Zootaxa 4780 (3): 594–600 ISSN 1175-5326 (print edition)

https://www.mapress.com/j/zt/
Copyright © 2020 Magnolia Press
Correspondence ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4780.3.11
http://zoobank.org/urn:lsid:zoobank.org:pub:55488386-C327-45E0-8481-0A6230CBAD09

Description of the tadpole of Dendropsophus haraldschultzi (Bokermann, 1962)

(Anura: Hylidae), with comments on reproductive biology
MARCELO MENIN1,2,*, ALEXANDRE PINHEIRO DE ALMEIDA2, FILLIPE PEDROSO-SANTOS3,4,
PATRICK RIBEIRO SANCHES3,5 & CARLOS EDUARDO COSTA-CAMPOS3,6
1
Universidade Federal do Amazonas, Instituto de Ciências Biológicas, Departamento de Biologia and Laboratório de Taxonomia e
Ecologia de Anfíbios e Répteis, Av. General Rodrigo Otávio Jordão Ramos 6200, 69.080-900, Manaus, AM, Brazil.
� menin@ufam.edu.br; http://orcid.org/0000-0002-7209-5083
2
Universidade Federal do Amazonas, Instituto de Ciências Biológicas, Programa de Pós-Graduação em Zoologia, Av. General Ro-
drigo Otávio Jordão Ramos 6200, 69.077-000, Manaus, AM, Brazil
� alexandre.dealmeida@hotmail.com; https://orcid.org/0000-0003-3938-4198
3
Universidade Federal do Amapá, Departamento de Ciências Biológicas e da Saúde, Laboratório de Herpetologia, Campus Marco
Zero do Equador, 68.903-419, Macapá, AP, Brazil
4�
fillipepedrosodossantos@gmail.com; https://orcid.org/0000-0001-7732-4482
5�
patricksanchs@gmail.com; https://orcid.org/0000-0002-6346-3817
6�
eduardocampos@unifap.br; https://orcid.org/0000-0001-5034-9268
*
Corresponding author

The genus Dendropsophus Fitzinger contains 108 recognized species (Frost 2020) distributed in nine species groups ac-
cording to Faivovich et al. (2005). However, recent phylogenetic analysis recognized the presence of nonmonophyletic
groups (e.g., Wiens et al. 2010; Fouquet et al. 2011; Motta et al. 2012; Jansen et al. 2019) suggesting that the relation-
ships among species of Dendropsophus require careful revision. Species of this genus are distributed in the tropical and
subtropical South America, including Trinidad, southward to northern Argentina and Uruguay and northward to Central
America and tropical southern Mexico (Duellman et al. 2016; Frost 2020). Dendropsophus haraldschultzi (Bokermann)
was described in the area of Santa Rita do Weill, municipality of São Paulo de Olivença, Amazonas State, Brazil. Adult
individuals of D. haraldschultzi are small sized frogs (males 18–22 mm and females 22–25 mm; Rodríguez & Duellman
1994) with tuberculate skin on dorsal surfaces, denser on the head. They are found near open ponds and permanent large
streams or in floating meadows along the Amazon river (Bokermann 1962; Rodríguez & Duellman 1994; Böning et al.
2017) and have been also found along the Amazon River Valley in the Brazilian states of Pará and Amapá (Missassi et al.
2017), and in Peru and Colombia. Bokermann (1962) and later authors (e.g., Lutz 1973) considered this species without
evident taxonomic affinities. Currently, D. haraldschulzi is not assigned to any species group within the genus (Faivovich
et al. 2005).
The tadpole of Dendropsophus haraldschultzi was first illustrated by Lynch & Suárez-Mayorga (2011) in a tadpole
identification key for species from eastern lowlands of Colombia, showing some morphological characteristics. However,
the tadpoles have not been described in detail and these authors do not show how they associated the tadpoles to this spe-
cies. In this work we present a formal description of the tadpole of D. haraldschultzi based on 12 individuals obtained
from a clutch and six individuals collected in a lake in two localities at northern Brazil. We also provide comments on
its reproductive biology. We then compare tadpoles of D. haraldschultzi with known tadpoles of the D. columbianus
(Boettger), D. leucophyllatus (Beireis), D. marmoratus (Laurenti), D. microcephalus (Cope), D. minutus (Peters), and D.
parviceps (Boulenger) groups (revisions in Faivovich et al. 2005; Caminer et al. 2017).
We obtained one egg clutch from an amplectant pair kept overnight in a plastic bag containing water collected
from their natural habitat. The frogs were found in a floodplain area of the municipality of Santana, Amapá State, Brazil
(0.0363°N, 51.1625°W, Datum: WGS84; 26 m.a.s.l.), in May 2018. The clutch was maintained in a glass container (8
cm3) in the laboratory and the tadpoles were fed with fish food. Twelve specimens were euthanized in solution of benzo-
caine diluted in water and fixed and preserved in a 1:1 solution containing ethanol 70% and formalin 15%. The tadpoles
and adults were deposited in the Herpetological Collection of the Universidade Federal do Amapá, Macapá, Amapá State,
Brazil (Tadpole lots CECCAMPOS 2655 to 2666; Adults CECCAMPOS 2386–2388, 0562, 0563, 0569–0571, 0573–

594 Accepted by F. Candioti: 28 Apr. 2020; published: 26 May 2020

0575, 0577). Additionally, this study also included six tadpoles collected on 08–09 February and 08 April 2013 in floating
meadows in the Catalão Lake (3.1667°S, 59.9142°W, Datum: WGS84), in the municipality of Iranduba, Amazonas State,
Brazil, and deposited in the Coleção Zoológica Paulo Bührnheim of the Universidade Federal do Amazonas, Manaus,
Amazonas State, Brazil (CZPB-LA tadpole lots 257/576, 258/580, 282/633, 282/634, 283/636, 285/648). Measurements
were obtained with use of an ocular micrometer (nearest 0.01 mm) in a stereoscopic microscope. Descriptive terminology
and morphometric measurements follow Altig & McDiarmid (1999): total length (TL), body length (BL), body width
(BW), body height (BH), tail length (TAL), maximum tail height (MTH), tail muscle height (TMH), tail muscle width
(TMW), internarial distance (IND), interorbital distance (IOD), eye diameter (ED), eye-nostril distance (END), nostril-
snout distance (NSD), and oral disc width (ODW). We also measured the spiracle length (SL) and the vent-tube length
(VTL). Developmental stages follow Gosner (1960).
Tadpole description. The description is based on two tadpoles at Stage 35 (CZPB-LA 258/580, 285/648). Body
depressed in lateral view (Fig. 1A) and violin-shaped with a slight constriction posterior to the orbits in dorsal and ven-
tral views (Fig. 1B and C); body length 31.8–33.3% of the total length. Maximum body height in the middle of the body
representing 42.0–46.9% of body length and 84.0–88.4% of body width. Body width at spiracle level near to body width
at eye level. Snout rounded in dorsal and lateral views. Eyes laterally positioned and directed; eye diameter representing
14.6–16.2% of body length. Nostrils oval and small, without a fleshy ring in its inner margins, anterolaterally positioned
and closer to the snout than to eyes, opening anterolaterally directed (Fig. 1D). Internarial distance representing 69.7–
75.0% of body width. Spiracle single, sinistral, long, and wide, opening in the posterior third of the body, located latero-
ventrally and posteriorly directed. Spiracle centripetal wall fused to the body wall; external wall longer than centripetal
wall and with irregular margin (Fig. 1D). Vent tube single, short, dextral, fused to the ventral fin. Tail length representing
66.7–68.1% of total length. Caudal musculature gradually tapering to a pointed tip in lateral view. Tail tip pointed. Tail
muscle height representing 67.5% of maximum tail height. Tail muscle width representing 47.5–48.1% of body width.
Dorsal fin shallow, originating on the anterior third of the tail. Ventral fin convex originating at the posterior extremity
of the ventral terminus and with linear margins following the tail axis. Lateral line system evident, with stitches of lat-
eral lines rounded; presence of oral, supraorbital, infraorbital, preorbital, angular, medial, dorsal, and ventral lines. Oral
disc (Fig. 1E) terminal, not emarginated, with wide gap on the anterior labium and rounded uniseriate marginal papillae
located in the ventral margin of the posterior labium. Lateral marginal papillae partially fused forming a dermal vertical
fold on both sides of oral disc. Submarginal papillae absent. Oral disc width representing 27.6–28.0% of body width. Up-
per and lower jaw sheaths large; upper jaw sheath arch-shaped and lower jaw sheath U-shaped; upper jaw sheath thinner
than the lower jaw sheath; both jaw sheaths black, keratinized, and finely serrated. Labial tooth row formula (LTRF) 0/1;
P1 medial and shorter than jaw sheaths. Posterior labium with a dermal ridge (likely a second labial ridge) between tooth
row and papillae.
Coloration. In life (Stage 25), dorsum, lateral and ventral body grayish-brown with small spots on the anterior por-
tion of the body and yellowish pigment spots from the posterior portion of the body to the tail. Digestive tract barely
visible ventrally. Tail musculature color similar to the body with small dark spots on lateral view. Fins translucent with ir-
regular marks grayish-brown on the posterior portion. Iris black. In preservative (Stage 35), body and caudal musculature
dark brown with small rounded spots on the dorsum, lateral of the body and caudal musculature. Fins translucent with
irregular dark brown marks on the posterior third. Iris dark brown. Glandular patches present on the dorsal and lateral
surface of the body.
Variation. Variation in 16 morphometric characteristics of tadpoles from Gosner Stages 25, 31, 34, 35 and 36 is given
in Table 1. There is no variation in LTRF along all development stages analyzed, but there is variation in the P1 row size,
usually short at Stage 25 and longer at older stages. Spiracle is more evident in the older stages analyzed and its size is
maximum at Stage 35 (2.05–2.25 mm). Dorsal fin originating at body-tail junction at stage 25 and on the anterior third of
the tail at older stages.
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).

TADPOLE OF DENDROPSOPHUS HARALDSCHULTZI Zootaxa 4780 (3) © 2020 Magnolia Press · 595
TABLE 1. Measurements (in mm) of 18 Dendropsophus haraldschultzi tadpoles at Gosner Stages 25, 31, 34, 35 and 36.
Values are means ± standard deviation and range except for stages with one or two individuals only, for which all values
are presented. Abbreviations according to the text.
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 Dendropso-
phus 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 re-
maining 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, Dendrop-
sophus 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 (Re-
inhardt 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

596 · Zootaxa 4780 (3) © 2020 Magnolia Press MENIN ET AL.

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. nahder-
eri (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 oc-
curring in Amazonia lay their eggs on the water, on the surface of leaves of emergent vegetation [D. bifurcus, D. boker-
manni (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).

FIGURE 1. Dendropsophus haraldschultzi tadpole at Stage 35. A) Lateral, (B) dorsal and (C) ventral views, (D) nostril (n) and spiracle
(sp), and (E) oral disc of an individual from CZPB-LA tadpole lot 258/580, collected at Catalão Lake, Iranduba municipality, Amazo-
nas, Brazil.

TADPOLE OF DENDROPSOPHUS HARALDSCHULTZI Zootaxa 4780 (3) © 2020 Magnolia Press · 597
 We are grateful to Maria Isabel da Silva for reviewing the English, Instituto Chico Mendes de Conservação da
Biodiversidade (ICMBio) for the collection permits (48102-2; 29197), SISBIOTA Program “Girinos do Brasil”—Con-
selho Nacional de Desenvolvimento Científico e Tecnológico (CNPq #563075/2010-4, 405640/2016-1), and Fundação
de Amparo à Pesquisa do Estado de São Paulo (FAPESP #10/52321-7)—for financial support. MM received a Research
Productivity Grant from CNPq.

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