Zootaxa 3753 (1): 047–058
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ISSN 1175-5326 (print edition)
Article
Copyright © 2014 Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3753.1.4
http://zoobank.org/urn:lsid:zoobank.org:pub:80782116-E419-4D5F-9286-AFA07C369291
A new species of spiny-tailed iguanid lizard (Iguania: Stenocercus)
from northwestern Peru
PABLO J. VENEGAS¹,², LOURDES Y. ECHEVARRIA¹ & SILVANA C. ALVAREZ¹
¹División de Herpetología-Centro de Ornitología y Biodiversidad (CORBIDI), Santa Rita N˚105 Of. 202, Urb. Huertos de San Antonio, Surco, Lima-Perú.
²Corresponding author. E-mail: sancarranca@yahoo.es
Abstract
We describe a new species of Stenocercus from the interandean valley of Río Chotano on the Amazonian slope of the
northern portion of the Cordillera Occidental of Peru (Cajamarca Region), at elevations of between 1997 and 2318 m.
Stenocercus arndti sp. nov. differs from other Stenocercus, except from S. bolivarensis, S. carrioni, S. chlorostictus, S.
crassicaudatus, S. empetrus, S. eunetopsis, S. simonsii, and S. torquatus, in having granular scales on the posterior surface
of the thighs, two caudal whorls per autotomic segment, mucronate caudal scales, and distinct longitudinal row of enlarged
vertebral scales. However, Stenocercus arndti sp. nov. is easily distinguished from these species in having a bold black
transversal band at midbody that extends ventrolaterally in adult males.
Key words: Cajamarca, Iguania, new species, Peru, Stenocercus, taxonomy
Resumen
Describimos una nueva especie de Stenocercus del valle interandino del Río Chotano en la pendiente amazónica de la porción norte de la Cordillera Occidental de Perú, Región de Cajamarca, entre los 1997 y 2318 m de altitud. Stenocercus arndti sp. nov. se diferencia de todas las demás especies de Stenocercus, a excepción de S. bolivarensis, S. carrioni, S.
chlorostictus, S. crassicaudatus, S. empetrus, S. eunetopsis, S. simonsii, and S. torquatus, por la combinación de los siguientes caracteres: escamas granulares sobre la superficie posterior de los muslos, vertebrales alargadas, dos anillos caudales por segmento autonómico, escamas caudales mucronadas y una fila longitudinal de escamas vertebrales distinta al
resto de escamas dorsales. Sin embargo, los machos adultos de Stenocercus arndti sp. nov., son fácilmente diferenciados
de las restantes especies, por tener una conspicua banda transversal negra a la mitad del dorso que se extiende hasta la
región ventrolateral.
Palabras clave: Cajamarca, Iguania, nueva especie, Perú, Stenocercus, taxonomía
Introduction
The iguanian lizard Stenocercus is currently composed of 63 species that occur at elevations of between 0–4000 m
in the Andes and adjacent lowland areas from northern Venezuela and Colombia to central Argentina, with some
species present in the Atlantic lowlands between southern Brazil and central Argentina, and other species present in
northeastern Brazil (Torres-Carvajal 2007a; Venegas et al. 2013). Members of this genus occupy a variety of
habitats, such as dry and humid lowland tropical forests, montane forests, cerrado, puna, and paramo (TorresCarvajal et al. 2006). In fact it is one of the most geographically— and ecologically— widespread reptile taxa
currently ranked as a genus in South America (Torres-Carvajal 2007b).
The major diversity of Stenocercus occurs in Peru, with currently 36 species, followed by Ecuador with 17,
Brazil 10, Colombia 7, Argentina 6, Bolivia 4, Uruguay 1, and Paraguay 1 (Torres-Carvajal 2007b, Torres-Carvajal
& Carvajal-Campos 2009, Torres-Carvajal & Mafla-Andora 2013, Venegas et al. 2010, Venegas et al. 2013).
Accepted by S. Carranza: 11 Nov. 2013; published: 3 Jan. 2014
47
Although the type species S. roseiventris was described more than 150 years ago (Dumeril & Bibron 1837),
approximately one-fourth of the species of Stenocercus have been described after 1990 (Avila-Pires 1995; Cadle
1991, 1998, 2001; Torres-Carvajal 2000, 2005 a, b, 2006, 2007c; Nogueira & Rodrigues 2006). Basically, one of
the main reasons for this dramatic rate of species discovery is that collections have recently been made in
previously unsampled areas throughout the Andes (e.g. Cadle 1991, 1998; Torres-Carvajal 2000). However, the
diversity of Stenocercus in Peru probably still remains underestimated due to the complex geography and diverse
terrain present, inadequate sampling in herpetological terms, and with some regions completely unexplored (Lehr
2002, Campbell & Lamar 2004).
Herein we describe a new species of Stenocercus that was discovered as a result of field trips to the interandean
valley of Río Chotano in Cajamarca on the Amazonian slopes of the Andes in northwestern Peru. This discovery
increases to 37 the number of Stenocercus species known from Peru.
Material and methods
All type specimens of the new species are deposited in the herpetological collections of the Centro de Ornitología y
Biodiversidad (CORBIDI) in Lima, Peru. Measurements of snout–vent length (SVL) and tail length (TL) were
taken with a ruler and recorded to the nearest 1 mm. All other measurements were made with digital calipers and
recorded to the nearest 0.1 mm. Sex was determined by dissection or by noting the presence of hemipenes. Data on
scutellation, such as the ranges, means and standard deviations, of all species of Stenocercus compared were taken
from Torres-Carvajal (2007b). Osteological characters were examined by dissection of two paratypes, one male
(CORBIDI 01682) and one female (CORBIDI 01685). We follow the terminology of Cadle (1991) and TorresCarvajal (2000, 2004, 2007b) for characters included in the description. For additional specimens examined, see
the Appendix.
Stenocercus arndti sp. nov.
(Figs. 1–4)
Proposed standard English name: La Granja whorltail lizard
Proposed standard Spanish name: Capón de La Granja
Holotype. CORBIDI 01680 (Figs. 1–3), an adult male from Quebrada Checos (6°21´12.63´´ S, 79°06´41.15´´ W),
at 1997 m elevation, La Granja District, Chota Province, Cajamarca Region, Peru, collected by P.J. Venegas on 22
August 2008.
Paratypes. CORBIDI 01681 and 01685, an adult male and adult female, respectively, collected with the
holotype; CORBIDI 01682 and 01688, adult males; CORBIDI 01692, adult female; CORBIDI 01686, 01687, and
01690, subadult males; CORBIDI 01683 and 01689, subadult females; CORBIDI 01691, a juvenile male;
CORBIDI 01684, a hatchling, all from Quebrada La Iraca, (6°22´09.9´´ S, 79°08´04.61´´ W), at 2213 m elevation,
La Granja District, Chota Province, Cajamarca Region, Peru, collected by P.J. Venegas on 9 August 2008.
Referred specimens (photo vouchers). Two adult females from Kañaris (6°03´26.18´´ S, 79°16´00.35´´ W),
at 2318 m elevation, Ferreñafe Province, Lambayeque Region, Peru, captured and released by P. J. Venegas on 25
May 2007.
Diagnosis. Stenocercus arndti differs from all other species of Stenocercus, except for S. bolivarensis Castro &
Ayala 1982, S. carrion Parker 1934, S. chlorostictus Cadle 1991, S. crassicaudatus Tschudi 1845, S. empetrus Fritts
1972, S. eunetopsis Calde 1991, S. simonsii Boulenger 1885, and S. torquatus Boulenger 1885, in having granular
scales on the posterior surface of the thighs, two caudal whorls per autotomic segment, mucronate caudal scales,
and a distinct longitudinal row of enlarged vertebral scales. However, the new species is easily distinguished from
these species in having a bold black transverse band at midbody that extends ventrolaterally in adult males.
Furthermore, S. carrioni, S. chlorostichus and S. euneptopsis differ from S. arndti (in parentheses) in having the
dorsal scales of the neck keeled and imbricate (slightly keeled and subimbricate) and the dorsal scales of the trunk
keeled (feebly keeled). Additionally, S. carrioni and S. euneptopsis have fewer scales at midbody (66–96, x =82.43
± 8.13 in S. carrioni and 60–80, x =70.62±5.38 in S. euneptopsis, versus 85–100, x =91.54 ± 6.32 in the new
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VENEGAS ET AL.
species), and S. chlorostichus has a strong sexual dichromatism, with the dorsal background color green in males
and brown in females (males and females gray or brown). Stenocercus empetrus differs from S. arndti in having a
venter of yellowish-orange with black reticulations (white, pale gray, or pale orange without reticulations), caudal
scales without strongly projected mucrons (strongly projected mucrons present), and attaining a larger size, with a
maximum SVL=103 mm in males and 90 mm in females (maximum SVL=90 mm in males and 77 mm in females).
Stenocercus simonsii differs from S. arndti in having the dorsal scales of the neck almost coarsely granular, while
in the new species these scales are slightly keeled and subimbricate. Stenocercus crassicaudatus and S. torquatus
differ from the new species in having the dorsal scales of the neck granular (slightly keeled and subimbricate) and
more scales around the midbody (97–121, x =108.87 ± 5.99 in S. crassicaudatus and 102–137, x =116.96 ± 8.21 in
S. torquatus, versus 85–100, x =91.54 ± 6.32 in S. arndti). Moreover, S. torquatus differs from S. arndti in having a
black antehumeral collar complete middorsally in adult males (incomplete), as well as two transverse bands
anterior to the antehumeral collar (absent), and the ability to change color between emerald green and dark brown
or gray (unable to change color). Stenocercus bolivarensis has strongly keeled and imbricate lateral body scales
(Torres-Carvajal 2007b), which are smooth in the new species, and fewer scales around the midbody (67–82,
x =73.08 ± 4.63 in S. bolivarensis, versus 85–100, x =91.54 ± 6.32 in S. arndti).
Characterization. (1) Maximum total length in males 90 mm (n = 7); (2) maximum total length in females 77
mm (n = 3); (3) vertebrals 74–94; (4) paravertebrals 87–108; (5) scales around midbody 85–100; (6) supraoculars
5–7; (7) internasals 3–4; (8) postrostrals 4; (9) loreals 2; (10) gulars 49–60; (11) lamellae on Finger IV 22–28; (12)
lamellae on Toe IV 23–30; (13) posthumeral mite pocket present as one or more vertical folds or ridges [Type 1 of
Torres-Carvajal (2007b)]; (14) postfemoral mite pocket present as a distinct pocket slightly depth with a
posteroventrally oriented slit-like opening [Type 2 of Torres-Carvajal (2007b)]; (15) parietal eye absent; (16)
occipital scales small, smooth, juxtaposed; (17) projecting angulate temporals absent; (18) enlarged supraoculars
occupying most of supraocular region in one row absent; (19) scales on frontonasal region smooth, juxtaposed;
(20) preauricular fringe absent; (21) antegular, antehumeral, gular, longitudinal, oblique, postauricular, supraauricular, and transverse antegular neck folds present; (22) lateral nuchals slightly smaller than dorsal nuchals; (23)
posterior gulars smooth, imbricate, apical pit absent; (24) lateral body scales smaller than dorsal body scales; (25)
vertebrals same size as dorsals, forming inconspicuous longitudinal row between fore- and hind-limbs; (26)
dorsolateral crests absent; (27) ventrals smooth, imbricate; (28) scales on posterior surfaces of thighs granular; (29)
prefemoral fold present; (30) inguinal groove present; (31) preanals not projected; (32) tail not compressed
laterally in adult males; (33) tail length 53–64 % of total length; (34) two caudal whorls per autotomic segment;
(35) caudals spinose; (36) dark stripe that extends anterodorsally from subocular region to supraciliaries absent;
(37) color pattern of gular region in adult females with clear marks, similar to ventral color pattern; (38) color
pattern of gular region in adult males with clear marks; (39) black blotch on ventral surface of neck in adult males
absent; (40) dark midventral stripe in adult males absent; (41) black patches on ventral surface of thighs in adult
males absent; (42) background color of dorsum grey or brown; (43) postxiphisternal inscriptional ribs not in
contact midventrally (Pattern 2B and 4A of Torres-Carvajal 2004).
Description of the holotype. Male (Figs. 1–3); SVL 79 mm; TL 101 mm; maximum head width 15.3 mm;
head length 18.9 mm; head height 11.5 mm; scales on parietal and occipital regions small, slightly wrinkled,
juxtaposed (Fig. 2); parietal eye not visible; supraoculars in five rows, smooth, with the lateral most three rows less
than half the size of the medial adjacent rows; distinct circumorbitals present; canthals two; anterior-most canthal
in contact with nasal; internasals four; postrostrals four, approximately as wide as long; supralabials seven;
infralabials six; loreals three; lorilabials in one row; preocular divided into four scales, the two dorsals in contact
with posterior canthal; lateral temporals granular; gulars in 49 rows between tympanic openings; all gulars cycloid,
smooth, imbricate, apical pit absent; second infralabial in contact with the second and third sublabials; first pair of
postmentals in contact medially; mental separated from the infralabials by the first pair of postmentals; dorsal and
lateral scales of neck granular; lateral scales of body conspicuously smaller than dorsals, but not granular, slightly
imbricate; scales around midbody 88; vertebrals enlarged, keeled, imbricate, in 74 rows, forming distinct vertebral
row; paravertebrals adjacent to vertebral row slightly enlarged, keeled, and imbricate; paravertebrals 92; ventrals
smooth, imbricate, more than twice the size of dorsals; preauricular fringe absent; antegular, antehumeral, gular,
longitudinal, oblique, postauricular, supra-auricular, and transverse antegular neck folds present; ventrolateral and
prefemoral folds present; dorsal scales of fore limbs imbricate, smooth; dorsal scales of hind limbs imbricate,
keeled, mucronate; ventral humeral scales smooth and imbricate, about one third the size of the dorsal humeral
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scales; ventral scales of forearms and hind limbs imbricate, smooth; palmars and plantars imbricate, keeled;
lamellae on Finger IV 26; lamellae on Toe IV 26; tail rounded; caudals strongly keeled, mucronate, imbricate;
basal subcaudals smooth, imbricate; posthumeral mite pocket present as one or more vertical folds [Type 1 of
Torres-Carvajal (2007b)]; postfemoral mite pocket present as a distinct pocket slightly depth with a
posteroventrally oriented slit-like opening [Type 2 of Torres-Carvajal (2007b)].
Color in life of holotype (Fig. 3): head dorsal surface green, loreal region and snout bright green, temporal
region and sides of head behind the eyes brown with white spots, thin dark brown temporal and two postocular
stripes present; dorsum brown with several pale green spots, diffuse dark brown bands, and a bold black transversal
band at midbody that extends ventrolaterally and is incomplete middorsally; black antehumeral collar present,
incomplete middorsally; flanks brown with several white spots on sides of neck and pale green flecks on sides of
trunk; tail brown; arms greenish brown and legs brown, both with dark brown flecks; throat greenish-white with
diffuse brown spots, chest and belly white with brown borders and gray flecks; ventral surface of thighs and cloacal
region cream.
Variation. Measurements, scutellation, and other morphological characters of Stenocercus arndti are presented
in Table 1. Loreals 1–5; supralabials 6–7; infralabials 5–7; second infralabials not in contact with third sublabials in
58% of specimens; first pair of postmentals not in contact medially in 25% of specimens. In two dissected
specimens the postxiphisternal pairs of inscriptional ribs were two in one specimen (both pairs long), and in the
other specimen (the first and second pair were long and the third pair short) (Pattern 2B and 4A of Torres-Carvajal
2004).
TABLE 1. Variation in scutellation and sexual dimorphism in snout–vent length (mm) of Stenocercus arndti sp. nov.
Range followed by mean ± standard deviation is given for quantitative characters if applicable.
CHARACTERS
n=13
Scales around midbody
85–100 (91.54 ± 6.32 )
Vertebrals
74–94 (83.23 ± 6.77)
Paravertebrals
87–108 (100.23 ± 5.83)
Gulars
49–60 (52.92 ± 3.65)
Supraoculars
5–7 (6)
Internasals
3–4 (4)
Subdigitals Finger IV
22–28 (24.15 ± 2.03)
Subdigitals Toe IV
23–30 (25.54 ± 1.94)
Tail length/total length
0.53–0.64 (0.58 ± 0.08)
Maximum SVL males
90
Maximum SVL females
77
Sexual dimorphism is evident in adult individuals. Males differ from females in having a conspicuous
anthehumeral black collar and a bold black transversal band at midbody, whereas females only have several diffuse
dark brown short transversal bands along the back. The other two collected adult males (CORBIDI 01681-82 and
01688) and one subadult male (CORBIDI 01690) differ from the holotype only by having the head completely
brown and the throat brown with white spots (Fig. 4: A & B). Subadult males (CORBIDI 01686-87) and a juvenile
male (CORBIDI 01691) differ from the aforementioned males in having the diagnostic mark on the dorsum dark
brown, while is conspicuous black in adult males.
In life, the adult female (CORBIDI 01692; Fig. 4: C & D) has a gray dorsum, while in the other collected
females the dorsum was brown as in the males (Fig. 4: E & G). The throat of females was gray with white spots
(CORBIDI 01692; Fig. 4 D), yellowish-green with gray spots (CORBIDI 01685; Fig. 4 F), or yellow with white
spots (CORBIDI 01689; Fig. 4 H). The two referred specimens, two adult females from Kañaris (Fig. 4: I–L),
differ from the adult females from the type locality only in having the throat and chest yellow without white or gray
spots on the throat (Fig. 4: J & L). Subadult females and the single hatchling collected have the transverse bands on
the dorsum more evident than in the adult female, especially the anthehumeral black collar.
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VENEGAS ET AL.
FIGURE 1. Stenocercus arndti sp. nov., holotype CORBIDI 01680, male, 79 mm SVL. Dorsal (A) and ventral (B) views.
Photographs by P.J. Venegas.
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FIGURE 2. Dorsal (A), lateral (B), and ventral (C) views of the head of Stenocercus arndti sp. nov. Holotype, CORBIDI
01680, male. Scale bar=3 mm. Photographs by P.J. Venegas.
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FIGURE 3. Dorsolateral (A), first half of body (B), and ventral (C) views of the holotype of Stenocercus arndti sp. nov.
(CORBIDI 01680) in life. Photographs by P.J. Venegas.
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FIGURE 4. Paratypes of Stenocercus arndti sp. nov. (A, B) Dorsolateral and ventral view of an adult male paratype
(CORBIDI 01688). (C, D) Dorsolateral and ventral view of an adult female (CORBIDI 01692). (E, F) Dorsal and ventral view
of an adult female (CORBIDI 01685). (G, H) Dorsolateral and ventral view of a subadult female (CORBIDI 01689). (I, J)
Dorsolateral and ventral view of an adult female from Kañaris (photo voucher). (K, L) Dorsolateral and ventral view of an adult
female from Kañaris (photo voucher). Photographs by P.J. Venegas.
Natural history observations. The habitat at the type locality and in the vicinity of Kañaris are croplands with
scattered large boulders and bushes. Croplands are embedded in a matrix of shorter-stature and drier forest.
However, the forest has been almost removed and only some small patches of secondary forest remain close to
ravines. All individuals of Stenocercus arndti were observed active on sunny days at between 10:00 and 15:00 h
basking on large rocks, fallen tree trunks, and fences of rock or wood. Stenocercus arndti is sympatric with S.
huancabambae and S. stigmosus in Quebrada La Iraca, and with only S. huancabambae in Quebrada Checos. Other
sympatric species of squamate reptiles collected with the new species in the type locality were Chironius
monticola, Dipsas peruana, Epictia teaguei, Liophis taeniurus, Micrurus peruvianus, and Pholidobolus sp. In the
vicinity of Kañaris, S. arndti was found sympatric only with Pholidobolus sp.
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VENEGAS ET AL.
FIGURE 5. Distribution map of S. arndti sp. nov. and its most similar species. Black star indicates the type locality and red
star indicates the locality of referred specimens (photo vouchers).
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Distribution. Stenocercus arndti is known only from two adjacent localities close to La Granja village in the
interandean valley of the Río Chotano and from one locality in the vicinity of Kañaris village in the interandean
valley of the Río Huancabamba on the Amazonian slope of the northern portion of the Cordillera Occidental in
northwestern Peru (Fig. 5). It occurs at elevations from 1997–2318 m in the regions of Cajamarca and
Lambayeque. The type locality lies within the Yungas (500–2300 m) ecoregions according to Brack (1986) and
Peñaherrera del Águila (1989).
Etymology. The specific name is a patronym for Dr. Rudolf G. Arndt of Pomona, New Jersey, USA, in
recognition of his financial support for the improvement of the herpetological collection of CORBIDI through the
BIOPAT-Programme.
Discussion
Due to the morphological similarity (e.g. spinose caudal scutellation and reduced number of caudal whorls) and
geographic proximity of Stenocercus arndti sp. n. with S. empetrus and S. eunetopsis, we assigned tentatively this
new taxon to the supraspecific clade Microphractoides, according with the phylogenetic classification of TorresCarvajal et al. (2006). The supraspecific clade Microphractoides is stemming from the most recent common
ancestor of S. empetrus, S. eunetopsis and S. imitator and is nested in the supraspecific clade Scelotrema together
with the supraspecific clades Microphractus and Saccodeira (Torres-Carvajal et al. 2006). However, tails armed
with strongly mucronate and spinose scales is a derived characteristic within Stenocercus, according to Frost and
Etheridge (1989), and Cadle (1991). Although the phylogenetic reconstruction of Torres-Carvajal et al. (2006) only
included four of the eight previously known species of Stenocercus characterized by a spiny tail and two caudal
whorls per autotomic segment, a latter phylogenetic hypothesis based on molecular and morphological characters,
that included more species showed that the species with spiny tails appeared independently in a clade composed by
species of the Scelotrema clade with no spiny tail and with three caudal whorls per autotomic segment (see TorresCarvajal 2007). Furthermore, other two species that have more spinose caudals, S. marmoratus and S. roseiventris,
than the aforementioned species also appear separated at the two different major clades in Torres-Carvajal (2007).
The definition of Microphractoides and Sceloterma is based on molecular data, which indicate that presenceabsence alone of spiny scales is not a useful character to define these groups.
Spinose caudal scutellation is also known in others lizards of several families, especially in rock-dwelling
species (Bellairs 1970). According to Bellairs (1970), Stenocercus with spinose caudal scutellation, including S.
arndti sp. n., frequent rock habitats (e.g. S. crassicaudatus, S. empetrus, S. eunetopsis, and S. simonsii), although a
few are also arboreal (e.g. S. chlorostichus and S. torquatus) (Fritts 1974; Cadle 1991; Torres-Carvajal 2000;
Torres-Carvajal et al. 2005). This suggests that our knowledge about the origin of the spiny tails and other
morphological characters, as well as the ecology of species in Stenocercus is unclear and needs to be improved for
an accurate definition of the supraspecific groups.
Acknowledgements
We thank J. Cordova and C. Torres for allowing access to the collection of the MUSM, and O. Torres-Carvajal for
allowing access to the reptile collection of the QCAZ. We also thank Rudolf G. Arndt, Pomona, New Jersey, USA,
for helpful comments on drafts of this manuscript. PJV is especially grateful to D. Rivera, C. Alvarado, K. Vinatea
and G. Tello of the environmental staff of Rio Tinto for logistic support in the field, and to F. Riva of CORBIDI.
The field work was funded by Rio Tinto.
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A NEW SPECIES OF STENOCERCUS FROM NORTHWESTERN PERU
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APPENDIX. Specimens examined.
Stenocercus carrioni— ECUADOR: Loja: Celica, Huajala QCAZ 10319, 10324, 10329.
Stenocercus chlorostichus— PERU: Cajamarca: Santa Cruz: Agua Azul, MUSM 25821.
Stenocercus crassicaudatus— PERU: Cuzco: Urubamba: Machu Picchu CORBIDI 09058, MUSM 4905, 4906, 8691.
Stenocercus empetrus— PERU: Cajamarca: Cajamarca CORBIDI 06532, 08638, MUSM 4909–10, La Colmena MUSM 8659,
San Vicente MUSM 8676; Departamento de la Libertad: Huamachuco MUSM 8658, 8675, 8677–8680.
Stenocercus eunetopsis— PERU: Cajamarca: Santa Cruz, Udima, MUSM 4018, 4022–4029.
Stenocercus simonsii—ECUADOR: Azuay: Giron, sendero El Salado, QCAZ 09636.
Stenocercus torquatus—PERU: Ayacucho: La Mar: carretera San Antonio-Chinquintirca, CORBIDI 06956; Cusco: La
Convención: Echarate, 06041, 06044, 08619; Junín: Alto Yurinaqui, MUSM 8617–8619; Pasco: Oxapampa: Huampal,
CORBIDI 07211; Bosque de Sholet, CORBIDI 09912.
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VENEGAS ET AL.