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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4114 (2): 171
–
181
http://www.mapress.com/j/zt/
Article
171
http://doi.org/10.11646/zootaxa.4114.2.6
http://zoobank.org/urn:lsid:zoobank.org:pub:B6EF32E1-641C-4349-BF8E-0CAABCEAC687
A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian
Brazil (Amphibia: Anura: Hylidae)
KARL-HEINZ JUNGFER
1
, VANESSA K. VERDADE
2
, JULIÁN FAIVOVICH
3,4
& MIGUEL T. RODRIGUES
5
1
Institute of Integrated Sciences, Department of Biology, University of Koblenz–Landau, Universitätsstr. 1, 56070 Koblenz, Germany.
E-Mail: khjungfer@aol.com
2
Centro de Ciências Naturais e Humanas, Universidade Federal do ABC, Av. dos Estados, 5001, CEP 09210-971, Santo André, São
Paulo, Brazil. E-mail: vanessa.verdade@ufabc.edu.br
3
División Herpetología, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”-CONICET, Angel Gallardo 470, 1405 Bue-
nos Aires, Argentina. E-mail: julian@macn.gov.ar
4
Departamento de Biodiversidad y Biologia Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires,
Argentina
5
Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11461, CEP 05508-090, São Paulo,
Brazil. E-mail: mturodri@usp.br
Abstract
A new species of treefrog of the genus Osteocephalus is described from the Rio Abacaxis, a southern tributary of the Am-
azonas in the state of Amazonas, Brazil. This member of the O. buckleyi group is characterized by green dorsal colouration
with irregular blotches of various shades of brown, light venter with tan spots and bold dark markings on the posterior
surfaces of the thighs. It can be distinguished from its closest relative, O. helenae from the same general area, by the lack
of an axillary membrane, a few indistinct tubercles on the proximal segment of Finger IV and single ulnar tubercles.
Key words: Amphibia, Anura, Hylidae; Amazonia, Brazil, Osteocephalus camufatus sp. nov.
Introduction
Spiny-backed treefrogs of the genus Osteocephalus are widespread in the Amazonian and Guianan regions of
South America. A molecular phylogenetic study (Jungfer et al. 2013) defined five species groups and revealed that
the number of species in the genus is largely underestimated and numerous species still await formal description.
One of those is a member of the Osteocephalus buckleyi species group from Central Amazonia in Brazil
distinguishable from their close relatives by molecular and morphological characters and referred to as Confirmed
Candidate Species (following the terminology of Vieites et al. 2009) in that paper. Frogs of the Osteocephalus
buckleyi group are stream-breeding, most of them occurring along the eastern Andean edge at elevations between
600 and 2300 m, namely O. carri (Cochran & Goin, 1970), O. duellmani Jungfer, 2011, O. festae (Peracca, 1904),
O. mimeticus (Melin, 1941), O. mutabor Jungfer & Hödl, 2002, and O. verruciger (Werner, 1901). Several species
also inhabit the lowlands eastward to the mouth of the Amazonas. These have been problematic from a taxonomic
point of view. They were considered one single species, O. buckleyi (Boulenger, 1882), by Trueb & Duellman
(1971), but several ones, also including an upland species (Lynch 2006), were later removed from its synonymy
(Duellman & Mendelson 1995, Jungfer 2010, Jungfer et al. 2013) or described as new (Ron et al. 2012). These
frogs from elevations mostly below 600 m currently comprise four species: Osteocephalus buckleyi from the
western Amazon Basin in eastern Ecuador and northern to central Peru (Ron et al. 2012, Jungfer et al. 2013), O.
cabrerai (Cochran & Goin, 1970) from southeastern Colombia (Cochran & Goin 1970; Lynch 2002), northeastern
Peru (Jungfer 2010; Ron et al. 2012) and northeastern Ecuador (Ron et al. 2011), O. cannatellai Ron, Venegas,
Toral, Read, Ortiz & Manzano, 2012 from lowlands, but also up to 1290 m asl. of the western Amazon Basin in
northern Colombia, eastern Ecuador and northern Peru (Ron et al. 2012, Jungfer et al. 2013), and O. helenae
(Ruthven, 1919) inhabiting a vast range from Santa Cruz, Bolivia, to the coastal Guianas (Jungfer et al. 2013).
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Among these lowland species there are frogs that are partly or predominantly green dorsally and laterally (with
individual variation), have posterior surfaces of the thighs light with dark spots or bars in preservative (though
sometimes uniform bluish or brownish in life) and are tuberculate on the head, especially on the canthus rostralis
and on the upper eyelids (Gorzula & Señaris “1988”, Duellman & Mendelson 1995, Lescure & Marty 2000, Lynch
2002, Lima et al. 2006, Kok & Kalamandeen 2008, Jungfer 2010, Ron et al. 2012). As these three characters are
absent both in the aforementioned upland members of the O. buckleyi group (Trueb & Duellman 1970, 1971,
Jungfer & Hödl 2002, Jungfer 2010, 2011, Ron et al. 2010) and all species of other groups (i.e. the O. alboguttatus,
O. leprieurii, O. planiceps and O. taurinus groups) (e.g. Trueb & Duellman 1971, Duellman 1978, Duellman &
Mendelson 1995, Jungfer et al. 2000, Jungfer & Hödl 2002), the lowland species can be distinguished relatively
easily from the rest of their congeners. But they are not so easily distinguishable morphologically from one another
(e.g. Ron et al. 2011). This also holds true for a new lowland species which has previously been identified on a
molecular basis (Jungfer et al. 2013) using tissues from specimens obtained along the Rio Abacaxis, a southern
tributary of the Amazonas in Amazonas, Brazil. Herein we proceed to its formal description.
Materials and methods
Measurements taken follow Duellman (2001), except that foot length is the distance from the bent tibiotarsal
articulation to the tip of the fourth toe. Snout length is calculated as the distance between eye and naris proportional
to head length (EN/HL). Terminology of skin texture is as described by Duellman & Lehr (2009) for terraranan
frogs. Webbing formula is that of Savage & Heyer (1967), as modified by Myers & Duellman (1982). Enumeration
of diagnostic characters follows Jungfer (2010). For ease of comparison unknown characters are also stated.
Abbreviations are as follows: ED: eye diameter; EN: distance from eye to naris; FD: diameter of finger disc on
third finger; FL: foot length; HL: head length; HW: head width; IN: internarial distance; SVL: snout-vent length;
TD: tympanum diameter; TE: distance between tympanum and eye; TL: tibia length. Measurements (all in mm)
were made using digital callipers or the ocular micrometer of a dissecting microscope if less than 5 mm. Museum
abbreviations follow Frost (2015). The numerical code used for the new species by Jungfer et al. (2013) refers to
“Ca” for candidate species, and the Genbank accession number of one of the sequences obtained from the holotype,
using the system established by Padial et al. (2010).
Results
Osteocephalus camufatus sp. nov.
(Figs. 1–4)
Osteocephalus buckleyi [Ca1_MTR12779_2748]—Jungfer et al. 2013
Holotype: MZUSP 142389 (field number MTR 12779), an adult male of 39.5 mm SVL with well-developed
nuptial pads from Brazil: Amazonas: Igarapé-açu (04°20'40''S 58°38'06''W), right bank of Rio Abacaxis, 30 m asl.,
collected by Miguel T. Rodrigues, Sergio Marques de Souza, José Cassimiro and José Mário Guellere, on 9 January
2007 (Figs. 1–4).
Paratype: MZUSP 157020 (field number MTR 13147), an adult male with well-developed nuptial pads from
Brazil: Amazonas: Areal, left bank of Rio Abacaxis (04°35'49''S 58°13'14''W), 39 m asl., collected by Miguel T.
Rodrigues, Sergio Marques de Souza, José Cassimiro and José Mário Guellere, on 20 January 2007.
Diagnosis and comparisons. Osteocephalus camufatus may be diagnosed as (1) a small to medium sized
species (as defined by Jungfer 2010: 29) in males (females unknown); (2) skin on dorsum of males shagreen with a
few irregular small tubercles, only some bearing keratinized tips; (3) skin on flanks coarsely areolate between limb
insertions; (4) rounded, tuberculate canthus rostralis curved inward; (5) frontoparietal ridges not visible externally;
(6) dentigerous processes of vomers angular; (7) thick tuberculate supratympanic fold from the posterior edge of
the orbit sloping in an arch towards the arm insertion, not reaching below tympanum posteroventrally; (8) webbing
on inner edge of third finger extending slightly beyond penultimate subarticular tubercle; (9) distal subarticular
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tubercle on Finger IV bifid; (10) dorsum boldly blotched irregularly; (11) throat, chest and venter creamy tan to
creamy white with numerous irregular tan spots; (12) supralabial area with irregular light and dark markings; (13)
flanks light with irregular dark markings; (14) vocal sacs paired, protruding ventral to angles of jaws; (15) juvenile
colouration unknown; (16) tadpole habitat (most likely in pools along streams) and labial tooth row formula
unknown; (17) colour of tibiofibular bones white in preservative.
O. camufatus is a member of the O. buckleyi species group as revealed by molecular data. A phylogenetic
analysis using both maximum parsimony (MP) and likelihood (ML) inference resulted in trees that showed that O.
camufatus was basal to a large and well-supported clade containing frogs from large areas of Amazonia and
Guiana. The oldest available name for this clade is O. helenae (Jungfer et al. 2013). Uncorrected p distances of the
mitochondrial 16S gene between O. camufatus and O. helenae from various localities were 1.3–2.6, while genetic
divergences within O. helenae, whose subclade relationships are not yet sufficiently resolved, were 0.0–2.2
(Jungfer et al. 2013: Appendix S4d).
The new species shares with several other species of the group the following character states: 1) greenish
ground colour, 2) strongly tuberculate canthus rostralis and upper eyelids and 3) posterior surfaces of thighs light
with large dark brown spots in preservative. These three characters distinguish it from all frogs in the O.
alboguttatus, O. leprieurii, O. planiceps and O. taurinus groups (Jungfer et al. 2013) in which the ground colours
are shades of tan or brown, the canthus rostralis is smooth to granulate and the posterior thigh surfaces are uniform.
From other species in the O. buckleyi group it differs as follows (characters of O. camufatus in parentheses): Six
upland species from the eastern Andean slopes between 600 and 2300 m differ as follows: in Osteocephalus carri,
a predominantly brown species (bold blotches of green and brown), the canthus rostralis and upper eyelids are
smooth (strongly tuberculate) and the iris is black with golden spots in life (light); O. duellmani is a tan species
with or without dark brown blotches (bold blotches of green and brown), with the posterior surfaces of the thighs
uniform tan (marbled) and the canthus and upper eyelid bearing a few low tubercles (strongly tuberculate); O.
festae exhibits brown ground colour (bold blotches of green and brown), a dark brown iris in life (light, Fig. 4) and
uniform tan posterior thigh surfaces (marbled). Males of Osteocephalus mimeticus, O. mutabor (the latter also
occurring as low as 150 m asl.) and O. verruciger have a brown ground color lacking green (blotches in shades of
green and brown), uniform posterior thigh surfaces (marbled dark brown), and heavily tuberculate dorsa with
numerous spinous tubercles bearing keratinized tips during breeding (a few irregular small tubercles, few of them
with keratinization). Species of the O. buckleyi group from the lowlands (< 600 m asl.), that in some cases share the
greenish ground colour of O. camufatus, differ as follows: Osteocephalus buckleyi has low tubercles or is smooth
on the upper eyelid (strongly tuberculate), and has low or lacks tarsal tubercles (prominent). Breeding males of O.
buckleyi exhibit dorsal tubercles, many with keratinized tips (few keratinized tips). Osteocephalus cabrerai has a
row of tubercles on the lower jaw (absent) and an irregular, deep fringe on the outer edge of Finger IV (low
tubercles on proximal segment of Finger IV). Osteocephalus cannatellai from lowlands, but also up to 1290 m asl.
(Ron et al. 2012), appears to be larger (38.5–57.2 mm SVL in males, mean, 46.8 mm, Ron et al. 2012) (to 40.4 mm,
but note that the sample size is n=2), and differs by a longer supratympanic fold that reaches the arm insertion (not
reaching lower level of tympanum); axillary membrane present (absent); usually a dark venter varying from light
grey to brown with or without dark markings (Ron et al. 2012) (yellowish white with small irregular brown spots).
Osteocephalus helenae (Ruthven) is a variable frog throughout its range. Specimens from localities closest to the
two sites known for O. camufatus, from about 230 km northwest of the type locality, near Manaus and other sites
just north and south of the Amazonas in Amazonas, Brazil, exhibit an axillary membrane (absent), a scalloped
fringe on the outer edge of Finger IV continued to the ulna (low, indistinct tubercles on proximal segment of Finger
IV and single tubercles on ulna), and more webbing on the hand, reaching beyond the distal subarticular tubercle
on Finger IV (to distal subarticular tubercle, Fig. 4a). Specimens from near Rio Branco, Acre, Brazil, roughly 1100
km WSW of the type locality of O. camufatus, have posterior thigh surfaces brown, finely mottled with cream
(bold dark brown markings on light ground) and lack tubercles on the eyelids (present).
Frogs in the closely related genus Dryaderces are similar in general appearance to some frogs of the O.
buckleyi group, but can be distinguished as follows: They differ in exhibiting bold dark ventral reticulation (dark
spotting) and uniform dark iris (light iris) in D. pearsoni (Gaige), a longer snout of EN/HL 0.27–0.33 (shorter, EN/
HL 0.25), non-tuberculate canthus rostralis (tuberculate) and light, uniform venter (venter with small dark spots) in
Dryaderces sp. (Dryaderces pearsoni [Ca1_MTR13158_2768] of Jungfer et al. 2013) (KHJ, unpublished data).
The inspection of a fresh specimen of “Osteocephalus” inframaculatus (Boulenger), a species until recently only
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known from the holotype and tentatively placed with the O. buckleyi group by Jungfer (2010), was placed in
Dryaderces by Hoogmoed (2013). It is easily distinguishable from O. camufatus by its bold dark markings on
throat and chest and has a longer snout with EN/HL 0.33 (0.25) and larger tympanum with TD/HL 0.26 (0.22–
0.23).
Description of holotype. Snout short (EN/HL 0.245) and blunt, head as long as wide. Nostrils raised, opening
laterally. Snout truncate in dorsal and lateral outline. Canthus rostralis well-developed, rounded, describing an
inwardly curved line, bearing large tubercles. Skin of head coarsely shagreen, orbits with a few additional small
tubercles, some bearing small keratinized tips. A few more scattered keratinized tips on some tubercles of the head
and anterior part of the body. No frontoparietal crests evident externally. Loreal region strongly concave bearing
low tubercles. Lips flared below the eye. Choanae oblique, elliptical, though slightly angular. Dentigerous
processes of vomers short, considerably shorter than choanae width, angular, not in contact with each other, bearing
4 teeth on the left and 3 on the right. The anterior edges of the dentigerous processes are in line with the posterior
third of the choanae, their posterior edges well behind the posterior margins of the choanae. Tongue elliptical,
slightly wider (by 6%) than long. The vocal sacs are paired, subgular and protrude ventral to the jaw articulation. A
thick glandular supratympanic fold from the posterior part of the orbit medially, covering tympanic annulus
dorsally from “11” (when tympanum is thought to be a clock face) to “3”, then sloping in a curve towards the arm
insertion, reaching no further than the lower one third level of the tympanum. The latter is conspicuous, rounded
(barely wider than high), with about 66% the diameter of the eye. Skin on dorsum coarsely shagreen with a few
low, irregularly spaced tubercles. Transversal furrows on the shoulders are absent. Skin smooth between eye and
tympanum and areolate posterior to the tympanum above the arm insertion, very coarsely areolate laterally between
the limb insertions. Throat and belly areolate. Cloacal opening situated at about half the level of thighs, surrounded
by large, irregular warts (except on the cloacal flap, which is smooth). Skin on arms shagreen with a few faint
tubercles dorsally, hidden surfaces finely shagreen. A row of low ulnar tubercles also encompasses the proximal
segment of Finger IV. Axillary membrane absent. Dorsal surfaces of hind legs shagreen, posterior surfaces of
thighs and ventral parts of shank smooth, proximal three fourths of thigh coarsely areolate ventrally. A row of low
outer tarsal tubercles from the heel to the proximal subarticular tubercle of Toe V. On the heel, three and four low
tubercles, respectively, directed dorsally and laterally. Finger and toe discs elliptical and wider than long. Diameter
of disc on Finger III is 74% of tympanum diameter. Relative finger length is I < II < IV < III. On the thumb dark
brown nuptial excrescences on the posterolateral side from its base to the level of the distal end of the subarticular
tubercle. A large elliptical thenar tubercle. Two rounded palmar tubercles. Proximal segments of Fingers II–IV
tuberculate, two supernumerary tubercles each on Finger II and Finger IV. Subarticular tubercles conical except the
distal ones on Finger III and Finger IV, which are bifid. Webbing formula is I basal II 1 ½—2 ¾ III 2 ⅓—2 IV.
On the foot a large elliptical inner metatarsal tubercle and a small rounded plantar tubercle. A larger rounded
conical outer metatarsal tubercle in line with the row of tarsal tubercles. The latter form a shallow fringe on the
distal part of the proximal toe segment that is continued to the toe disc. A few indistinct supernumerary tubercles
on the proximal segments of Toes III–V. Subarticular tubercles single and conical. Relative lengths of appressed
toes I < II < III ≤ IV < V. Webbing formula: I 1—2
-
II 1—2 III 1—1
+
IV 1
+
—1 V.
Measurements and proportions. SVL 39.5; HL 14.3; HW 14.3; TL 22.2; FL 26.8; ED 4.7; TD 3.1; FD 2.3;
EN 3.5; IN 3.4; TE 2.0. HL/SVL 0.36; HW/SVL 0.36; TL/SVL 0.56; FL/SVL 0.68; TD/ED 0.66; TD/FD 1.35; EN/
HL 0.25; EN/SVL 0.09; HL/HW 1.00; TE/TD 0.65; TD/HL 0.22.
Colour in preservative. The ground colour of head and dorsum is light (green in life) with numerous, more or
less irregular blotches in tan and dark brown (various shades of brown in life). This pattern extends to the lateral
sides of the head and body. One ill-defined dark brown mark somewhat more conspicuous than others has two arms
between the eyes (including the orbits), two between the tympana and on midbody the posterior two arms diverging
like the lower part of an X and are continued on the flanks. Two additional ill-defined dark brown marks posteriorly
to the proximal part of the sacrum, continued to the flanks. Throat and belly with small irregular brown spots.
Although there are light areas on the upper lip, there are no discrete labial marks distinguishable from the rest of
the pattern. Tympanum tan. Arms dorsally and laterally with irregular dark brown crossbars, with irregular brown
spotting ventrally. Legs dorsally with irregular dark brown crossbars, posterior surfaces of thigh and calf marbled
dark brown. Ventral surfaces of thigh tan with lighter areolae, of femur tan peppered with darker markings.
Webbing tan, slightly marbled. Subcloacal warts are in creamy white and different shades of tan.
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NEW SPINY-BACKED TREEFROG OSTEOCEPHALUS
FIGURE 1. a. Dorsal, b. ventral view of the preserved holotype of O. camufatus sp. nov.
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FIGURE 2. Lateral view of the head, circumcloacal area and posterior surface of thigh of the preserved holotype of O.
camufatus sp. nov.
Colour in life. Colours were taken from photographs (Fig. 4). Dorsal surfaces covered by bold blotches of
different shades of green and brown. A light mark posteroventral to the eye is light green, the posterior part of the
upper lip dark brown, of the lower lip white. The tympanum is tan. The iris is light tan in its upper half and creamy
white in its lower one, the latter bearing a diffuse dark brown median vertical streak. Upper and lower halves are
separated by a weakly demarcated reddish brown horizontal streak and are both finely reticulated black. Some light
blue coloration can be seen on the hidden surface of the thigh. Outer tarsal and heel as well as ulnar tubercles are
tipped white.
Variation in the paratype. An adult male slightly larger than the holotype (40.4 mm SVL) with well-
developed nuptial pads. The snout is bluntly rounded in dorsal aspect. The dentigerous processes bear five teeth on
each side. Keratinized tips on tubercles are lacking entirely. Coloration is similar, but the large tan dorsal mark is
less conspicuously outlined. The belly is spotted evenly. There is slightly more webbing on the hand with I basal II
1
2
/
3
—2
2
/
3
III 2—2 IV and little variation on the foot: I 1
+
—2 II 1—2
-
III 1
—
1
1
/
3
IV 1
1
/
3
—1 V. There is also little
variation in proportions.
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FIGURE 3. a. Right hand and forearm and b. right foot of the preserved holotype of O. camufatus sp. nov. Not to scale.
Measurements and proportions. SVL 40.4; HL 15.0; HW 14.9; TL 21.2; FL 25.9; ED 5.0; TD 3.4; FD 2.1;
EN 3.7; IN 3.3; TE 1.9. HL/SVL 0.37; HW/SVL 0.37; TL/SVL 0.53; FL/SVL 0.64; TD/ED 0.68; TD/FD 1.62; EN/
HL 0.25; EN/SVL 0.09; HL/HW 1.01; TE/TD 0.56; TD/HL 0.23.
Distribution and habitat. The new species is so far only known from two sites about 54 km apart along the
terra firme forests of the right bank of the Rio Abacaxis, a black water river south of the Amazonas that discharges
into Paraná do Arariá, a white water channel delimiting the southern border of Ilha de Tupinambarana (Fig. 5). This
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is a huge island (about 300 km long) extending between Nova Olinda do Norte on the lower Rio Madeira to
Parintins on the Amazonas. Habitat at Igarapé-açu consisted of typical terra firme forest with higher trees reaching
up to 40 m and a diameter at breast height of around 80–100 cm. The understory was particularly rich in low palm
trees and the leaf litter was dense. The specimen was obtained close to a small stream perched on a tree about 1 m
high. At Areal, a site further upstream (also referred to as Paca or Pacamiri locally) the local habitat consisted of an
extensive white sand area (campinarana) with scattered vegetation near the margin of the river with abundant
clumps of ground bromeliads, gradually replaced by denser areas covered by palm trees and then terra firme forest.
The terra firme forest with large trees became more evident farther from the river margin, where the second
specimen was obtained.
Etymology. The species name camufatus is the Latinized past participle of Italian camuffare, to disguise or
mask, from which the word camouflaged is derived, in allusion to the shape-dissolving pattern of greens and
browns of the new species.
FIGURE 4. Adult male holotype of O. camufatus n. sp. from Igarapé-açu, Rio Abacaxis, Estado Amazonas, Brazil. Enclosure:
close-up of the eye.
Acknowledgements
We are grateful to to Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP), Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq) and Coordenação de Aperfeiçoamento de Pessoal de Ensino
Superior (CAPES) for funding. For help in field we thank Sergio Marques de Souza, José Cassimiro and José
Mário Guellere. JF thanks ANPCyT 2011–1895, 2013-404, Grants 2012/10000-5, and 2013/50741-7, São Paulo
Research Foundation (FAPESP), and CONICET PIP 11220110100889. For access to comparative material we are
grateful to Ana Almendáriz (EPN), Franco Andreone (MZUT), Rafe Brown, Linda Trueb and William E.
Duellman (KU), Barry T. Clarke and Colin McCarthy (BMNH), Jesús Córdova and César Aguilar (MUSM),
Raffael Ernst (MTD), Darrel R. Frost and David Kizirian (AMNH), Alexander Haas and Jakob Hallermann
(ZMH), W. Ronald Heyer (USNM), Gunther Köhler (SMF), John D. Lynch (ICN), Göran Nilson (GNM), Ronald
Nussbaum and Greg Schneider (UMMZ), Dennis Rödder and Wolfgang Böhme (ZFMK), Andreas Schlüter
(SMNS), Andreas Schmitz (MHNG), Harold K. Voris (FMNH), Hussam Zaher, Carolina Castro-Mello and Alberto
Barbosa de Carvalho (MZUSP). We also thank Albertina Lima for photos of additional material from Central
Amazonia. Two anonymous reviewers made helpful comments.
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NEW SPINY-BACKED TREEFROG OSTEOCEPHALUS
FIGURE 5. Map of the collecting sites of Osteocephalus camufatus sp. nov. (red dots) in Estado Amazonas, Brazil. The type
locality is Igarapé-açu.
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APPENDIX. Material examined of frogs of the Osteocephalus buckleyi species group.
Osteocephalus buckleyi: ECUADOR: Napo: Jatun Sacha: KU 217746, SMNS 13714-5. Orellana: Hacienda Primavera, 40 km
S Coca: MHNG 2556.58; Yasuní (S 00°41’00’’; W 76°24’00’’): SMNS 13713. Pastaza: Canelos: BMNH 1947.2.13.40–
1947.2.13.41, 1947.2.13.43, 1947.2.13.44 (lectotype), 1947.2.13.45. Sucumbíos: San Pablo de Kantesiya: MHNG
2366.39–2366.40; Santa Cecilia: KU 105208–9, 150492–3, 152477, 175500; Shushufindi: MHNG 2560.61.
PERU: Loreto: 28 km S Iquitos: MUSM KHJ-F 067.
Osteocephalus cabrerai: COLOMBIA: Amazonas: Caño Guacayá: USNM 152759 (holotype). Leticia, Río Pure: ICN 46691–
701. PERU: Loreto: Quebrada Negra, Río Iauasiyacu: MUSM KHJ-082–3. Ucayali: 65 km ENE Pucallpa, SE slope Cerro
Tahuayi: MUSM 649. Parque Nacional Sierra del Divisor (08º12'16.4''S;73º52'58.3''W): MUSM 24411, 24413–4.
Osteocephalus camufatus: BRAZIL: Amazonas: Igarapé-açu, right bank of Rio Abacaxis, 30 m (04°20'40''S; 58°38'06''W):
MZUSP 142389. Areal, left bank of Rio Abacaxis, 39 m (04°35'49''S; 58°13'14''W). MZUSP 157020.
Osteocephalus carri: COLOMBIA: Caquetá: Parque Nacional Cordillera Los Picachos, vereda Cristo Rey, 1500–1600 m: ICN
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NEW SPINY-BACKED TREEFROG OSTEOCEPHALUS
MC-9037–40, MC-9101–2, MC-9440–1, MC-944357, MC-9459, MC-9460–3, MC-9465–81, MC-9483–9, MC-9649–50,
MC-9661, MC-9794. Parque Nacional Cordillera Los Picachos,vereda La Esperanza, 1380–1440 m: ICN MC-9802–5,
MC-9845, MC-9871–2. Huila: Acevedo, Río Suaza, Río Aguas Claras, near San Adolfo, 1400 m: FMNH 69702
(holotype). Meta: Villavicencio, parte alta Caño Maizazo: ICN 26988.
Osteocephalus duellmani: ECUADOR: Morona-Santiago: Cordillera del Cóndor, Río Piuntza, 1910 m asl, about 3°25'S,
78°27'W: KU 147171 (paratype), 147172 (holotype).
Osteocephalus festae: ECUADOR: Morona-Santiago: Valle de Santiago (= lower Río Zamora): MZUT An. 208 (holotype).
Napo: Archidona: MHNG 2560.60. Loreto, Ávila, subcentro Caimítoyacu: EPN 5577–8. Sucumbíos: Cuyabeno,
Campamento Concienti: EPN AA-5611, AA-5607.
PERU: Cajamarca: San Ignacio: El Sauce: MUSM 19224. San Martín: 1 km NW Venceremos, 1600 m: KU 217302. 14
km W Venceremos, 2000 m: KU 217303.
Osteocephalus helenae: BOLIVIA: Santa Cruz: Buena Vista: MZUSP 95529. BRAZIL: Acre: Catuaba (S10°04'00'';
W63°37'00''): SMNS 14190-1, 17502, 17520. Amapá: Rio Maracá, boca Igarapé Camaipí: MZUSP 99990-99999; Seringal
Boa Fortuna: MZUSP 74253. Amazonas: Reserva Adolpho Ducke: MZUSP 59516, 74253, 75648, 84581; Reserva INPA-
WWF (Rio Preto): MZUSP 58055. Rondônia: Cachoeira de Nazaré, Rio Machado: MZUSP 63805. UHE Samuel: MZUSP
76921. GUYANA: Cuyuni-Mazaruni: Kartabo: AMNH H-13492. Demerara-Mahaica: Marudi Creek: AMNH A-46233.
Essequibo: Shudikar-wau: AMNH A-49252. Essequibo Islands-West Demerara: Dunoon: UMMZ 52681 (holotype).
Potaro-Siparuni: Iwokrama Forest: MTD 48360-2; Magdalen's Creek, NW bank of Konawaruk River, ca. 40 km WSW
Mabura Hill: AMNH A- 166220-4, 166229. VENEZUELA: Amazonas: SW sector of Cerro Yapacana: AMNH-A 100598.
Osteocephalus mimeticus: PERU: Ayacucho: San José on Río Santa Rosa, 1005 m: KU 196994–5. Sivia, Río Apurimac, 760 m:
FMNH 39853. Cuzco: Pozo Pagoreni, Comunidad Kirigueti: MUSM 21872. Zona Reservada Nahua Kugapakori: MUSM
23179–80, 23188, 23207, 23218, 23226. Huánuco: Fundo Flor, Río Pachitea, 300 m: SMNS 6515. San Martín: W slope
Abra Tangarana, 7 km NE San Juan de Pacaysapa, 1080 m: KU 212189–90. Cataratas Ahuashiyacu, 14 km NE Tarapoto,
730 m: KU 212191. Río Cainarache, 33 km NE Tarapoto on road to Yurimaguas: KU 209454–5. Río Cumbaza Valley, 9.4
km N Tarapoto, 390 m: KU 212182–5. Roque: GNM 469 (lectotype). Pongo de Shilcayo, ca. 4 km NNW Tarapoto, 470 m:
KU 212196–9. 14 km ESE Shapaja, 360 m: KU 212193. 22.7 km NE Tarapoto, 810 m: KU 212200. 28 km NE Tarapoto,
600 m: KU 212201–3. 29 km NE Tarapoto, 550 m: KU 212181. Tocache, Río Huallaga: MUSM 10845. Venceremos, 89
km NW Rioja, 1650 m: KU 212186–7. Pasco: Pozuzo, 770 m: MUSM 20351. Puerto Bermudez: MUSM 17801. Santa
Isabel: MUSM 17847–52, 18022, 18024. Yaupi, Río Paucartambo, 1600 m: KU 136312. Ucayali: Boquerón del Padre
Abad: ZFMK 33352, 39614–5, 39748–50, 39752–3, 40152–3.
Osteocephalus mutabor: ECUADOR: Napo: South slope of Volcán Sumaco on Río Pucuno, 1000 m, between Guamaní and
Guagua Sumaco (0° 42' 24'' S; 77° 35' 54'' W): EPN H-6658 (holotype), EPN H-6659 (paratype). Comuna Veinticuatro de
Mayo: EPN H-5695-9 (paratypes). Río Chaloyacu on Carretera Narupa – Coca: ZFMK 66237 (paratype). San Pablo de
Kantesiya: MHNG 2260.4-6, 2260.97, 2366.43, 2366.45, 2366.48, 2366.53, 2366.61-63, 2366.67, 2373.84, 2373.86-87,
2556.57 (paratypes). Pastaza: Canelos, 530 m: KU 120915. Locación Petrolera Garza 1, NE Montalvo, 300 m: KU
217747-9. Vicinity of Arutam Biological Field Station: SMF 79721-2 (paratypes). Sucumbíos: Cascales: EPN-H5716-9
(paratypes). Limoncocha, 200 m: KU 99210-6. Santa Cecilia, 340 m: KU 105210-20, 109509-11, 111971, 122964-87,
123169, 150494-5, 152277.
PERU: Loreto: San Jacinto, 175-190 m: KU 221928. 1.5 km N Teniente Lopez, 310-340 m: KU 221929-32.
Osteocephalus verruciger: COLOMBIA: Caquetá: Municipio de Florencia: Escuela Tarqui, carretera Altamira-Florencia km
48-49: ICN 23648. Municipio de Florencia: 35.2 km. arriba de Florencia: ICN 23943. Municipio de Florencia: Vereda
Tarqui 38.8-39.0 km: ICN 23944-5, ICN 23948, ICN 23952, ICN 23954. Municipio de Florencia: Vereda Tarqui 13.2 km
arriba de Florencia: ICN 23946. Municipio de Florencia: 39.3 km arriba de Florencia: ICN 23947. Huila: Acevedo, Río
Suaza, Río Aguas Claras near San Adolfo, 1400 m: FMNH 69709-10. Palestina: ICN 01542. Putumayo: 10.3 km W El
Pepino, 1440 m: KU 169586-7, 169589-93, 169595-6,169599, 169601-4, 169606-7. ECUADOR: No specific locality:
ZMH-A946 (syntype). Cotopaxi: Las Pampas (in error): MHNG 2259.20, 2560.62, 2560.64-68. Napo: El Reventador:
MHNG 2259.18, 2273.28, 2485.65-70, 2560.63. 2 km SSW Río Reventador, 1700 m: KU 164408, 164414, 164416-9,
164421, 164423-4, 164426. 3.2 km NNE Oritoyacu, 1910 m: KU 178839-44. Río Azuela, 1740 m: KU 143210-2, 143215-
7, 143219-24, 164434, SMNS 14197. Río Azuela, 9.5 km W of Reventador, 1630 m: KU 217750-1. Río Salado, 1 km
upstream from Río Coca, 1420 m: KU 164437, 164442, 178844, 178846-7. 0.7 km NE Río Salado bridge on Lago Agrio
road, 1380 m: KU 190054. San Rafael: MHNG 2259.19, 2272.98-99. S slope Cordillera del Due, 1150 m: KU 123181,
123186. 11.1 km NE Santa Rosa, 1900 m: KU 19004953. 16.5 km NNE Santa Rosa, 1700 m: KU 143209. Pastaza: 9.5 km
NW Mera, 1270 m: KU 178848. Tungurahua: 11 km E Río Negro, 1170 m: KU 146469-70.
