NORTH-WESTERN JOURNAL OF ZOOLOGY 10 (2): 365-373 ©NwjZ, Oradea, Romania, 2014 Article No.: 141505 http://biozoojournals.ro/nwjz/index.html

Reproductive modes, use, and richness of anurans from Chapada dos Veadeiros, central Brazil

Guilherme Ramalho Chagas Cataldi SANTORO1,* and Reuber Albuquerque BRANDÃO2

1. Programa de Pós-graduação em Zoologia, Instituto de Biologia, Universidade de Brasília – DF. Campus Universitário Darcy Ribeiro, CEP 70910-900, Brasília, DF, Brazil. 2. Universidade de Brasília, Laboratório de Fauna e Unidades de Conservação, Departamento de Engenharia Florestal, Caixa Postal 04357. CEP 70919-970. Brasília, DF, Brazil. *Corresponding author, G.R.C.C. Santoro, E-mail: [email protected]

Received: 06. February 2014 / Accepted: 14. March 2014 / Available online: 16. October 2014 / Printed: December 2014

Abstract. Anurans have the largest number of reproductive modes among vertebrates, and are especially diverse in New World Tropical Forests. However, species inhabiting open and drier biomes present reproductive modes adapted for breeding in these environments. Despite recent studies, the diversity of reproductive modes, species richness, and distribution of anurans in the Cerrado is still poorly known. The Chapada dos Veadeiros, in Central Brazil, harbors high species richness and endemism. This region is located in the central portion of the Cerrado and presents various physiognomies, especially those associated with high altitude plateaus (above 900 m a.s.l.). Herein, we evaluated the diversity of reproductive modes, habitat use, and conservation status of anurans from the Chapada dos Veadeiros. We found 54 species from eight families, which presented 14 reproductive modes, including a new mode for the Cerrado. Anuran richness in the Chapada dos Veadeiros is the highest in the Cerrado, highlighting its importance for the biome biodiversity. Among the species, 66% are Cerrado endemics, occurring mainly in open areas. Moreover, five species are known only from the region, and six have not been described yet. Most species occur in typical Cerrado formations, are terrestrial, and breed in lentic environments. Leptodactylids had the highest diversity of reproductive modes, followed by Hylids. In general, species showed adaptations for breeding in dry environments, being foam nests the most characteristic breeding strategy. Anuran conservation in the Cerrado depends on the maintenance of environmental heterogeneity of open areas.

Key words: Cerrado, breeding, reproductive biology, richness, phytophysiognomies.

Introduction Provete et al. 2011, Conte et al. 2013). The constraints generated by the specific re- Anurans have the greatest reproductive diversity quirements related to the reproductive modes of all tetrapods (Duellman & Trueb 1986), with 39 causes high habitat specificity in anurans (Ber- different reproductive modes (Haddad & Prado toluci & Rodrigues 2002). Therefore, regions with 2005). The classification of these categories of re- high structural complexity (Oda et al. 2009) and productive strategies is based on oviposition site, environmental heterogeneity (Diniz-Filho et al. egg characteristics, and type of parental care 2005) tend to present higher amphibian richness. (Salthe & Duellman 1973, Toledo et al. 2012). Dif- Previous studies suggested that anurans tend to ferent populations of the same species may exhibit segregate their reproductive periods temporally distinct reproductive modes due to competition or and spatially, so they can coexist and diminish the changes in oviposition site availability (Touchon & effects of competition (Oda et al. 2009, Kopp et al. Warkentin 2007, Toledo et al. 2012). 2010, Brandão et al. 2013). Nevertheless, historical The New World wet forests have the highest factors may also influence breeding activities and diversity of reproductive modes for anurans site preferences (Kopp & Eterovick 2006). (Wells 2007). The Brazilian Atlantic forest, for ex- The anuranfauna of open formations devel- ample, harbors 27 out of 39 known modes oped a series of evolutionary adaptations, primar- (Haddad & Prado 2005). Several studies con- ily related to breeding in dry environments with a ducted in open and seasonal biomes showed a well-defined seasonality (Duellman & Trueb smaller diversity of reproductive modes compared 1986). The majority of their reproductive strategies to forested biomes, and most species limit its re- involve adaptations for desiccation and sun expo- productive activity to the rainy season and use sure (Haddad & Prado 2005). in sea- mainly lentic water bodies for egg laying (Haddad sonal biomes usually have a short reproductive & Prado 2005, Prado et al. 2005, Kopp et al. 2010, period and utilize accumulated water or some mi-

366 G.R.C.C. Santoro & R.A. Brandão

Figure 1. The Chapada dos Veadeiros National Park (black line). White circles represent surveyed localities.

crohabitats that retain humidity for egg deposi- conservation priority for Cerrado amphibians due tion, such as burrows, rock crevices, tree holes, to its high richness and endemicity (Diniz-Filho et and fallen logs; Duellman & Trueb 1986, Eterovick al. 2005). Its climatic stability during the late Pleis- & Fernandes 2001, Wells 2007, Kopp et al. 2010). tocene provided a refuge for Cerrado fauna, ex- Thus, even terrestrial breeders depend on water plaining its high richness (Werneck et al. 2012). A for eggs development (Gomez-Mestre et al. 2012). portion of this region (about 65,000 ha) is pro- The Cerrado is the most diverse savanna in tected by the Chapada dos Veadeiros National the world (Klink & Machado 2005), showing a Park and other small private reserves, but most of strongly seasonal climate, with a wet and warm the region lacks effective conservation policies, de- (October to April) and a dry and cold seasons spite its biological relevance. (May to September). Its original area comprised Herein, we provide data on habitat use, re- 23% of the Brazilian territory and is characterized production modes, and conservation status of the by a complex landscape with great horizontal het- anuranfauna from the Chapada dos Veadeiros Na- erogeneity in the distribution of open, savanic, tional Park and surrounding areas. and forested (Ribeiro & Walter 1998). As a consequence, species distribution differs among those areas (Colli et al. 2002). About 52% of the 213 Materials and Methods anurans recorded for the Cerrado are endemic (Valdujo et al. 2012). Despite the recognized anu- Study area Fieldwork was conducted in the Chapada dos Veadeiros, ran richness in the central and northern portions Northern Goiás, central Brazil (between the latitudes 13° of the Cerrado, most of the surveys were focused and 15° S; and longitudes 47° to 49° W; Felfili 2007). The on the southern region of the Cerrado or near ur- Chapada dos Veadeiros National Park comprises four ban areas (Colli et al. 2002). Some studies showed municipalities: Alto Paraíso de Goiás, Cavalcante, Colinas that the diversity of Cerrado amphibians is greater do Sul, and Teresina de Goiás (Fig. 1). This region is char- in open habitats (Brandão & Araújo 2001, Valdujo acterized by highlands (up to 1670m a.s.l.), covered et al. 2011, 2012). Although some species use for- mainly by fields and open Cerrado physiognomies (Fig. 2). The soil is shallow and seasonally wet. The climate is ested habitats, the majority is adapted to open tropical highland (Köppen’s Cwa), with monthly mean habitats (Brandão & Araujo 2001), which are ex- temperatures ranging from 21°C to 22°C and 1,550 mm of tremely threatened (Silva & Bates 2002). mean annual rainfall (Ramalho et al. 2013). Forested habi- The region of the Chapada dos Veadeiros is a tats correspond to less than 5% of landscape (Felfili 2007).

Diversity and reproduction of Cerrado anurans 367

Figure 2. Examples of physiognomies in the Chapada dos Veadeiros. A – Grassland (Campo limpo), B – Bush grassland (Campo sujo), C – Wet grassland (Campo alagável), D - Cerrado sensu strictu, E – Valley with gallery Forest, F – Riparian forest.

Deep valleys are present in the surrounding and have We classified species by their habitats, reproduction more dense forests (Fig. 2). Grasslands are less repre- habitats, typical perch, distribution, and reproductive sented at low altitudes. The region has complex and het- modes (Table 1). The reproductive mode classification erogeneous vegetation, including open grasslands (campo (Haddad & Prado 2005) was based on natural history ob- limpo), bush grasslands (campo sujo), gallery forests, wet servations and collection information. grasslands (veredas) with Buritis (Mauritia flexuosa) palms, wet grasslands (campos alagáveis), and highland grass- lands (campos de altitude; Fig. 2). Results

During the last 20 years, several researchers from the We recorded 54 anuran species from eight families Universidade de Brasília, including us, surveyed anurans and 19 genera. The most diverse family is , in the Chapada dos Veadeiros National Park and its sur- roundings (Fig. 1). We used these records, along with an- followed by (Table 1). More than urans deposited in the Herpetological Collection of the half (66%) of the species are Cerrado endemics Universidade de Brasília (CHUNB) to establish the list of and five of them are known to occur only in the species occurring in the region. We also included de- study area (Adenomera juikitam, Hypsiboas ericae – scribed species from recent publications associated to the Fig. 3A; Hypsiboas phaeopleura – Fig. 3B; Procerato- Chapada dos Veadeiros. phrys rotundipalpebra; and tapiti – Fig.

368 G.R.C.C. Santoro & R.A. Brandão

Table 1. Anuran species recorded in the Chapada dos Veadeiros, showing habitat use (Of: open grasslands, including campo limpo, campo rupestre, vereda, and campo sujo; Ce: cerrado, including campo cerrado, cerrado aberto, cerrado rupestre, cerrado strict sense; Fh: Forested habitats, including gallery forests, ciliar forests, dry forests, and cerradão; Aa: Altered areas, including artificial dams, cattle dams, artificial ); breeding habitat (Lo: lentic, Lo: lotic, or Tr: terrestrial reproduction), typical perch (Te: Terrestrial, Gr: grasses; Lb: low bushes, Hb: high bushes; Br: bromeliad), and distribution (E: Cerrado Endemic, NE: narrow endemic, W: wide distribution, “?”: no information on species distribution).

Repro- Repro- Tipical calling Distri- Species Habitat ductive ductive perch bution habitat mode goianus (Bokermann, 1975) Fh Tr Te E 20 BUFONIDAE Rhaebo guttatus (Schneider, 1799) Fh Le Te W 1 mirandaribeiroi (Gallardo, 1965) Ce, Aa Le Te E 1 Rhinella rubescens (Lutz, 1925) Of, Ce, Fh, Aa Lo, Le Te E 1, 2 Rhinella schneideri (Werner, 1894) Of, Ce, Fh, Aa Lo Te W 1 ternetzi (Miranda-Ribeiro, 1937) Ce, Fh Tr Te E 23 DENDROBATIDAE Ameerega flavopicta (Lutz, 1925) Of, Ce Lo Te E 20 HYLIDAE Aplastodiscus sp. Fh Lo Hb, Lb E 5 Bokermannohyla pseudopseudis (Miranda-Ribeiro, 1937) Of, Ce, Fh Lo Te, Lb E 2 jimi (Napoli & Caramashi, 1999) Of, Ce Le Gr E 1 Dendropsophus minutus (Peters, 1872) Of, Ce, Aa Le Lb, Gr W 1 Dendropsophus rubicundulus (Reinhardt & Lütken,1862) Of, Ce, Aa Le Gr E 1 Hypsiboas albopunctatus (Spix, 1824) Of, Ce, Aa Le Lb, Hb, Gr W 1 Hypsiboas ericae (Caramashi & Cruz, 2000) Fh, Ce Lo Hb, Lb NE 2 Hypsiboas goianus (Lutz, 1968) Fh Lo Hb, Lb E 2 Hypsiboas lundii (Burmeister, 1856) Fh Lo Hb, Lb E 4 Hypsiboas phaeopleura (Caramashi & Cruz, 2000) Fh Lo Lb, Hb NE 2 Hypsiboas raniceps Cope, 1862 Of, Fh, Aa Le Lb W 1 Hypsiboas sp. Of Lo Lb, Gr ? ? Phyllomedusa azurea Cope, 1862 Of, Ce, Fh, Aa Le Lb, Hb,Te, Gr W 24 Phyllomedusa oreades Brandão, 2002 Of, Ce Lo Lb, Te, Gr E 25 aff. squalirostris (Lutz, 1925) Of, Ce Le Gr ? 1 Scinax fuscomarginatus (Lutz, 1925) Of, Ce, Aa Le Gr, Lb W 1 Scinax fuscovarius (Lutz, 1925) Of, Ce, Fh, Aa Le Lb, Hb, Gr, Te W 1 Scinax rogerioi Pugliese, Baêta & Pombal, 2009 Of Le Gr,Te E 1 Scinax similis (Cochran, 1952) Of, Ce, Fh, Aa Le Te, Lb, Hb, Gr W 1 Scinax skaios Pombal, Carvalho, Canelas & Bastos, 2010 Of, Ce, Fh Lo Te, Lb E 2 Scinax tigrinus Nunes, Carvalho & Ferreira, 2010 Of, Ce Le Gr E 1, 2 Scinax sp. Of, Ce Lo Gr,Te E 2 Trachycephalus typhonius (Linnaeus, 1758) Fh, Aa Le Hb, Lb W 1 LEPTODACTYLIDAE Adenomera cotuba Carvalho & Giaretta 2013 Of, Ce Le Te E 32 Adenomera juikitam Carvalho & Giaretta 2013 Of, Ce Le Te NE 32 Adenomera saci Carvalho & Giaretta 2013 Of, Ce Le Te E 32 Leptodactylus furnarius Sazima & Bokermann, 1978 Of Le Te E 30 Leptodactylus fuscus (Schneider, 1799) Of, Ce, Fh, Aa Le Te W 30 Leptodactylus labyrinthicus (Girard, 1853) Of, Ce, Fh, Aa Le Te W 13 Leptodactylus mystaceus (Spix, 1824) Of, Ce, Fh Le Te W 30 Leptodactylus mystacinus (Burmeister, 1861) Of, Ce Le Te W 30 Leptodactylus sertanejo Giaretta & Costa, 2007 Of, Ce Le Te E 30 Leptodactylus syphax Bokermann, 1969 Of, Ce Le Te E 29 Leptodactylus tapiti Sazima & Bokermann, 1978 Of, Ce Lo Te NE 31 Leptodactylus latrans (Steffen, 1815) Of, Ce, Fh, Aa Le Te W 11

continued on next page Diversity and reproduction of Cerrado anurans 369

Table 1. (continued).

Repro- Repro- Tipical calling Distri- Species Habitat ductive ductive perch bution habitat mode Leptodactylus sp. Of,Ce Lo Te E 31 Physalaemus centralis Bokermann, 1962 Of, Ce Le Te E 11 Physalaemus cuvieri Fitzinger, 1826 Of, Ce, Fh, Aa Le Te W 11 mystacalis (Cope, 1887) Of, Ce Le Te W 1 Pseudopaludicola saltica (Cope, 1887) Of, Ce, Fh Le Te E 1 Pseudopaludicola ternetzi Miranda-Ribeiro, 1937 Of, Ce, Aa Le Te E 1 MICROHYLIDAE Elachistocleis cesarii (Miranda-Ribeiro, 1920) Of, Ce, Aa Le Te E 1 Elachistocleis sp. Of Le Te E 1 Odontophrynus cultripes Reinhardt & Lütken,1862 Ce, Fh Le Te E 1, 2 Odontophrynus salvatori Caramashi, 1996 Of, Ce Lo Te E 2 goyana (Miranda-Ribeiro, 1937) Of, Ce, Fh Lo Te E 2 Proceratophrys rotundipalpebra Martins & Giaretta 2013 Of, Ce Lo Te NE 2

3H) and were considered narrow endemics (sensu (mode 2) water bodies (Table 1). Another common Valdujo et al. 2012; Table 1). Some populations strategy involves the use of foam nests for egg represent undescribed species (Aplastodiscus aff. deposition out of water (modes 29 to 32) presented perviridis; Hypsiboas sp.; Scinax sp.; Scinax aff. by the majority of the leptodactylids, and aquatic squalirostris (Fig. 3D), Elachistocleis sp., and Lepto- foam nests (modes 11 and 13; Table 2). Three spe- dactylus sp.). Moreover, 17 endemic species only cies presented reproduction independent of water occur in open Cerrado formations (Table 1). We (Modes 20 and 23). Allobates goianus and Ameerega recorded seven species that occurred exclusively flavopicta (Fig. 3J) carry tadpoles to water bodies, inside forests, being five of them Cerrado endem- and Barycholos ternetzi (Fig. 3F) has direct devel- ics (Table 1). Most species used field and savanic opment of terrestrial eggs. Only two species pre- formations (sensu Ribeiro & Walter 1998; 81% and sented arboreal modes (24 and 25), both of Phyl- 79%, respectively), whereas 46% occurred in forest lomedusa genus (Phyllomedusa azurea and P. oreades formations, and 33% also used altered areas (Table – Fig. 3C). 1). Additionally, most species are exclusively ter- restrial (59%), followed by arboreal species that use grasses and small herbs for perching (five spe- Discussion cies; Table 1). About one third perch on bushes (with 31% use low bushes, and 18% use tall The diversity of reproductive modes found in the bushes), but most call on the ground during the Chapada dos Veadeiros is the greatest recorded reproductive period (74%, N = 40). About 64% of for a Cerrado locality or even open formations in the species lay eggs on lentic water bodies, 35% South America (Table 2), being comparable to use lotic watercourses, and less than 4% have ter- some sites in the Amazon (Haddad & Prado 2005). restrial reproduction (Table 1). The high horizontal heterogeneity of habitats and We recorded 14 reproductive modes. One of physiognomies in the Cerrado (Garda et al. 2013) them recorded for the first time for the Cerrado should play an important role in this diversity. biome (mode 5). This mode is presented by Aplas- The most frequent reproductive mode at the study todiscus aff. perviridis that deposits eggs in subter- area is eggs and exotrophic tadpoles in lentic wa- ranean flooded nests, with exotrophic tadpoles in ter bodies (mode 1). This is also the most common ponds or streams (mode 5; Table 1). This increases mode in other open formations of South America, to 15 the reproductive modes known for the Cer- such as the Pantanal floodplain (Prado et al. 2005), rado (Table 2). We could not determine the repro- Caatinga (Vieira et al. 2009), Pampas (Moreira et ductive strategy of Hypsiboas sp. (Table 1). al. 2007, Santos et al. 2008), Chaco (Perotti 1997), The most common reproductive modes were 1 and other Cerrado areas (Kopp et al. 2010, Conte and 2, with aquatic reproduction and eggs and et al. 2013). The variety of reproductive modes in- exotrophic tadpoles in lentic (mode 1) and lotic volving foam nests (terrestrial and aquatic) con-

Figure 3. Some of the Cerrado endemic anurans from the Chapada dos Veadeiros. A- Hypsiboas ericae, B - Hypsiboas phaeopleura, C - Phyllomedusa oreades, D – Scinax aff. squalirostris, E – Scinax rogerioi, F – Barycholos ternetzi, G – Leptodactylus furnarius, H – Leptodactylus tapiti, I – Adenomera saci, J – Ameerega flavopicta, K – Rhinella rubescens, L – Odontophrynus salvatori.

Diversity and reproduction of Cerrado anurans 371

Table 2. Reproductive modes (sensu Haddad & Prado 2005) of the anuran species of the Chapada dos Veadeiros and other South American biomes.

Localities Reproductive modes TOTAL References Present study 1,2,4,5,11,13,20,23 - 25,29-32 14 Atlantic forest 1-6,8,10,11,13-15,18-25,27,28,30-32,36,37 27 Haddad & Prado 2005 Amazon 1,2,4,6,8,11,13,15,16,18,20,21,23-27,30,32,35-37 22 Haddad & Prado 2005 Cerrado 1-4,11,13,20,23-25,29-32 14 Brandão 2002, Vasconcelos & Rossa-Feres 2005, Costa et al. 2006 Kopp et al. 2010, Provete et al. 2011, Rodrigues et al. 2011, Conte et al. 2013 Caatinga 1,4,11,13,15,17,24,30,32 9 Vieira et al. 2009 Pantanal 1,3,8,18,21,22 6 Prado et al. 2005 Chaco 1-4 4 Perotti 1997 Pampas 1,11,24,30 4 Moreira et al. 2007, Santos et al. 2008

tributed largely to the number of reproductive entiation between these species in strategies for strategies (Table 1), and seems to be an interesting egg laying. pattern for Cerrado anuran communities. Terrestrial reproduction in anurans is usually In general, we found the same pattern of pre- associated with a high rate of predation in perma- vious studies on Cerrado, with species presenting nent water bodies in forested habitats (Magnusson high frequency of reproductive modes involving & Hero 1991, but see Haddad & Prado 2005, and eggs on lentic water with aquatic exotrophic lar- Gomez-Mestre et al. 2012). However, they can also vae and of terrestrial eggs with exotrophic aquatic be influenced by phylogenetic relationships (Et- larvae (Prado et al. 2005, Santos et al. 2008, Kopp erovick & Fernandes 2001, Kopp & Eterovick 2006, et al. 2010, Conte et al. 2013). Some authors sug- Gomez-Maestre et al. 2012), suggesting that re- gest that this pattern could be produced by lower productive modes could exhibit phylogenetic sig- availability of microhabitats for reproduction in nal in some lineages. In our study, leptodactylids open areas (Magnusson & Hero 1991, Haddad & showed three aquatic reproductive modes (modes Prado 2005). However, this pattern is also ex- 1, 11 and 13), or variations of terrestrial reproduc- pected for open and drier environments (Gomez- tion (modes 29 to 32). Even with lesser number of Mestre et al. 2012). Garda et al. (2013) suggested reproductive modes (6), hylids showed more plas- that the horizontal heterogeneity of the Cerrado, ticity in microhabitats for egg deposition as well as when compared to the vertical heterogeneity of perching preferences (Table 1). Tadpoles of hylid forested biomes, would maintain a high diversity have higher morphological variation than of habitats and ecological niches, and the high that of leptodactylids (Eterovick & Fernandes number of species with aquatic and “transitional” 2001), which can be related to the higher plasticity reproductive modes (i.e. arboreal and terrestrial in the use of aquatic habitats. The great variations eggs with aquatic exotrophic larvae; such as phyl- in external morphology of adult hylids enable lomedusinaes and Leptodactylus fuscus species them to utilize a greater variety of microhabitat for group, respectively). This represents a quite inter- perching and eggs deposition (Kopp & Eterovick esting situation for studies concerning the evolu- 2006, Wells 2007). Furthermore, despite the con- tion of reproductive modes in Amphibia (Gomez- servatism of reproductive modes displayed by Mestre et al. 2012). some genera or families (Gomez-Maestre et al. The horizontal heterogeneity of the Cerrado 2012), reproductive strategies may vary between could also explain the high diversity of closely re- populations of the same species (Toledo et al. lated Leptodactylus species presenting terrestrial 2012). breeding in the study area, such as Leptodactylus Recent studies showed that central and north- fuscus, L. furnarius (Fig. 3G), L. mystaceus, L. ern Cerrado regions are richer than previously mystacinus, L. sertanejo, L. tapiti (Fig. 3H), and Lep- suggested (Valdujo et al. 2011, 2012). Our results todactylus sp. Although these species present show that the Chapada dos Veadeiros have the variations on their terrestrial reproductive modes, highest richness recorded for any Cerrado locality. most of them are not found syntopically (Brandão Moreover, there are still some poorly sampled ar- et al. 2013), suggesting that habitat heterogeneity eas in this region, mainly in its northeastern por- can create some degree of diversification on re- tion (Fig. 1). Cerrado highlands exhibit high rates source use, which could account for some differ- of herpetofauna endemism (Nogueira et al. 2011, 372 G.R.C.C. Santoro & R.A. Brandão

Valdujo et al. 2012). However, the extent of the Acknowledgments. We would like to thank Renata D. Chapada dos Veadeiros National Park (about Françoso and Erika C. Gonzaga for the map. Carlos 65,000 ha) is not enough to protect species and Cândido provided several photos. Guarino R. Colli and Isis Arantes allowed access to the herpetological habitat diversity (Felfili et al. 2007). Like other collection from the Brasília University. Instituto Cerrado areas, several common threats for anuran PROAVES/IBAMA provided funding for some fieldwork conservation occurs in the region, such as wild- during the studies for the management plan of the fires (Rocha et al. 2008), agriculture (Mann et al. Chapada dos Veadeiros National Park in 1996. The 2009), deforestation for cattle ranching (Gallant et owners of the Fazenda Miraflores allowed access to their al. 2007) and charcoal production, hydroelectric property. dams (Brandão & Araujo 2008), and growing cities (Smallbone et al. 2012). Other possible threats in- clude the fungus Batrachochytrium dendrobatidis, References which was recently recorded in the region (Ra- malho et al. 2013). Bastos, R.P., Motta, J.A.O., Lima, L.P., Guimarães, L.D. (2003): Since several anuran populations recorded Anfíbios da Floresta Nacional de Silvânia, GO. Stylo Gráfica e Editora, Goiânia. here may represent undescribed cryptic species, a Bertoluci, J., Rodrigues, M.T. (2002): Utilização de habitats deeper taxonomic understanding is necessary to reprodutivos e micro-habitats de vocalização em uma known their real distributions and determine their taxocenose de anuros (Amphibia) da Mata Atlântica do Sudeste do Brasil. Papéis Avulsos de Zoologia 42: 287–297. conservation status (Dubois 2003). For example, Bokermann,W.C.A. (1975): Uma nova espécie de Colostethus do Allobates goianus is no longer found in its type lo- Brasil Central (Anura, Dendrobatidae). Iheringia Série Zoologia cality (Chapada dos Veadeiros) since its descrip- 46: 13–16. Brandão, R.A. (2002): A new species of Phyllomedusa Wagler, 1830 tion (Bokermann 1975). Therefore, the taxonomic (Anura: Hylidae) from Central Brazil. Journal of Herpetology identity of populations outside this region (Bastos 36: 571–578. et al. 2003, Brandão & Araújo 2008) is unclear. Fur- Brandão, R.A., Araújo, A.F.B. (2001): A herpetofauna associada às matas de galeria no Distrito Federal. pp. 561–604. In: Ribeiro, thermore, Scinax aff. squalirostris (Fig. 3D) from the J.F., Fonseca, C.E.L. & Sousa-Silva, J.C. (eds), Cerrado: study area and other Cerrado regions seems to Caracterização e Recuperação de Matas de Galeria. Planaltina, represent a still undescribed species (Faria et al. Embrapa Cerrado. Brandão, R.A., Araújo, A.F.B. (2008): Changes in anuran species 2013). richness and abundance resulting from hydroeletric dam Open physiognomies are quickly altered be- flooding in central Brazil. 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