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NUMBER 10, PAGES 1-21 DECEMBER 6, 1972

NEW SPECIES OF LIZARDS OF THE

GENUS STENOCERCUS FROM PERtJ

(SAURIA: IGUANIDAE)

By

Thomas H, Fritts^

In the course of an evolutionary study of the genus Stenocercus
seven unnamed species were studied in the field and laboratory. In
order to facilitate discussion of the evolutionary relationships of
the genus, these new species are described in the present paper.
Although several other taxonomic problems exist, the nomenclature
of Etheridge (1970) is used herein. A complete taxonomic review
will be presented in a paper cun'ently in preparation.

It has become apparent that the generic status of all species in
the genera Ophnjoessoides Dumeril, 1851, and Stenocercus Dumeril
and Bibron, 1837, warrants consideration. My work indicates that
many of the species presently included in Ophnjoessoides are more
closely related to species of Stenocercus than to other Ophnjoes-
soides. All species named in this paper are members of the genus
Stenocercus as it is viewed by me.

ACKNOWLEDGMENTS

I am grateful to Richard E. Etheridge for information on type
specimens of Stenocercus and Ophnjoessoides in European muse-
ums. He has generously shared specimens in his care and given
freely of his time. William E. Duellman has encouraged me in my
work with Stenocercus and provided energies in laboratory and
field aspects of the study. Many persons in South America assisted

1 Research Fellow, Museum of Natural History, University of Kansas. Pres-
ent Address: Department of Biology, St. Edward's University, Austin, Texas
78704.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

in field work; I am particularly grateful to Oscar Ochoa M., Manuel
Moro, Luis Inope, James Muir, and Hugo Paz. Access to the her-
petological collections of several American institutions is gratefully
acknowledged.

The committee on Systematics and Evolutionary Biology spon-
sored by the National Science Foundation (GB-4446X, George W.
Byers, principal investigator and GB-8785, J. Knox Jones, Jr., prin-
cipal investigator) provided funds for field and laboratory studies.
Funds for travel to South America were also provided from the
F. William Saul Fund of the Museum of Natural History, Univer-
sity of Kansas.

MATERIALS AND METHODS

The new species reported in the present report are represented
in a collection of nearly 1000 specimens of the genus Stenocercus
collected in the period of 1968-1972 and deposited in the University
of Kansas Museum of Natural History (KU). Taxa are delineated
on the basis of morphological and geographical discordance. The
characters discussed in the diagnoses are numbered to facilitate
comparison of character states between taxa; to avoid redundancy
not all character states are given in each diagnosis. The ranges of
characters presented in the diagnoses are based on all known speci-
mens. The description is based on specimens from the type locaHty
only. All other samples of the taxa are discussed in the "Remarks"
section when pertinent. Geographic variation is examined through
use of the Sum of Squares-Simultaneous Test Procedure (SS-STP).
The SS-STP is an a posteriori test of significance of means (Sokal
and Rohlf, 1969) and defines nonsignificant subsets of samples after
significant variance is demonstrated by an overall analysis of
variance. The test is used for samples of two means realizing that
the probability level obtained from the analysis of variance is of
an a priori test, not an a posteriori one. Significance in analyses
with only two samples is therefore tentative.

TAXONOMIC ACCOUNTS

Stenocercus apurimacus new species
Figure IB

Holotype.—KU 134273, an adult male, from Puente Pachachaca,
15 km west of Abancay, 1800 m, Departamento Apurimac, Peru,
one of a series collected on 20-22 November 1969, by Thomas H.
Fritts, Patricia R. Fritts, and Oscar Ochoa M.

Paratopotypes.—KU 134270-134272, 134274-134309, same data
as holotype.

Diagnosis.— I) Posthumeral pocket deep; 2) postfemoral pocket
absent; 4) scales posterior to tympanum equal in size to dorsal

NEW SPECIES OF LIZARDS FROM PERtJ 3

neck scales, keeled, imbricate; 5) scales on posterior thigh keeled,
imbricate; 6) vertebral row continuous, forming high dorsal crest
extending onto tail; 7) caudal scales equal in size to ventrals, three
caudal whorls per autonomic segment; 8) antehumeral fold absent;
9) neck fold absent; 10) scales around middle of body less than 62;
11) scales across gular region between tympana less than 27; 13)
venter of males gray-beige to gray with a blue hue; 17) posterior
head scales large, wider than long, rounded posteriorly, a pair of
large parietals in broad contact posterior to small interparietal.
Stenocercus apurimaciis is the only species of Stenocerciis occurring
in southern Peru which lacks antehumeral and neck folds. Steno-
cercus apurimaciis differs from species of Ophnjoessoides in lacking
keeled ventrals and an antehumeral fold and in having enlarged
scales on the posterior margin of the head.

Description. — N=35; scales of parietal and occipital regions of
head large, with curved posterior margins; head scales smooth and
only slightly convex; interparietal small, bordered laterally by pair
of large parietal scales which meet posterior to it; scales across
occipital region between angulate temporals 6-9 (7.1±0.1804);
largest supraoculars nearly equal in size to scales of interorbital
region; supraoculars at greatest width of semicircles 4-6 (5.4±0.094).
Scales of lateral neck imbricate and keeled, slightly smaller than
dorsal neck scales; antehumeral and neck folds absent; scales across
gular region between tympana 18-26 (23.0+0.362).

Vertebral row continuous, forming high dorsal crest from occiput
to tail; dorsal scales large, imbricate, keeled; scales around middle
of the body 48-61 (52.3±0.519); lateral and ventral scales approxi-
mately equal in size to dorsals; ventrals smooth; caudal scales
normal; three caudal whorls per autonomic segment. Deep post-
humeral pocket present; postfemoral pocket absent; scales of ventral
surface of proximal forelimb large, imbricate, keeled; scales of
posterior thigh smaller, imbricate and keeled; subdigital lamellae
of the fourth finger 15-22 (17.9±0.256); subdigital lamellae of
fourth toe 21-28 (25.9+0.264).

Coloration of males: Dorsum red-brown to medium brown with
few indistinct black chevrons; narrow black collar edged posteriorly
with white across lateral and dorsal neck; labial scales near the
angle of the jaw black; chin white with few gray flecks; venter
beige- white with faint blue or pink tint in some specimens; ventral
pelvic region, ventral base of tail and ventral thighs dull yellow.

Coloration of Females: Dorsum brown with gold-bronze tint;
dorsal and lateral surfaces of the neck gold-brown; chin and gular
region black posterior to level of insertion of forelimb; chin white
medially in some specimens; black collar absent; dorsal chevrons
less distinct than those of males; venter gray- white.

Distribution. — The species is known in Peru from elevations of

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 1. A. Stetwcercus empetrus, KU 134390, $ 92 mm in snout-\ent length.
B. S. apurimacus, KU 134285, $ 84 mm. C. S. ochoai, KU 133888, 6 95 mm.

NEW SPECIES OF LIZARDS FROM PERO

FiG. 2. A. Stenocercus praeomatm, ct. KU 134224-134232, S approximate
snout-vent length 100 mm. B. S. orientalis, KU 134456, 6 65 mm. C. S.
7iiibicola, KU 134107, 6 72 mm.

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

1800 and 2700 m in the interandean valleys of the Rio Apurimac
and Rio Pachachaca in the departments of Cuzco and Apurimac
( Fig. 3 ) . The upper valley of the Rio Apurimac divides into several
smaller valleys, of which the Rio Pampas, Rio Pachachaca and
Rio Apurimac valleys are the largest. Although Stenocercus
apurimacus is known only from the last two valleys it is likely to
occur in the former. Mountain ranges above 4000 m in elevation
separate upper regions of the Rio Pampas from the adjacent Rio
Mantaro drainage. The Cordillera Vilcabamba is largely above
5000 m and separates the Rio Apurimac from the Rio Urubamba.

Fig. 3. Distribution in Perii of Stenocercus apurimacus (closed triangles),
S. ivitus (open triangle), S. nubicola (closed circle), S. ochoai (open circles),
S. orientalis (closed square), S. praeornatus (open squares), and S. empetrus
( stars ) .

NEW SPECIES OF LIZARDS FROM PERtJ 7

The Rio Apurimac courses between high cordilleras until its junc-
tion with the Rio Ene and Rio Mantaro in central Perii.

Ecological Observations. — The species is found in xeric habitats
throughout its range. At localities below 2400 m. Acacia and other
thorn shrubs and cactus form the predominant plant cover. The
lizards are active on the ground at the bases of shnjbs and on
rock piles. Localities at 2700 m are more mesic and have more
grasses and shrubs. At these higher elevations individuals are ob-
served in rows of Agave. Stenocercus ochoai occurs in sympatry
with S. apurimucus at Curahuasi, 2700 m, Departamento Apurimac.

Etymology. — The trivial name for this lizard, which occurs
throughout the drainage of the Rio Apurimac, is a latinized noun
from the Quecchua language, apu meaning the great, and rimac
meaning the talker or one who speaks. Guardia (1967) translated
the word apurimac as meaning "el gran hablador" or great speaker.
The species is dedicated to Dr. Philip S. Humphrey, who has al-
lowed use of the facilities of the University of Kansas Museum of
Natural History.

Remarks. — Specimens from the main valley of the Rio Apurimac
in the Departamento Apurimac (Puente Cunyac and Curahuasi)
have more scales around the middle of the body than those from
the type locality. The populational means for the two main valley
localities (56.8 and 56.0 respectively) are included in a nonsig-
nificant subset significant from the mean of the type locality ( 52.3 )
in SS-STP analysis, p<0.05.

The extent of white coloration on the medial chin of females
varies with altitude in the main valley of the Rio Apurimac. Females
from elevations below 2000 m have a solid black chin, whereas those
from between 2000 and 2700 m on both sides of the valley have a
gray- white or white region in the middle of the chin.

Stenocercus empetrus new species
Figure lA

Holotype. — KU 134394, an adult male from Huamachuco, 3350
m, Departamento La Libertad, Perii, one of a series collected on 16-
17 April 1970 by Thomas H. Fritts and Patricia R. Fritts.

Paratopoty pes. —KU 134379-134393, 134395-134400, same data
as the holotype.

Diagnosis. — 1) Posthumeral pocket absent; 2) postfemoral pocket
large; 3) dorsals imbricate, smooth or weakly keeled; 4) scales
posterior to tympanum granular, smaller than dorsal neck scales;
5) scales of posterior thigh granular; 6) vertebral row discon-
tinuous, only a few scales on neck and anterior body morphologically
distinct from adjacent dorsal scales; 7) caudals spinose, enlarged,
subequal, two caudal whorls per autonomic segment; 8) ante-
humeral fold present; 9) neck fold well developed; 10) scales

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

around middle of body more than 86; 13 ) dorsal body dark brown or
black, numerous yellow flecks and spots on dorsum in some pop-
ulations.

Three species of Stenocercus have spinose caudal scutellation
and lack a continuous vertebral row. Stenocercus empetrus differs
from S. marmoratus (D'Orbigny, 1837) in having more than 86
scales around the middle of the body and in having more than 38
scales across gular region between tympana. Stenocercus empetrus
differs from S. carrioni Parker, 1934, in having caudal whorls which
are subequal in size and in having a more robust body. It differs
fr >m S. shnonsii Boulenger, 1899, in lacking a vertebral row forming
a distinct dorsal crest, in having smooth or weakly keeled dorsals,
and in having a less spinose caudal scutellation.

Description. — N=21; scales of dorsal head small, subequal in
size, convex; scales of posterior head similar to dorsal neck scales;
posterior margin of head indistinct; temporal region bulbous; inter-
parietal scale not distinct, parietal eye lacking; scales across occipital
regon between angulate temporals 14-17 ( 15.3±0.211); supraocular
s ales small subequal in size, lacking rugosities and keels; supra-
o ulars at greatest width of semicircle 5-7 (5.8±0.118); head scales
along medial line between occiput and rostral 16-20 ( 17.9±0.217).
Scales of lateral neck granular, smaller than dorsal neck scales;
anteh-'meral fold present, lined with granular scales; neck folds
well devel )ped, connected to antehumeral fold by longitudinal fold
n some specimens; largest neck fold extending ventrally across
gular region in some specimens; scales of gular region between
tympana small, 39-49 (43.1±0.599).

Vertebral row discontinuous; vertebral scales distinct from ad-
jacent scales on neck only, forming an incomplete dorsal crest in
some specimens; dorsal scales imbricate, smof)th or weakly keeled,
scales around middle of body 90-112 (99.9±1.140); scales of lateral
body less than one-half size of dorsals, granular, meeting dorsals
abruptly in dorsolateral region; caudal scales large, with heavy
keels which project slightly beyond posterior margins of scales;
two caudal whorls per autonomic segment; dorsomedial row of
caudal scales lacking enlarged keels, not forming high crest. Post-
humeral pocket absent; postfemoral pocket formed by large fold;
scales of ventral surface of proximal forelimb granular; scales of
posterior thigh granular; subdigital lamellae of fourth finger 19-25
(21.9±0.379); subdigital lamellae of fourth toe 25-32 (29.0±0.495).

Coloration: Little sexual dimorphism in color pattern; dorsal
ground color black, heavily flecked and spotted with pale yellow;
lateral neck and body black with prominent yellow spots in irregular
longitudinal and transverse rows; head black with numerous yellow
flecks; prominent red spots in interorbital and occipital region;
venter of males orange-yellow with heavy black reticulations; chin

NEW SPECIES OF LIZARDS FROM PERtJ 9

of males orange-yellow with black or bright yellow with black
reticulations; venter of females yellow with black reticulations.

Distribution. — In addition to the type locality the species is
known from Laguna Sacsacocha, 12 km E Huamachuco, 3200 m,
Departamento La Libertad; Otuzco, 2730 m, Departamento La
Libertad; Cajamarca, 2800 m, Departamento Cajamarca; and 3 km
E Celendin, 2650 m, Departamento Cajamarca (Fig. 3). All of
the localities, except Otuzco, are in the drainage of the Rio Crisneja
and adjacent tributaries of the Rio Marafion. The single locality in
the Pacific drainage, Otuzco, is in the upper valley of the Rio Moche.

Ecological Observations. — Individuals of both sexes are found
in large rock piles and rock exposures. Stenocercus rnelanopygus
Boulenger, 1900, occurs in sympatry with S. empetrus throughout
the distribution of the latter species in the Atlantic drainage; S.
ineJanopijgus is common on the ground at the bases of shrubs, on
solitary rocks, and at the periphery of large rock piles. Stenocercus
empetrus is common in the mesic habitats near Huamachuco and
Celendin but rare on the dry slopes near Cajamarca, despite the
availability of large rocks and rock piles preferred at other localities.
The black coloration of individuals from Celendin closely matches
the black volcanic rocks on which the lizards occur. Stenocercus
empetrus is also sympatric with S. ornatissimus (Girard, 1857)
on the Pacific slopes near Otuzco. The latter species was abundant
in agave rows and at the bases of shrubs; habitat partitioning be-
tween the two species similar to that observed on Atlantic slopes
is probable.

Stenocercus empetrus diff^ers from the two species with which
it is sympatric in several notable aspects. It differs in lacking
marked sexual dimorphism in the ventral coloration and in having
enlarged and spinose caudal scutellation, a bulbous temporal region,
smooth or weakly keeled dorsals, granular scales on lateral body
and neck, and a larger body size.

Etymology. — The trivial name is from the Greek empetros mean-
ing rock-dwelling and refers to the preferred habitat of the species.

Remarks. — Individuals of this species from Celendin, 2650 m,
Departamento Cajamarca, differ from those from the type locality
in having greater sexual dimoiphism in coloration. Adult males
are entirely black, except for a few small white flecks on the lateral
body and posterior surface of the thighs. Subadult males are black
to brown-black with transverse rows of small white spots extending
from the dorsum to the lateral edge of venter. The antehumeral
fold is black and outlined on anterior and posterior margins with
white scales. The venter is white with gray reticulations whereas
the chin is black with white spots. The posterior one-half of the
venter has a pink sufll^usion. Adult females are medium gray with
faint transverse rows of white flecks. The venter is gray- white with

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

a pink suffusion on the posterior one-half. Juveniles are similar to
females, except in having more prominent white flecks on lateral
body.

Individuals from Cajamarca, 2800 m, Departamento Cajamarca,
are somewhat similar in coloration to subadult males from Celendin,
except in having equal numbers of yellow-beige and black scales
on the dorsal and lateral body. The occipital region of the head
is black whereas the interorbital region has a few scales which are
dull red-brown. The venter is orange-yellow with prominent black
spots and reticulations.

The number of paravertebral scales is higher in the specimens
from Celendin than in those of other localities (p<0.05; SS-STP).
The dorsal scales are smooth in the majority of specimens from
Celendin, whereas they are smooth laterally and weakly keeled
medially in specimens from other localities.

Stenocercus ivitus new species

HoIotype.—K\J 134654, an adult female from summit of
Cordillera Huancabamba between Canchaque and Huancabamba,
3100 m, Departamento Piura, Peru,. obtained on 11 May 1970, by
Thomas H. Fritts and Patricia R. Fritts.

Diagnosis.— 1) Posthumeral pocket absent; 2) postfemoral
pocket present; 4) scales posterior to tympanum nearly equal in
size to dorsal neck scales, imbricate, keeled; 5) scales of posterior
thigh granular; 6) vertebral row discontinuous, vertebral scales not
forming dorsal crest; 7) caudals normal, three caudal whorls per
autonomic segment; 8) antehumeral fold absent; 9) neck fold ab-
sent; 10) scales around middle of body 44.

Stenocercus ivitus differs from all species of Ophryoessoides and
Stenocercus in lacking antehumeral and neck folds, in lacking a
continuous vertebral scale row which forms a dorsal crest, and in
having granular scales on the posterior surface of the thigh.

Description.— ^=1; head scales moderate in size, subequal,
prominently keeled; 16 scales on middorsal line between occiput
and rostral; supraoculars subequal, keeled. Interparietal scale lack-
ing unpigmented spot of parietal eye; scales of occipital region
equal in size to dorsal neck scales, posterior margin of head indis-
tinctly demarcated from dorsal neck; a longitudinal row of angulate
temporal scales from posterior margin of eye to posterior margin of
head scales; 11 scales between angulate temporals across occipital
region. Scales of anterior margin of tympanic opening slighdy
denticulate, projecting into opening; scales of lateral neck large,
equal in size to dorsal neck scales, imbricate, keeled; scales posterior
to tympanum imbricate, keeled; no antehumeral fold; no neck
folds; 21 scales across the gular region between tympana.

Vertebral scales not in continuous row, similar to adjacent dor-

NEW SPECIES OF LIZARDS FROM PERtJ 11

sals, not forming distinct dorsal crest; dorsal scales imbricate, keeled,
mucronate, lanceolate; lateral scales equal in size to dorsals; ventral
scales imbricate, smooth, lanceolate; 44 scales around middle of
body. Scales of posterior thigh granular; scales of ventral surface
of proximal forelimb imbricate, keeled; no posthumeral pocket;
postfemoral pocket moderate in size; 20 subdigital lamellae on
fourth fingers; 26 subdigital lamellae on fourth toe. Caudal scales
normal, imbricate, keeled; three caudal whorls per autonomic

segment.

Coloration of Female: Dorsum light brown with large black
blotches in two longitudinal rows between yellow dorsolateral
stripes; light yellow stripe on lateral neck and body from ventral
margin of tympanum to base of hind limb; venter white with pinkish
bronze tint; dark flecks on gular region; indistinct black lines on

lateral venter.

Distribiition.—Stenocercus ivitus is known only from the type
locality at the summit of the Cordillera Huancabamba (Fig. 3).
This Cordillera forms the continental divide between the Rio Piura
(Pacific drainage) and the Rio Huancabamba (Atlantic drainage).

Ecological Observations.— The adult female was observed bask-
ing on a pile of dead branches at the margin of a low shrub-forest.
The moist forest at this locality is produced by moisture-saturated
winds coming from the Pacific. The western side of the summit has
a low cloud forest, whereas the eastern side is drier with only local
patches of forest and extensive tall grasses. Stenocercus ivitus is
known to occur in sympatry with S. nuhicola.

Etymologi/.—The trivial name is a latinized acronym of the
Instituto Veterinario de Investigaciones Tropicales y de la Altura
a cooperative research institute of the Universidad San Marcos and
the United Nations FAO. The faculty and staff of IVITA assisted
field studies through provision of facilities and logistical support.

Stenocercus nubicola new species
Figure 2C

Holotijpe.—KU 134107, an adult male, one of a series from sum-
mit of Cordillera Huancabama between Canchaque and Huanca-
bamba, 3100 m, Departamento Piura, Peru, collected on 11 May
1970 by Thomas H. Fritts and Patricia R. Fritts.

Paratopotijpe.—KU 134108, a juvenile male with same data as
theholotype.

Diagnosis.— I) Posthumeral pocket absent; 2) postfemoral
pocket present; 4) scales posterior to tympanum less than one-halt
size of dorsal neck scales, imbricate and smooth or granular; 5)
posterior thigh granular; 6) vertebral row complete, forming dorsa
crest; 7) caudal scales equal in size to ventrals, keeled, three caudal
whorls per autonomic segment; 8) antehumeral fold distinct; 9)

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

neck fold present; 10) scales around middle of body more than 90;
11) scales across gular region between tympana fewer than 40;
13 ) black midventral stripe in males, bifurcate anteriorly, extending
onto ventral thighs posteriorly.

Description. — N=2; head scales smooth, sHghtly convex, larger
than supraoculars, subequal; scales along medial line between
occiput and rostral 17; supraoculars subequal, smooth; supraoculars
in transverse line at greatest width of semicircles 5-6; scales of
occipital region rough, not keeled, distinct from dorsal neck scales;
scales across occipital region between angulate temporals 10-12;
parietal eye absent; anterior margin of auricular opening with three
denticulate scales; antehumeral fold present; neck folds present;
scales posterior to tympanum imbricate and smooth, smaller than
dorsal neck scales; scales across gular region between tympana 38-39.

Vertebral row continuous, forming distinct dorsal crest; dorsal
scales small, imbricate, keeled; laterals slightly smaller than dorsals;
scales around middle of body 93-94; scales of posterior thigh gran-
ular; scales of ventral humerus imbricate, smooth; posthumeral
pocket absent; postfemoral pocket distinct; subdigital lamellae of
fourth finger 17-22; subdigital lamellae of fourth toe 24-28; caudal
scales equal in size to ventrals, keeled; three caudal whorls per
autonomic segment.

Coloration of Male: Chin, gular region and ventral pectoral
region white with numerous black spots; wide charcoal-gray mid-
ventral stripe, bifurcating anteriorly to axilla, bifurcating onto
ventral thighs posteriorly; lateral body and lateral venter light
green with numerous red-orange spots and flecks; dorsum medium
brown with small dark brown or black blotches; beige dorsolateral
stripes connected with white loreal region; supralabials and infra-
labials black. Numerous light blue-green transverse markings on
dorsum; collar fold bordered posteriorly by black bar.

Coloration of Juvenile: Dorsum gray-brown; dorsolateral stripes
beige.

Distribution. — Stenocercus nubicola is known only from the type
locality (Fig. 3). The Cordillera Huancabama forms the con-
tinental divide in extreme northern Peru and esparates the Rio
Huancabama Valley (Atlantic drainage) and the Rio Piura Valley
( Pacific drainage ) .

Ecological Observations. — Stenocercus nubicola occurs in sym-
patry with Stenocercus ivitus. The holotype was observed basking
on a fallen tree limb in an t)pen areas of the low montane shrub-
forest. Stenocercus nigromaculatus Noble, 1924, is known at lower
elevations in the same region.

Etymology. — The trivial name is from the Latin nubes meaning
cloud and cola meaning dweller. The name refers to the association

NEW SPECIES OF LIZARDS FROM PERtJ 13

of the species with the heavy fogs characteristic of the Pacific
slopes in northern Peru.

Stenocercus ochoai new species
Figure IC

Holotype. — KU 133888, an adult male, one of a series from
Chiica, 10 km NW Ollantaytambo, 2700 m, Departamento Cuzco,
Peru, collected on 30 November 1969 by Thomas H. Fritts, Patricia
R. Fritts and Oscar Ochoa M.

Paratypes.—KU 133874-133887, 133889, 139263. Topoparatypes
with same data as holotvpe, except KU 133874-133883 collected
15-16 November 1969, KU 133889 collected 17 January 1970, and
KU 139263 collected 9 January 1971.

Diagnosis. — 1) Posthumeral pocket absent; 2) postfemoral
pocket deep; 3) dorsal scales but slightly larger than lateral scales,
imbricate, keeled; 4) scales posterior to tympanum one-half size of
dorsal neck scales, imbricate, usually keeled; 5) scales of posterior
thigh imbricate, keeled; 6) vertebral row continuous, forming raised
dorsal crest; 7) caudal scales approximately equal in size to ventrals,
three caudal whorls per autonomic segment; 8) antehumeral fold
present; 9) neck fold present; 10) scales around body fewer than
71; 11) scales across gular region between t>'mpana fewer than 27;
13) venter of adult males black.

Stenocercus ochoai is most similar to Ophryoessoides arenarium
(Tschudi, 1845) from which it differs in having larger dorsal scales,
black ventral coloration in males, and only three caudal whorls
per autonomic segment.

Description. — N=12; head scales convex, not keeled, subequal;
scales of lateral occipital region largest of head scales; parietal eye
indistinct or absent; head scales distinct from dorsal neck scales;
scales across occipital region between angulate temporals 8-12
(10.3±0.333); scales on medial line between occiput and rostral
16-19 (17.1±0.285). Entire anterior margin of tympanic opening
with denticulate scales; antehumeral fold prominent; neck folds
present, occasionally extending onto ventral neck, not forming
strong gular fold; scales posterior to tympanum approximately one-
half size of dorsal neck scales, imbricate, usually keeled; scales
across gular region between tympana 20-26 (22.7±0.582); dorsals
imbricate, keeled; lateral scales slightly smaller than dorsals; verte-
bral row continuous, forming distinctly raised dorsal crest; keels
of dorsal scales forming longitudinal lines which converge only
slightly on posterior one-half of body; scales around middle of body
56-67 (61.7±0.882). Caudals approximately equal in size to ventrals,
keeled, three caudal whorls per autonomic segment; scales of ven-
tral surface of upper arm large, imbricate, weakly keeled; scales of
posterior thigh imbricate, keeled; subdigital lamellae of fourth finger

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

18-21 (19.8±0.241); subdigital lamellae of fourth toe 24-31 (27.5±
0.634); posthumeral pocket absent; postfemoral pocket deep.

Coloration of Males: Dorsum medium to copper brown with
occasional small black blotches; lateral body, limbs and tail brown
with lime-green suffusion; ventrolateral body greenish yellow; chin
light yellow with extensive black suffusion; venter duU black with
some yellow laterally; anal area, ventral base of tail and ventral
thighs dull yellow, occasionally with black suffiusion.

Coloration of Females: Dorsum gray-brown to charcoal with
occasional beige or light gray scales; lateral body light brown;
\'entrolateral body gray-brown with bronze or light red-orange
suffusion; venter beige-white; chin beige-white with gray suffusion;
lateral neck same as dorsal ground color. Juveniles similar with
dark brown blotches on dorsum.

Distribution. — Stenocercus ochoai (Fig. 3) is known between
2000 and 3000 m in the valleys of the Rio Urubamba ( Burt and Burt,
1930) and Rio Apurimac ( Mertens, 1952). The two rivers are sepa-
rated by the Cordillera Vilcabamba in excess of 4000 m elevation.

Ecological Observations. — Stenocercus ochoai occurs in sym-
patry with Stenocercus crassicaudatus (Tschudi, 1845) between
2000 m and 2600 m in the Rio Urubamba Valley. It occurs in
sympatry with Stenocercus apurimacus at Curahuasi, 2700 m, De-
partamento Apurimac, in the Rio Apurimac Valley. In areas of
allopatry, individuals can be observed using the ground substrate,
bases of shrubs, isolated rocks and rock piles for feeding and basking
activity. Males tend to occupy rocks whereas females are more
frequently found on the ground. In sympatry with the rock-dwelling
Stenocercus crassicaudatus, S. ochoai is confined to the ground,
isolated rocks and small rock piles.

Etymology. — The trivial name is a patronym for Dr. Oscar Ochoa
M., who generously provided assistance and support with field work
in southern Peru.

Remarks. — Specimens of Stenocercus ochoai have been errone-
ously reported by Barbour and Noble (1920) and Burt and Burt
(1930) under the names Liocephalus ervingi (Stejneger, 1913)
and Liocephalus arenarius (Tschudi, 1845), respectively.

Stenocercus orientalis new species
Figure 2B

Holotype. — KU 134466, an adult male, one of a series from
Chachapoyas, 2340 m, Departamento Amazonas, Peru, collected on
2 May 1970, by Thomas H. Fritts and Patricia R. Fritts.

Paratopotypes.—KU 134447-134465, 134467-134473 collected 1-2
May 1970.

Diagnosis. — 1) Posthumeral pocket absent; 2) postfemoral
pocket present; 4) scales posterior to tympanum half size of dorsal

NEW SPECIES OF LIZARDS FROM PERtJ 15

neck scales, imbricate, keeled; 5) scales of posterior thigh granular;
6) vertebral row continuous, not forming dorsal crest; 7) caudal
scales approximately equal in size to ventrals, keeled, three caudal
whorls per autonomic segment; 8) antehumeral fold present; 9)
neck fold present; 10) scales around the middle of the body 47-58;
15) head scales prominently keeled.

Stenocercus orientalis, S. ornatissimiis (Girard, 1857), S. moestus
Boulenger, 1885, S. chrysopygus Boulenger, 1900, and Ophryoes-
soides haenschi (Werner, 1901) differ from other species of Steno-
cercus and Ophryoessoides in lacking a dorsal crest and spinose
caudal scutellation, and in having antehumeral and neck folds.
Stenocercus orientalis is uniquely characterized by prominently
keeled scales on the snout and interorbital region. It differs from
S. ornatissimus in having a well-developed postfemoral pocket, and
from S. moestus in having more than 47 scales around the middle
of the body. Stenocercus orientalis is distinguished from S. haenschi
by the absence of green dorsal coloration and presence of fewer
than 30 scales across the gular region. It differs from S. chrysopygus
in having imbricate scales posterior to the tympanum and keels of
dorsals in continuous longitudinal series.

Description. — N=27; head scales subequal, keeled; supraoculars
subequal, keeled; parietal eye only occasionally visible as unpig-
mented spot on small interparietal scales; head scales distinct from
those of dorsal neck; 10-13 (12.0±0.173) scales across occipital re-
gion between angulate temporals; 14-18 (16.3±0.267) scales on
middorsal line between occiput and rostral. Anterior margin o£
auricular opening with 1-2 denticulate scales projecting into open-
ing; antehumeral fold prominent; neck fold present; scales posterior
to dorsal one-half of tympanum imbricate, keeled, nearly granular,
less than one-half size of dorsal neck scales. Scales on edges of
antehumeral and neck fold larger than adjacent scales, keeled,
imbricate; 23-38 (25.4±0.289) scales across gular region between
tympana. Dorsals imbricate, keeled; vertebral row continuous, not
forming dorsal crest; keels of dorsal scales forming longitudinal
lines converging slightly posteriorly; 47-58 (51.9±0.552) scales
around middle of body. Caudals approximately equal in size to
ventrals, keeled; three caudal whorls per automatic segment. Scales
of ventral humerus keeled, imbricate; scales of posterior thigh
granular; 17-22 (19.7±0.275) subdigital lamellae on fourth finger;
25-31 (27.7±0.287) subdigital lamellae on fourth toe; posthumeral
pocket absent; postfemoral pocket deep.

Coloration of Males: Head brown with a few light spots; light
stripe from under eye to dorsolateral region of shoulder; dorsum
brown with irregular dark brown or black blotches; blotches oc-
casionally forming chevrons; ventral pelvic region and ventral thighs
yellow; chin and gular region beige, similar to venter.

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Coloration of Females: Dorsum brown; gray-white stripe from
under eye extending posteriorly dorsal to tympanum and onto dorso-
lateral body; two rows of irregular dark blotches on dorsum; verte-
bral stripe absent; chin and venter gray-beige.

Distrilntfion. — The species is known in Peru only from the type
locality in the valley of the Rio Utcubama (Fig. 3). The Rio
Maraiion Valley has an elevation of less than 1000 m at this latitude
and bounds this range of the Andes on the west and north. The
most eastern ridge of the Andes, reaching above 3000 m at this
latitude, is immediately east of Chachapoyas. The extremely xeric
valley of the Rio Maraiion is in extreme contrast with the moist
eastern slopes.

Ecological Observations. — In the vicinity of Chachapoyas
patches of open chaparral are intermixed with low montane forest.
The species was collected in open areas at the bases of shrubs.
There are few exposed rocks in the immediate area; lizards seek
refuge in holes at the bases of shrubs and in tall grasses. No other
iguanid species is known in the area.

Etymology. — The trivial name is from the Latin oriens meaning
east and refers to the distribution of the species east of the Rio
Maraiion valley.

Stenocercus praeornatus new species

Figure 2A

Holotype. — KU 134231, an adult male, one of a series from
Comas 3220 m, Departamento Junin, Peru, collected on 26-27
September 1969, by Thomas H. Fritts and Patricia R. Fritts.

Paratopotypes.—KU 134224-134230, 134232, same data as holo-
type.

Diagnosis. — 1) Posthumeral pocket consisting of weak fold; 2)
postfemoral pocket large; 4) scales posterior to tympanum granular,
less than one-half size of dorsal neck scales; 5) scales of posterior
thigh granular; 6) vertebral row continuous, forming high dorsal
crest; 7) caudals approximately equal in size to ventrals, keeled,
three caudal whorls per autonomic segment; 8) antehumeral fold
present; 9) neck fold present; 10) scales around middle of body
more than 8(S; 13) ventral coloration of males rose pink; 14) gular
region of males black.

Of the species having a black ventral collar in males, Stenocercus
praeornatus is the only one having more than 88 scales around the
middle of the body and having pink ventral coloration in males.

Description. — N=9; scales of head subequal, convex, not keeled;
supraoculars smooth, not greatly enlarged; supraoculars at greatest
width of semicircles 5-7 (6.3±0.236); scales of medial region of
occipital area granular, smaller than more lateral scales, scales
across occipital region between angulate temporals 12-14 ( 13.3±

NEW SPECIES OF LIZARDS FROM PERtJ 17

0.236); parietal eye absent, interparietal indistinct; scales of dorsal
snout slightly larger than other head scales; scales along midline
between occiput and rostral 17-20 ( 1(S.(S±0.324 ) . Scales of anterior
margin of auricular opening only weakly projecting into ear; ante-
humeral fold prominent; neck folds often connected with ante-
humeral fold; scales posterior to tympanum granular, less than one-
half size of dorsal neck scales; neck fold continuing ventrally, meet-
ing its fellow, forming a weak gular fold in some specimens; scales
across gular region between tympana 40-57 (48.2± 1.786).

Vertebral row continuous; vertebral scales forming distinct
raised dorsal crest; skin of lateral neck folded, forming slight dorso-
lateral fold from dorsal margin of tympanum to point posterior to
scapular region; dorsal scales imbricate, keeled; scales around
middle of body 100-121 (106.8+2.29); lateral scales less than one-
half size of dorsal scales, granular or slightly imbricate; lateral scales
sharply meeting dorsals at dorsolateral region; shallow fold in
axillary region forming shallow posthumeral pocket; postfemoral
pocket large; dorsal crest continuing onto tail; caudal scales ap-
proximately equal in size to ventrals, imbricate, keeled; three caudal
whorls per autonomic segment. Scales of ventral humerus imbricate,
keeled or smooth; scales of posterior thigh granular; subdigital
lamellae of fourth finger 20-22 (20.8+0.324); subdigital lamellae
of fourth toe 28-31 ( 29.4+0.324 ) .

Coloration of Males: Head dull chocolate brown with scattered
yellow-beige spots; nape dull brown posterior to level of shoulders
with diagonal yellow markings and spots; dorsum gray-brown with
faint irregular darker brown diamond blotches; lateral areas light
yellow-brown with dull yellow spots; chin gray-brown peripherally;
central chin blue-green with yellow spots; gular region rose pink
with faint gray reticulation; chest anterior to pectoral girdle jet
black; ventral pectoral region, ventral fore- and hindlimbs and
narrow mid ventral line pale yellow; lateral venter rose pink; base
of tail yellow with orange tint.

Coloration of Females: Dorsum gray-beige with middorsal
brown blotches; dorsal blotches paired, nearly connecting medially;
venter gray-white; ventral thighs with slight yellow tint; chin beige
with gray-brown spots or reticulations.

Distribution. — The species is known in central Peru from the
higher elevations of the Rio Tulumayo in Departamento Junin ( Fig.
3). Additional specimens of this species are known from northern
Peru, Abra Porculla, 2144 m, Departamento Piura. Although both
localities from which the species is known are Amazonian, the
major river valleys of the Rio Maranon and Rio Huallaga are be-
tween them. A similar distribution is exhibited by Stenocercus
hoettgeri Boulenger, 1911.

Ecological Observations. — No other iguanid species is known to

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

occur in sympatry with S. praeornattis. Populations of S. variabilis
Boulenger, 1901, and S. hoettgeri are known from geographically
close areas of Departamento Junin. Stenocercus praeornatus was
collected in large rock piles and in terraces made of rocks. Because
S. variabilis is largely confined to terrestrial habitats and small rocks,
sympatry with S. praeornatus is more probable than with the rock-
dwelling species, S. boettgeri.

Etymology. — The trivial name is from the Latin prae meaning
front or anterior part and ornatus meaning ornate; it refers to the
ornate coloration of the head and shoulders in contrast to that of
the remainder of the body.

Remarks. — Five specimens from Abra Porculla in northern Peru
have fewer scales across the gular region between tympana, 32-35
(33.0+0.632), than topotypic specimens 40-57 (48.2+1.786). The
scales around the body in specimens from the former locality range
from 89 to 112 (97.8±3.219). Females from Abra Porculla lack
dark blotches on the chin. Although differences in scutellation
between the two populations are suggested by the small samples
available, acquisition of more material is necessary to assess geo-
graphic variation. The close similarity in most scutellational aspects,
as well as in coloration, is the basis for assignment of the northern
population to this species.

RESUMEN

Mientras un estudio evolucionario del genero Stenocercus cor-
riente, siete especies nuevas estaban siendo estudiadas. Las especies
nuevas que estan siendo descritas son de zonas poco conocidas en
los Andes Peruanos. Discordancia de dato morfologicos, geograficos,
y ecologicos es la base de la delimitacion taxonomica. El presente
conocimiento necesita consideracion de la posicion generica de
todas las especies en Ophryoessoides y Stenocercus con motivo de
formacion de grupos naturales que conformen con relaciones
evolucionarias. Algunas especies de Ophryoessoides son mas afines
con Stenocercus que otras especies de Ophryoessoides. La variacion
morfologica, distribucion geografica, y preferencias ecologicas son
presentados para cada especie.

SPECIMENS EXAMINED

The following abbreviations are used to indicate the museums
housing specimens examined in this study: AMNH, American Mu-
seum of Natural History; FMNH, Field Museum of Natural History;
KU, University of Kansas Museum of Natural History; MCZ, Mu-
seum of Comparative Zoology; MVZ, Museum of Vertebrate Zool-
ogy— Berkeley; SM, Senckenbergische Natur-Museum; UMMZ,
University of Michigan Museum of Zoology; USNM, United States

NEW SPECIES OF LIZARDS FROM PERtJ 19

National Museum. Specimens are listed alphabetically by species,
country, state and locality.

Stenoceicus apxirimactis
PERU: Apuriniac: Curaluiasi, 2700 in, KU 134240, 134242-134249,
134251-134252; Hacienda Matara, Dist. Matara, MCZ 62253; Puente
Pachachaca, 1800 m, KU 134270-134309; Rio Apurimac, Puente Cunyac, 1830
ni, KU 134235-134239, 134253-134257. Ciizco: Limatambo, 2700 ni, KU
134263-134269; 3 km E Puente Cunyac, 1830 m, KU 134258; 8 km E Puente
Cunyac, 2300 m, KU 134259-134262.

Stenocercus empetrus

PERU: Cajamarca: Banos, MCZ 8084; Cajamaica, 2800 m, FMNH 3941-
3942, 5710-5711, KU 134411-134413; 3 km E Celendin, 2650 m, KU 134414-
134427; Hacienda Huagac, UMMZ 60094-60095. La Libcitad: Huamachuco,
3350 m, KU 134379-134400; Laguna Sacsacocha, 12 km E Huamachuco, 3200
m, KU 134401-134410; Otuzco, FMNH 5708.

Stenocercus ivitus
PERU: Piurci: Summit Cordillera between Canchaque and Huancabamba,
3100 m, KU 134653.

Stenocercus mihicola

PERU: Piura: Summit of Cordillera between Canchaque and Huanca-
bamba, 3100 m, KU 134107-1,34108.

Stenocercus ochoai
PERU: Apurimac: Curahuasi, 2700 ni, KU 134241, 1342,50. Cuzco:
Chilca, 10 km N Ollantaytambo, 2760 m, KU 1,33874-13,3889, 139263;
Chospvoc, Rio Huarocondo, USNM 60705-60706; Ruinas de Machu Picchu,
2400-24,50 m, KU 117108, 1342,33-134234, 1,39267-139268; Ollantaytambo,
USNM 60810, MCZ 41984; Rio Huaracondo, 3048 m, MCZ 12410; Torontoy,
USNM 60811.

Stenocercus orientalis
PERU: Amazonas: Chachapoyas, 2340 m, KU 134447-134473.

Stenocercus praeornatus
PERU: Junin: Comas, ,3220 m, KU 134224-134232. Piura: Abra
Porculla, 2144 m, SM 65252, 65254, 65257-65259, MVZ 82370.

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

LITERATURE CITED

Barbour, T. and G. K. Noble

1920. Amphibians and reptiles from southern Peru collected by the
Peruvian expedition of 1914-19L5 under the auspices of Yale Uni-
versity and the National Geographic Society. Proc. U.S. Natl. Mus.,
58:609-620.

BOULEXGER, G. A.

1885. Catalogue of the lizards in the British Museum (Natural History).
2nd Ed. London, 2:xiii+497 pp.

1899. Descriptions of new reptiles and batrachians collected by Mr. P. O.
Simons in the Andes of Ecuador. Ann. Mag. Nat. Hist., (7)
4:454-457.

1900. Descriptions of new batrachians and reptiles collected by Mr. P. O.
Simons in Peru. IhicL, (7) 6:181-186.

1901. Further descriptions of new reptiles collected by Mr. P. O. Simons
in Peru and Bolivia. Ibid., (7) 7:546-549.

1911. Descriptions of new reptiles from the Andes of South America,
preserved in the British Museum. Ibid., (8) 7:19-25.
Burt, C. E. and M. D. Burt

1930. The South American lizards in the collection of the United States
National Museum. Proc. U.S. Natl. Mus., 78(6): 1-52.
D'Orbigny, a. D.

1837. 350-353, 356-359 pp. In Dumeril, A. M. C. and G. Bibron, 1837.
Erpetologie generale. Vol. 4, 572 pp.
Dumeril, A. M. C. and G. Bibrox

1837. Eiptologie generale. Vol. 4, 572 pp.
Dumeril, M. A.

1851. Catalogue methodique de la collection des reptiles. Museum
D'Histoire Naturelle de Paris. Liv. 1, 128 pp.
Etheridge, R. E.

1970. Stcnocercus. pp. 254-257. In Peters, J. A. and R. Donoso-Barros,
Catalogue of neotropical Squamata: Part II. Lizards and am-
phisbaenians. Bull. Smithsonian Inst., 297:1-293.

GiRARD, C.

1857. Descriptions of some new reptiles collected by the United States
exploring expedition, under the command of Capt. Charles Wilkes,
U.S.N. Fourth part including the species of saurians exotic to
North America. Proc. Acad. Nat. Sci. Philadelphia, 1857:195-198.

GUARDIA MaYORGA, C.

1967. Diccionario Kechwa-Castellano, Castellano-Kechwa. Editora Los
Andes, Lima, Peru. 219 pp.
Mertexs, R.

1952. Amphibien und Reptilien I. pp. 257-266. In Titschack, E. Beitrage
zur Fauna Perus. Verlag Gustav Fischer, Jena. Bd. III.
Noble, G. K.

1924. New lizards from northwestern Peru. Occas. Papers Boston Soc.
Nat. Hist., 5:107-113.
Parker, H. W.

1934. Reptiles and amphibians from southern Ecuador. Ann. Mag. Nat.
Hist., (10) 14:264-273.
SoKAL, R. R. and F. J. Rohlf

1969. Biometry. W. H. Freeman and Company, San Francisco. 776 pp.
Stejxeger, L.

1913. Results of the Yale Peruvian expedition of 1911. Batracians and
reptiles. Proc. U.S. Natl. Mus., 45:541-547.

NEW SPECIES OF LIZARDS FROM PERtJ 21

TSCHUDI, J. J-

1845. Reptiliuni conspectuni quae in republica Peruana reperiuntur et
pleraque obsenata \el collecta sunt in itinera. Arch. Naturg., 11
(1):150-170.
Werner, F.

1901. Ueber Reptilien und Batrachier aus Ecuador und Neu-Guinea.
Verb. Zool.-Bot. GeselL Wien, 51:593-614.