Zootaxa 4375 (2): 296–300 ISSN 1175-5326 (print edition)

http://www.mapress.com/j/zt/
Copyright © 2018 Magnolia Press
Correspondence ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4375.2.10
http://zoobank.org/urn:lsid:zoobank.org:pub:071DCC19-CEF5-4F22-A90C-2597EAC0F221

The tadpole of Dendropsophus novaisi (Bokermann, 1968) (Anura: Hylidae),

with comments on natural history

DANILO SILVA RUAS1, CAIO VINICIUS DE MIRA-MENDES2,3,6, MARIA LÚCIA DEL-GRANDE4,

JULIANA ZINA5& MIRCO SOLÉ2
1
Universidade Estadual do Sudoeste da Bahia, Campus Itapetinga, Praça Primavera, 40 - Bairro Primavera, CEP 45700-000,
Itapetinga, Bahia, Brasil
2
Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, km 16, 45662-900 Ilhéus,
Bahia, Brasil
3
Programa de Pós-Graduação em Sistemas Aquáticos Tropicais, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, km 16,
45662-900 Ilhéus, Bahia, Brasil.
4
Departamento de Ciências Naturais, Universidade Estadual do Sudoeste da Bahia, Estrada do Bem Querer, km 4, 45083-900 Vitória
da Conquista, Bahia, Brasil
5
Departamento de Ciências Biológicas, Universidade Estadual do Sudoeste da Bahia, Rua José Moreira Sobrinho, s/n, 45206190
Jequié, Bahia, Brasil
6
Corresponding author. E-mail: caioviniciusmm@gmail.com

Several frog species descriptions from this century provide larval morphology, acoustical, molecular, ecological and
behavioral information that, together, provide a better species characterization (e.g., Glos et al. 2005; Toledo et al. 2010;
Vassilieva et al. 2016). For the majority of anuran species, basic information is still lacking, despite being fundamental to
understand the evolution of a taxonomic unit as well as phylogenetic relationships established between taxa. Such
information can be particularly useful for species groups with unresolved taxonomic issues, such as the Dendropsophus
marmoratus group (Bokermann 1964; Heyer 1977; Caramaschi & Jim 1983; Gomes & Peixoto 1996).
The Dendropsophus marmoratus species group currently includes eight species occurring throughout South
America (Faivovich et al. 2005; Frost 2017). Species in this group show features like warty skin around the margin of the
lower lip, limbs with crenulated margin, a marbled pattern on the dorsum, a large vocal sac, and a thick sheath of tissue at
the basis of the tail musculature and fins of the tadpole that may represent possible morphological synapomorphies of the
group (Bokermann 1964; Altig & McDiarmid 1999; Faivovich et al. 2005; Orrico et al. 2009; Hepp et al. 2012). Species
of this group show an explosive breeding pattern, calling mainly after heavy rainfall (Bertoluci 1998; Canelas &
Bertoluci 2007). These life history traits make the collecting of new data on natural history difficult, and as a result,
species of this group are poorly known. For example, none of the eight species have available data on natural history and
the larval external morphology of Dendropsophus acreanus, D. dutrai and D. novaisi remains unknown.
Dendropsophus novaisi (Bokermann 1968) is a species described from Maracás, Bahia State, Brazil. Its currently
known distribution encompasses northern Minas Gerais and Bahia State (Frost 2017). Herein, we describe the tadpole of
D. novaisi, and provide comments on natural history. Data were collected at the Reserva Ecológica Michelin (-13.824076,
-39.184791; Datum = WGS84; 136 m.a.s.l.), municipality of Igrapiuna, southern Bahia, eastern Brazil. We collected 10
tadpoles (MZUESC 18538) in a temporary pond at the study site on 10 September 2011. Tadpoles were euthanized with
lidocaine, preserved in 10% formalin and deposited in the zoological collection of the Universidade Estadual de Santa
Cruz, Ilhéus, Bahia. We took the following measurements: total length (TL); body length (BL); body width (BW); body
height (BH); tail length (TAL); tail height (TAH); tail musculature width (TMW); tail musculature height (TMH); dorsal
fin height (DFH); ventral fin height (VFH); internostril distance (IND); interorbital distance (IOD); eye diameter (ED);
nostril diameter (ND); snout-spiracle distance (SSD); snout-eye distance (SED) and oral disc width (ODW).
Measurements and terminology follow Altig & McDiarmid (1999), Altig (2007), and Conte et al. (2007). Measurements
and illustrations were obtained with a stereoscopic microscope (Olympus®) coupled to a camera lucida.
Description of larval external morphology. Description was based in 10 individuals between Stages 33–37
(Gosner 1960). The tadpole has an elliptic body in dorsal and ventral view and triangular in lateral view (Fig. 1), with
body length about 36.1% of TL. The maximum height of the body is at the beginning of the dorsal fin, and represents
59.5% of BL. The maximum width of the body is near the spiracle line, about 60.4% of BL. Snout is rounded in dorsal

296 Accepted by F. Candioti: 28 Nov. 2017; published: 24 Jan. 2018

view and sloped in lateral view. Nostrils are oval, small, oriented anteriorly and near to the tip of the snout, being the IND
about 44.6% of BW. The eyes are laterally oriented with ED representing about 14.8% of BL; IOD about 91.4% of BW.
Spiracle is sinistral, short, attached to the body without inner wall and located below the midline of the body. Vent tube is
short, located at the beginning of the tail musculature. Tail is about 63.9% of TL; TAH is about 36% higher than BH,
with tail tip ending in a flagellum. Dorsal and ventral fins are arc-shaped. Dorsal fin originates at the posterior third of
the body and it is about 6% higher than ventral fin. There is a thick sheath of tissue in the basal portion of the tail muscle
and fins. Oral disc is anterior, not emarginated, its width representing approximately 30% of the BW (Fig. 2). Upper
labium is thick, partially covering the upper jaw sheath, with four to five small marginal papillae on the upper lateral
portion. Lower labium is also thick, with a medial row of uniseriate marginal papillae; in some cases, papillae of this
region are alternate and have varied shapes: papillae are short and triangular in tadpoles at advanced stages (Fig. 2A) and
longer at tadpoles at early stages (Fig. 2B). Upper and lower jaw sheaths are robust, slightly arc-shaped, and serrated;
serrations are short with a wide base. Labial tooth row formula (LTRF) is 0/1, with row P1 usually very short and medial.
Small and short labial teeth in P1 are present in all tadpoles analyzed.

FIGURE 1. Dendropsophus novaisi tadpole at Stage 36 of Gosner (1960): (A) dorsal view; (B) lateral view; (C) ventral view.

Measurements [in mm, expressed as mean±SD (range)]. TL 34.3±1.68 (31.54–36.90); BL 12.4±0.67 (11.26–
13.48); BW 7.5±0.95 (6.10–8.81); BH 7.4±0.7 (6.50–8.58); TAL 21.9±1.26 (19.50–23.50); TAH 10.1±0.94 (9.00–
11.88); TMW 2.9±0.21 (2.60–3.33); TMH 4.0±0.36 (3.62–4.60); DFH 3.9±0.45 (3.30–4.58); VFH 3.7±0.46 (3.00–4.40);
IND 3.3±0.26 (2.96–3.85); IOD 6.0±0.45 (5.44–6.94); ED 1.8±0.12 (1.69–1.98); ND 0.3±0.07 (0.25–0.40); SSD 8.0±0.6
(7.10–9.20); SED 5.05±0.3 (4.68–5.69); ODW 2.3±0.25 (1.86–2.68).
Coloration. In life (Fig. 3A), dorsum grayish-brown with some regions having more pigmented spots than others;
belly uniformly gray. Tail musculature color similar to the dorsum; fins varying from reticulated to marbled; fins contour
reddish; flagellum unpigmented. Iris black. In reproductive environments, where there was little vegetation and clayey

TADPOLE OF DENDROPSOPHUS NOVAISI Zootaxa 4375 (2) © 2018 Magnolia Press · 297
soil, the tadpoles were less pigmented with a light gray dorsum, and translucent fins. In formalin, the color pattern is
similar to life, but without the red pigmentation in the contour of the fin. Iris whitish.

FIGURE 2. Oral disc of Dendropsophus novaisi: (A) Tadpole at Stage 37 of Gosner (1960); (B) Tadpole at Stage 33 of Gosner (1960).

FIGURE 3. Dendropsophus novaisi (A) tadpole in life; (B) calling male; (C) couple in amplexus; (D) egg clutch in a temporary pond.

Natural history traits. Adult Dendropsophus novaisi were found only during days with heavy rainfall when they
were encountered in calling activity (Fig. 3B) at temporary ponds. The amplexus is axillary (Fig. 3C) and a gelatinous
mass is laid on the water surface of lentic water bodies (Fig. 3D). Egg clutches have in mean 813±164.2 eggs (697–1001
eggs; n = 3). Mean egg size was 1.76±0.10 mm (1.6–1.19 mm; n = 15). We observed predation attempts on tadpoles of R.

298 · Zootaxa 4375 (2) © 2018 Magnolia Press RUAS ET AL.

crucifer by tadpoles of D. novaisi, but due to water turbidity we were not able to follow the event until the end. Under
laboratory conditions, we offered tadpoles of R. crucifer (Stages 25 and 26; ~15 mm long) to D. novaisi tadpoles (Stage
34; ~34 mm length). Tadpoles of D. novaisi attacked the ventral region of the tadpoles of R. crucifer while they were
moving, and all prey tadpoles were completely ingested.
The tadpole of Dendropsophus novaisi is morphologically similar to other tadpoles described in the D. marmoratus
group. According to Peixoto & Gomes (1999), common characteristics include anterior oral disc with reduced mouth
parts, body ovoid in dorsal view and triangular in lateral view, and a high tail attenuated distally into a flagellum. A labial
tooth row formula 0/1 is described in all species except in D. marmoratus (Duellman 1978), although the illustration by
Lynch & Suárez Mayorga (2011) of what appears to be a P1 ridge in this latter makes suspect some intraspecific
variation. The tadpole of D. novaisi differs from that of D. seniculus by its taller fin relative to the body, and from D.
seniculus, D. nahdereri and D. melanargyreus by dorsal fin higher than ventral fins (Gomes & Peixoto 1991; Peixoto &
Gomes 1999; Schulze et al. 2015). Dendropsophus novaisi differs from D. soaresi by having circular nostrils and the
posterior spiracle opening, while D. soaresi has elliptical nostrils and spiracle open posterodorsally oriented (Gomes &
Peixoto 1991).
Tadpoles of Dendropsophus are frequently referred in the literature as macrophagous feeders, eating both plants and
animal resources (e.g., Altig & McDiarmid 1999; Peixoto & Gomes 1999; Vera Candioti 2007; Schulze et al. 2015).
Carnivorous diets often consist of insect larvae, worms, and small crustaceans. To our knowledge, predatory behavior on
heterospecific tadpoles is only known for Dendropsophus minutus, which attack and ingest Physalaemus larvae (Peixoto
& Gomes 1997). In addition, ours would be a new record of predation on anuran tadpoles, the first within the D.
marmoratus group. This observation and the wide resemblance of larval mouthparts in the group suggest that this
predaceous behavior may be a common feature within this clade.

Acknowledgments. We thank Francisco Fonseca Ribeiro de Oliveira, Carla Santana Cassini, Ricardo Lourenço de
Moraes and Diogo B. Provete, and especially Florencia Vera Candioti for helpful comments and suggestions on the
manuscript. Victor G. Dill Orrico for validating the identification of species. We also thank Center for Biodiversity
Studies of the Reserva Ecológica Michelin for logistical and financial support during field survey. DSR thanks CAPES
for a graduate scholarship. C.V.M.M. thanks FAPESB for a graduate scholarship and Fernanda Tonolli for her patience,
affection and eternal loving support. This study is part of the project "Girinos do Brasil", funded by the SISBIOTA
program (CNPq 563075/2010-4 and FAPESP 2010/52321-7). Tadpoles were collected under the permit 13708-1 issued
by the ICMBio to M. Solé.

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