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The tadpole of the endemic poison frog Ameerega pulchripecta (Silverstone,

1976) with the description of its chondrocranium (Anura: Dendrobatidae:
Colostethinae)

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 Zootaxa 5115 (2): 295–300 ISSN 1175-5326 (print edition)
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Copyright © 2022 Magnolia Press
Correspondence ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5115.2.9
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The tadpole of the endemic poison frog Ameerega pulchripecta (Silverstone,

1976) with the description of its chondrocranium (Anura: Dendrobatidae:
Colostethinae)
TIAGO LEITE PEZZUTI1, RAÍLA BRENA ARAÚJO1, PATRICK RIBEIRO SANCHES2, FILLIPE PEDROSO-SAN-
TOS3, MARCOS ROBERTO DIAS-SOUZA4 & CARLOS EDUARDO COSTA-CAMPOS5,*
1
Universidade Federal de Minas Gerais, Instituto de Ciências Biológicas, Departamento de Zoologia, Programa de Pós-Graduação
em Zoologia, Laboratório de Herpetologia, 31270-90, Belo Horizonte, MG, Brazil.
2
Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Ecologia (Biologia), Manaus, Amazonas, Brazil.
3
Universidade Federal do Amapá, Programa de Pós-Graduação em Biodiversidade Tropical, Macapá, Amapá, Brazil.
4
Universidade Federal do Pará, Instituto de Ciências Biológicas, Laboratório de Biologia Celular e Helmintologia “Prof. Dra Rein-
alda Marisa Lanfredi”, Belém, Pará, Brazil.
5
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.
*
Corresponding author: � [email protected]; https://orcid.org/0000-0001-5034-9268
TIAGO LEITE PEZZUTI— https://orcid.org/0000-0002-1162-3954
RAÍLA BRENA ARAÚJO— https://orcid.org/0000-0001-8507-8576
PATRICK RIBEIRO SANCHES— https://orcid.org/0000-0002-6346-3817
FILLIPE PEDROSO-SANTOS— https://orcid.org/0000-0001-7732-4482
MARCOS ROBERTO DIAS-SOUZA— https://orcid.org/0000-0001-5378-912X

The genus Ameerega Bauer comprises 29 species distributed geographically in Central Brazil and Eastern Amazon region,
as well as west of the Andes, Bolivia, Venezuela, Ecuador, and Peru (Grant et al. 2017; Frost 2021). This genus is a well-
supported clade characterized by some morphological and coloration aspects of adults, as the dorsal skin strongly granular,
finger IV of males with weak preaxial swelling, a typical marbled pattern of abdomen, among other characters (Grant et al.
2017). However, the internal relationships of the genus have differed greatly among the distinct phylogenetic hypotheses
proposed, depending on the dataset and optimality criteria used (e.g., Grant et al. 2017; Guillory et al. 2020).
Currently, tadpoles of only 12 species are described in the genus: A. altamazonica (Twomey & Brown 2008), A.
braccata (Cope 1887; Haddad & Martins 1994), A. bilinguis (Poelman et al. 2010), A. flavopicta (Haddad & Martins 1994;
Costa et al. 2006; Dias et al. 2018; Pezzuti et al. 2021), A. hahneli (Haddad & Martins 1994; Rodríguez & Duellman 1994;
Duellman 2005; Menin et al. 2017), A. macero (Rodríguez & Myers 1993), A. petersi (as A. petersi and A. smaragdina,
Silverstone 1976), A. parvula (Poelman et al. 2010), A. picta (Lescure 1976; Silverstone 1976; Haddad & Martins 1994;
Duellman 2005; Schulze et al. 2015), A. rubriventris (Lötters et al. 1997), A. silverstonei (Silverstone 1976; Myers &
Daly 1979), and A. trivittata (Wyman 1859; Silverstone 1976; Rodríguez & Duellman 1994).
Ameerega pulchripecta (Silverstone 1976) is an endemic species only known from Serra do Navio, Amapá state,
Eastern Amazon, inhabiting elevations between 100 and 400 m, East of Amapari River (Silverstone 1976). Its phylogenetic
position has been largely controversial due to the distinct methods and datasets used in the phylogenetic proposals. It has
been recovered as a sister species of Ameerega hahneli (see Grant et al. 2017), as a sister species of the clade formed by
A. trivittata and A. picta, or as sister species of a large clade containing the A. macero, A. simulans, A. petersi, and A.
hahneli species groups (sensu Guillory et al. 2020). The last complementary information about the natural history of A.
pulchripecta was the description of its advertisement call (Costa-Campos et al. 2016) and helminth parasites (Tavares-
Costa et al. 2019). Recently, the tadpole of A. pulchripecta was used for a comparison in the description of A. flavopicta
tadpoles (Dias et al. 2018), and as one of the outgroup taxa in an evolutionary analysis of Silverstoneia larvae, another
Colostethinae genus (Dias et al. 2021). Some characters of external and buccopharyngeal morphology of A. pulchripecta
were optimized in this study but focusing mainly on the evolution of the Silverstoneia characters. Thus, a formal description
of the tadpole of A. pulchripecta is still lacking, as well as more detailed data on internal morphology. Here we describe
the tadpoles of A. pulchripecta including chondrocranial morphology, aiming to increase the knowledge about the species
and to provide larval data that could be useful to systematic studies of Ameerega.

Accepted by F. Candioti: 16 Feb. 2022; published: 14 Mar. 2022 295

 Tadpoles of Ameerega pulchripecta (Stage 25; Gosner 1960) were collected from the back of a male individual
found in activity, at Parque Natural Municipal do Cancão (0.9066°N, 52.0073°W; Datum WGS 84), municipality of Serra
do Navio, Amapá state, northeastern Brazilian Amazon, in March 2018. We euthanized nine tadpoles in 5% lidocaine,
then preserved in solution containing 50% alcohol (diluted to 70%) and 50% formalin (diluted to 15%). Voucher lots are
housed in the Herpetological Collection of the Universidade Federal do Amapá (CECC), Macapá, Amapá State, Brazil
and in the Tadpole Collection of Centro de Coleções Taxonômicas at the Universidade Federal de Minas Gerais (UFMG),
Belo Horizonte, Minas Gerais State, Brazil. External morphology descriptions, measurements and proportions were based
on nine tadpoles at Stage 25 (CECC 2622–2627, 2675–2677; Gosner 1960). Terminology followed Altig & McDiarmid
(1999). Measurements were obtained with a stereoscopic microscope, along with a micrometric ocular (nearest 0.1 mm)
and follow Altig & McDiarmid (1999) for total length (TL), body length (BL), tail length (TAL), maximum tail height
(MTH), internarial distance (IND), interorbital distance (IOD), tail muscle height (TMH), tail muscle width (TMW);
Lavilla & Scrocchi (1986) for body height (BH), body width (BW), body width at eyes level (BWE), eye-nostril distance
(END), nostril-snout distance (NSD), eye diameter (ED), oral disc width (ODW); Lins et al. (2018) for spiracle length
(SL) and vent-tube length (VTL). For the analysis of the chondrocranium, two specimens (UFMG 2445) were cleared and
stained following protocol of Wassersug (1976). Drawings of skeleton were made based on photographs taken in Leica
M205 stereomicroscope. Chondrocranium terminology follows Haas (2003) with English names when available.

FIGURE 1. Tadpole Ameerega pulchripecta at Stage 25 in lateral (A), dorsal (B), and ventral (C) views (CECC 2675), collected
in the Parque Natural Municipal do Cancão, municipality of Serra do Navio, state of Amapá, Brazil. Open oral disc (D) and
frontal view (E) of another specimen at Stage 25 (UFMG 2445). Scale bars, A–C = 3 mm; D = 0.5 mm; E = 1 mm.

External morphology. Body depressed (BH/BW = 0.64–0.86; Fig. 1A–C), 0.31–0.35 times TL; oval in dorsal and
ventral views, elongated in lateral view; ventral contour convex in gular and abdominal regions, with a well-defined
constriction anterior to the spiracle level (Fig. 1A). Snout rounded in lateral view and truncated in dorsal view. Eyes dorsal

296 · Zootaxa 5115 (2) © 2022 Magnolia Press PEZZUTI et al.

(IOD/BWE = 0.50–0.63), laterally directed. Nostrils elliptical, small, dorsolaterally located, anterolaterally directed (Fig.
1E); presence of an elevated ring in the inner margin of the marginal rim, with a small fleshy projection on the medial
portion. Spiracle sinistral, located below lateral midline, visible in dorsal and ventral views, posterodorsally directed,
opening at the end of middle third of the body; aperture smaller than the diameter of tube, with its inner wall free from
the body. Vent tube short, opening dextrally, posteriorly directed, fused to the ventral fin and positioned at its ventral
margin. Tail low, slightly higher than the body (MTH/BH = 1.04–1.24); tail musculature robust (TMH/BH = 0.48–0.78),
tapering homogeneously until the acuminated tail tip; dorsal fin higher than ventral fin (DFH/VFH = 1.09–1.33). Dorsal
fin emerging on the posterior third of the body at a low slope, being straight until the middle third of the tail and convex
posteriorly; its maximum height at the posterior third of the tail. Ventral fin slightly convex; its maximum height at the
posterior third of the tail. Oral disc small (ODW/BW = 0.30–0.36, measured with oral disc closed), laterally emarginated,
anteroventrally directed (Fig. 1D, E); anterior labium with groups of three or four rounded and poorly-defined papillae
forming crenulated lateral flaps on each side of the anterior labium, interrupted by a large medial anterior gap, which
corresponds from 45 to 55% of oral disc width; posterior labium projected anteriorly, surrounded by 17 to 20 short and
conical papillae aligned or alternated, in a single row. Submarginal papillae absent. Upper jaw sheath W-shaped with
a deep medial notch and well-developed lateral processes. Lower jaw sheath V-shaped, wider than upper jaw sheath;
serrations on upper and lower sheaths extend the entire lengths of sheaths. Labial tooth row formula 2(2)/3; A2 gap from
18 to 35% of oral disc width; A1 and A2 approximately equal in length; P1 and P2 approximately equal in length, longer
than P3.
Measurements. All measures in mm, values expressed as mean ± SD (range). TL 16.48 ± 0.74 (15.2–17.5); BL
5.47 ± 0.17 (5.2– 5.7); TAL 10.81 ± 0.91 (9.5–12.1); MTH 2.88 ± 0.1 (2.7–3.0); IND 1.13 ± 0.28 (0.72–1.45); IOD 1.65
± 0.41 (1.12–2.4); TMH 1.61 ± 0.14 (1.4–1.87); TMW 1.27 ± 0.32 (0.8–1.84); BH 2.73 ± 0.27 (2.4–3.12); BW 3.62 ±
0.36 (2.8–4.0); BWE 2.77 ± 0.37 (2.32–3.21); END 0.91 ± 0.3 (0.56–1.52); NSD 0.51 ± 0.21 (0.24–0.8); ED 0.59 ± 0.2
(0.24–0.75); ODW 1.25 ± 0.11 (1.11–1.44); SL 0.42 ± 0.25 (0.15–0.88); VTL 0.65 ± 0.19 (0.32–0.96).
Color in preservative. Body translucent with small dark brown spots and black dots irregularly scattered on dorsum,
laterally, and ventrally, including the vent tube (Fig. 1 A, B, C); a lighter transversal area ventrally, with few dark spots,
anterior to the abdominal region; spiracle translucent. Tail musculature cream, fins translucent, both with angular large
dark spots; ventral fin more pigmented than dorsal fin.
Chondrocranium. Chondrocranium longer than wide; greatest width at the plane of the subocular bar (Fig. 2A, B).
Suprarostral cartilage consists of two central and separated corpora and two lateral alae; ala and corpus proximally fused
(Fig. 2C). Trabecular horns short, slightly divergent, curved ventrally; originated from the ethmoid plate and articulated
with the suprarostral cartilage in the point of articulation of the corpora and alae; lateral process of trabecular horn
present, small, triangular. Adrostral tissue mass formed of connective tissue, not chondrified. Cranial floor cartilaginous,
pierced by two pairs of foramina; the craniopalatine foramen, smaller and anteriorly positioned, and the carotid foramen,
which are larger and posterior. Lateral walls of the braincase formed by the orbital cartilage, which posteriorly does not
connect to the otic capsules; optic, oculomotor, and prootic foramina identifiable. Chondrocranium open dorsally through
a frontoparietal fenestra delimited anteriorly by ethmoid plate, laterally by taeniae tecti marginales, and posteriorly by
tectum synoticum. Otic capsules rhomboid in shape. Palatoquadrate C-shaped in dorsal view, with a short and narrow
articular process; anteriorly, the palatoquadrate connects to the braincase through the quadratocranial commissure which
has a small triangular quadratoethmoid process; triangular-shaped and short pseudopterygoid process present; posteriorly,
the palatoquadrate connects to the braincase at the level of oculomotor foramen through the ascending process, which
consists of a thin rod-like cartilage. Muscular process triangular, tall, and thin; dorsally, the edges of the muscular process
curve, approaching and connecting to the conspicuous antorbital process by ligament. Lower jaw formed by the infrarostral
and Meckel’s cartilages; infrarostral cartilages rectangular, arranged describing a V shape, joined medially by connective
tissue; Meckel’s cartilage sigmoid in shape, located ventral to the trabecular horns. Ceratohyal cartilages wide, flat,
subtriangular (Fig. 2D); each ceratohyal bears a conspicuous triangular anterior process, a small anterolateral process,
and a well-developed, triangular posterior process; ceratohyal bonded together by the pars reuniens, which is shorter
than the basibranchial. Urobranchial process small. Basihyal absent. Hypobranchial plates triangular and articulated
anteromedially. Branchial basket with four curved ceratobranchials with numerous lateral projections; ceratobranchial
distally joined by terminal commissures; proximally, ceratobranchials attach to the hypobranchial plates; four spicules,
curved, long, arise dorsally from each ceratobranchial.
Comparison of external morphology in the genus Ameerega. The tadpoles of Ameerega pulchripecta are similar
to those of other species of Ameerega by presenting a globular/depressed body, low tail and fins, an oral disc laterally

Tadpole of Ameerega pulchripecta Zootaxa 5115 (2) © 2022 Magnolia Press · 297
emarginated with a large anterior gap on marginal papillae, and a large A2 gap. Some important variations mainly
concerning oral disc characters have been illustrated and described in the literature (see Menin et al. 2017; Dias et al.
2018). The marginal papillae on the anterior labium, for example, have been described as 1) absent on the anterior labium
in A. parvula, A. bilinguis, and A. hahneli (Poelman et al. 2010; Haddad & Martins 1994; Menin et al. 2017); in these
species the lateral edges of anterior labium are elevated and smooth (i.e., without defined papillae), extending from the
oral disc emargination until the level of lateral extremes of A1 tooth row; 2) as a crenulated flap (described in some studies
as flap free from the body wall; Twomey & Brown 2008) in A. altamazonica, A. macero, and A. pulchripecta (Rodrigues
& Myers 1993; Twomey & Brown 2008; this study); 3) disposed in row of defined papillae in A. braccata, A. flavopicta,
A. picta, and A. silverstonei (Myers & Daly 1979; Haddad & Martins 1994; Schulze et al. 2015; Dias et al. 2018; Pezzuti
et al. 2021). This condition seems to be also present in A. petersi, A. rubriventris, and A. trivittata, according to the
schematic drawings presented (Silverstone 1976; Lötters et al. 1997). The tadpoles of Ameerega pulchripecta can be
distinguished from A. bilinguis, A. hahneli, A. flavopicta, and A. parvula by the LTRF 2(2)/3, which is the most common
configuration in the genus (LTRF 2(2)/3(1) in A. bilinguis, A. flavopicta, and A. parvula; 0/0, 1/1, or 1/2 in A. hahneli;
Haddad & Martins 1994; Duellman 2005; Poelman et al. 2010; Menin et al. 2017). It is worth noting that these species
were comparatively analyzed by Dias et al. (2018) and only A. flavopicta and a Peruvian population of A. hahneli were
reported as presenting 2(2)/3(1). Whether differences between the previous descriptions and the specimens analyzed in
Dias et al. (2018) are related to intraspecific variation (e.g., ontogenetic or populational variation, as already reported
for A. hahneli; Menin et al. 2017) remains to be investigated. Additionally, A. pulchripecta is distinct from A. flavopicta
by the W-shaped anterior jaw sheath (arc-shaped in this species; Dias et al. 2018; Pezzuti et al. 2021). This character
may have an underestimated variation, as the medial notch of the anterior W-shaped jaw can be conspicuously deep as
in A. pulchripecta, A. parvula, A. bilinguis, A. hahneli, or discreet, as figured in Ameerega altamazonica, A. macero,
A. picta (Rodriguez & Myers 1993; Twomey & Brown 2018; Schulze et al. 2015). However, this character cannot be
unequivocally compared using all schematic drawings currently available, requiring a more specific and comparative
analysis to evaluate this variation. Dias et al. (2018) characterized the tadpoles of A. pulchripecta as presenting the dorsal
fin originating at the tail, differing from the condition found here, in which the tail originates at the body tail junction.
Comparison of chondrocranium in the genus Ameerega. Data on chondrocranium of Ameerega are available
only for A. bassleri, A. bilinguis, A. parvula, and A. trivittata (Dias et al. 2021). Dias et al. (2021) used these species
as outgroup for optimizations, aiming to account for the variation among Silverstoneia larvae. Thus, few characters are
available for a detailed comparison between A. pulchripecta and other Ameerega species. Several suprarostral cartilage
configurations have been described for Dendrobatidae (Haas 2003; Dias et al. 2021). The condition of completely free
suprarostral corpora (state 0; character 13 of Dias et al. 2021) described here for A. pulchripecta was also found for A.
trivittata, A. bassleri, A. parvula, and A. bilinguis, as well for other Colostethinae genera (Dias et al. 2021). This character
state differentiates these species from Ameerega hahneli and A. petersi, in which the corpora are proximally fused (Dias
et al. 2021). Dias et al. (2021) suggested that the proximal fusion between the corpora could be a synapomorphy for the
A. petersi group (sensu Grant et al. 2017), requiring the knowledge of this character in other species, including Ameerega
pulchripecta. Considering phylogenetic hypotheses that retrieve A. pulchripecta as close related to A. hahneli (see Grant
et al. 2017), the distinct character states in both species, together with the lack of information in the close related clades,
brings an ambiguity in the optimization of this character in the genus (Dias et al. 2021). Ameerega pulchripecta, like
all species of Ameerega and most Colostethinae (except Colostethus and Leucostethus; Dias et al. 2021) present the
suprarostral alae and corpora fused proximally (state 1; character 14 of Dias et al. 2021). This feature has been considered
a synapomorphy of the subfamily (Dias et al. 2021). The condition of adrostral tissue mass not chondrified, as reported
here for A. pulchripecta, is also found in most Colostethinae, including Ameerega (Dias et al. 2021). The absence of a free
basihyal cartilage in the hyobranchial apparatus, as described here for A. pulchripecta, was also described for A. hahneli,
while it is present in A. trivittata and A. parvula (Dias et al. 2021). The variation in chondrocranium features among the
few known species of Ameerega, points to a possible underestimated skeletal diversity in the genus.
We are grateful to M. F. Vera Candioti and an anonymous reviewer for their comments and suggestions that improved
the manuscript. We are thankful to Albertina Lima for his constructive comments on early drafts of this manuscript and
for sharing her enormous experience in anurans with us. We thank Jeni Magnusson, Iasodhara Rodrigues for providing
photographic plates of tadpoles used in this manuscript, Paulo C. A. Garcia (UFMG) for providing laboratory workspace,
and Christoph Jaster (Parque Nacional Montanhas do Tumucumaque) for hospitality and generous logistic support and
assistance in the field. TLP acknowledges CAPES for his PNPD fellowship (8887.468027/2019-00). Collections were
made with permits provided by Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio, 48102-2). This

298 · Zootaxa 5115 (2) © 2022 Magnolia Press PEZZUTI et al.

study was carried out as part of the Project “Natural history of anurans of eastern Amazon” of the Departamento de
Pesquisa of the Universidade Federal do Amapá.

FIGURE 2. Chondrocranium (UFMG 2445, Stage 25) in dorsal (A) and ventral (B) views, suprarostral in frontal view (C), and
hyobranchial apparatus (D). Abbreviations: a, ala; alpc, anterolateral process of ceratohyal; apc, anterior process of ceratohyal;
app, articular process of palatoquadrate; asp, ascending process; c, corpora; cb I–IV, ceratobranchial I–IV; cf, carotid foramen;
ch, ceratohyal; cpf, craniopalatine foramen; hp, hypobranchial plate; ic, infrarostral cartilage; ff, frontoparietal fenestra; lt,
lateral process of trabecular horn; mc, Meckel’s cartilage; mp, muscular process; oc, otic capsule; pp, pseudopterygoid proc-
ess; ppc, posterior process of ceratohyal; pq, palatoquadrate; pr, pars reuniens; qcc, quadratocranial commissure; s, spicule; sc,
suprarostral cartilage; th, trabecular horns; ts, tectum synoticum; ttm, taenia tecti marginalis. Scale bars, A and B = 1.0 mm; C
and D = 0.5 mm.

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