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Reproductive Biology of the Amazonian Treefrog Dendropsophus Walfordi (Bokermann, 1962) (Anura, Hylidae)

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Reproductive Biology of the Amazonian Treefrog Dendropsophus Walfordi (Bokermann, 1962) (Anura, Hylidae) Herpetology Notes, volume 14: 973-976 (2021) (published online on 07 July 2021) Reproductive biology of the Amazonian treefrog Dendropsophus walfordi (Bokermann, 1962) (Anura, Hylidae) Aline Emanuele Oliveira-Souza1, Maria Madalena Salviano Santana1, Patrick Ribeiro Sanches2,*, Patrick de Castro Cantuária3, and Carlos Eduardo Costa-Campos1 The Neotropical treefrog genus reproductive biology of D. walfordi from the Amazon Dendropsophus Fitzinger, 1843 is one of the most Forest in northern Brazil. taxonomically complex genera of hylids due to its We sampled D. walfordi in the Toca da Raposa high intraspecific variation and high morphological Farm (0.1500°N, 51.0384°W), located in the River similarity among most species (Fouquet et al., 2011). Curiaú Environmental Protection Area, municipality Dendropsophus comprises 109 species (Frost, 2020) of Macapá, state of Amapá. The region belongs to divided into in nine species groups (Orrico et al., 2021) the Amazon rainforest, and has an average annual that exhibit a diversity of reproductive modes with either precipitation of 2,600 mm/year and temperature of 27 aquatic (Touchon and Warkentin, 2008) or terrestrial °C. The driest period is between June to November and nests (Crump, 1974; Hartmann et al., 2010; Mageski et the highest precipitation occurs between December and al., 2014; Silva et al., 2019), and aquatic or terrestrial May (Souza and Cunha, 2010). spawns (Miranda et al., 2008; Zina et al., 2014). Between May 2013 and April 2014, we made 12 Studies on the reproductive biology of Dendropsophus field trips, each lasting three days (36 observation days have been conducted in the Atlantic Forest biome (e.g., in total), to a temporary pond of 120 m2 in size. Pond Miranda et al., 2008; Silva et al., 2019), Araucaria forest vegetation are predominately Poaceae and Cyperaceae (Leivas et al., 2018), and in Panama (Touchon and species and shrubby vegetation reaching up to 1.5 m. Warkentin, 2008), but such information is scarce for We conducted the surveys between 18:30-20:30 h and species of the Amazon Forest biome. Dendropsophus recorded all individuals heard or sighted during the walfordi (Bokermann, 1962) is one of the 33 members search period. The sampling procedure consisted of of the Dendropsophus microcephalus group (Faivovich visual and auditory searching (Crump and Scott Jr., et al., 2005), with a distribution restricted to the 1994). Amazon basin (Teixeira, 2018). The lack of natural Satellite males were recognised by their peripheral history information for D. walfordi from less-explored positions to calling males, adopting alternate mating regions, such as northern Brazil, precludes a better tactics in an attempt to intercept females that are understanding of its biology, ecology and taxonomy attracted to calling males (Wells, 2007). To verify (Fouquet et al., 2011). Here, we present data on the spawning characteristics, we placed pairs in amplexus in water-filled plastic bags (a single pair per plastic bag) with a water depth of ~3 cm, and an upper air layer of ~20 cm. From this method, we obtained five clutches, which were preserved in 5% formalin. The eggs of 1 Laboratório de Herpetologia, Departamento de Ciências clutches were counted and the diameters of 30 eggs Biológicas e da Saúde, Universidade Federal do Amapá, per spawn were measured using a stereomicroscope Campus Marco Zero do Equador, Macapá, Amapá 68903- and lens micrometre. We measured snout-vent length 419, Brazil. 2 Instituto Nacional de Pesquisas da Amazônia, Avenida André (SVL) of the females corresponding to the clutches with Araújo, Manaus, Amazonas 69011-970, Brazil. a Mitutoyo digital calliper (0.01 mm precision). 3 Instituto de Pesquisas Científicas e Tecnológicas do Estado do The climate data were obtained from Núcleo de Amapá, Avenida Juscelino Kubitschek, Fazendinha, 68.903- Hidrometereologia e Energias Renováveis (NHMET), 970, Macapá, AP, Brazil. Instituto de Pesquisas Científicas e Tecnológicas do * Corresponding author. E-mail: [email protected] Estado do Amapá (IEPA) database. The collected © 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. treefrogs were euthanised by topical application of a 2% 974 Aline Emanuele Oliveira-Souza et al. liquid solution of lidocaine hydrochloride, fixed in 10% formalin, preserved in 70% ethanol and deposited in the Herpetology Collection of Universidade Federal do Amapá under the acronomy CECC (0635-0681; 0901- 0919). The collection permits were provided by Sistema of Autorização and Informação in Biodiversidade– SISBIO (license number # 41586-1). We used Pearson’s correlation coefficient to test the possible relationship between SVL and the number and mean diameter of the eggs observed in each female, and male calling activity and rainfall (Zar, 1999). Sex-specific differences in SVL were tested with the Student’s t-test. Presented data concern means and Figure 1. Adult males, females, satellite males and gravid standard deviations. All data were tested for normality females of Dendropsophus walfordi recorded between May via a Kolmogorov–Smirnov test, and analyses were 2013 and August 2014 in Toca da Raposa Farm, River Curiaú performed using the Bioestat 5.0 software (Ayres et al., Environmental Protection Area, state of Amapá, Brazil, in relation to total monthly rainfall. Numbers above the months 2007). For all tests, the significance threshold was p ≤ represent number of eggs from clutches obtained. The number 0.05. of eggs for February represent a mean of two clutches, the We collected a total of 64 D. walfordi individuals: 28 remaining values are single clutches. females and 36 males. Eight of these individuals were gravid females, five pairs in amplexus and five satellite males. We observed gravid females and spawns of D. walfordi from February to May (Figure 1). Females were larger in SVL (21.1 mm ± 1.4; range = 19.1–22.9; n = 28) than males (18.2 mm ± 0.4; range = 17.4–18.9; n = 36; t = 12.6; p = <0.0001). Males used the vegetation bordering temporary ponds as calling sites: Cyperaceae (n = 19; 52.8%), Poaceae (n = 12; 33.3%) and Rubiaceae (n = 5; 13.9%). The perch height ranged from 25 to 78 cm above the water surface. We observed females (n = 11; 39.3%) and satellite males (n = 5; 13.9%) near of vocalising males on perches. Satellites males (SVL = 17.2 mm ± 0.4; range 16.5– Figure 2. Minimum distance (cm) between Dendropsophus 17.4) were not significantly larger (t = 0.99; p = 0.35) walfordi males during the calling activity recorded in Toca than vocalising males. Despite the spatial distribution da Raposa Farm, River Curiaú Environmental Protection of the calling sites around the temporary ponds, 56% Area, state of Amapá, Brazil. Black bars represent distances of the calling males were found separated by 20–40 to satellite males and grey bars represent distances to nearest cm from each other. The spatial distribution reflects calling male. the social dominance of the males in the population: territories located at the edge of the pond were occupied by calling males, while satellite males were stationed on reproductive mode and spawning similar in appearance. the peripheral or centrally located territories in the pond. The number of eggs per clutch ranged from 352 to 462 At these distances, male-satellite male interactions were (x̄ = 403 mm ± 41.6; n = 5 clutches), and the diameter observed (Fig. 2). of eggs varied from 0.8 to 1.1 mm (x̄ = 0.9 mm ± 0.1; Five clutches were obtained in plastic bags (n = 5 n = 30 eggs). There was a positive and non-significant pairs in amplexus) and placed in the water. Clutches relationship between female SVL and the number of were not collected in the field due to other species eggs (r2 = 0.83; p = 0.07) and diameter of eggs (r2 = with reproductive activity in the same pond: Scinax 0.43; p = 0.47), was observed. The number of calling ruber (Laurenti, 1798) and Elachistocleis helianeae males was correlated with daily rainfall (r2 = 0.84; p = Caramaschi, 2010; these species have the same 0.001). Reproductive biology of the Amazonian treefrog Dendropsophus walfordi 975 Individuals of D. walfordi reproduced from February (ICMBio) for the research license (41586-1 #). And we are to May, characterising a prolonged breeding pattern grateful to Mateus Ramos who permitted accesses to his farm. (sensu Wells, 1977) in which females attend the choruses in an extremely brief and asynchronous, while References males remain in reproductive activity (calling) during Ayres, M., Ayres Jr., M., Ayres, D.L., Santos, A.S. (2007): BioEstat the whole reproductive season. Males were territorial 5.0. Aplicações estatísticas nas áreas das Ciências Biológicas e during the breeding activity and defended calling sites Médicas. Sociedade Civil de Mamirauá, Belém, Brazil. through acoustic and physical interactions. However, Bokermann, W.C.A. (1962): Nova espécie de Hyla de Rondônia, the number of these interactions was low and may be Brasil (Amphibia, Salientia). Atas da Sociedade de Biologia do Rio de Janeiro 6: 52–55. related to the abundant availability of calling sites, and Crump, M.L. (1974): Reproductive strategies in a tropical anuran low population density in the temporary pond. community. University of Kansas Museum of Natural History The reproductive period of anurans from tropical Miscellaneous Publications 61: 1–68. seasonal habitats is strongly influenced by rainfall Crump, M.L., Scott Jr., N.J. (1994): Visual encounter surveys. (Gottsberger and Gruber, 2004; Wells, 2007). Male adult In Heyer, W.R., Donnelly, M.A., McDiarmid, R.W., Hayek, D. walfordi were found in vocal activity throughout the L.A.C., Foster, M.S. (Eds), Measuring and Monitoring months of January to May (rainy season). No individuals Biological Diversity, Standard Methods for Amphibians, pp. 84–92. Smithsonian Institution Press, Washington DC. were observed during the period of June to December, Crump, M.L.
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