Herpetology Notes, volume 11: 671-673 (2018) (published online on 24 August 2018)

Combat behaviour in the glassfrog uranoscopa (Müller, 1924)

Igor Yuri Fernandes1,*, José Flávio Cândido-Junior2, and Igor Luis Kaefer1,3

The family Centrolenidae is composed by 155 anuran Thus, the authors emphasized the need for behavioural (Frost, 2018), popularly known as glassfrogs. observations from additional species. Territoriality of males occasionally involves combat On 10 February 2017, we observed two males of behaviours, but these have been poorly documented (Müller, 1924) in combat for this family (Rojas-Runjaic and Cabello, 2011). To behaviour on the vegetation at the banks of a stream near date, this behaviour was reported for only 15 species a waterfall. The site is located in the Centro de Educação (Rojas-Runjaic and Cabello, 2011; Dautel et al., 2011, Ambiental Suely Festugatto, locally known as Parque Cardozo-Urdaneta and Señaris, 2012), including one Ambiental de Cascavel (24.5949ºS, 53.1733ºW), in the case of interspecific combat (Sorokin and Steigerwald, Municipality of Cascavel, Paraná State, southern . 2017). Ruiz-Carranza and Lynch (1991) proposed two Vitreorana uranoscopa is a centrolenid of the subfamily distinct types of combat behaviours for centrolenids: a , usually found in Atlantic Forest habitats basal (known as primitive) in which the fight takes place in eastern Brazil from Minas Gerais and Espírito Santo through amplexus-like (pseudoamplectant) positions on States to the north of Rio Grande do Sul State, and also the leaf surface, and a second type (known as derived), in northeastern Argentina (Frost, 2018). characterized by males grappling venter-to-venter, The sequence of behaviours reported here lasted 16 dangling upside down from the vegetation. min (20:41 h to 20:57 h) and occurred between a larger This generic classification of combat behaviours was male (male A), which was vocalising on the vegetation later proposed as a taxonomic character at the subfamily next to an egg clutch (ca. 50 cm; Fig. 1A) and a smaller level, the basal type being restricted to the subfamily male (male B), which was approximately 3 meters away and the derived type to the subfamily from male A, both on a tree branch hanging ca. 250 cm Centroleninae (Guayasamin et al., 2009). A later review over a stream (males were not measured, but their sizes indicated that both types of combat behaviour occur were visually different). When male B started vocalising, in at least four species of centrolenids, suggesting that male A turned his body towards male B, and displayed the taxonomic and evolutionary value of this character an upright posture of forearms and hindlimbs on the leaf should be interpreted with caution considering the few (Fig. 1B). After 3 min of vocalisation emitted by male species observed (Rojas-Runjaic and Cabello, 2011). B, male A started to climb the branches towards male B. When both males met, they grappled venter-to-venter with their forearms and dangled upside down from the twig stuck by their feet (Fig. 1C). Such behaviour lasted 1 Programa de Pós-Graduação em Ecologia, Instituto Nacional for about 1 min, when the males fell from the vegetation, de Pesquisas da Amazônia, Av. André Araújo 2936, 69060- became briefly separated and then grappled each other 001, Manaus, Amazonas, Brazil. over the lamp of one of the observers (Fig. 1D). After 2 Laboratório de Ecologia e Biologia da Conservação, Centro about 30 s, male B vocalised, and male A climbed over de Ciências Biológicas e da Saúde, Universidade Estadual male B, performing an inverted pseudoamplectant do Oeste do Paraná, Rua Universitária 2069, 85819-110, position (Fig. 1E), probably attempting to subjugate Cascavel, Paraná, Brazil. male B. Shortly after, male A released male B, and both 3 Instituto de Ciências Biológicas, Universidade Federal do Amazonas, Av. General Rodrigo Octávio 6200, 69080-900, jumped back to a leaf. Male B did not vocalise after Manaus, Amazonas, Brazil. that moment, and male A vocalised while chasing his * Corresponding author. ������� ������������������������� opponent, which moved through the vegetation in the 672 Igor Yuri Fernandes et al.

Figure 1. Sequence of combat behaviour stages between two males of the glassfrog Vitreorana uranoscopa in southern Brazil. (A- B) Male A at rest and emitting advertisement vocalisations, respectively. (C) Males A and B grappled venter-to-venter dangling from the vegetation. (D-E) Males in pseudoamplectant position after falling from the vegetation on the observer’s lantern. (F) Winner male (male A) calling after the end of the combat. Combat behaviour in the glassfrog Vitreorana uranoscopa 673 opposite direction of the nest and male A, leaving the glassfrogs (Amphibia: Centrolenidae) and their sister taxon dispute. After a period of ca. 10 min, male A resumed Allophryne ruthveni. Zootaxa 2100: 1–97. calling near the nest and close to its initial position (Fig. Rojas-Runjaic, F.J.M., Cabello, P. (2011): daidaleum (Ruiz-Carranza & Lynch, 1991) (Anura, Centrolenidae): A 1F). glassfrog with primitive and derived combat behavior. ������� Vocalisations during male-male combats are usually 2833: 60-64. reported for Centrolenidae (Cardozo, 2012), when males Ruiz-Carranza, P.M., Lynch, J.D. (1991): Ranas Centrolenidae emit low-frequency, territorial calls similar to a “preep”, de Colombia I. Propuesta de una nueva clasificación genérica. unlike their advertisement calls which are characterized Lozania 57: 1–30. by shorter or sharper notes similar to a “peep” (Rojas- Sorokin, A., Steigerwald, E. (2017): Interspecific combat between Runjaic and Cabello, 2011). In our observations, aff. grandisonae and prosoblepon. Herpetology Notes 10: 283-285. males of Vitreorana uranoscopa vocalised only when physically separated, both side-by-side and during the chase. The lack of calls or vocal sac inflations during both venter-to-venter and pseudoamplectant positions may be one of the factors that make combat behaviour in this family inconspicuous and rarely observed in the field. Our observation constitutes the first record of combat behaviour for the widely distributed Vitreorana uranoscopa, and the fifth record of both types of combat behaviour for a centrolenid , besides the four species reported by Rojas-Runjaic and Cabello (2011). However, knowledge on this topic is still scarce compared to the 155 species that currently compose the family. Hence, records from additional taxa within Centrolenidae are needed to gain insight about the evolutionary and taxonomic significance of this character.

Acknowledgments. Special thanks to Luís F. Andrietti for his fellowship and help during field observations. To Marcelo V. B. Garcia for the assistance in translating the manuscript and for his precious friendship, and for all the friends of the Laboratory of Ecology of Unioeste for the support and discussions on ecology and behaviour.

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