi, P.V. Scherrer, and A.P. Almeida for helping with field and laboratory work. L.P. Costa provided laboratory support at Mastozoology and Biogeography Lab. Z. Spanos, B. Ferreira, R.C. Carvalho, and two reviewers M. Vences and D. Rödder provide valuable coments and revised the text. Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) provided master scholarship to JFRT.

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Appendix 1. Voucher list (MNRJ – Museu Nacional, Rio de Janeiro, RJ; MBML – Museu de Biologia Prof. Mello Leitão, Santa Teresa, ES).

Allobates olfersioides (MBML7384-85), Aparasphenodon brunoi (MBML7386), Arcovomer cf. passarellii (MNRJ68069-70), Chiasmocleis cf. capixaba (MBML7387-89), Dendropsophus berthalutzae (MNRJ68061-62), Dendropsophus bipunctatus (MNRJ68063-64), Dendropsophus branneri (MNRJ68046-50), Dendropsophus cf. decipiens (MNRJ68045), Dendropsophus elegans (MBML7390-91), Dendropsophus haddadi (MNRJ68051-52), Dendropsophus seniculus (MNRJ68041), Hypsiboas albomarginatus (MNRJ68028), Hypsiboas pombali (MNRJ68026), Hypsiboas semilineatus (MNRJ68030, MNRJ68057-58), Leptodactylus fuscus (MNRJ68031), Leptodactylus latrans (MNRJ68037), Leptodactylus natalensis (MNRJ68073), Leptodactylus thomei (MNRJ68027), Physalaemus crombiei (MNRJ68071-72), Phyllomedusa burmeisteri (MBML7392), Proceratophrys laticeps (MBML7393-94), Rhinella crucifer (MNRJ68034), Rhinella granulosa (MNRJ68088), Scinax alter (MNRJ68029, MNRJ68065-68), Scinax argyreornatus (MNRJ68074-77), Scinax cuspidatus (MNRJ68043), Scinax eurydice (MNRJ68035), Scinax fuscovarius (MNRJ68033), Sphaenorhyncus planicola (MNRJ68038, MNRJ68055-56), Stereocyclops incrassatus (MNRJ68036), Trachycephalus sp. (MNRJ68087).

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Accepted by Miguel Vences; Managing Editor: Dennis Rödder