ZooKeys 69 | : | 49-| 62 (20 | 7) A peer-reviewed open-access journal
doi: 10.3897/zookeys.69 1.13595 RESEARCH ARTICLE #ZooKey S

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Formal recognition of the species of Oreosaurus
(Reptilia, Squamata, Gymnophthalmidae) from the
Sierra Nevada de Santa Marta, Colombia

Santiago J. Sanchez-Pacheco'’, Pedro M. Sales Nunes?*, Sergio Marques-Souza’,
Miguel T. Rodrigues?, Robert W. Murphy'*

| Department of Ecology and Evolutionary Biology, University of Toronto, 25 Willcocks Street, Toronto, ON
M5S 3B2, Canada 2 Department of Natural History, Royal Ontario Museum, 100 Queens Park, Toronto,
ON MDS 2C6, Canada 3 Departamento de Zoologia, Instituto de Biociéncias, Universidade de Sto Paulo, Sao
Paulo, CEP 05508-090, SP Brazil 4 Departamento de Zoologia, Centro de Biociéncias, Universidade Federal
de Pernambuco, Av. Professor Moraes Rego S/n, Cidade Universitdria 50670-901, Recife, PE, Brazil

Corresponding author: Santiago J. Sdnchez-Pacheco (santiago.sanchezpacheco@mail.utoronto.ca)

Academic editor: A. Herrel | Received 9 May 2017 | Accepted 11 July 2017 | Published 17 August 2017
http://z0obank.ore/1897E08F-76 15-4470-8586-55B3EAAF74FA

Citation: Sanchez-Pacheco SJ, Nunes PMS, Marques-Souza S, Rodrigues MT, Murphy RW (2017) Formal
recognition of the species of Oreosaurus (Reptilia, Squamata, Gymnophthalmidae) from the Sierra Nevada de Santa
Marta, Colombia ZooKeys 691: 149-162. https://doi.org/10.3897/zookeys.69 1.13595

Abstract

Oreosaurus is one of the two genera extracted from the former Riama sensu lato, which was recently rec-
ognized as polyphyletic. Oreosaurus is a small clade (five named and two undescribed species) of montane
gymnophthalmid lizards and exhibits an exceptional distributional pattern. Its nominal and undescribed
species are discontinuously distributed on the Cordillera de la Costa of Venezuela, the tepuis from the
Chimanta massif in Venezuela, the highlands of the island of Trinidad, and the Sierra Nevada de Santa
Marta in Colombia (SNSM). Herein, we describe the species of Oreosaurus that is endemic to the SNSM.
Historically, this species associates with two names that are currently nomina nuda: Proctoporus serranus
and P specularis. Formal nomenclatural recognition of Oreosaurus serranus sp. n. renders specularis a per-
manently unavailable name for this taxon. Oreosaurus serranus sp. n. is the sister of all remaining conge-
ners, and differs primarily from them in having only one pair of genial scales, as well as a unique pattern

of scutellation. We provide an identification key to the species of Oreosaurus.

Copyright S.J. Sanchez-Pacheco et al. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

150 Santiago J. Sanchez-Pacheco et al. / ZooKeys 691: 149-162 (2017)

Resumen

Oreosaurus es uno de los dos géneros que fueron extraidos de Riama sensu lato, el cual fue reconocido re-
cientemente como polifilético. Oreosaurus es un clado pequefio (cinco especies nominales y dos indescritas)
de gimnoftalmidos de montafia y presenta un patron de distribucién excepcional. Sus especies nominales
e indescritas se distribuyen discontinuamente sobre la Cordillera de la Costa de Venezuela, los tepuyes del
macizo de Chimanta en Venezuela, las tierras altas de la isla de Trinidad, y la Sierra Nevada de Santa Marta
en Colombia (SNSM). Describimos la especie de Oreosaurus que es endémica de la SNSM. Histéricamente,
esta especie se asocia con dos nombres que son nomina nuda: Proctoporus serranus y P specularis. El recono-
cimiento formal de Oreosaurus serranus sp. n. hace que specularis sea un nombre permanentemente indis-
ponible para este taxén. Oreosaurus serranus sp. n. es la especie hermana de todos los congéneres restantes, y
se diferencia de ellos principalmente por tener un tinico par de escamas geneiales, asi como por presentar un

patron unico de escamacién. Proveemos una clave de identificacién para las especies de Oreosaurus.

Keywords
Microteiid lizard, Oreosaurus serranus, nomenclatural recognition, Proctoporus serranus, Proctoporus specularis,

nomina nuda, South America, taxonomy

Introduction

Oreosaurus Peters, 1862 (Reptilia: Gymnophthalmidae) contains five named species of
montane lizards that have discontinuous distributions on the Cordillera de la Costa
and tepuis from the Chimanta massif in Venezuela, and the Aripo northern range in
the Caribbean island of Trinidad (Sanchez-Pacheco et al. 2017). An additional spe-
cies that is the sister of all remaining congeners and is endemic to the Sierra Nevada
de Santa Marta in Colombia (SNSM) remains undescribed. SAnchez-Pacheco et al.
(2017) referred to it as “Sierra Nevada”.

Over 30 years ago, Ayala and Castro reviewed the Colombian lizard fauna in their
unpublished but widely distributed book “Lizards of Colombia”. Their work included
brief descriptions of several species and they referred to informal specific epithets as-
sociated with authors to indicate that formal descriptions were not yet published, but
were forthcoming. Among these species, Ayala and Castro included “Proctoporus” “ser-
ranus’, a gymnophthalmid lizard from the Serrania de San Lorenzo, SNSM, and they
provided a reference for the description (Harris, dated to 1984). However, Harris’
formal description of this taxon was never published. Although Ayala and Castro in-
cluded a brief description (based on an undetermined number of specimens), the name
“serranus isa nomen nudum because it does not have a reference, and therefore fails to
conform to ICZN (1999) Art. 11. Similarly, Ayala (1986) published a list of Colom-
bian lizards, which included undescribed species referred to names within quotes (“”)
and associated with authors to indicate imminent formal descriptions. Most of these
names were the same ones provided by Ayala and Castro (unpublished data), the ex-
ception being “Proctoporus” “specularis”, also from San Lorenzo, SNSM. Nevertheless,
both the locality and the given reference (Harris, but this time dated to 1986—also
never published) were strongly suggestive that “serranus” and “specularis” referred to

Formal recognition of the species of Oreosaurus (Reptilia, Squamata, Gymnophthalmidae)... 151

the same species. However, in accordance with ICZN (1999) Art. 13, the absence of a
description for “specularis’ (Ayala 1986) renders this name a nomen nudum.

While carrying out field work in the SNSM, we had the opportunity to collect a se-
ries of specimens that conform to the unpublished description of “serranus’. Two termi-
nals labeled “Sierra Nevada” 1 and 2 were included in a recently published phylogenetic
analysis of Riama Gray, 1858 sensu lato (Sanchez-Pacheco et al. 2017), which recovered
this species as part of the resurrected Oreosaurus. Although “serranus” and “specularis”
are currently nomina nuda, and by definition unavailable names (i.e., they fail to con-
form to ICZN Arts. 11 and 13), both of them have reached the modern literature
(Rueda-Almonacid et al. 2012 and de Albuquerque et al. 2012, respectively). A nomen
nudum can be made available (or validated) if it is published again in a way that meets
the criteria of availability (ICZN 1999). Anadia altaserrania Harris & Ayala, 1987, an-
other endemic gymnophthalmid lizard from the SNSM, is a pertinent example. It was
included in Ayala and Castro’s unpublished book (with reference to Harris, Ayala and
Castro, 1984) and listed by Ayala (1986; this time with reference to Harris and Ayala,
1986), but finally published formally by Harris and Ayala (1987). The situation with
Oreosaurus is not unlike that of Anolis in which Poe et al. (2009) provided examples of
nomen nudum species of Anolis lizards listed by Ayala (1986). Below we provide a name
and a description for the species of Oreosaurus from the SNSM.

Materials and methods

For comparative purposes, specimens of Oreosaurus achlyens (Uzzell, 1958), O. luctuo-
sus Peters, 1862, O. shrevei (Parker, 1935) and the undescribed O. “Venezuela” were ex-
amined (Appendix 1). Data for O. mcdiarmidi (Kok & Rivas, 2011) and O. rhodogaster
(Rivas et al., 2005) were taken from the literature (Kok and Rivas 2011 and Rivas et
al. 2005, respectively). Measurements (snout-vent length [SVL] and tail length) were
taken to 0.1 mm with a digital caliper. Sex was determined by noting the presence
of hemipenes in males and/or secondary sex characters, such as the number of femo-
ral pores. To facilitate comparisons with other species of Oreosaurus, scutellation and
head-scale terminology follows Kizirian (1996). Bilateral variation is reported as left/
right. Hemipenes were prepared following the procedures described by Manzani and
Abe (1988) as modified by Pesantes (1994) and Zaher (1999). The retractor muscle
was severed manually and an everted organ was filled with stained petroleum jelly. Fol-
lowing Uzzell (1973) and Nunes et al. (2012), calcareous hemipenial structures were
stained in an alcoholic solution of alizarin red. Terminology follows Dowling and Sav-
age (1960), Savage (1997) and Nunes et al. (2012).

The following collection abbreviations are used herein: AMNH (American Mu-
seum of Natural History, New York), EBRG (Museo de la Estacion Bioldgica de Ran-
cho Grande, Maracay, Venezuela), MCZ (Museum of Comparative Zoology, Harvard
University, Cambridge, USA), ROM (Royal Ontario Museum, Toronto, Canada), and
USNM (National Museum of Natural History, Washington D.C., USA).

152 Santiago J. Saénchez-Pacheco et al. / ZooKeys 691: 149-162 (2017)

Species description

Oreosaurus serranus sp. 0.
http://zoobank.org/5BBOFBOB-47E8-4788-BD79-2784FF9 1 F63F
Figures 1-3

Holotype. ROM 53608 (field number JJS 548; Fig. 1), an adult female collected
by S.J.S-P., P.M.S.N., S.M.S, Liliana Saboyd-Acosta, Jhon Jairo Ospina-Sarria, Sandy
B. Arroyo, and Mariane Targino Rocha in Colombia, Sierra Nevada de Santa Marta,
Departamento de Magdalena, headwaters of the Rio Guachacos, Corregimiento de
Minca, finca Vista Hermosa, approximately 2156 m, June 2013. This locality is situ-
ated at approximately 11°05'N, 74°01'W.

Paratypes. ROM 53609 (adult female, Fig. 2), ROM 53610 (subadult male),
ROM 53611 (subadult female), ROM 53612-13 (juvenile females), and ROM 53614
(juvenile male), all with same data as holotype.

Diagnosis. Oreosaurus serranus sp. n. can be distinguished from all its congeners by
the number of genial pairs (1 in O. serranus sp. n. versus 2 in the other species). It also
differs from all other species of Oreosaurus, except O. mcdiarmidi, by the number of
supraoculars (3 in O. serranus sp. n. and O. mcdiarmidi versus 4 in the other species),
and dorsal scale relief (smooth in O. serranus sp. n. and O. mcdiarmidi versus keeled
or slightly keeled in the other species). Oreosaurus serranus sp. n. also differs from O.
mcdiarmidi by the absence of prefrontal scales (present in O. mcdiarmidi).

Description. Oreosaurus serranus sp. n. possesses the following characteristics:
(1) maximum known SVL in males 60 mm (7 = 2), in females 70.4 mm (z = 5);
(2) frontonasal equal to or longer than frontal; (3) prefrontal scales absent; (4) naso-
loreal suture complete [= loreal present]; (5) supraoculars three, all in contact with
ciliaries; (6) superciliary series incomplete, formed only by the anteriormost supercili-
ary scale; (7) supralabial-subocular fusion absent; (8) postoculars two; (9) postparietals
two; (10) supratympanic temporals two; (11) genials in one pair; (12) dorsal scales
rectangular, juxtaposed, smooth; (13) nuchal scales smooth; (14) longitudinal dorsal
scale rows 10—11; (15) transverse dorsal scale rows 33—36; (16) ventral scales smooth,
in 21—22 transverse scale rows; (17) lateral scale rows (oval, non-granular scales) 4—6;
(18) femoral pores per hind limb in males 7-9, in females 2—3 (located proximal-
ly); (19) scales between medialmost femoral pores two; (20) subdigital scales on toe
I four; (21) anterior cloacal plate scales four or six; (22) posterior cloacal plate scales
seven; (23) dorsum dark brown to black with fine brown mottling; distinct dorsolat-
eral stripes absent; lateral ocelli (i.e., white spots surrounded by dark blotches) absent
(white or cream spots instead); venter black with conspicuous whitish spots mostly on
scale sutures; (24) hemipenial body globose, slightly bilobed, ornamented by 14—15
chevron-shaped flounces on each side.

Description of holotype. Adult female (Fig. 1), SVL = 70.4 mm, tail length =
72.4 mm; head scales smooth, glossy; rostral scale wider than long, higher than adjacent
supralabials, in contact with frontonasal, nasals, and anteriormost supralabials posteri-

Formal recognition of the species of Oreosaurus (Reptilia, Squamata, Gymnophthalmidae).... 153

Figure |. Oveosaurus serranus sp. n. (holotype, ROM 53608 [70.4 mm SVL]). Dorsal, lateral and ventral

views of the head, and ventral view of the pelvic region.

orly; frontonasal roughly quadrangular, longer than wide, widest posteriorly, equal in
length to frontal, in contact with nasals and loreals laterally, and frontal posteriorly; pre-
frontals absent; frontal longer than wide, anterior suture convex, lateral sutures concave,
posterior suture angular with point directed posteriorly, in contact with anteriormost

154 Santiago J. Sdénchez-Pacheco et al. / ZooKeys 691: 149-162 (2017)

=
«*

Figure 2. Oreosaurus serranus sp. n. (paratype, ROM 53609 [68.6 mm SVL]) in life. Photos: $.M.S (top)
and Jhon Jairo Ospina-Sarria (bottom).

Formal recognition of the species of Oreosaurus (Reptilia, Squamata, Gymnophthalmidae).... 155

supraoculars and superciliaries posterolaterally, and frontoparietals posteriorly; fron-
toparietals pentagonal, in contact anterolaterally with all supraoculars on the left side
and second and third supraoculars on the right side, and posteriorly with parietals and
interparietal; interparietal hexagonal, longer than wide, lateral sutures concave, in con-
tact with parietals laterally, postparietals posteriorly; parietals in contact with third su-
praoculars anterolaterally, dorsalmost temporal and postocular scales laterally, and post-
parietals posteriorly; postparietals pentagonal, two, in broad contact; supraoculars three,
all in contact with ciliaries. Nasoloreal suture complete, nasal quadrangular; loreal quad-
rangular, not in contact with second supralabial; superciliary series incomplete, formed
only by the anteriormost superciliary scale, which barely extends onto dorsal surface of
head, and lies between loreal, frontal, first supraocular, and anteriormost ciliaries; palpe-
bral disc of lower eyelid divided into three large, unpigmented scales; frenocular quad-
rangular, in contact with loreal and nasal anteriorly; circumorbital scales between pos-
teriormost supraocular and frenocular five; postoculars two; temporals smooth, glossy,
polygonal; supratympanic temporals two; supralabials seven; infralabials four. Mental
wider than long, in contact with anteriormost infralabials and postmental posteriorly;
postmental roughly pentagonal, posterior suture angular with point directed posteriorly,
in contact with first and second infralabials laterally; genials in one pair, roughly quad-
rangular, in contact with second and third infralabials; scale rows between genials and
collar fold (along midventral line) eight, medialmost scales of posteriormost scale row
distinctly enlarged, smooth; posteriormost gular row enfolded posteriorly, concealing
one small scale row; lateral neck rounded, smooth.

Dorsal scales rectangular, longer than wide, juxtaposed, smooth, in 35 transverse
rows; longitudinal dorsal scale rows at fifth transverse ventral scale row nine, at 10"
transverse ventral scale row 10, at 15" transverse ventral scale row 11; lateral scale rows
at fifth transverse ventral scale row 6/5, at 10° transverse ventral scale row four, at 15"
transverse ventral scale row four; lateral scales on body near insertion of forelimb small
to granular; ventral scales quadrangular, smooth; complete transverse ventral scale rows
22; longitudinal ventral scale rows at midbody 10; anterior cloacal plate scales six; pos-
terior cloacal plate scales seven, medialmost scale with a horizontal suture; scales on tail
rectangular and juxtaposed; midventral subcaudals smooth, wider than adjacent scales,
nearly square. Femoral pores per hind limb two, located proximally; scales between
medialmost femoral pores two.

Coloration of holotype. In life, dorsal ground color dark brown to black with
fine brown mottling; dorsal surfaces of head, body and tail with an iridescent bluish
shine. White or cream spots laterally from neck to posterior portion of body, becoming
less distinct posteriorly. Ventral surfaces of head and body predominantly black, with
conspicuous whitish spots mostly on scale sutures; subcaudally black without spots.
In preservative (70% ethanol), dorsal ground color brown with fine light brown mot-
tling; dorsal surfaces of head, body and tail without the iridescent bluish shine. Ventral
surfaces of head and body brown with cream spots on scale sutures.

Hemipenial morphology. Right organ of subadult male ROM 53610 (Fig. 3) was
partially everted and filled. Basal and lobular regions are partially damaged. Hemipe-

156 Santiago J. Sdénchez-Pacheco et al. / ZooKeys 691: 149-162 (2017)

Figure 3. Oreosaurus serranus sp. n. Sulcate (left), lateral (center) and asulcate (right) views of the right
hemipenis of ROM 53610 (paratype).

nial body is roughly globose, ending in two small and partially everted, barely visible
lobes. Partial eversion and some damages precluded the detection of folds, or any other
ornamentation, on the lobes.

The sulcus spermaticus, central in position, originates at the base of the organ and
proceeds in a straight line towards the lobes. It is bordered by two parallel nude areas,
and divided by a fleshy fold. Branches of the sulcus spermaticus are not visible. Two
columns of at least 14 chevron-shaped flounces ornament the sides of the organ and
the borders of the sulcate and asulcate faces of the hemipenial body. Although these
flounces do not present calcified comb-like spicules, it is possible that such absence is
due to the age of the specimen. ‘These calcified structures are present in adults of most
species of Cercosaurinae that have their hemipenial morphology described, includ-
ing species of Oreosaurus (e.g., Kok and Rivas 2011, Nunes 2011, Rivas et al. 2005).
A broad nude area occupies at least 50% of the asulcate face. Some damages at the
basis of the organ precluded the detection of the isolated horizontal flounces on the
proximal-central region of the asulcate face that are often present in species of Cerco-
saurinae (e.g., Kok and Rivas 2011, Nunes 2011, Rivas et al. 2012, Sanchez-Pacheco
et al. 2011).

Variation. Paratypes consist of four females (SVL = 41.4-68.6 mm) and two males
(SVL = 40.4-60 mm). The paratypes are similar to the holotype with the follow-
ing noteworthy exceptions. Frontonasal longer than frontal in ROM 53609-12 and
53614; loreal scale in contact with second supralabial in ROM 53612-13; ventralmost
postocular fused with posteriormost subocular on the right side in ROM 53613; me-
dialmost scale of the posterior cloacal plate not divided horizontally in ROM 53610-11
and 53614; palpebral disc of the lower eyelid divided into two large, pigmented scales
in ROM 53609; femoral pores per hind limb in female ROM 53612 three. Femoral

Formal recognition of the species of Oreosaurus (Reptilia, Squamata, Gymnophthalmidae).... 157

se ail Caribbean Sea

4

\ 5 ne ¥
Fe oo

Figure 4. Distribution of Oreosaurus serranus sp. n. in the northwestern slopes of the Sierra Nevada de

Santa Marta, northern Colombia. Pentagon indicates type locality.

pore number is the most evident sexually dimorphic character, with males having 7—9
pores per hind limb (ROM 53610 8/9, ROM 53614 9/7) and females having 2-3.

Distribution and natural history. Oveosaurus serranus sp. n. is known exclusively
from the type locality (Figs 4, 5) and San Lorenzo (Ayala and Castro unpublished data,
Ayala 1986), two adjacent cloud forest localities on the northwestern slopes of the
Sierra Nevada de Santa Marta (SNSM) at elevations of about 1800-2156 m (Fig. 4).
This forest-dwelling lizard is often found under fallen, rotten trunks or logs. Holotype
and paratypes were collected manually during the day. The new species was found at
the type locality in sympatry with Anadia pulchella, another gymnophthalmid endemic
to the SNSM.

Etymology. ‘The specific epithet serranus, which is an adjective derived from the
Spanish adjective serrano (meaning from the sierra), refers to the location of the spe-
cies’ type locality in the Sierra Nevada de Santa Marta, and preserves the original ety-
mological intent of Harris, as stated by Ayala and Castro (unpublished data).

Comments. Formal nomenclatural recognition of Oreosaurus serranus sp. n. ren-
ders specularis (Ayala 1986) a permanently unavailable name for this taxon. Specimens
reported by Ayala and Castro (unpublished data) were not included herein because
they are presumably lost (S.J.S-P. personal observation).

158 Santiago J. Saénchez-Pacheco et al. / ZooKeys 691: 149-162 (2017)

Figure 5. Type locality (top) and habitat (bottom) of Oreosaurus serranus sp. n. in the Sierra Nevada de
Santa Marta, Colombia. Photos: Jhon Jairo Ospina-Sarria (top) and S.M.S (bottom).

Formal recognition of the species of Oreosaurus (Reptilia, Squamata, Gymnophthalmidae).... 159

O. serranus sp.n. SNSM
O. mcdiarmidi TCM
O. achlyens CCC

O. shrevei IT

O. “Venezuela” CCO

Figure 6. Summary of the phylogeny and geographic distribution of Oreosaurus (Sanchez-Pacheco et al.
2017). SNSM = Sierra Nevada de Santa Marta, Colombia; TCM = tepuis from the Chimanta massif,
Venezuela; CCC = Cordillera de la Costa Central, Venezuela; IT = island of Trinidad; CCO = Cordillera
de la Costa Oriental, Venezuela. Oreosaurus luctuosus, from the CCC, and O. rhodogaster, from the CCO,
were included in this genus due to the presumed close relationships of these species and O. achlyens and
O. shrevei, respectively. Data taken from Sanchez-Pacheco et al. (2017).

Oreosaurus is one of the two genera extracted from the former Riama sensu lato,
which was recently found to be non-monophyletic (Sanchez-Pacheco et al. 2017). The
other clade, Andinosaura Sanchez-Pacheco et al., 2017, includes 11 Andean species and
Riama sensu stricto is also an exclusively Andean radiation of 16 named species.

Sanchez-Pacheco et al. (2017) discussed the disjunct geographic distributions of
species of Orveosaurus, as well as their phylogenetic relationships. Figure 6 summarizes
these findings. All species of Oreosaurus share the absence of a narrow band of differen-
tiated granular lateral scales (present in species of Andinosaura and Riama).

Key to the species of Oreosaurus

i One pairot Genial SCAless, fevatdaheges anu hisieneenan’ Oreosaurus serranus sp. n.
- AO SpAITS Gl CTIA) SCAG rok Sad cries Sddudrohendedenhyoheechoomvanetnlssnabiera nab ielesseh voted 2
2 Pretrontal:scalés:p resent 3.5. ites nee been O. mcdiarmidi
- Pretrorbal scales ase mtie ts... Scales taba lesiasiy. uatethsusstihs lia haitasnnae laden seat Meio 3
3 Morell Scales Se Tbe es ty pense ale Se lice banutsc idl iret idee MU eet an nc hatte btn rad -.
— Moreal Seales present 2: cers eens ee geney a orertavaceesveaewan temew alanis eee e alee meecerenmucedl 5
- Anterior cloacal plate row composed of a small scale... eee O. shrevei
- Anterior cloacal plate row composed of two large scales....... O. “Venezuela”
5 Borsal body scales Nexa coil’. ...cwccataeedelsaatesnventennsoavccowabi ate uounstongaheateBpauenlnes 6
- Dorsaltbodyisealesirecta new lar’, sc.) ectowosenntnessusdivonantnesd ocuntopoeiatah O. luctuosus
6 42-44 transverse dorsal scale rOWS........sceceeseesseeeeeseeseesseeseeaes O. rhodogaster

— 37—40 transverse dorsal scale rowS..........ccssssccccccesseccccccssecccccesscees O. achlyens

160 Santiago J. Saénchez-Pacheco et al. / ZooKeys 691: 149-162 (2017)

Acknowledgments

Funding for S.J.S-P. was provided by a COLCIENCIAS doctoral fellowship (Becas
Francisco José de Caldas), an Ontario Graduate Scholarship at the University of To-
ronto, and an AMNH Collection Study Grant. NSERC Discovery Grant 3148 sup-
ported the research. P.M.S.N., S.M.S and M.T.R. are grateful to Fundacao de Amparo
a Pesquisa do Estado de Sao Paulo (FAPESP) and Conselho Nacional de Desenvolvi-
mento Cientifico e Tecnolégico (CNPq) and P.M.S.N. is grateful to Fundacao de
Amparo a Ciéncia e Tecnologia do Estado de Pernambuco (FACEPE) for financial
support. S.J.S-P. thanks D. Frost and D.A. Kizirian for work space and other facilities
provided during his stay at the AMNH. For specimen loans and access to collections
we are grateful to D.A. Kizirian (AMNH), J. Rosado (MCZ), and K. de Queiroz and
T.D. Hartsell (USNM). We thank P. Pulido-Santacruz, S.B. Arroyo, J.J. Ospina-Sarria
and M. Targino Rocha for arranging logistics and collaborating on fieldwork at Sierra
Nevada de Santa Marta. The manuscript benefited from comments and suggestions
from O. Torres-Carvajal and T.M. Doan. J. Hsiung provided the excellent drawings
of the holotype, and P. Pulido-Santacruz helped to make the figures. J.J. Ospina-Sarria
provided photographs.

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Appendix |

Comparative material examined

Oreosaurus achlyens: VENEZUELA: Aragua: Rancho Grande (AMNH 137260,
137267-69, 137271-76, 137278-82, 137297). O. luctuosus. VENEZUELA:
Aragua: Rancho Grande (AMNH 137270, 137277, MCZ 100410, USNM
196336), Parque Nacional Henry Pittier, Rancho Grande (USNM 259170).
O. shrevet: TRINIDAD & TOBAGO: Horne Tucuche (MCZ 62506-07); El
Teluche [in error, probably Tucuche] (MCZ 100466-68); Mt. Tucuche (MCZ
160065-66). O. “Venezuela”: VENEZUELA: Anzoategui: Cerro El Guamal,
Macizo del Turimiquire, municipio Freites, 2150 m (EBRG 5962).