Bioacoustics of Dendropsophus haraldschultzi

Natural history. The couple of Dendropsophus haraldschultzi in amplexus was found in a floodplain, perched on aquatic macrophytes ( Montrichardia sp.) 69 cm from the water surface. Snout-vent length and body mass were 19.72 mm and 0.4 g for the male, and 22.62 mm and 0.6 g for the female. The amplexus is axillary. Clutch size was 70 eggs (one clutch); eggs were white and clumped in a gelatinous mass in the water surface. We observed males of D . haraldschultzi , D. leucophyllatus and Scarthyla goinorum (Bokermann) in calling activity at night (between 18:30 to 20:00 h) in the same site where the couple was found. The advertisement call of D. haraldschultzi (not recorded) is composed of two types of pulsed notes with similar dominant frequencies emitted separately (only one individual was observed). Characters Stages 25 (N=12) 31 (N=1) 34 (N=1) 35 (N=2) 36 (N=2) TL 7.38±0.23 (6.80–7.60) 20.96 20.16 23.52–25.12 25.28–26.72 BL 2.69±0.09 (2.56–2.88) 6.88 7.04 7.84–8.00 8.64–8.80 BW 1.70±0.08 (1.52–1.84) 3.68 3.84 4.00–4.16 4.80–5.12 BH 1.60±0.04 (1.52–1.68) 3.20 3.20 3.36–3.68 4.16-4.16 TAL 4.68±0.23 (4.16–4.96) 14.08 13.12 15.68–17.12 16.64–17.92 MTH 1.27±0.08 (1.20–1.44) 3.20 3.30 4.00–4.00 4.00–4.50 TMH 0.72±0.04 (0.64–0.80) 2.50 2.50 2.70–2.70 2.60–3.00 TMW 0.62±0.06 (0.56–0.72) 1.70 1.80 1.90–2.00 2.10–2.10 IND 1.07±0.04 (1.04–1.12) 2.50 2.75 2.90–3.00 3.00–3.30 IOD 1.46±0.09 (1.20–1.52) 3.50 3.65 4.10–4.30 4.25–4.60 ED 0.48 1.00 1.15 1.15–1.30 1.30–1.35 END 0.48±0.06 (0.40–0.56) 1.00 1.05 1.20–1.35 1.35–1.50 NSD 0.40±0.04 (0.32–0.48) 0.80 0.80 0.75–0.80 0.70–0.80 VTL 0.25±0.05 (0.16–0.32) 0.75 0.80 1.10–1.10 1.00–1.00 SL 0.77±0.07 (0.72–0.96) 1.50 1.60 2.05–2.25 1.80–1.85 ODW 0.47±0.04 (0.40–0.56) 1.07 1.07 1.12–1.15 1.17–1.27 The tadpoles of Dendropsophus haraldschultzi at stages 31 and 34 to 36 are morphologically similar to what was illustrated by Lynch & Suárez-Mayorga (2011) as the tadpole of D. haraldschultzi from Colombia . Forty-six Dendropsophus tadpole species from six species groups were described and/or figured previously ( Bokermann 1963 ; Duellman & Fouquette 1968 ; Kenny 1969 ; Duellman 1970 , 1972 , 1978 , 2005 ; Duellman & Crump 1974 ; Weygoldt & Peixoto 1987 ; Hero 1990 ; Heyer et al. 1990 ; Lavilla 1990 ; Cruz & Dias 1991 ; Gomes & Peixoto 1991a , 1991b ; Wild 1992 ; Santos et al. 1998 ; Peixoto & Gomes 1999 ; Cruz et al. 2000 ; Wogel et al. 2000 ; Pugliese et al. 2000 , 2001 ; Carvalho-e-Silva et al. 2003 ; Lynch 2006 ; Rossa-Feres & Nomura 2006 ; Lynch & Suárez-Mayorga 2011 ; Lourenço-de-Moraes et al. 2012 ; Abreu et al. 2014 , 2015 ; Fouquet et al. 2015 ; Schulze et al. 2015 ; Ruas et al. 2018 ). Tadpoles of D. haraldschultzi can be easily differentiated from all of them by its long, wide, and lateroventral spiracle (spiracle is sinistral and short in the remaining species). An elongate spiracle was reported in tadpoles of other hylids, including species of Boana albopunctata Spix group ( Wild 1992 ; Rossa-Feres & Nomura 2006 ) and Sphaenorhynchus Tschudi (e.g., Nunes et al. 2007 ; Araujo- Vieira et al. 2015), and in the microhylid genus Otophryne Boulenger (e.g., Pyburn 1980 ). Like in Sphaenorhynchus species, tadpoles of D. haraldschultzi are found in the root of aquatic macrophytes in floating meadows (Bönning et al. 2017), but functional aspects of the long spiracle are unknown ( Araujo-Vieira et al. 2015 ). In most general features, e.g., small size, snout rounded, oral disc anterior with mouthparts usually reduced, Dendropsophus haraldschultzi larvae are similar to other tadpoles of the genus. In some individual traits however these tadpoles resemble larvae of different species groups. For instance, the post-ocular constriction that gives the body a violin-shape occurs also in most tadpoles of the D. leucophyllatus group [ D. arndti Caminer, Milá, Jansen, Fouquet, Venegas, Chávez, Lougheed, and Ron , D. bifurcus (Anderson) , D. elegans (Wied-Neuwied) , D. leucophyllatus , D. reticulatus (Jiménez de la Espada) , D. salli Jungfer, Reichle, and Piskurek , D. sarayacuensis (Shreve) , and D. triangulum (Günther) ], species of the D. microcephalus group [i.e., D. branneri (Cochran) , D. microcephalus , D. nanus (Boulenger) , D. rubicundulus (Reinhardt and Lütken) , D. sanborni (Schmidt) , D. studerae (Carvalho-e-Silva, Carvalho-e-Silva and Izecksohn)], within the D. marmoratus group [i.e., D. melanargyreus (Cope) ], the D. parviceps group [i.e., D. timbeba (Martins and Cardoso) ], and in D. minutus . The tail tip without a flagellum is also characteristic of D . decipiens (Lutz) , D . haddadi , D . oliveirai (Bokermann) , and D. riveroi (Cochran and Goin) of the D. microcephalus group, and D. microps (Peters) and D. ruschii (Weygoldt and Peixoto) of the D. parviceps group. The configuration of the oral disc is widely variable in Dendropsophus , and varied extents of oral structure reductions are reported in several groups (e.g., Duellman 1978 , 2005 ; Schulze et al. 2015 ). On one hand, marginal papillae reduced to lateral commissures on upper lateral portion and lower labium appear in D. haraldschultzi and most species of the D. leucophyllatus group ( D. arndti , D. ebraccatus (Cope) , D . elegans , D. rossalleni (Goin) , D. sarayacuensis , D. salli ), and D. parviceps group [ D. giesleri (Mertens) , D. microps ]. The fusion of lateral marginal papillae forming a dermal vertical fold on both sides of oral disc has been described only in D. counani Fouquet, Orrico, Ernst, Blanc, Martinez, Vacher, Rodrigues, Ouboter, Jairam, and Ron. On the other hand, the number of tooth rows in the genus varies from 0/0 in many species of the D. leucophyllatus , D. marmoratus , D. minutus , D. parviceps and D. microcephalus species groups to 2/ 2 in D . anceps (Lutz) . Besides in D . haraldschultzi , a LTRF 0/1 has been reported in D . elegans , D . rossalleni , D. salli ( D. leucophyllatus group), D. giesleri , D. microps , and D. ruschii of the D. parviceps group; in D. melanargyreus , D. nahdereri (Lutz and Bokermann) , D. novaisi (Bokermann) , D. seniculus (Cope) , and D. soaresi (Caramaschi and Jim) of the D. marmoratus group, and in D. minutus . In addition, further labial ridges without labial teeth are found also in some species of the D. microcephalus group (one ridge in D haddadi , two ridges in D. berthalutzae and D. decipiens ). This combination of characters indicates that larval morphology is ambiguous to assign D. haraldschultzi to any species group. As a final comment, clutch size of Dendropsophus haraldschultzi found in our study is smaller than those reported in the literature for gravid females with mature ovarian eggs ( 216 eggs ; Hödl 1990 ). Most species of Dendropsophus occurring in Amazonia lay their eggs on the water, on the surface of leaves of emergent vegetation [ D. bifurcus , D. bokermanni (Goin) , D. brevifrons (Duellman and Crump) , D. reticulatus , D. sarayacuensis , D. timbeba and D. reticulatus ]. Conversely, D. haraldschultzi is similar to D. koechlini (Duellman and Trueb) , D. leali (Bokermann) , D. marmoratus , D. minutus , and D. parviceps in that they deposit the eggs directly in the water ( Duellman 1978 , 2005 ; Rodríguez & Duellman 1994 ).