Society for the Study of Amphibians and Reptiles

Two New Sympatric Species of Leaf-Toed Geckos (Gekkonidae: Phyllodactylus) from the Balsas
Region of the Upper Maraon Valley, Peru
Author(s): Pablo J. Venegas, Josiah H. Townsend, Claudia Koch and Wolfgang Bhme
Source: Journal of Herpetology, Vol. 42, No. 2 (Jun., 2008), pp. 386-396
Published by: Society for the Study of Amphibians and Reptiles
Stable URL: http://www.jstor.org/stable/40060525
Accessed: 01-11-2015 09:57 UTC
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Journalof Herpetology,Vol. 42, No. 2, pp. 386-396, 2008

Copyright 2008 Society for the Study of Amphibians and Reptiles

Two New Sympatric Species of Leaf-Toed Geckos (Gekkonidae:

Phyllodactylus) from the Balsas Region of the Upper Maranon
Valley, Peru
Pablo J. Venegas,1 Josiah H. Townsend,2'3Claudia Koch,4 and Wolfgang BOhme4
1Centrode Ornitologiay Biodiversidad(CORBIDI), Santa Rita 117, Huertos de San Antonio, Surco, Lima, Peru;
E-mail:sancarranca@yahoo.es
2Schoolof Natural Resourcesand Environment,and FloridaMuseum of Natural History, University of Florida,
Gainesville,Florida 32611-7800, USA
4ZoologischesForschungsmuseumAlexanderKoenig,Adenauerallee160, D-53113 Bonn, Germany

Abstract.- Two new species of the genus Phyllodactylusare described based on materialcollected in the
southern portion of Departamentode Amazonas, Peru. Both species are well differentiated from all other
South AmericanPhyllodactylusbased on characteristicsof their morphology.In the case of one species, its
large adult size and lack of well-defined rows of strongly keeled scales differentiate it from other
Phyllodactylus,whereas in the second species, the presenceof an enlargedpostanal scale is diagnostic.Both
species were collected in the xeric Balsas region of the upper MaranonValley, and exhibit some similarities
to other assemblages of sympatric Phyllodactylus in South America. The Balsas region is an area of
endemism that warrantsfurtherattention from systematists and conservationbiologists.
- Dos nuevas especies del genero Phyllodactylusson descritasbas&ndoseen materialcolectado
Resumen.
en la porci6nsur del Departamentode Amazonas,Peru.Ambas especies poseen caracteresmorfologicosbien
definidos que los diferencian de las dem4s especies de Phyllodactylus de Sudamerica.Parael caso de la
primera especie, esta se diferencia por su talla adulta y carecer de filas bien definidas de escamas
fuertementekilladas, en comparaci6na otros Phyllodactylus,y en la segunda especie, el caiicter diagnostico
es la presenciade una escamapostanal alargada.Ambas especies fueron colectadasen la aridaareade Balsas
en la cuenca alta del Maran6n,y exhiben cierta similitud a otros grupos de Phyllodactylussimpatricosde
Sudamerica.El reade Balsas,es una zona de endemismos con gran potencial en el estudio de la sistematica
y biologia de la conservaci6n.

The New World Leaf-Toed Geckos, genus

Phyllodactylus
Gray 1828,occuralong the Pacific
coast from southern California,USA, through
Mexico and CentralAmerica to western Panama and from southern Ecuador to northern
Chile, including the Galapagos Islands (Dixon
and Huey, 1970;Bauer et al., 1997).The genus
also has representatives along the Caribbean
coast of the Yucat&nPeninsula, the Islas de la
Bahia of Honduras, the Caribbean coasts of
Colombiaand Venezuela,the OrinocoBasin,the
Caribbeanislands of Aruba,Barbados,Bonaire,
Curacao,Puerto Rico, and Hispaniola,and Isla
Malpelo off the Pacific coast of Colombia
(Dixon, 1964; Dixon and Huey, 1970; Huey,
1975;Baueret al., 1997).Becauseof the form of
the terminallamellaeof theirtoes, characterized
by having the distal phalanges symmetrically
dilated with two ventral terminalplates (Peters
and Donoso-Barros,1970; Rosier, 1995), these
3

Corresponding Author.

reptiles are commonly referredto as the LeafToed Geckos.

In Peru, there are 11 recognized species

of Phyllodactylus: P. angustidigitus, P. clinatus,

P. gerrhopygus, P. inaequalis, P. interandinus, P.
johnwrighti, P. kofordi, P. lepidopygus, P. microphyllus, P. reissi, and P. sentosus (Dixon and

Huey, 1970). The majorityof these species are

distributedalong the arid Pacificcoastal plain,
although P. interandinus,P. johnwrighti,and P.
reissiinhabitdry inter-Andean valleys in northwestern Peru (Dixon and Huey, 1970). In
the interval since Dixon and Huey (1970)
provided the only majortreatmentof the South
American Phyllodactylus,
no new species have
been described in this genus from mainland
South America. Moreover,the complex physiography of the Andes in northern Peru has
limited herpetological collecting in many potentially diverse regions (Duellman and Pramuk, 1999), and some regions have not been
explored at all (Lehr, 2002; Campbell and
Lamar,2004).

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TWO NEW PERUVIAN PHYLLODACTYLUS

387

from Peru.
Table 1. Comparisonof diagnosticcharacteristicsin species of Phyllodactylus
Max SVL
(mm)

P. delsolari

81

P. thompsoni
P. angustidigitus

42
57

P. clinatus
P. gerrhopygus

46
56

P. inaequalis

42

P. interandinus
P. johnwrighti

47
44

Dorsal tubercles

small, round,
smooth
large, trihedral
Dorsal scales
subequal
large, trihedral
Dorsal scales
subequal
small, round,
smooth
large, trihedral
large, trihedral

Number
of 4th toe
lamellae

Terminal
digital pads

Abdominal
plaque

^ 10, irregular

13-16

large

no

> 10
None

11-13
15-20

small
small

no
yes

> 10
None

13-15
12-15

large
small

no
yes

< 10, irregular

10-12

large

no

> 10
> 10

10-12
10-12

large
large

no
no

Rows of
dorsal tubercles

To contribute to the knowledge of the Andean

herpetofauna, two of the authors (PJV and CK)
surveyed the inter-Andean valleys of the upper
Maranon basin in July 2005, and one author (PJV)
returned for a second survey in June 2006 to
collect additional information on the amphibians
and reptiles of this region. The investigations
resulted in the discovery of a comparatively
huge new species of Phyllopezus (Koch et al.,
2006) and two apparently new species of
Phyllodactylus.Simultaneously, JHT was working
in the Herpetology Collection at the Florida
Museum of Natural History (UF), Gainesville,
Florida, and became aware of material possibly
representing two species of undescribed Phyllodactylus from the upper Maranon basin. These
geckos were collected in 1972 by F. G. Thompson,
Curator of Malacology at the Florida Museum of
Natural History and, together with the material
recently collected by PJV and CK, represent two
distinctive new species of Phyllodactylus,which
are described below.
Materials and Methods
All measurements were taken to the nearest
0.1 mm using dial calipers and a stereomicroscope with an optical micrometer; some tail
measurements were taken with a tape measure.
Terminology in the diagnoses and descriptions
follows that used by Dixon and Huey (1970) and
Bauer et al. (1997), with the term "cloacal spurs"
used in favor of "postanal tubercles." Comparative data for other South American species of
Phyllodactyluswere taken from Dixon and Huey
(1970); in addition, we examined specimens of
South American Phyllodactylus housed in the
Museo de Historia Natural San Marcos, Peru
(MUSM), Florida Museum of Natural History,
USA (UF), and Zoologisches Forschungsmuseum Alexander Koenig, Germany (ZFMK).

Diagnostic characteristics for Peruvian Phyllodactylus are compared in Table 1. Measurements and morphological classifications generally follow that used by Dixon (1964).
Measurements of type specimens are always
in millimeters unless otherwise specified, and
measurements are abbreviated as follows:
snout-vent length (SVL), axilla-groin length
(AGL), length of leg (LLE), length of arm
(LAR), length of tail (LTA), length of head
(LHE), depth of head (DHE), width of head
(WHE), length of snout (LSN), diameter of eye
(DEY), diameter of ear (DEA), distance from eye
to ear (DEE). Sex was determined by the
presence of enlarged cloacal spurs and by
probing the base of the tail. For additional
specimens examined, see Appendix 1.
Descriptions of New Species
Phyllodactylus thompsonisp. nov.

Figures 1, 2, 3
Museum of NaturalHistoFlorida
Holotype.
ry (UF) 126943, a female from 7 km east of
Balsas, 1,400 m elevation, Departamento de
Amazonas, Peru (6.847S,77.986W),collected
29 April 1972by F. G. Thompson,original field
number FGT1511.
- Six; two adult males (ZFMK
Paratypes.
84998, 85000) and four adult females (MUSM
19561,19563 [Fig. 1], 19564,ZFMK84999)from
Quebrada Honda in the vicinity of Balsas,
approximately 900-1,000m above sea level,
Provincia de Chachapoyas, Departamento de
Amazonas,Peru (6.817S,78.00W);collectedby
P. J. Venegas and C. Koch on 10 July 2005.
Diagnosis. Of the species of Phyllodactylus

found in mainlandSouth America,P. thompsoni

is the only species with an enlarged postanal
scale; this species can be furtherdifferentiated
from P. angustidigitus, P. gerrhopygus, and P.

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388

P. VENEGAS ET AL.

Fig. 1. Femaleparatypeof Phyllodactylus

thompsonisp. nov. (MUSM19563).
heterurus by lacking an abdominal plaque
anterior to the cloaca (abdominal plaque present
in aforementioned species), from P. inaequalis
and P. microphyllusby having well-defined rows
of enlarged, trihedral tubercles (dorsal tubercles
small, rounded, not arranged in regular rows),
from P. dixoni, P. reissi, P. delsolari, and P.
ventralis by not exceeding 42 mm SVL (those
four species regularly exceed 70 mm SVL), from
P. johnwrighti by having internasals in contact
along the medial suture (internasals usually
separated in P. johnwrighti), from P. sentosus by
having enlarged medial subcaudals (medial
subcaudals not enlarged in P. sentosus), from
P. clinatus, P. interandinus,and P. lepidopygusby
having enlarged trihedral tubercles on the
dorsal surface of tibia (dorsal surface of tibia
with very small granular scales in P. clinatus or
with rounded, elevated scales in P. interandinus
and P. lepidopygus), from P. pumilis by having
homogenous scales on the proximal one-fourth
of the tail (scales heterogeneous on the proximal
one-fourth of the tail in P. pumilis), and from P.
kofordiby lacking rows of enlarged tubercles on
the tail (enlarged tubercles on the tail present in
P. kofordi).
DescriptionofHolotype.- A small female gecko
(33.8 mm SVL) with a broad, depressed head,
1.75 times as long as it is wide; head distinct
from neck; snout with concave granular scales

slightly larger than granular dorsal scales;

temporal and posterior dorsal surface of head
with granular scales interspersed with enlarged,
round, smooth scales; snout length 3.5 mm; 19
scales across the snout at the level of the third
supralabial; rostral twice as wide as it is high,
with a medial groove extending from the dorsal
edge approximately halfway down the rostral;
two internasals; 9/9 supralabials, with supralabials 5-9 entirely below the orbit; six supralabials to a point below the center of eye; eye
large, pupil vertical, orbit width 2.8 mm, 3-4
denticulate scales bordering posterior-dorsal
one- third of orbit; 11 scales from anterior edge
of orbit to nostril; seven granules between
orbits, interorbital distance 0.9 mm; eye to ear
distance 3.0 mm; ear opening oval shaped,
diagonally compressed, inside of ear opening
strongly denticulate on the anterior and posterior edges; ear diameter about one-third the
diameter of the eye; mental larger than infralabials, bell-shaped, 2.0 mm long, 1.1 and
1.7 mm wide at its narrowest and widest points;
infralabials 7/7, with the first pair largest and
each subsequent pair smaller than the last; five
infralabials to a point below the center of eye;
three irregular, enlarged medial chinshields
contact the mental; scales on chin largest
anteriorly, becoming smaller, almost granular
toward the gular region. Body essentially

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TWONEW PERUVIANPHYLLODACTYLUS

Fig. 2. Ventral view of left hind foot of a paratype

of Phyllodactylusthompsonisp. nov. (ZFMK 85000).

cylindrical;dorsum covered with small granules, with eight complete rows of greatly
enlarged, trihedral, posteriorly projecting
keeled dorsal tubercles and at least four more
partial rows of same scales on lateral surfaces
between forelimbsand hind limbs;maximumof
39 enlargedtuberclesper row between the head
and level of the vent; about 108 dorsal granules
between level of the axilla and level of the vent;
about 86 scales around the midbody; lateral
scales enlarged,imbricate,keeled or "pointed,"
projecting outward, becoming smaller and
gradinginto ventralscales;ventralscales larger
than laterals, smooth, imbricate, becoming
larger posteriorly, 44 ventral scales along
midventral line between axilla and cloaca;
abdominal plaque anterior to cloaca absent;
enlarged plate present posterior to cloaca and
anteriorto base of tail, 1 mm long by 2.2 mm
wide, with the anterioredge continuous with
the posterior interior surface of the cloaca;
enlarged,narrow scale present borderinganterior edge of cloaca; leading edges of fore and
hind limbs covered with enlarged, imbricate
scales, largest on the forearms and thighs;
dorsal surfaces of tibia with enlarged trihedral
tubercles;fourthtoe with 11 subdigitallamellae,
terminal lamellae enlarged, longer than wide,
with the tip of the claw primarily concealed.
Tailcomplete,0.82 times the length of the body,
stout, tapering distally; dorsal and lateral
surfaces covered with keeled, imbricatescales
that are largest at the base and becoming
gradually smaller distally; medial subcaudals

389

enlarged, somewhat irregular,platelike, imbricate, more than twice as wide as long.

- Snout-vent length,
Measurements
ofHolotype.
33.8 mm; tail length, 28.1 mm; head length,
10.9 mm; and head width, 6.2 mm.
- Coloration of the
Color (in Preservative).
after
35
holotype,
yr preserved in formalinand
stored in ethanol, is as follows: dorsal ground
color of head and body pale brown; head and
snout with faint pale brown mottling; brown
line visible from posterioredge of orbitrunning
along lateral surface of body, breakingup and
becoming diffuse posteriorly; dorsal surfaces
with irregular,brown mottling;exceeding small
dark brown spots are scattered across most
scales of the body, most profuse laterally;
ventral scales on head and body immaculate
or marked with one or two microscopic dark
brown spots; tail pale brown with three pairs
of brown spots, becoming paler and closer
togetherdistally.
Colorin Life.- Colorationin life of an adult
female paratype (Fig. 1; MUSM 19563) is as
follows: ground color pinkish-tan with brown
indistinct thin bars reaching to the middle of
dorsum, tail with seven brownish bands and
yellowish-browninterspaces,bands a bit smaller than interspaces, head dirty yellowishbrown, dark brown line from nostril through
eye to above arminsertion,venterwhite, ventral
surfaceof tail dirty whitish.
- Male paratypes range from 40Variation.
42 mm SVL, and female paratypes range from
36-41 mm SVL.Tail length ranges from 42-54%
of total length. Postmentals vary from two to
three, always contacting first infralabial,often
contactingsecond infralabial;number of scales
immediatelyfollowing postmentalsvaries from
5-7. Interorbitalscales range from 20-26, scales
across snout at level of third labialsfrom 17-21.
Usually two internasalsborderedby five scales,
only one specimen with four internasals bordered by four scales; internasals always in
contact.Scales from nostril to eye number from
10-13. Twenty-five to 29 scales across venter,
from gular region to vent from 57-68; dorsal
tubercularrows always 10;paravertebraltubercles from head to base of tail vary from 36-40,
with 20-24 between axilla and groin; enlarged
rows of tuberclesreachingbase of tail number6
in five specimens and 4 in one specimen.
Tubercles absent from tail. Lamellae beneath
fourthtoe 12 or 13 in all paratypes.Dorsalcolor
patternwith double row of indistinct8-10 spots
or transversebars or lines, dorsal surfaceof tail
usually banded with 4-9 brownish bands, with
ground color interspacesof almost equal width;
bands not present on regeneratedtails and only
very weak developed in some individuals.
Ventralcolorationdirty white in all specimens.

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390

P. VENEGAS ET AL.

Fig.3. Close-up of dorsal scalationin a paratypeof Phyllodactylus

thompsonisp. nov. (UF 126943).
Etymology.- The specific name thompsoni is a
patronym honoring F. G. Thompson, collector of
the holotype of this new species. Thompson has
served as a curator of Malacology at the Florida
Museum of Natural History since 1966, during
which time he has conducted biological surveys
in 18 countries worldwide that have led him to
describe two subfamilies, 35 genera, and over
200 species of mollusks, as well as new species
of Thamnophis, Xenosaurus, and Leptotyphlops.
Thompson's collections have also led to the
description of many other amphibian and
reptile taxa, including Eleutherodactylusaraiodactylus Duellman and Pramuk 1999 and Phrynopus
thompsoni Duellman 2000, which are only
known from material collected by F. G. Thompson in Peru in May 1972.
Distribution and Natural History.- Phyllodactylus thompsoniis known only from the mountains
east of Balsas and the Rio Maranon in vicinity of
the type locality, from 900-1,400 m elevation on
the western slope of the northern portion of the
Cordillera Central (see Duellman and Pramuk,
1999). The vicinity of the type locality receives
less than 500 mm of rain annually (Duellman
and Pramuk, 1999), and the vegetation is
characterized as dry deciduous tropical forest
and thorn scrub forest with a short, sparse
overstory that includes the plants Acacia macracantha, Aeschynomene scoparia, Caesalpinia sp.
nov., Capparissp., Coursetia cajamarcana,Croton
sp., Dalea exilis, Dalea carthagenensisvar. brevis,
Eriothecasp., Leucaena trichodes,Maraniona lavi-

nii, Mimosa incarum,Mimosa ctenodes,Parkinsonia

praecox, Pithecellobium excelsum, and Tecoma
rosifolia (Hughes et al., 2004). This habitat is
within the Ecuatorial Dry Forest ecoregion
(Brack, 1986).
Paratypes collected in 2005 and 2006 were
also found at night, either perched on low rocks
or actively moving on the ground. Air temperature during these nights averaged 27.3C;
substrate temperature of the rocks averaged
28.8C.
Phyllodactylus delsolari sp. nov.
Figures 4, 5, 6
Museum of Natural HistoFlorida
Holotype.
an
adult
female from 7 km east of
50059,
(UF)
ry
Balsas, 1,400 m elevation, Departamento de
Amazonas, Peru (6.847S, 77.986W), collected
29 April 1972 by F. G. Thompson, original field
number FGT 1508.
Paratypes.- -UF 50060, 50062 adult males, UF
50061 juvenile male, collected with the holotype
on 29 April 1972, by F. G. Thompson; MUSM
26327, adult male and MUSM 26328-26330,
adult females, from Quebrada Honda in the
vicinity of Balsas (0649'S, 7800'W, approximately 900-1,000 m above sea level), Provincia
de Chachapoyas, Departamento de Amazonas,
Peru, collected by P. J. Venegas on 18 June 2006;
MUSM 19566, 26410 and ZFMK 85002 adult
males, MUSM 26411, ZFMK 85001, 85003 adult
females from Quebrada Honda; collected by P.
J. Venegas and C. Koch on 10 July 2005.

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TWONEW PERUVIANPHYLLODACTYLUS

Fig. 4.

391

Male paratype of Phyllodactylus delsolari sp. nov. (MUSM 26410).

Fig. 5. Ventral view of left hind foot of a paratype

of Phyllodactylusdelsolari sp. nov. (ZFMK 85001).

- Of the Phyllodactylusfound in
Diagnosis.
western South America, P. delsolariis one of
two "giant" species, the other being P. reissi.
BothP. delsolariand P. reissiexceed 70 mm SVL,
with no other species from the region exceeding
60 mm SVL. Phyllodactylusdelsolari can be
readily distinguished from P. reissi by having
fewer than 10 poorly defined rows of small,
smooth, round tubercles (12-18 well-defined
rows of enlarged, trihedral, strongly keeled
tubercles in P. reissi), and by having broad,
well-defined dark dorsal cross-bands(no crossbands or sometimes incomplete,dark,irregular,
narrow cross-bandsin P. reissi).There are two
recorded
other species of "giant"Phyllodactylus
from mainland South America:P. dixoniand P.
ventralis. Both of these species may exceed
70 mm SVLand are found in centralVenezuela
and along the Caribbeancoast of Colombiaand
Venezuela, respectively. Phyllodactylusdelsolari
can be distinguished from P. dixoni and P.
ventralisby having poorly defined, irregular
rows of small, smooth, round tubercles (welldefined, regular rows of enlarged, trihedral,
strongly keeled tubercles in P. dixoni and P.
ventralis).This species can be furtherdifferentifound in South
ated from all otherPhyllodactylus

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392

P. VENEGASET AL.

Fig. 6. Close-up of dorsal scalation of the holotype of Phyllodactylusdelsolarisp. nov. (UF 50059).

Americaas follows (Table1): from P. angustidiand P. heterurusby lacking

gitus, P. gerrhopygus,
an abdominal plaque (abdominal plaque present in aforementionedspecies); from P. clinP. johnwrighti,P. kofordi,P.
atus, P. interandinus,
P. pumilis,and P. sentosusby having
lepidopygus,
fewer than 10 poorly defined rows of enlarged,
smooth, round tubercles (14-22 rows of en10larged trihedraltuberclesin P. interandinus,
16 rows of enlargedkeeled or trihedraltubercles
in the other aforementionedspecies); from P.
microphyllus
by having large terminal lamellae
and
(small terminallamellae in P. microphyllus);
fromP. inaequalis
by having ear denticulationon
the anteriorand posterioredges (ear denticulation absent in P. inaequalis).There is one other
species of South American Phyllodactylusthat
has broad, dark dorsal cross-bands,P. transversalis,which is endemic to Isla de Malpelo,but it
lacks ear denticulation (inside of ear strongly
denticulateon the anteriorand posterioredges
in P. delsolari)and is not reported to exceed
57 mm SVL.
- A large female gecko
Descriptionof Holotype.
mm
with
a
broad, depressed head,
(73.2
SVL)
1.3 times longer than it is wide; head distinct
from neck, covered in small granules that are
largest on snout and smallest on back of head;
snout granules concave, larger than dorsal
granules, snout length 7.6 mm; 22 scales across
the snout at the level of the third supralabial;
rostral broad, rectangular, lacking a medial
groove; two internasals; supralabials 10/10,
with supralabials6-10 entirely below the orbit;
seven supralabialsto a point below the centerof
eye; eyes large, pupil vertical, orbital width

5.3 mm, three rows of small, somewhat trihedral supraciliaryscales bordering the anterior
edge; one row of enlarged platelike scales
bordering the eye, and 2-3 rows of small
granules between platelike scales and upper
edge of orbit; 13 scales from anterioredge of
orbit to nostril; 18 granules between orbits,
interorbitaldistance2.6 mm; eye to ear distance
6.7 mm; ear opening oval shaped, vertically
compressed,1.5 mm high by 1 mm wide, inside
of ear opening strongly denticulateon anterior
and posterior edges; mental larger than infralabials, bell-shaped, 3.7 mm long, 1.7 and
3.4 mm wide and its narrowest and widest
points; infralabials9/9, first pair largest and
each subsequentpair smaller;six infralabialsto
a point below the center of eye; one pair of
enlarged medial chinshields contacts the mental; scales on chin largest where they contact
infralabialsand chinshields, becoming smaller,
almost granular toward the midventral line.
Body depressed; dorsal surface of neck with
small granules that are somewhat smaller than
those on the head; dorsum covered with small
granules, with about nine poorly defined
longitudinal rows of enlarged, smooth scales
that are 2-3 dorsal granules in width (Fig. 2); a
maximum of 28 enlarged scales per row
between level of the axilla and level of the vent;
enlarged rows most evident on posterior onethird of the dorsum, where enlarged scales are
also interspersed with granules; about 134
dorsal granules between level of the axilla and
level of the vent; about 115 scales around the
midbody; ventral surface with smooth, platelike, subimbricatescales that graduallybecome

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TWO NEW PERUVIAN PHYLLODACTYLUS

larger posteriorly; 51 ventral scales along
midventral line between axilla and vent; preanal plate absent; leading edges of fore and hind
limbs covered with enlarged, platelike scales
that are largest on the forearms and shins;
fourth toe with 13 subdigital lamellae, terminal
lamellae greatly enlarged, truncate, with the tip
of the claw exposed. Tail complete, 1.26 times
the length of the body; dorsal surface of tail
covered with squarish, platelike scales, without
whorls of enlarged scales; medial subcaudal
scales broadly expanded, two or more times as
wide as they are long, larger than any other
scales found on the body.
MeasurementsofHolotype.- Snout-vent length,
73.2 mm; tail length, 92.4 mm; head length,
21.4 mm; and head width, 16.5 mm.
Color (in Preservative).- Coloration of holotype after 35 yr preserved in formalin and
stored in ethanol, is as follows: dorsal ground
color of head and body pale brown, with the
snout and supralabials heavily mottled brown
and irregular brown flecks and blotches on the
dorsal surface of the head; dorsum with broad
brown bands, which are 1-2 times as wide as
the pale intervening areas; dorsal bands interrupted by a narrow pale brown middorsal
stripe; lateral surfaces of the body yellowishcream with pale brown mottling, lateral scales
marked with minute dark brown spots; dorsal
surfaces of limbs tan with brown flecks and
mottling; ventral surface of head and body
immaculate cream; dorsal surface of tail with
about five poorly defined brown bands, subcaudal surface immaculate cream.
Color in Life.- Coloration of an adult male
paratype (MUSM 26327) is as follows: ground
color grayish-white, dorsal surface from rear of
head to vent with 5-6 broad brown transversal
bands, which are cut vertically by a slender
grayish vertebral stripe with the margin of each
band undulated (Fig. 4). Head is grayishbrown, limbs with brown flecks and blotches
(Fig. 4), and the venter white.
Variation.- Male paratypes range from 54.580 mm SVL, and female paratypes range from
47.5-81 mm SVL. In two adult male paratypes,
UF 50062 possesses 3/3 cloacal spurs, and UF
50060 has 4/4 cloacal spurs. All paratypes have
a single pair of postmental chinshields, and
usually a single pair of internasals in broad
contact. Supralabials 11/10 in UF 50660 (7/8 to
a point below the center of eye), 10/11 in UF
50062 (8/8 to a point below the center of eye),
and 11/10 in UF 50061 (8/8 to a point below the
center of eye). Infralabials 7/9 in UF 50060 (6/7
to a point below the center of eye), 8/8 in UF
50061 (6/6 to a point below the center of eye),
and 9/9 in UF 50062 (6/6 to a point below the
center of eye). Among all paratypes, there are

393

11-14 scales between anterior edge of orbit and

nostril; dorsal tubercles intermixed between
granules, only sometimes forming short indistinct rows ranging from 6-12. All male and
female paratypes lack precloacal or femoral
pores. The color pattern exhibited in the
paratypes is very similar to that of the holotype,
with broad transversal bands from rear of head
to vent numbering five or six, with 6-8 dark
bands on tail. One juvenile male (UF 50061)
differs from all other specimens in lacking a
thin, pale middorsal stripe interrupting the
transverse dorsal bands.
Etymology.- The specific name is a patronym
honoring Gustavo del Solar, in recognition of his
continued and unattenuated work in the conservation of the White Winged Guan, Penelope
albipennis, a critically endangered species of
Cracid bird that is endemic to northwestern Peru.
Distribution and Natural History.- The known
geographic and ecological distribution of P.
delsolarimirrors that of P. thompsoni(see species
account above). The holotype and paratypes of
P. delsolaricollected in 1972 were found between
2100 and 2230 h on a damp, west-facing
exposed limestone wall from 3-5 m above the
ground. Paratypes collected in 2005 and 2006
were also found at night, usually perched
between 30 cm and 3 m above the ground on
tall, nearly vertical boulders in Quebrada
Honda, a tributary of Rio Maranon. Air temperature during these nights averaged 27.3C;
substrate temperature of the rocks averaged
28.8C. Two individuals of P. delsolari were
observed running on the ground a few meters
away from verticals rock walls. This new
species appeared abundant, along with the
sympatric giant gecko Phyllopezus maranjonensis
(Koch et al., 2006), in the zones where the ravine
is very narrow and forms a small canyon
surrounded by completely vertical rocky walls.
Along with P. maranjonensis, the congener
Phyllodactylus reissi is also sympatric with P.
delsolari and P. thompsoni in the Balsas region,
giving that area a unique assemblage of
sympatric geckos (especially giant geckos)
notable among Neotropical herpetofaunas.
Three female paratypes of P. delsolari
were gravid. Two of these females (MUSM
26328, 26330) contained two oviductal eggs with
the following sizes: 3.8 X 3 mm and 3.6 X
2.7 mm; 8.5 X 6.9 mm and 8.6 X 7.3 mm,
respectively; the other female (MUSM 26328)
contained a single oviductal egg measuring 5.5
X 5 mm.
Discussion
The New World Phyllodactylus remains a
relatively little studied yet significant group of

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394

P. VENEGASET AL.

gekkonids, and virtually no systematic work nodes, Parkinsoniaperuviana,two undescribed

has appeared since Dixon and Huey (1970) species of Caesalpinia,and the endemic genus
presented their exhaustive review of the South and species Maranionalavinii (Hughes et al.,
American taxa. In that work, all revisionary 2003, 2004;Hughes, 2005). Many of these taxa,
systematics were based on comparative mor- as well as endemic species from other plant
phological data, and no attempt was made to families,exhibithighly localizeddistributionsin
resolve higher-level relationships within the the Balsasarea, some of which (like Maraniona)
South American taxa. Bauer et al. (1997) are restrictedto a narrow elevational belt that
restricted the genus Phyllodactylusto New also encompassesthe known distributionof the
World taxa, naming five new genera and two new Phyllodactylus.
The MaranonValley is also an area of bird
retrieving a sixth from synonymy to represent
monophyletic clades of geckos from Africa, endemism, with 22 species restricted to the
Asia, and Europethat were previously referred valley, and Balsasand its immediatesurroundto as Phyllodactylus.
As indicatedby Baueret al. ings up to 2,900 m elevation is considered an
(1997), the monophyly of the New World area of importancefor endemic bird conservaPhyllodactylushas not been tested, and phylo- tion (Frankeet al., 2005). This region and the
genetic relationships among species of Phyllo- surrounding highlands is potentially a center
for land snail endemism, with some endemics
dactylusremain unknown.
The P. gerrhopygusgroup is defined by the already described (Thompson, 1982) and over
presence of abdominal plaques in the group's 70 new species awaiting description (F. G.
three constituent species: P. angustidigitus,P. Thompson, pers. comm.), all of which were
gerrhopygus,and P. heterurus.An abdominal collected on the same trip that produced the
plaque (referredto as "preanalplate" by Dixon type material of Phyllodactylusdelsolariand
and Huey, 1970) is a novel sensory organ Phyllodactylusthompsoni,as well as that of E.
resembling an enlarged, roundly triangular araiodactylus(Duellman and Pramuk, 1999),
plate or scale located anterior to the cloacal Phrynopus spectabilis (Duellman, 2000; later
opening on the ventral surface and is most synonymized with Pleurodemamarmorataby
(Duelllikely a heat sensing organ used to actively Lehr,2006),and Phyllodactylus
thompsoni
sample substratetemperature(Krolland Dixon, man, 2000).
Eleven of the 15 species of Phyllodactylus
1972). Members of the P. gerrhopygusgroup
inhabitsome of the driestand coldest habitatsof reviewed or described by Dixon and Huey
any South American species of Phyllodactylus, (1970) are sympatric with at least one other
making the evolution of the heat-sensing ab- memberof the genus. Thereare four species of
dominalplaque potentiallyadvantageous(Kroll sympatric geckos in the Balsas region: Phylloand Dixon, 1972).Phyllodactylus
thompsonipos- dactylusdelsolari,Phyllodactylus
thompsoni,Phylsesses an enlarged scalelike plate or pad lodactylusreissi, and Phyllopezusmaranjonensis
extending from the posterior inside edge of (Koch et al., 2006). In their discussion of
the cloacal lip posteriorly onto the body. The sympatry in Phyllodactylus,Dixon and Huey
extremely dry habitat in the upper Maranon (1970)noted two generalcommonalitiesamong
first, sympatric
Valley, not unlike that of the arid Andean sympatricsets of Phyllodactylus:
foothills inhabitedby P. gerrhopygus,
may have species typically exhibit a wide divergence in
fueled evolution in P. thompsoniof an organ body size, often typified by the presence of one
homologous to the abdominal plate in the P. relativelylarge and one relativelysmall species;
gerrhopygusgroup. Externalexaminationof the second, sympatricspecies usually demonstrate
postcloacalplate does not allow for determina- clear differences in their respective degrees of
tion of whether this plate is a sensory plaque arboreality.Although P. delsolariand P. thomplike those found in the P. gerrhopygusgroup, soniobviously conformto the first of Dixon and
some other sort of sensory organ, or perhaps Huey's (1970) characterizations,the ecological
simply cloacal ornamentation.
relationships among the two giant species of
After being overlooked by biologists until Phyllodactylus
(P. delsolariand P. reissi)and the
relativelyrecently,the upper MaranonValley in even larger sympatric gecko Phyllopezusmarthe area of Balsasis beginning to be recognized anjonensisremains unclear. Clearly, focused
as an important center for endemism within examinationof the unique gekkonidassemblage
the MaranonValley, particularlyamong plants in the Balsas region is warranted,particularly
(Bregman, 1996; Bridgewater et al., 2003; given the apparent ecological overlap among
Hughes et al., 2004; Pendry, 2004; Hughes, the three species of giant geckos.
In lower parts of the Maranonbasin around
2005). The vicinity of the type locality for P.
delsolariand P. thompsonialso encompasses the 120-150 km north of Balsas, in the Huancadistributionof the endemic legume (Legumino- bamba Depression near Bagua and the confluMimosacte- ence of the Maranon and rios Chinchipe and
sae) species Coursetiacajamarcana,

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TWONEW PERUVIANPHYLLODACTYLUS
Utcubamba,the giant species P. reissi is syman
patricwith the small species P. interandinus,
endemic to the Bagua region. The HuancabambaDepressionis both an importanthotspot
for endemism across a variety of taxonomic
groups (Cadle, 1991;Duellman and Wild, 1993;
Reeder, 1996; Duellman and Pramuk, 1999;
Sancho, 2004) and one of the most significant
biogeographic barriers in the entire Andes,
separatingthe northernand southern cordilleras of the Andes and formingthe southernedge
of the distributionfor taxa found to the north
and the northernedge for species found in the
southernCordilleras(Duellman,1979;Weigend,
2002). Suites of plant genera found in both the
Bagua and Balsas regions of the Maranon
Valley, such as the legumes Mimosa and
Coursetia,each contain species endemic to that
respective region (Weigend, 2002; Hughes,
2005), a pattern not unlike that seen in Phyllodactylus.Evidence from the fossil record indicates that some dry forest-inhabiting plant
species in northwestern Peru have remained
remarkablyconserved in terms of morphology
for at least 10 million years, indicatingthat dry
forests in this region have been present continuously for the intervening period and became
graduallymore fragmented,shapingthe current
phytogeographyof the region (Burnham,1995;
Burnhamand Barranco,2004;Hughes, 2005).
Despite the identification of a number of
locally endemic species in the Balsas area, this
region remains poorly known and undersampled biologically. Even taxonomic groups
that are relatively well studied in this region,
such as plants, have thus far only received
limited attention from systematists, and P.
delsolariand P. thompsoniare the second and
third endemic reptile species to be described
from the immediate vicinity of Balsas (Koch et
al., 2006). The potential for discovery of additional endemic, undescribed species in the
Balsas region is high, and a concertedeffort to
characterizethe composition and conservation
status of the herpetofaunaof the upper Maranon Valleyshould be undertakenat the earliest
possible opportunity.
- JHT is grateful to F. G.
Acknowledgments.
for
Thompson bringinghis attentionto the type
series of P. delsolari,for supporting efforts to
describe these taxa, and for providing comments on an earlierversion of this manuscript.J.
D. Austin and L. D. Wilson also provided
helpful commentson a draftof this manuscript.
J. Cordovaand C. Aguilar grantedaccess to the
collectionsof the Museo de HistoriaNaturalde
la UniversidadNacional Mayorde San Marcos,
Lima,Peru, and facilitatedthe loan of material.
M. A. Nickerson, F. W. King, and K. Krysko

395

provided access to the UF herpetology collection and database.While preparingthis manuscript, JHT was supported by a Tropical
Conservationand Development Graduate Fellowship, a University of Florida Foundation
GrinterFellowship,and grants from the Reptile
and Amphibian Conservation Corps (RACC).
PV is indebted to C. Elerafor allowing access to
the laboratoryof the Museo de Sican.CKthanks
the Deutscher Akademischer Austauschdienst
(DAAD) for financial support and A. Schliiter,
StaatlichesMuseum fur Naturkunde,Stuttgart,
for sharing his experience with Peruvian authorities.
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Appendix 1
Additional SpecimensExamined
Phyllodactylus kofordi.-FERU: Piura: UF 34719,
34722-34733, 34745-34749.
Phyllodactylus microphyllus.-PERU: Piura: UF
34718, 34720-34721, 34738, 34743, 126944-126956.
Phyllodactylus reissi.- ECUADOR: Esmeraldas: UF
71934, Guayas: UF 39458-39459, 71502-71503, 9062O90631, 99334, 126957-126958, PERU: Cajamarca: UF
39454-39457, Piura: UF 34717, 34734-34737, 3473934742, 34744, 126959-126967.
Phyllodactylus ventralis- VENEZUELA: Falcon: UF
40739^0741, Guarico: UF 40709-40710, 40729^0737,
60946, Portuguese UF 40671^0708, 40742^0743.

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