(Anura: Hylidae) in the Brazilian Eastern Amazon
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Herpetology Notes, volume 13: 599-602 (2020) (published online on 05 August 2020) Physical combat of two species of Phyllomedusidae (Anura: Hylidae) in the Brazilian eastern Amazon Marcos R. Dias-Souza²,*, Vinícius A. M. B. de Figueiredo1, Lorena F. Souza Tavares-Costa²,³, and Carlos Eduardo Costa-Campos¹ Territorially is defined as a place where a species Phyllomedusa and Pithecopus (Duellman et al., 2016; has access to primary resources. The defence of these Frost, 2020). Previous studies registered the aggressive areas has two benefits: the first is protecting resources behaviour between males in the genera Agalychnis, and breeding success (Kaufmann, 1983; Shi-Ryong for Agalychnis callidryas (Pyburn, 1970; Gonzalez and Cheong, 2002). Territorial behaviour has observed and Briggs, 2011), A. moreletii (Serrano et al., 2018); in distinct animal groups (e.g. insects: Baker, 1983; Phyllomedusa, for Phyllomedusa boliviana (Jansen and fishes: Barlow, 1974; amphibians: Wells, 1977; reptiles: Köhler, 2008), P. sauvagii (Rodrigues et al., 2003), P. Stamps, 1983; birds: Brown, 1964; mammals: Ostfeld, burmeisteri (Wogel et al., 2004) and P. iheringii (Dias et 1990). al., 2017); and Pithecopus, for Pithecopus azureus (Costa In frogs, most territories are defended against intruder et al., 2010), P. hypochondrialis (Matos et al., 2000), P. males that utilised the aggressive behaviour for a nordestinus (Vilaça et al., 2011), P. rohdei (Abrunhosa protecting resource. These interactions are determined and Wogel, 2004), P. megacephalus (Oliveira et al., by size, residence and body condition, and involve 2011), and P. ayeaye (Oliveira, 2017). Here, we report a combination of physical combat and acoustic and two physical combat events by Phyllomedusa bicolor visual displays (Shi-Ryong and Cheong, 2002; Wogel (Boddaert, 1772) and Pithecopus hypochondrialis et al., 2004; Méndez-Narváez and Amézquita, 2014; (Daudin, 1800) in the Amapá state, Brazilian eastern Reichert and Gerhardat, 2014). In general, intraspecific Amazon. interaction is more common due to the high overlap in Phyllomedusa bicolor (Boddaert, 1772) is found in the the use of resources and has already been observed in Amazonian forests of Brazil, the Guianas, Venezuela, the Phyllomedusidae family (Jansen and Köhler, 2008; Colombia, Peru and Bolivia (Frost, 2020). It is one of the Peiman and Robinson, 2010). largest tree frogs of the Amazon (males measure 91-103 The Phyllomedusidae family comprises 63 species mm snout-vent length [SVL] and females 111-119 mm). distributed in eight genera: Agalychnis, Callimedusa, The species is arboreal and nocturnal, characterised by Cruziohyla, Hylomantis, Phasmahyla, Phrynomedusa, dorsum green with lateral region and the belly varying from white to yellow-white or cream, a prominent gland extends from behind the eye over the tympanum (Lima et al., 2006). Pithecopus hypochondrialis (Daudin, 1800) occurs in pond areas with grasses and widely distributed in 1 Universidade Federal do Amapá, Departamento de Ciências Colombia, Venezuela, Guianas, Amazonian Brazil from Biológicas e da Saúde, Laboratório de Zoologia, Macapá, states: Pará, Mato Grosso, Goiás, and Tocantins (Cole et Amapá 68.903-419, Brazil. al., 2013; Frost, 2020). The males measure 31-46.5 mm ² Universidade Federal do Pará, Instituto de Ciências SVL and females 40-53 mm. This species are identified Biológicas, Laboratório de Biologia Celular e Helmintologia by dorsum green, interior portion of members with “Prof. Dra Reinalda Marisa Lanfredi”, Belém, Pará 66.075- cream stripe; posterior half of flanks, upper arms, things 110, Brazil. ³ Museu Paraense Emílio Goeldi, Programa de Pós-graduação an feet bright orange with dark blue or purple bars; em Zoologia, Avenida Perimetral, 1901, Belém, Pará, 66077- abdomen and ventral surfaces of thighs is pale yellow 830, Brazil. to dark orange (Caramaschi, 2006; Cole et al., 2013; * Corresponding author. E-mail: [email protected] Duellman et al., 2016). 600 Marcos R. Dias-Souza et al. On 25 May 2017, two males of P. bicolor were The physical interaction observed in P. bicolor found in aggressive behaviour (Fig. 1) in the Cancão differs from that recorded in P. boliviana from Bolivia, Municipal Natural Park, municipality of Serra do which has been described by Jansen and Köhler Navio, Amapá state (0.9008°N, 52.0134°W; WGS84). (2008) as a combination of slow movement and visual The males were observed at the top of a shrub, 1.5 m communication, resulting in an axillary amplexus. This away from us. Male A grabbed the other by the hind type of amplexus has also been observed by Najar and limbs, while male B threw its arms into the bush to Ferrante (2018) in C. tomopterna. On the other hand, our escape. The individuals moved slowly and both emitted observations are quite similar to the physical interactions calling during the interaction. The physical combat observed in P. burmeisteri by Wogel et al. (2004) and resulted in an amplexus in the inguinal region and the which resulted in amplexus in the inguinal region and males remained in this position for approximately 10 physical encounters lasting from 20 to 80 min. minutes, until they separated. We did not interfere in the The physical combat by P. hypocondrialis has been interaction, we just taking the photography. previously described by Matos et al. (2000) in males On 29 April 2018, we observed two males of P. suspended from branches and supported only by the hypochondrialis in physical combat on the side edge of finger of one of the males, with the loser falling to abandoned pool located in the urban area, municipality the ground. However, our observation differs from of Serra do Navio (0.8984°N, 52.0018°W; WGS84; the authors because the individuals were found in 144 m a.s.l.), Amapá state. During observation, the physical combat already on the ground. In anurans from individuals were found embraced ventrally in the Phyllomedusidae family, the proximity between males ground. The dorsum of one male was on the ground is caused by competition for territory (Wogel et al., with the legs clasped with the other male, a similar 2004; Toledo and Haddad, 2005; Vilaça et al., 2011). movement to guillotine at the jiu-jitsu (Fig. 2). During Therefore, physical interactions have consequences for this physical combat both frogs did not vocalise. The males because it demanding energy expenditure, cause specimens remained in the same position for 30 minutes injuries and increase vulnerability to predation (Vilaça after initial observation, until they separate. et al., 2011). According Haddad and Prado (2005), the reproductive In the study area from May 2017 to April 2018, modes of Phyllomedusidae family fit into Modes 18, shorter distances were recorded between individuals 24 and 25, which are characterised by oviposition in of P. bicolor and P. hypocondrialis, as well as a shorter lentic or lotic water environments. It is exactly close to distance between the calling sites and the nearest male. these environments that males defend their territories, This observation suggests that there is a brief breeding increasing the chances of amplexus, oviposition and period in which males gather around temporary combats (Vilaça et al., 2011). ponds and defend territories, which would increase Figure 1. Males of Phyllomedusa bicolor in physical combat Figure 2. Males of Pithecopus hypochondrialis in physical in the Cancão Municipal Natural Park, municipality of Serra combat in the urban area, municipality of Serra do Navio, do Navio, Amapá state, Brazilian Eastern Amazon. Amapá state, Brazilian Eastern Amazon. Physical combat of two species of Phyllomedusidae, Brazilian eastern Amazon 601 the occurrence of physical interactions. Thus, these Gonzalez, S.C., Briggs, V.S. (2011): Aggression in froglets of red- results suggest the possible existence of a reproductive eyed treefrogs, Agalychnis callidryas. Herpetology Notes 4: seasonality, corroborating Villaça et al. (2011) for 315–318. Haddad, C.F.B., Prado, C.P.A. (2005): Reproductive modes in Pithecopus nordestinus (Caramaschi, 2006). frogs and their unexpected diversity in the Atlantic forest of The physical interactions in different species of Brazil. BioScience 55: 207–217. Phyllomedusidae are similar, except in P. boliviana, Jansen, M., Köhler, J. (2008): Intraspecific Combat Behavior of indicating that this behaviour is a synapomorphy of Phyllomedusa boliviana (Anura: Hylidae) and the Possible the group (Abrunhosa and Wogel, 2004; Wogel et al., Origin of Visual Signaling in Nocturnal Treefrogs. Herpetological 2004; Costa et al., 2010). Therefore, our observations Review 39: 290–292. contribute for the knowledge of reproductive biology Kaufmann, J.H. (1983): On the definitions and functions of dominance and territoriality. Biological Reviews 58: 1–20 and agonistic interactions of P. hypochondrialis and P. Lima, A.P., Magnusson, W.E., Menin, M., Erdtmann, L.K., bicolor from Brazilian eastern Amazon. Rodrigues, D.J., Keller, C., Hodl, W. (2006): Guia de sapos da Reserva Adolpho Ducke, Amazônia Central, 2nd Edition. Acknowledgements. We thanks to Gilda Vasconcelos for Manaus, Áttema Design Editorial. valuable suggestions on the manuscript. Additionally, we grateful Matos, R.H.R., Andrade, G.V., Hass, A. (2000): Reproductive to Instituto Chico Mendes de Conservação da Biodiversidade Biology and Territoriality of Phyllomedusa hypochondrialis in (ICMBio #48102-2) for providing collection permits and Northeastern Brazil. Herpetological Review 31: 84–86. Christoph Jaster