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Three new species of Proceratophrys Miranda-Ribeiro 1920 from Brazilian

Cerrado (Anura, Odontophrynidae)

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DOI: 10.11646/zootaxa.3750.4.2 · Source: PubMed

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 Zootaxa 3750 (4): 321–347 ISSN 1175-5326 (print edition)
www.mapress.com /zootaxa /
Copyright © 2013 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3750.4.2
http://zoobank.org/urn:lsid:zoobank.org:pub:AF5F57AF-8F2D-413B-9B98-82484835555F

Three new species of Proceratophrys Miranda-Ribeiro 1920 from Brazilian Cer-

rado (Anura, Odontophrynidae)

REUBER ALBUQUERQUE BRANDÃO1,6, ULISSES CARAMASCHI2,5, WILIAN VAZ-SILVA3 &

LEANDRO AMBRÓSIO CAMPOS4
1.
Departamento de Engenharia Florestal – EFL, Faculdade de Tecnologia – FT, Universidade de Brasília – UnB, 70910-900 Brasília,
DF, Brasil. E-mail: reuber@unb.br
2.
Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Quinta da Boa Vista, São Cristóvão,
20940-040 Rio de Janeiro, RJ, Brasil. E-mail: ulisses@acd.ufrj.br
3.
Departamento de Biologia, Pontifícia Universidade Católica de Goiás - PUC Goiás. Rua 235, n. 40, Bloco L, Setor Universitário,
74605-010 Goiânia, GO, Brazil. E-mail: herpetovaz@gmail.com
4.
Laboratório de Anatomia Comparativa de Vertebrados, Departamento de Ciências Fisiológicas, Instituto de Biologia, Universidade
de Brasília – UnB, 70910-900 Brasília, DF, Brasil. E-mail: leandro.ambrosio@gmail.com
5.
Fellow of Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq).
6.
Corresponding author: Reuber Albuquerque Brandão: E-mail: reuber@unb.br

Abstract

Based on the analyses of specimens collected at several areas in the Cerrado domain from Central Brazil and others housed in
scientific collections and on specimens collected at the type-locality, herein we describe three new species belonging to the P.
cristiceps group: Proceratophrys bagnoi sp. nov., from UHE Serra da Mesa power plant (13°49’47.5”S, 48°19’17”W; 570 m
a.s.l.; WGS84 datum), Municipality of Minaçu, State of Goiás; Proceratophrys branti sp. nov., from Jalapão, Municipality of
Mateiros (05o15’25”S, 48o12’00”W; 109 m a.s.l.; WGS84 datum), State of Tocantins; and Proceratophrys dibernardoi sp. nov.,
Municipality of Mineiros (17o33’52”S, 52o33’20”W; 803 m a.s.l.; WGS84 datum), State of Goiás. The diversity of
Proceratophrys in Brazilian Cerrado is still underscored and several species will be described in the following years.

Key words: Amphibia, Proceratophrys goyana, Proceratophrys bagnoi sp. nov., Proceratophrys branti sp. nov.,
Proceratophrys dibernardoi sp. nov., Taxonomy

Introduction

Despite lacking a formal definition based on synapomorphies, the genus Proceratophrys Miranda-Ribeiro, 1920 is
considered homogeneous based on overall morphology and it is likely a monophyletic taxon, with Odontophrynus
Reinhardt & Lütken, 1862 + Macrogenioglottus Carvalho 1946 as its sister group (Frost et al. 2006; Pyron &
Wiens 2011). All species in the genus share the following combination of characters: body without enlarged glands,
absence of nuptial pads on thumbs, toes not webbed but laterally fringed, supernumerary tubercles present on
hands and feet, dorsal surfaces of fingers and toes rugose, zygomatic ramus of squamosal bone close to or in sutural
contact with maxilla, and cervical cotylar arrangement type II (cotyles closely approximated) (Savage and Cei
1965; Lynch 1971; Prado and Pombal 2008; Martins and Giaretta 2011). Currently, the genus is composed of 29
species (Ávila et al. 2012; Cruz et al. 2012; Frost 2013), distributed in four putative species groups based mainly
on morphological traits (Kwet and Faivovich 2001; Giaretta et al. 2000; Prado and Pombal 2008; Cruz and Napoli
2010; Dias et al. 2013). These groups have not been accessed in a comprehensive phylogenetic analysis, and
unfavorable evidence based on molecular data has been presented by Amaro et al. (2009), Pyron and Wiens (2011),
Teixeira Jr. et al. (2012), and Dias et al. (2013). Herein we keep the traditional taxonomic arrangement for the
genus, providing comparisons when necessary.
The Proceratophrys bigibbosa species group, characterized by the presence of developed postocular swellings
and absence of prominent palpebral appendages, was reviewed by Kwet and Faivovich (2001) and includes P.

Accepted by M. Vences: 18 Nov. 2013; published: 20 Dec. 2013 321

avelinoi Mercadal de Barrio & Barrio 1993, P. bigibbosa (Peters 1872), P. brauni Kwet & Faivovich 2001, and P.
palustris Giarettta & Sazima 1993. These species are distributed in southern Brazil, Argentina, and Paraguay (Frost
2013), except by P. palustris, treated as relictual species in southeastern Brazil (Kwet and Faivovich 2001).
The second and more speciose species group, the Proceratophrys boiei group (Izecksohn et al. 1998; Prado
and Pombal 2008), is composed of species with prominent, long horn-like palpebral appendages, and absence of
postocular swellings. Later, Cruz and Napoli (2010), Teixeira Jr et al. (2012), and Dias et al. (2013) considered that
the species of the former Proceratophrys boiei group having long rostral appendages and triangular palpebral
appendages must be grouped in the Proceratophrys appendiculata group. Considering this, the Proceratophrys
boiei group is composed by P. boiei (Wied-Neuwied 1824), P. paviotii Cruz, Prado & Izecksohn 2005, and P.
renalis (Miranda-Ribeiro 1920), whereas the Proceratophrys appendiculata involves P. appendiculata (Günther
1873), P. belzebul Dias, Amaro, Carvalho-e-Silva & Rodrigues 2013, P. izecksohni Dias, Amaro, Carvalho-e-Silva
& Rodrigues 2013, P. laticeps Izecksohn & Peixoto 1981, P. melanopogon (Miranda-Ribeiro 1926), P. moheringi
Weigoldt & Peixoto 1985, P. phyllostomus Izecksohn, Cruz & Peixoto 1999, P. rondonae Prado & Pombal 2008, P.
sanctaritae Cruz & Napoli 2010, P. subguttata Izecksohn, Cruz & Peixoto 1999, and P. tupinamba Prado &
Pombal 2008. The species in these groups are distributed mainly throughout the eastern Brazilian Atlantic Forest,
except P. rondonae, occurring in the southwest Amazon Rainforest. Additionally, besides included in the P.
appendiculata group, P. rondonae presents a short multi-cuspidate instead of a unique long palpebral appendage.
The approximately same condition, but with slightly shorter multi-cuspidate palpebral appendage, is present in P.
schirchi (Miranda-Ribeiro 1937) and P. minuta Napoli, Cruz, Abreu & Del Grande 2011; however, these species
were not associated to any group due to their distinct morphological characters (Prado and Pombal 2008, Napoli et
al. 2011). On the other hand, Amaro et al. (2009) suggested the relationship of P. schirchi with P. cristiceps, but
also did not clearly propose its inclusion in any species group. Recently, Teixeira Jr. et al. (2012) described P.
redacta and suggested that this species is closely related to P. minuta, and composing a sister clade to P. cristiceps
+ P. schirchi.
Finally, the Proceratophrys cristiceps group in the sense of Giaretta et al. (2000) comprises species lacking
both palpebral appendages and postocular swellings, but also lacking any recognized shared derived character. It
currently contains P. aridus Cruz, Nunes & Juncá 2012, P. caramaschii Cruz, Nunes & Juncá 2012, P. carranca
Godinho, Moura, Lacerda & Feio 2013, P. concavitympanum Giaretta, Bernarde & Kokubum 2000, P. cristiceps
(Müller 1884), P. cururu Eterovick & Sazima 1998, P. goyana (Miranda-Ribeiro 1937), P. huntingtoni Ávila,
Pansonato & Strüssmann 2012, P. moratoi (Jim & Caramaschi 1980), P. vielliardi Martins & Giaretta 2011, P.
rotundipalpebra Martins & Giaretta 2013, and P. strussmannae Ávila, Kawashita-Ribeiro & Morais 2011 (Giaretta
et al. 2000; Martins and Giaretta, 2011; Ávila et al. 2011). The species of the P. cristiceps group are mainly
associated to seasonally open habitats in Brazilian Cerrado and Caatinga morphoclimatic domains (sensu Ab’Sáber
1977), with exception of P. concavitympanum, found at the southwest Amazon Rain Forest in Rondônia State and
on transitional areas in Cerrado and Amazon Rain Forest, in Mato Grosso and Tocantins States (Ávila et al. 2012,
Teixeira Jr. et al. 2012).
Until now, seven species of Proceratophrys, all belonging to the P. cristiceps species group, are considered
Cerrado endemic species (P. goyana, P. cururu, P. moratoi, P. vielliardi, P. huntingtoni, P. carranca, and P.
rotundipalpebra) (Valdujo et al. 2012; Brandão et al. 2012; Teixeira Jr. et al. 2012; Dias et al. 2013). Based on the
analyses of several specimens of Proceratophrys collected in several areas of the Cerrado domain housed in
scientific collections, we describe three new species related to the P. cristiceps species group occurring in Central
Brazil.

Material and methods

Specimens used for descriptions and comparisons (Appendix 1) are housed in the Coleção Herpetológica da
Universidade de Brasília, Brazil (CHUNB) and in the Museu Nacional, Rio de Janeiro, Brazil (MNRJ). We also
used data from the formal descriptions of the members of the Proceratophrys cristiceps species group available in
the literature (Eterovick and Sazima 1998; Giaretta et al. 2000; Santana et al. 2010; Ávila et al. 2011; Martins and
Giaretta 2011; Napoli et al. 2011; Cruz et al. 2012; Ávila et al. 2012; Teixeira Jr. et al. 2012, Dias et al. 2013,
Godinho et al. 2013, Martins & Giaretta 2013).

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 The following measurements were taken to the nearest 0.1 mm with digital calipers, modified from Prado and
Pombal (2008) and Teixeira Jr. et al. (2012): snout-vent length (SVL), head length (HL), head width (HW), head
height (HH, maximum head height, measured behind the eyes), eye–nostril distance (END), eye diameter (ED),
interorbital distance (IOD), forearm length (FAL, from the elbow to the hand basis), hand length (HAL, from the
hand basis to the third finger tip), thigh length (THL), shank length (SHL), tarsus length (TL, from the knee to the
basis of the inner metatarsal tubercle), and foot length (FL, from the basis of the inner metatarsal tubercle to the
fourth toe tip). Web formula is based on Savage and Heyer (1967).

FIGURE 1. Dorsal view of Proceratophrys bagnoi sp. nov. showing the continuous ocular-dorsal ridge of warts (ODR)
and the different portions used in this study. A) Interocular ridge (IOR); B) Supra-scapulae region; C) Pre-sacral
constriction; D) Sacral-urostyle region; and E) Coccyx end. Bars equal 10 mm.

Morphological Characters Description. The external morphology of the species belonging to the
Proceratophrys cristiceps species group is extremely complex. The patterns formed by the combination of
tubercles sizes, formats, and distribution, make the species descriptions and identifications a hard task. Herein we
describe the morphological traits used for species descriptions and propose formulae aiming to help comparisons
between species in the group.
Ocular-dorsal ridge of warts. This is one of the most noticeable dorsal traits of Proceratophrys. Despite, this
trait has been called by different terms, as “sagittal ridges” or “sagittal crest” (p.ex., Godinho et al. 2013; Martins
and Giaretta 2013), “rows of warts” (Cruz and Napoli 2010), and “cutaneous dorsal crest” (Ávila et al. 2011). We

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decide to use the term “ocular-dorsal ridge of warts” (ODR) due to the following reasons: 1) allow a prone
visualization of the structure; 2) sagittal is an adjective, denoting the spur-like format of the ridge; however, it also
denotes the suture between parietal bones in mammals and reptiles. This double meaning turns the term confusing
and inappropriate; and 3) this term has been recently used by other authors (e.g., Teixeira Jr. et al. 2012; Prado and
Pombal 2008).
The ODR is formed by two simple rows of enlarged warts (but not larger than other scattered larger dorsal
tubercles), that extend from the medial or posterior edge of the eyelid to the sacral region, where these two rows
joint at the extreme tip of coccyx, forming a spur-like pattern. We divided the ODR in the following sections: A)
eyelid contact; B) supra-scapulae region; C) pre-sacral constriction; and D) sacral-urostyle portion (Figure 1).
We found three patterns of ODR in the Proceratophrys cristiceps species group: A) Continuous, observed in P.
goyana and P. bagnoi sp. nov.; B) Interrupted, observed in P. huntingtoni, P. branti sp. nov., and P. carranca; and
C) Absent, as observed in P. vielliardi and P. moratoi (Figure 2). The ODR interruption often is placed on the sacral
region or on the pre-sacral constriction.

FIGURE 2. Different patterns of ocular-dorsal ridge of warts (ODR). A) complete in Proceratophrys bagnoi sp. nov.; B)
interrupted at supra-sacral region in Proceratophrys branti sp. nov.; and C) absent in Proceratophrys vielliardi. Bars
equal 10 mm.

Eyelid tubercles. We observed a common pattern on external margin eyelid tubercles of all examined species.
There is always a longer medial tubercle on eyelid, that can cover almost all eyelid border in some species. The
ODR is joined at the posterior end of this medial tubercle. We also noted that often smaller anterior and posterior
tubercles can be present to this medial elongated one. The number and format of these anterior and posterior
tubercles varies between different species. We considered these tubercles as fused when they are poorly
differentiated from others, showing only discrete groove between them. Not fused tubercles are clearly discrete
from neighbor ones. These tubercles can be short (higher than long) or elongated (longer than high), round or
conical, pointed or not pointed. Although the tubercle format is similar within the same species, the number of
tubercles can vary between right and left eyelids in the same individual. We propose here a formulae for describe
the eyelid pattern, taking into account the number of anterior and posterior tubercles in each eyelid (right and left),
and the extension of the medial tubercle, expressed by the side (L or R), the number of anterior tubercles, the
extension of the elongated medial tubercle (expressed in proportion), and the number of posterior tubercles. Taking
one example: L 1,4/5,2; R 1,4/5,2. This example denotes that the individual have four tubercles on left eyelid,
being one anterior, the medial covering 4/5 of the eyelid margin, and two posteriors. The same number and pattern
is observed at right side. This example was picked in a sorted P. vielliardi individual. We choose to keep all medial
tubercle proportions divided by five for practical purposes. Figure 3 presents other examples of eyelid formulae.
We tested this formula with several individuals and we are convinced that this can be useful for accounting
differences between individuals and species (by expressing the differences on tubercles numbers using dashes).
Using P. vielliardi as example again (N=9), the pattern could be expressed as L 0–2, 3/5–4/5, 1–3; R 0–2, 2/5–4/5,
2–4.

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FIGURE 3. Eyelid tubercles and eyelid formulae of left eye. A) round and fused anterior (1) and posterior tubercles (3),
with medial one covering almost all eyelid external margin and without points in Proceratophrys branti sp. nov.
(formulae 1, 4/5, 1); B) conical slightly fused anterior and posterior eyelid tubercles, with medial tubercle showing small
points in Proceratophrys dibernardoi sp. nov. (formulae 1,3/5,1); C) conical, not fused, and pointed anterior and
posterior tubercles in Proceratophrys bagnoi sp. nov. (formulae 2, 3/5, 6). Bars equal 1 mm.

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FIGURE 4. Different patterns of interocular ridge (IOR). A) complete in Proceratophrys bagnoi sp. nov.; B) warts
present but not organized in a row in Proceratophrys dibernardoi sp. nov.; C) IOR absent in Proceratophrys vielliardi.
Bars equal 1 mm.

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FIGURE 5. Two magnified forearm tubercles of Proceratophrys bagnoi sp. nov., showing the absence of pointed
spicules and without pigmentation. Bar equal 2 mm.

FIGURE 6. Different patterns of rows of enlarged forearm tubercles. A) two rows of very close and enlarged forearm
tubercles in Proceratophrys dibernardoi sp. nov. (bar = 3 mm); B) one row of enlarged tubercles, but showing some
scattered ones, not organized in row, in Proceratophrys goyana (bar = 4 mm); C) two spaced rows of enlarged forearm
tubercles, not closely disposed in Proceratophrys bagnoi sp. nov. (bar = 4 mm).

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FIGURE 7. Palmar surface without spicules between tubercles in Proceratophrys bagnoi sp. nov. (A), and with several
spicules between tubercles in Proceratophrys branti sp. nov. (B). Bars equal 2mm.

328 · Zootaxa 3750 (4) © 2013 Magnolia Press BRANDÃO ET AL.

FIGURE 8. Contact point between the ocular-dorsal ridge of warts and the external eyelid margin tubercles. A) posterior
third in Proceratophrys branti sp. nov.; B) medial in Proceratophrys bagnoi sp. nov.; and C) posterior edge in
Proceratophrys dibernardoi sp. nov.

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 Interocular ridge. The interocular ridge is joined at the anterior edge (not at the anterior tip) of the eyelid and
is formed by a single row of enlarged tubercles. This row is not continuous to the eyelid tubercles. We found four
states of this character (Figure 4): A) continuous; B) with medial gap; C) tubercles not organized in row; and D)
absent.
Enlarged forearm tubercles. An interesting morphological trait of Proceratophrys is the presence of several
enlarged smooth tubercles placed on several portions of the body, as at the corner of the mouth, close to the
cloacae, and at the external or ventrolateral surfaces of forearms. These tubercles are often not pigmented and lack
the typical spicules found on others body tubercles (Figure 5), but its function remains unknown.
In the forearm, these tubercles are organized in rows (one row in P. vielliardi and P. carranca, two in P. bagnoi
sp. nov., P. strussmannae, and P. rotundipalpebra, or three, in some specimens of P. goyana). In some individuals
of P. goyana the tubercles are also organized in one row plus some scattered tubercles (Figure 6).
Presence of spiculae in palmar surfaces. We found two states in this character. In Proceratophrys bagnoi sp.
nov. spicules are absent in palmar surfaces, but present in other species (Figure 7).
Eyelid and ODR contact point. In most species, the ODR touch the eyelid tubercles at the posterior third of
eyelid, often in the posterior tip of the medial eyelid tubercle. More two states were observed within the analyzed
specimens. In Proceratophrys bagnoi sp. nov. the ODR touch the eyelid in its medial position, whereas in P.
dibernardoi sp. nov. the ODR touch the eyelid in its posterior tip (Figure 8).

Results

The high number of syntopic species of Proceratophrys in Cerrado localities is surprising. Since all species
belongs to the same phenetic group, they looks very similar between them and a correct identification relies on the
combination of several characteristics. Based on the specimens observed, it was possible to recognizes some
populations that represents new species, described below. Considering it, Proceratophrys goyana, the first species
described for central Brazil, occurs syntopically with several species along its distribution. For example, at
Chapada dos Veadeiros it could occurs with P. rotundipalpebra and P. branti, whereas in Serra da Mesa, Minaçu, it
could occurs along with P. bagnoi and P. branti. However, these species could be taken apart based on some
characteristics, as the pattern of eyelid tubercles and forearm tubercles (see below), and we are convinced that all
species described where are independent evolutionary lineages. Whereas Proceratophrys goyana seems to show
large distribution and can use different habitats along it range, P. bagnoi is related to rivulets placed on slopes on
the Alto Tocantins river valley region, P. branti was found on sand substrate areas, and P. dibernardoi is a species
related to Chapadas (typical Central Brazil relief, similar to Mesas) on areas belonging to the Alto Araguaia river
region.

Accounts on species

Proceratophrys bagnoi sp. nov.

(Figs. 9–11)

Proceratophrys cf. cristiceps—Oda et al. 2009.

Holotype. CHUNB 35803, adult male, collected at the UHE Serra da Mesa power plant (13°49’47.5”S;
48°19’17”W; 550 m a.s.l.; WGS84 datum), Municipality of Minaçu, State of Goiás, Brazil, on 20 November 1999,
by R.A. Brandão.
Paratypes. CHUNB 35801, 35804 (adult females), 35802 (adult male), collected with the holotype; CHUNB
69223 (adult male), 69224, 69225 (adult females), collected at “Mata do Alex”, Serra da Mesa region
(13°45’03.5”S; 48°18’08”W, 845m a.s.l.; WGS84 datum), Municipality of Minaçu, State of Goiás, Brazil, in
September 1998, by A.F.B. Araújo.
Diagnosis. A Proceratophrys species belonging to the P. cristiceps group. Proceratophrys bagnoi sp. nov. is
characterized by: (1) small to medium size (31.4–47.2 mm SVL in males; 43.7–58.9 mm SVL in females; Table 1);
(2) a pointed eyelid tubercle, forming an small flesh point (Figure 9); (3) snout rounded from above, vertical in

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profile; (4) continuous ocular-dorsal ridge of warts; (5) tympanic region slightly concave; (6) presence of dark gray
spots on the outer margin of the ocular-dorsal ridge of warts, forming a characteristic pattern; (7) ocular-dorsal row
from the medial portion of the eyelid; (8) absence of spicule on the palmar surface; (9) eyelid tubercles formulae L
0-2, 3/5 -4/5, 1-6; R 1-3, 2/5-4/5, 2-4; (10) forearm tubercles formulae L I(4-9), II (3-6); R I (5-8), II (3-6).

FIGURE 9. Detail of the eyelid ornamentation of Proceratophrys bagnoi sp. nov., showing warts belonging to the
dorsal-ocular ridge of warts and tubercles of the upper eyelid. Note the development of the medial eyelid appendage.

Comparison with other species. Proceratophrys bagnoi sp. nov. differs from other species in the P. cristiceps
group by having conical and pointed, not fused, anterior and posterior tubercles on the border of the upper eyelid
(anterior and posterior tubercles rounded and not pointed in P. cristiceps, P. minuta, P. moratoi, P. strussmannae,
and P. vielliardi; pointed but not fused in P. cururu; fused with short points in P. goyana; slightly fused but without
appendages in P. huntingtoni); by having an elongated, fused tubercle at medial portion, showing a conspicuous
flesh extremity (not present in P. carranca, P. cururu, P. huntingtoni, P. moratoi, P. rotundipalpebra, and P.
vielliardi; present but smaller in P. caramaschii, P. cristiceps, P. goyana, and P. aridus); by continuous longitudinal
ridge of warts from the medial portion of the eyelid to the coccyx (interrupted in the supra-sacral region or in the
pre-sacral constriction in P. carranca, P. cristiceps, P. cururu, P. huntingtoni, P. concavitympanum, and P.
rotundipalpebra; absent in P. moratoi and P. vielliardi); by the vertical snout in profile (obtuse in P. aridus, P.
caramaschii, P. carranca, P. cristiceps, P. cururu, P. huntingtoni, P. moratoi, P. vielliardi, and P. strussmannae); by
having SVL ranging from 31.4–47.2 mm in males and from 43.7–58.9 mm in females (SVL in males of P. moratoi,
P. minuta, and P. redacta ranging from 19.2–30 mm; females SVL ranging from 25.9–34.9 mm), and by having the
snout round in dorsal view (snout triangular in dorsal view in P. aridus). Proceratophrys bagnoi sp. nov. occurs
sintopically with the morphologically similar P. goyana at Serra da Mesa and Niquelândia, but can be distinguished
by having palpebral formula (L 0–3, 4/5, 1–4; R 0–3, 4/5, 1–4 in P. goyana), by the absence of spicule on the
palmar surface (spicule present on P. goyana), by having the ODR touching the eyelid margin on its medial portion
(touching in the posterior third in P. goyana), and by prolonged medial eyelid extremity.
Description of holotype. Adult male; general aspect robust (Figures 10, 11 A, and 11 B); head 1.7 times wider
than high, 1.3 times wider than long; snout round in dorsal view, vertical in profile (Figure 11 C); nares simple,
elongated, dorsal, small; upper eyelid tubercles short along all eyelid margin, with one or more pointed; medial
tubercle with developed, sharply and flesh pointed, occupying about 3/5 of the eyelid border (Figure 9); eyelid
tubercles formula L 2, 3/5, 6; R 3, 3/5, 2; longitudinal ocular-dorsal ridge of warts complete, without gaps or

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FIGURE 10. Unvouchered adult male of Proceratophrys bagnoi sp. nov. from Serra da Mesa, Municipality of Minaçu,
State of Goiás, Brazil. Photo: Antonio Sebben.

interruptions; canthal crest present, evident, from the eyelid border to the anterior margin of nares, without
granules; interocular region concave, without granules, except for few ones anterior to the interocular ridge of
warts; canthus rostralis evident, slightly concave; tympanic region slightly concave; inferior border of eyes
without granules; maxillae margin poorly barred, virtually unbarred in ventral view (Fig. 11 B); tongue cordiform
covered by small round papillae, with two discrete grooves close to the basis; choanae round, evident; two
vomerine teeth patches, close to choanae; vocal slit just behind the tongue basis, and covered by a thin membrane;
ostium of Eustaquian tube elliptic, placed on the palate sides, posteriorly to the eye globes, close to the oral
corners; forelimbs short, robust; fingers free, without vestiges of web; relative fingers size IV>II>III>V (Fig. 11
D); subarticular tubercles on phalange 2 in fingers IV and V; fingers tips not blackened, with discrete fringe;
palmar surface with conical tubercles not spiculated; absence of spicules in the space between palmar tubercles;
inner metacarpal tubercle large, round; outer metacarpal tubercles elongated, widely fused at median portion; two
series of tubercles on forearms; conical tip of the forearm tubercles without spicules; forearm tubercles formula L I
6, II 4; R I 5, II 3; hindlimbs small; foot web formula I 1 – 1 II 21/2 – 1 III 2 – 21/2 IV 2 – 2 V; interdigital web paler
than plantar surface; relative toes size IV>III>V>II>I (Fig. 11 E); subarticular and plantar tubercles without
spicules; spicule present in the space between plantar tubercles; subarticular tubercles present at the base of all
toes; subarticular tubercles in phalange 2 of toes III, IV, and V; subarticular tubercles in phalange 3 of toe IV; toes
tips cream light, with discrete fringe; inner metatarsal tubercle developed, keratinized, similar to a shovel; outer
metatarsal tubercle absent; ocular-dorsal ridge of warts complete, presenting constrictions close to the presacral
and urostyle regions, forming a spear-like drawn posteriorly oriented; dorsal region delimited by the ocular-dorsal
ridge of warts tan; rest of the body with ochre background; warts of the ocular-dorsal ridge very close and more
elongated than other granules on the body; tubercles at dorsal surface densely covered by diminute spicules; some
large conical tubercles on the lateral side of the body, forming discrete rows antero-posteriorly oriented; flanks and
dorsal surface of limbs covered by small, irregular spiculated granules; ventral region covered by small, regular

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FIGURE 11. Holotype of Proceratophrys bagnoi sp. nov. (CHUNB 35803) from Serra da Mesa, State of Goiás, Brazil.
A) dorsal view; B) ventral view; C) head profile; D) palmar view; E) plantar view. Bars equal 10 mm.

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FIGURE 12. Dorsal view of individuals from series type of Proceratophrys bagnoi. Females on upper row and males on
inferior row.

spiculated granules; small dark gray spots on ventral region, darker on belly than on gular region, forming a
vermiculate pattern; right foot deformed; an evident row of granules from the base of toe I to the median portion of
the shank; presence of one large tubercle on the oral corner, lighter than others; very large hands.
Measurements of holotype (mm). SVL 41.2; HW 20.2; HH 12.1; HL 15.2; ED 5.2; EMD 4.3; IOD 8.5; FAL
10.3; HAL 13.6; THL 19.3; SHL 16.8; TAL 7.4; FOL 20.5.
Color in life. Dorsal background gray, with some brownish regions in flanks; dorsal area delimited by the
ocular-dorsal ridge of warts light brown, without spots or blotches; dark gray spots beginning on the outer margin
of the ocular-dorsal ridge of warts. Venter light cream, with grayish vermiculations.
Color in preservative. Dorsal background light brown; dorsal spots and bars on limbs brown; dorsal area
delimited by the ocular-dorsal ridge of warts cream; venter light cream with faded vermiculations.
Variation. Females are larger than males (Table 1); some individuals have the region delimited by the ocular-
dorsal ridge of warts grayish, but only one female presented blotches. The tympanic region is concave in all
individuals observed. About 80% of all individuals have small gaps at the medial portion of the IOR. Most of
individuals (70%) show pale and isolated blotches at the venter, the ventral surface is immaculate in 10% of the
specimens, and 20% present pale and anastomosed small spots. The ODR is continuous in all individuals analyzed.
Most individuals were collected using pitfall traps and some individuals died in the traps. Due to this, some
individuals in the type series became darkened and some characteristics were not so evident at a first glance.
Etymology. The specific name is a homage to Marcelo Araújo Bagno, an amazing ornithologist and great

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friend untimely deceased in a boat accident in the Caiapó River, State of Goiás, during a field work, on 11 January
2001.
Natural history notes. Males were found calling in campo sujo and campo limpo physiognomies (sensu
Ribeiro and Walter 1998), close to gallery forests. One individual was found calling in a rivulet placed on a valley
forest. Most individuals were captured in pitfall traps, including an amplectant pair. The traps were placed close to
a stream, where probably occurs the egg laying and larval development, suggesting that couples are formed in open
areas, moving posteriorly to streams. This species seems to be uncommon, since only one specimen was found in a
sample of 829 individuals by Oda et al. (2009) and in a sample of 388 individuals by Brandão and Araujo (2008).
Geographic distribution. Proceratophrys bagnoi is known from the municipalities of Minaçu and
Niquelândia (Oda et al. 2009), State of Goiás, Brazil (Figure 13).

FIGURE 13. A) Distribution of Proceratophrys goyana (circle) and Proceratophrys branti sp. nov. (square), showing
their type localities (dot center circle and dot center square, respectively); B) Distribution of Proceratophrys bagnoi sp.
nov. (square) and Proceratophrys dibernardoi sp. nov. (circle), showing their type localities (dot center square and dot
center circle, respectively). The area selected in the inset map corresponds to the magnified region in each map.
Abbreviations for Brazilian states are Bahia (BA), Goiás (GO), Maranhão (MA), Minas Gerais (MG), Mato Grosso
(MT), Mato Grosso do Sul (MS), Piauí (PI), Tocantins (TO), and Distrito Federal (DF).

Proceratophrys branti sp. nov.

(Figs. 14–15)

Proceratophrys sp. – Vitt et al. 2005.

Holotype. CHUNB 41977, adult male, collected at the Estação Ecológica do Jalapão, Municipality of Mateiros
(10o30’18”S, 46o26’W; 540 m a.s.l.; WGS84 datum), State of Tocantins, on 21 November 2004, by G.R. Colli.
Paratypes. CHUNB 41978–41979 (males), collected with the holotype; CHUNB 27366, 27368–27373,
27375–27378, 27382, 27610 (males), collected at Mateiros, State of Tocantins, on 13 February 2002, by G.R. Colli;
CHUNB 38585 (male), collected at Mateiros, State of Tocantins, on 21 November 2004, by G.R. Colli.

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TABLE 1. Mean, standard deviation, and range (in parenthesis) of measurements (in mm) of males and females of
Proceratophrys bagnoi sp. nov., Proceratophrys branti sp. nov., and Proceratophrys dibernardoi sp. nov.
Measurements Proceratophrys bagnoi Proceratophrys branti Proceratophrys dibernardoi
Males Females Males Females Males Females
(N=4) (N=4) (N=29) (N=7) (N=7) (N=4)
Snout-vent length (SVL) 38.1±7.5 52.6±6.4 36±4.8 47.5±4 32.1±2.1 41.5±2
(31.4–47.2) (43.7–58.9) (25.8–48.5) (42.4–55) (28.8–34.6) (39.7–43.6)
Head width 18.7±3.4 25.2±4 16.4±2.2 21.6±1.1 14±1.2 18±0.9
(HW) (15.8–22.8) (21–29.7) (12.9–22.4) (20.4–23.6) (12.3–15.5) (17.3–19.2)
Head high 11.2±2.1 13.5±2.2 9.8±1.1 12.7±0.8 8.1±0.5 10.1±0.6
(HH) (8.3–13.3) (11.4–16.6) (6.6–12.1) (11.5–13.8) (7.2–8.6) (9.5–10.7)
Head length 14.1±2.7 19.5±3.2 12.9±1.8 17±1.4 11.4±0.8 14.1±1.5
(HL) (11.8–17.4) (16.1–23.3) (10.3–17.5) (15.3–19.8) (10.4–12.5) (13.2–16.4)
Eye diameter 4.6±0.6 5.9±0.9 4.2±0.6 5.1±0.2 3.6±0.7 4±0.8
(ED) (4.1–5.2) (5–7.2) (3–5.4) (4.9–5.3) (3.2–4.7) (3.2–5.1)
Eye-nostril distance (END) 3.4±0.7 4±0.5 3±0.5 3.5±0.3 2.9±0.4 3±0.3
(2.7–4.3) (3.6–4.7) (2.3–3.8) (3.2–3.9) (2.3–3.4) (2.8–3.4)
Interorbital distance (IOD) 7.4±1.1 9.2±1.1 6±0.8 7.5±1.1 5.4±0.3 6.5±1.1
(6.1–8.5) (8.2–10.6) (4.7–7.6) (5.7–8.6) (4.9–5.8) (5.2–7.7)
Forearm length (FAL) 9±1.9 12.1±1.5 7.6±1.2 9.8±0.9 7±0.7 8.5±1.4
(7–11) (10.4–13.9) (5.6–10.8) (8.3–11.3) (6–8.2) (7.4–10.4)
Hand length (HAL) 12.1±1.7 15.1±2.4 9.4±1.1 12.1±0.6 9±0.5 10.4±0.4
(10.4–13.6) (13.3–18.6) (7.8–12.3) (11.3–12.8) (8.2–9.6) (10.1–10.8)
Thigh length (THL) 16.1±2.4 19.9±4.6 13.1±1.6 17.6±1.1 12±1 15.5±1.2
(13.6–19.3) (15.2–25) (10.7–17.4) (16.3–19.1) (11–13.7) (14.4–17)
Shank length (SHL) 15.4±2.2 20±2.9 12.5±1.5 (9.9– 16.2±0.7 11.8±0.6 (11– 14.7±0.8
(13.3–17.8) (16.5–23.7) 16.2) (15.6–17.6) 12.6) (13.8–15.9)
Tarsus length (TAL) 7±1.6 9.8±1.8 5.8±0.8 7.7±0.7 5.5±0.8 7.3±0.8
(5.3–9) (7.1–10.8) (4.4–8) (7.2–9.1) (4.5–6.5) (6.7–8.4)
Foot length 18.1±2.8 22.2±3.5 14±1.8 18.3±1.3 (17– 13±1 16.8±1
(FT) (15.1–20.5) (19.3–26.6) (11–19.6) 20.9) (11.8–14.3) (15.8–18.2)

Referred specimens. CHUNB 16120 (male), collected at Palmas (10°7’8”S; 48°13’20”W; 580 m a.s.l.;
WGS84 datum), State of Tocantins, in October 2000, by R.A. Brandão and A.K. Péres Jr.; CHUNB 24007 (male),
collected at Palmas, State of Tocantins, on 01 March 2001, by R.A. Brandão and G.C. Costa; CHUNB 24618–
24619 (males), collected at Alto Paraíso de Goiás (14°4’38”S; 47°30’38”W; 1340 m a.s.l.; WGS84 datum), State
of Goiás, no date, by G.R. Colli; CHUNB 33953 (male), collected at Chapada Gaúcha (15°23’30”S; 45°34’34”W;
735 m a.s.l.; WGS84 datum), State of Minas Gerais, on 04 November 2002, by P.H. Valdujo and G.C. Costa;
CHUNB 36659 (male), collected at Pedro Afonso (9°2’43”S; 48°2’43”W; 250 m a.s.l.; WGS84 datum), State of
Tocantins, no date, by M.B. Lion; CHUNB 38129–38130, 38140 (males), collected at Paranã (12°35’22”S;
47°50’37”W; 315 m a.s.l.; WGS84 datum), State of Tocantins, on 25 March 2004, by G.R. Colli; CHUNB 50246
(male), collected at Serra da Mesa (13°47’S; 48°17’40”W; 810 m a.s.l.; WGS84 datum), Minaçu, State of Goiás, in
November 1998, by R.A. Brandão; CHUNB 50343 (male), collected at Colinas do Sul (14°12’34”S; 48°3’16”W;
485 m a.s.l.; WGS84 datum), State of Goiás, on 10 October 2006, by P.H. Valdujo and D. Velho; CHUNB 50946
(female), collected at Minaçu (13°36’29”S; 48°10’38”W; 500 m a.s.l.; WGS84 datum), State of Goiás, on 04
October 2007, by C. Cândido; CHUNB 52398 (female) and CHUNB 52408 (male), collected at Monte Alegre de
Goiás (13°19’2”S; 46°53’54”W; 480 m a.s.l.; WGS84 datum), State of Goiás, no date, by G.R. Colli; CHUNB
62730 (female), 62731, 62736–62737 (males), collected at Combinado (12°52’40”S; 46°40’11”W; 545 m a.s.l.;
WGS84 datum), State of Tocantins, on 20 October 2009, by C. Cândido; CHUNB 62732 (male), collected at
Conceição do Tocantins (12°14’45”S; 47°16’29”W; 410 m a.s.l.; WGS84 datum), State of Tocantins, on 09
October 2009, by C. Cândido; CHUNB 62733 (male), collected at Paranã, State of Tocantins, on 08 October 2009,
by C. Cândido; CHUNB 72516 (male), collected at Parque Estadual da Serra do Lajeado, Palmas (10°9’38”S;
48°10’2”W; 580 m a.s.l.; WGS84 datum), State of Tocantins, on 15 December 2012, by R.F. Magalhães and R.N.
Costa.

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 Diagnosis. A Proceratophrys species belonging to the P. cristiceps group. The species is characterized by: (1)
small to medium size (25.8–48.5 mm of SVL in males; 42.4–55.0 mm of SVL in females; Table 1); (2) anterior and
posterior eyelid tubercles short, round, medial slightly pointed, being the medial one without appendages; (3) snout
rounded in dorsal view, vertical in profile; (4) presence of a ocular-dorsal ridge of warts poorly evident,
incomplete, interrupted at the sacral region; (5) tympanic region slightly concave; (6) black anastomosed dorsal
spots absent; (7) a conspicuous lighter stripe between eyes; white (or very lighter) spots on the anterior and
posterior region of eyes; lighter stripes bordering the inner edge of the ocular-dorsal ridge of warts, forming a “X”
like pattern; (8) dorsal granules densely covered by spicules, conferring a rough aspect; (9) one row of forearm
tubercles; (10) forearm formula L I 3–7, R I 3–8; (11) eyelid formula L 0–3, 3/5-5/5, 1–4; R 0–3, 3/5-5/5, 0–5.

FIGURE 14. Unvouchered male of Proceratophrys branti sp. nov. from Estação Ecológica do Jalapão, Municipality of
Mateiros, State of Tocantins, Brazil. Photo: Janalee P. Caldwell.

Comparison with other species. Proceratophrys branti sp. nov. differs from other species of the P. cristiceps
group by showing anterior and posterior tubercles on border of upper eyelid slightly fused, but without appendages
(rounded, not fused in P. cristiceps, P. huntingtoni, P. moratoi, P. redacta, and P. vielliardi; fused and short ocular
appendages in P. goyana, P. concavitympanum, and P. strussmannae; pointed but not fused in P. cururu; not fused
and conical in P. bagnoi); by medial eyelid tubercle very long, covering 3/5 to 5/5 of the eyelid border, without
points (medial eyelid tubercle never covering all eyelid border and with points in P. goyana, P. cururu, P. aridus, P.
caramaschii, P. cristiceps, P. strussmannae, and P. concavitympanum; medial tubercle with conspicuous flesh
appendage in P. bagnoi); by presence of the ocular-dorsal ridge of warts interrupted at sacral region (continuous in
P. goyana, P. bagnoi, P. concavitympanum, and P. strussmannae; absent in P. moratoi and P. vielliardi); by having
vertical snout profile (obtuse in P. aridus, P. caramaschii, P. carranca, P. cristiceps, P. cururu, P. huntingtoni, P.
moratoi, P. vielliardi, and P. strussmannae); by females presenting SVL 42.4–55 mm (SVL in females varying
from 25.9–34.9 mm in P. minuta, P. redacta, and P. moratoi).

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FIGURE 15. Holotype of Proceratophrys branti sp. nov. (CHUNB 41977) from Estação Ecológica do Jalapão,
Municipality of Mateiros, State of Tocantins, Brazil. A) dorsal view; B) ventral view; C) head profile; D) palmar view;
E) plantar view. Bars equal 10 mm.

338 · Zootaxa 3750 (4) © 2013 Magnolia Press BRANDÃO ET AL.

 Description of holotype. Adult male; aspect robust (Figures 14 and 15), head 1.6 times wider than high, 1.3
times wider than long; snout round in dorsal and ventral views (Figures 15 A, B), vertical in profile (Figure 15 C);
nares simple, elongated, dorsal, with small opening; anterior and posterior tubercles of upper eyelid, short, conical,
slightly pointed; medial tubercle covering 4/5 of upper eyelid border, without point; eyelid formulae L 1, 4/5, 3; R
1, 4/5, 1,dorsum covered by round tubercles strongly spiculated; ocular-dorsal ridge of warts discontinuous on
sacral region; inner edge of the ocular-dorsal ridge of warts with light gray irregular stripes; evident light gray
stripe between eyes; white spot on anterior and posterior edge of eyes; interocular region concave; all area between
eyes and nares covered by spiculated tubercles; canthus rostralis evident; canthal crest present, poorly evident;
absence of spicules on the inferior border of eyes; single, white, large tubercle on mouth corner; maxillae
presenting dark bars, covered only by spicules, without tubercles; jaw without bars; tympanum hidden, tympanic
region slightly concave; two small, round vomerine teeth patches, placed between choanae; choanae large, round;
tongue cordiform, with dorsal surface smooth, covered by small, round papillae; absence of grooves on tongue;
large vocal slit, single, placed on the posterior margin of the tongue basis; ostium of Eustaquian tube elliptical,
placed on the sides of palate, posteriorly to ocular globes, close to the oral corner; forelimbs short, robust; fingers
free, without vestiges of interdigital web; relative fingers size IV>V>III>II (Figure 15 D); palmar surface covered
by light conical granules, that highlights against the darker background; palmar surface densely covered with
darkened spicules in the spaces between palmar tubercles; inner metacarpal tubercle round; supernumerary
tubercles on sole and fingers, two outer metacarpal tubercles, being the proximal larger than the distal; subarticular
tubercles on basis of all fingers; subarticular tubercle on phalange 2 of fingers IV and V; fingers tips not blackened,
round, with discrete fringe on fingers III, IV, and V; forearm row tubercles conical, pointed, organized in a single
row, containing five tubercles on left forearm, and seven tubercles on right forearm (L I 5; R I 7); tip of forearm
row tubercles without spicules; forearm row tubercles slightly larger than other tubercles on anterior limb;
hindlimbs small; foot with plantar surface covered by small, conical, spiculated tubercles (Figure 15 E);
subarticular tubercles evident on the basis of all toes; interdigital web poorly developed; foot web formula I 2 – 1 II
2½ – 1 III 2 – 1 IV 1 – 1 V; toe tips not blackened, with discrete fringe; inner metatarsal tubercle large, forming a
shovel with external margin keratinized; outer metatarsal tubercle absent; subarticular tubercles on phalange 1 of
toes I and II, on phalanges 1 and 2 of toes III and IV, and phalanges 1, 2, and 3 of toe IV; relative toes size
IV>III>V>II>I; dorsum with complex color pattern formed by irregular brown, gray, black, cream, and strawy
blotches; most of the warts in the ridge are more elongated and larger than other ones on the dorsum; flanks and
dorsal surface of limbs with smaller granules than those of dorsum; dorsal surface of limbs with dark gray
transversal bars; ventral surface covered by diminute, regular, spiculated granules; vocal sac evident, darker than
ventral region; ventral region marbled by dark spots; two larger tubercles close to vent.
Measurements of holotype (mm). SVL 35.3; HW 16.9; HH 10.3; HL 12.6; ED 4.3; END 3.2; IOD 6.8; FAL
7.2; HAL 8.8; THL 12.5; SHL 11.5; TAL 5.1; FOL 14.
Color in life. Dorsal background gray to dark gray, with a complex pattern of brown, tan, cream, and strawy
blotches. Light gray spots on anterior and posterior margins of eyes; irregular light gray or light brown stripes
bordering the inner edge of the ocular-dorsal ridge of warts; irregular cream blotches on dorsum; evident light gray
stripe between eyes. Palmar and plantar tubercles pink, especially in younger individuals; external margin of
maxillae lighter than dorsum.
Color in preservative. Most of specimens present light gray background on dorsum; some exhibit yellowish
coloration on dorsal tubercles (especially those from northeastern State of Tocantins). The lighter stripes became
light gray, and ventral black spots became faded.
Variation. Variation in measurements is presented in Table 1. Females are larger than males (Figure 16). Some
individuals presents fingers tips more elongated than others. Dorsal coloration varies from dark gray background to
light gray. In some individuals, the interocular stripe is gray instead of light gray. Most individuals (80%) presents
the ODR discontinuous from supra-sacral region, but the ODR interruption could be observed on the presacral
region (12%), or on supra-scapulae (8%). The IOR is not organized on most individuals (58%), not organized in
row in 21% or with medial gap in 21% of them.
Etymology. The specific epithet is a homage to our friend Eng. Agronomist Sérgio Brant Rocha, by his relevant
contribution for the creation and establishment of several areas for biodiversity protection in Brazil. Throughout his
brilliant career, Sérgio Brant was official of the Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais
Renováveis (IBAMA), Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio), and Ministério do

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Meio Ambiente (MMA), where he developed a painstaking work for the creation of some protected areas where this
new species could be found, as the Grande Sertão Veredas National Park and Nascentes do Parnaíba National Park.
Besides the creation of some of the most relevant protected areas in Brazilian Cerrado, his major contribution was
claim for more rigor and earnest policies for nature conservation and public property in Brazil.
Natural History. Proceratophrys branti males call directly from soil, in open Cerrado physiognomies, often
related to sand or rocky substrate.
Geographical distribution. Proceratophrys branti occurs in a wide region in central-north portion of Cerrado
biome, especially in areas of sandy or rocky soil. Besides its type locality (Mateiros, State of Tocantins), the species
was recorded in the states of Tocantins (municipalities of Dianópolis, Porto Alegre do Tocantins, Combinado,
Conceição do Tocantins, Paranã, Palmas, Taquaralto, and Pedro Afonso), Goiás (municipalities of Minaçu, Alto
Paraíso de Goiás, Colinas do Sul, Monte Alegre de Goiás, Sítio d’Abadia, and São Domingos), and Minas Gerais
(Municipality of Chapada Gaúcha) (Figure 13).

FIGURE 16. Dorsal view of individuals from series type of Proceratophrys branti. Females on upper row and males on
inferior row.

Proceratophrys dibernardoi sp. nov.

(Figures 17–18)

Proceratophrys sp.—Vaz-Silva et al. 2007.

Holotype. CHUNB 28131, adult male, collected at the Municipality of Mineiros (17o37’09”S, 52o30’59”W; 715m
a.s.l; WGS84 datum), State of Goiás, Brazil, on 01 April 2002, by P.H. Valdujo.
Paratypes. CHUNB 28132 (male), 28133–28134 (females), collected with the holotype; CHUNB 50305
(male), collected at the Municipality of Rio Verde (17°48’29”S; 51°22’26”; 855 m a.s.l.; WGS84 datum), State of
Goiás, no date, by D.A. Velho; CHUNB 65844 (male), collected at the Municipality of Alto Araguaia (17°02’30”S;
53°07’17”W; 520 m a.s.l.; WGS84 datum), State of Mato Grosso, on 23 March 2000, by A.F.B. Araújo and R.A.
Brandão; CHUNB 69222 (female), collected at the Municipality of Aporé (18º39’48”S; 51º52’17”W; 600 m a.s.l.;
WGS84 datum), State of Goiás, on 16 April 2003, by W. Vaz-Silva and P.V.C. Mathias; CHUNB 69220 (female),
collected at the Municipality of Alto Taquari (17°41’51”S; 53°27’45”W; 725 m a.s.l.; WGS84 datum), State of
Mato Grosso, no date, by P.H. Valdujo; CHUNB 69219, 69221 (males), collected at the Parque Estadual da Serra
Azul (approx. 15°51’24”S; 52°15’20”W, 520 m a.s.l.), Municipality of Barra do Garça, State of Mato Grosso, no
date, by P.H. Valdujo.

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 Diagnosis. A Proceratophrys species belonging to the P. cristiceps group. The species is characterized by: (1)
small to medium size (28.4–34.6 mm SVL in males; 39.7–43.6 mm SVL in females, Table 1); (2) anterior and
posterior tubercles on external margin of upper eyelids conical, slightly fused; (3) medial eyelid tubercle covering
4/5 of eyelid margin, with small points; (4) snout rounded from above, vertical in profile; (4) ocular-dorsal ridge of
warts poorly defined, discontinuous close to sacral region; (6) ocular-dorsal ridge of warts touching the eyelid in its
posterior end (7) tympanic region indistinct; (8) dorsal background grayish, small scattered ocelli on dorsal surface;
a marked “X” like pattern formed by two light gray stripes that borders the inner edge of the ocular-dorsal ridge;
(9) presence of spots on anterior and posterior regions of eyes; (10) dorsal granules round, regular in size; (11)
eyelid formula L 0–2, 4/5, 1–2; R 0–2, 4/5, 1–2; (11) forearm tubercles formula L I(4–9), II(0–4); R I(4–7), II(0–5).

FIGURE 17. Unvouchered male of Proceratophrys dibernardoi sp. nov. from the Municipality of Aporé, State of Goiás,
Brazil. Photo: Sheila Andrade.

Comparison with other species. Proceratophrys dibernardoi sp. nov. shows anterior and posterior tubercles
on upper eyelid border not fused and round to conical, with small points (fused and showing with small points in P.
goyana, P. strussmannae, P. carranca, P. branti, and P. concavitympanum; pointed but not fused in P. cururu;
slightly fused, without appendage in P. huntingtoni, P. vielliardi, and P. moratoi; conical and pointed in P. bagnoi);
medial eyelid tubercle without ocular appendages (with short ocular appendages in P. goyana, P. cururu, P.
strussmannae, P. caramaschii, P. cristiceps, P. aridus, and P. concavitympanum; long and without appendage in P.
huntingtoni, P. vielliardi, P. moratoi, and P. branti; with an elongated and pointed appendage in P. bagnoi; with
several cusps in P. minuta); by presenting an interrupted ocular-dorsal ridge of warts (continuous in P. goyana, P.
strussmannae, and P. bagnoi; absent in P. moratoi and P. vielliardi); by showing vertical snout profile (obtuse in P.
aridus, P. caramaschii, P. carranca, P. cristiceps, P. cururu, P. moratoi, P. vielliardi, and P. huntingtoni); head
round in dorsal view (triangular in dorsal view in P. aridus); males SVL 28.8–34.6 mm (males SVL from 39.5–56.3
mm in P. cristiceps, P. caramaschii, P. concavitympanum, and P. strussmannae); females SVL 39.7–43.6 mm
(female SVL 43.7–65 mm in P. bagnoi, P. caramaschii, P. concavitympanum, and P. strussmannae); dorsum with
small rounded spots (absent in P. aridus, P. cristiceps, P. minuta, P. goyana, and P. concavitympanum); presence of
irregular white stripes on the internal edge of the ocular-dorsal ridge of warts, forming a “X” shaped pattern
(present in P. moratoi; incomplete in P. branti and P. carranca); dorsal rows of tubercles interrupted in sacral

NEW SPECIES OF PROCERATOPHRYS FROM CENTRAL BRAZIL Zootaxa 3750 (4) © 2013 Magnolia Press · 341
FIGURE 18. Holotype of Proceratophrys dibernardoi sp. nov. (CHUNB 28131) from Municipality of Mineiros, State of
Goiás, Brazil. A) dorsal view; B) ventral view; C) head profile; D) palmar view; E) plantar view. Bars equal 10 mm.

342 · Zootaxa 3750 (4) © 2013 Magnolia Press BRANDÃO ET AL.

FIGURE 19. Dorsal view of individuals from series type of Proceratophrys dibernardoi. Females on upper row and
males on inferior row.

region (continuous to the coccyx in most individuals of P. caramaschii, P. aridus, P. minuta, P. bagnoi, and P.
goyana); absence of light interocular stripe (present in P. branti and P. carranca) and of a white spot on the anterior
and posterior margins of eyes (present in P. branti, P. carranca, and P. cururu); ocular-dorsal ridge touching the
eyelid on its posterior margin (ODR touching eyelid on its posterior third in most species; touching the medial
portion of eyelid in P. bagnoi).
Description of holotype. Body small, robust (Figures 17, 18 A, 18 B). Head 1.6 times wider than high, 1.1
times wider than longer; head length 0.45 times body length, head width 0.48 times total length; snout short, round
in dorsal and ventral views (Figures 18 A, 18 B), vertical in profile (Figure 18 C); nostrils directed dorsolaterally;
nostrils aperture circular; internarial distance smaller than eye to nostril distance (54%), and 0.13 times the head
width; eyes large, located anterolaterally (Figure 18 C), eye diameter 0.27 times head length; upper eyelids well
developed covering eyes in dorsal view; tubercles of the eyelid external margin not protuberant; eyelid formula L
1, 4/5, 1; R 0, 4/5, 2; loreal region gently sloping, showing two protuberant tubercles; canthus rostralis distinct;
canthal crest present, evident, from eye to nostril; interocular region concave; granules on the interocular ridge
region not organized on a row; vomerine teeth in two small patches, transverse, well-separated, located behind the
choanae; choanae widely separated; tongue large, cordiform; tympanum hidden, tympanic region indistinct; a clear
spot between upper eyelids; skin on dorsum granular, warty, ornamented with tubercle ridges of variable
arrangement and size; dorsal surface of arms and legs presenting tubercles with minute keratinous spines; distinct
longitudinal ocular-dorsal ridge of warts, discontinuous on the sacral region. One row of tubercles on forearms.

NEW SPECIES OF PROCERATOPHRYS FROM CENTRAL BRAZIL Zootaxa 3750 (4) © 2013 Magnolia Press · 343
Forearm tubercles formula L I(6), R I(7). Skin on venter granulated with small, homogeneous tubercles. Fingers
without webbing (Figure 18 D); tips of fingers not dilated or darkened; relative lengths of fingers IV>II>V>III;
subarticular tubercles distinct, rounded; palmar tubercles small, distinct; spiculae present between palmar
tubercles; inner carpal tubercle oval, unique; outer carpal tubercles elongated, similar in size, separated by a deep
groove. Toes slightly webbed (Figure 18 E), tips not expanded, dorsally with small granules with keratinous spines;
webbing formula I 1 – 1 II 2 – 11/2 III 2 – 1 ½ IV 1 – 2 V; subarticular tubercles small, distinct, rounded; inner
metatarsal tubercle enlarged, elliptical, projecting, with length 0.16 times foot length; outer metatarsal tubercle
very small, rounded; discrete row of tubercles extending from inner metatarsal tubercle to half of the tarsus.
Measurements of holotype (in mm). SVL 28.8, HW 12.7, HH 7.8, HL 11.2, ED 3.3, END 3.4, IOD 5.7, FAL
6.0, HAL 8.5, THL 11.4, SHL 11.0, TAL 4.5, FOL 12.1.
Color in life. Dorsal coloration ranging from tan to gray. Dark gray spots on dorsum. Plantar and palmar
tubercles white. Ventral region strawy, without spots.
Color in preservative. Dorsal background light gray, with brownish or dark gray blotches.
Variation. Variation in measurements is presented in Table 1. The specimens of the type series are variable in
the dorsal pattern coloration (Figure 19). The specimen CHUNB 28132 shows small spots on dorsum. The flanks
show large tubercles in females and small tubercles in males. Little variation was observed in the dorsal pattern
with “X” shape. About half of the available specimens presented the ocular-dorsal ridge interrupted at the presacral
constriction.
Etymology. The specific name is a noun in the genitive case in honor of Dr. Marcos Di Bernardo (in
memoriam), for his friendship and extensive contributions for the knowledge of the Brazilian herpetofauna.
Natural History. Proceratophrys dibernardoi was always found in open areas covered by Cerrado and
“veredas” (sensu Ribeiro and Walter 1998), with small streams, where probably the tadpoles develop.
Geographical distribution. Proceratophrys dibernardoi is known in the municipalities of Mineiros, Jataí, Rio
Verde, and Aporé in southwestern State of Goiás, and in the municipalities of Alto Taquari, Alto Araguaia, and
Barra do Garças, in the State of Mato Grosso, Brazil (Figure 12).

Discussion

Although we used external morphology characters to describe three new Proceratophrys of the Cerrado biome, we
strongly suggest that other set of characters must also be used for future taxonomic studies with this genera. Some
morphological characters are variable within the same species, making very difficult to find appropriate diagnostic
(synapomorphies) characters.
The rapid increment on new Proceratophrys for the Cerrado biome suggests that the diversity of the genus in
the Cerrado is underscored. In the last five years, seven species in the genus have been described within the
Cerrado biome or from contact areas between Cerrado and other biome limits (e.g., P. carranca, P. huntingtoni, P.
minuta, P. redacta, P. rotundipalpebra, P. strussmannae, and P. vielliardi). The central portion of the Cerrado seems
to be a hotspot of Odontophrynidae diversity in the biome, due to the occurrence of high local diversity. Some
localities, as Chapada dos Veadeiros, Serra da Mesa, and Brasília, harbor at least three Proceratophrys species, plus
two Odontophrynus species, typically O. cultripes and O. salvatori, a species morphologically similar to P. moratoi
(Caramaschi 1996), and that may belong to the genus Proceratophrys (Brandão and Batista 2000; Amaro et al.
2009). The astonish number of syntopic species is not surprising, since this pattern of species distribution was
previously observed (Teixeira Jr. et al. 2012; Martins and Giaretta 2013). This sintopy could be possible due to the
high horizontal habitat heterogeneity observed on Cerrado landscape (Garda et al., 2013). The presence of different
habitat spots placed on the Cerrado landscapes create conditions for species establishment, especially for small
vertebrates (Alho 1981), as frogs.
Although recognized as one of the world’s biodiversity hotspots (Myers et al. 2000), due to various threats
(e.g., increasing deforestation, charcoal production, large hydroelectric dams, livestock, and agricultural activities),
most areas in Cerrado are still poorly sampled. Based only on IUCN Criterion B of geographical distribution of the
species (area of occupancy and extension of occurrence), we suggest P. branti and P. goyana to be ‘Least Concern’
(LC) species, and P. dibernardoi and P. bagnoi as ‘Data Deficient’ (DD) in IUCN Red List. All records of P. bagnoi
come from Serra da Mesa hydroelectric power plant, raising concerns about the conservation status of this species.

344 · Zootaxa 3750 (4) © 2013 Magnolia Press BRANDÃO ET AL.

Acknowledgements

We thank Antonio Sebben, Janalee P. Caldwell, and Sheila Andrade for providing photos of adult individuals. To
Ana Beatriz Costa and Taynã Rapp Py-Daniel for help with some photos and specimens measurements. To Miguel
Vences, Natan Maciel and Adrian Garda for relevant comments on the manuscript and language improvements. UC
acknowledges the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for financial support.

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Appendix 1

Additional specimens examined

Odontophrynus salvatori.—BRAZIL: State of Goiás: Municipality of Alto Paraíso de Goiás, Parque Nacional da Chapada dos
Veadeiros, MNRJ 15870 (holotype), MNRJ 15871 (paratype).
Proceratophrys aridus.—BRAZIL: State of Ceará: Milagres, Minador farm, MNRJ 55782 (holotype), MNRJ 55349, 55778–
55781, 55783–55822, 75156–75157, 75158-75168 (paratypes); Jardim, CHUNB 56961–56962; Crato, CHUNB 64738–
64745.
Proceratophrys caramaschii.—BRAZIL: State of Ceará: Fortaleza, Mucuripe, MNRJ 16592 (holotype), MNRJ 1419–1420,
1680, 16470–16484, 16487–16591, 16593–16600 (paratypes). State of Piaui: Piripiri, Parque Nacional de Sete Cidades,
CHUNB 61185–61218.
Proceratophrys cristiceps.—BRAZIL: State of Pernambuco: Caruaru, CHUNB 35793; Arco Verde, MNRJ 58030; Casinhas,
MNRJ 307. State of Bahia: Feira de Santana, MNRJ 47839. State of Paraíba: Areia, MNRJ 50378; Maturéia, MNRJ
54756–54758. State of Rio Grande do Norte: Macaíba, MNRJ 1775, 1777. State of Sergipe: Brejo Grande, MNRJ 50376;
São Cristóvão, MNRJ 50377.
Proceratophrys goyana.—BRAZIL: State of Goiás: Minaçu, Serra da Mesa, MNRJ 17309–17313, 17314, 20209–20212;
Colinas do Sul, MNRJ 66224–66225, 66527.
Proceratophrys concavitympanum.—BRAZIL: State of Tocantins: Palmas, CHUNB 15263, 16121, 24260. State of Pará: Novo
Progresso, CHUNB 44890–44891; Parauapebas, Serra dos Carajás, MNRJ 58854–58855.
Proceratophrys cururu.—BRAZIL: State of Minas Gerais: Parque Nacional da Serra do Cipó, MNRJ 17905 (paratype).
Proceratophrys huntingtoni.—BRAZIL: State of Mato Grosso: Chapada dos Guimarães, Parque Nacional da Chapada dos
Guimarães, CHUNB 35975.
Proceratophrys moratoi.—BRAZIL: State of São Paulo: Botucatu, Rubião Júnior, MNRJ 14248–14249; 15872 (paratypes).
Proceratophrys rotundipalpebra.—BRAZIL: State of Goiás: Colinas do Sul, CHUNB 50338 –50342; 50344; 50347 –50348;
50351 –50353.
Proceratophrys vielliardi.—BRAZIL: State of Minas Gerais: Paracatu, CHUNB 26655, 38967–38971. Distrito Federal:
Brasília, CHUNB 48361, 57484, 57501.

NEW SPECIES OF PROCERATOPHRYS FROM CENTRAL BRAZIL Zootaxa 3750 (4) © 2013 Magnolia Press · 347

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