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Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [Special Section]: 1-11 (e96).

A new species of Andean lizard Proctoporus (Squamata:

Gymnophthalmidae) from montane forest of the Historic

Sanctuary of Machu Picchu, Peru

^uis Mamani, 2 Noemi Goicoechea, and 3 Juan C. Chaparro

1 ? Museo cle Historki Natural, Universidad Nacional de San Antonio Abad del Cusco, Plaza de Armas s/n (Paraninfo Universitario), Cusco, PERU

2 Department of Biodiversity and Evolutionary Biology, Museo Nacional cle Ciencias Naturales-CSIC, C/ Jose Gutierrez Abascal 2, 28006 Madrid,

SPAIN

Abstract .— We describe a new species of lizard assigned to the genus Proctoporus from the Historic

Sanctuary of Machu Picchu in the Department of Cusco (southeastern Peru) where it inhabits a

montane forest region at an elevation between 2,760-2,800 m. The new species is distinguishable

from all other species of Proctoporus by a unique combination of morphometric, scalation, and

color pattern characteristics.

Resumen . — Describimos una nueva especie de lagartija asignada al genero Proctoporus, proveniente

del Santuario Historico de Machu Picchu en el Departamento del Cusco (Sureste de Peru), habita la

region de bosques montanos entre los 2,760-2,800 m de altitud. La nueva especie se distingue de

todas las demas especies de Proctoporus por la combinacion unica de caracteres morfometricos,

escamacion y caracteristicas en los patrones de coloracion.

Key words. Oriental Cordillera, Cusco, Peru, South America, Andean lizard, Proctoporus, Natural Protected Area,

Cercosaurinae

Palabras clave. Cordillera Oriental, Cusco, Peru, America del Sur, Lagartija andina, Proctoporus, Area Natural Pro-

tegida, Cercosaurinae

Citation: Mamani L, Goicoechea N, Chaparro JC. 201 5. A new species of Andean lizard Proctoporus (Squamata: Gymnophthalmidae) from montane

forest of the Historic Sanctuary of Machu Picchu, Peru. Amphibian & Reptile Conservation 9(1) [Special Section]: 1-11 (e96).

NoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation', official journal website <amphibian-

reptile-conservation. org> .

Received: 11 February 2015; Accepted: 27 April 2015; Published: 30 April 2015

Introduction

Gymnophthalmid lizards of the genus Proctoporus in-

cludes eleven species that occur in central and southern

Peru, Bolivia, and northern Argentina, and an addition-

al two unnamed species known from Peru (Doan et al.

2005; Goicoechea et al. 2012). These small, semi-fosso-

rial lizards occur in habitats characterized by cloud for-

est, steppes, cacti, shrubs, and wet puna habitats along

the eastern slopes of the central Andes (Doan and Cas-

toe 2003; Doan et al. 2005). The highest diversity of the

genus Proctoporus occurs in Peru, which includes ten

species: P. bolivianus Werner 1910; P. carabaya; P. iri-

descens; P. kiziriani Goicoechea, Padial, Chaparro, Cas-

tro viejo -Fisher, and De la Riva 2013; P. chasqui (Chavez

et al. 2011); P. guentheri (Boettger 1891); P. lacertus

(Stejneger 1913); P. pachyurus Tschudi 1845; P. sucul-

lucu Doan and Castoe 2003; and P. unsaacae Doan and

Castoe 2003. Three species, P. guentheri, P. bolivianus,

and P. xestus (Uzzell 1969), occur in Bolivia and one,

P. xestus, reaches northern Argentina and probably Peru

(Goicoechea et al. 2013). Taxonomic works published

in this century include revisions (e.g., Doan and Castoe

2003; Doan et al. 2005; Goicoechea et al. 2012; Goicoe-

chea et al. 2013) and the description of several new spe-

cies (Doan et al. 2005; Chavez et al. 2011; Goicoechea

et al. 2013). However, the actual diversity of this genus

is far from known well, and new species continue to be

Correspondence. Email: l luismamanic@ gmail.com (Corresponding author); 2 n.goicoechear@ gmail.conr, ) jchaparroauza@yahoo.com

April 2015 | Volume 9 | Number 1 | e96

Amphib. Reptile Conserv.

Mamani et al.

found as herpetological surveys are carried out in previ-

ously unexplored or poorly known areas.

Recent biological exploration in the southern Peru-

vian Cordillera Oriental of the Andes has revealed the

existence of a new species of gymnophthalmid lizard in

the montane forest region within the national protected

area of the Historic Sanctuary of Machu Picchu. The spe-

cies is described herein and assigned to the genus Proc-

toporus.

(Table 1) used in previous studies on gymnophthalmid

systematics were examined for 120 specimens (Ap-

pendix I). Character definition and usage follow Uzzell

(1970) and Doan and Castoe (2003). Drawings were

elaborate using a stereo microscope with camera lucida..

Coloration in life is based on the field notes and photog-

raphy by LM. Geographic coordinates were taken using

a global positioning system (GPS) device and geodetic

datum WGS 84.

Materials and Methods

Specimens were collected by hand, euthanized with

Halatal, fixed in 10% formalin, and later transferred to

70% ethanol for long-term museum storage. The speci-

mens were deposited at the Museo de Historia Natural

de la Universidad Nacional de San Antonio Abad de

Cusco (MHNC) in Peru. Morphological data were ob-

tained from preserved specimens of all known species

of Proctoporus. Because only two specimens (one adult

male and one juvenile) of Proctoporus cliasqui , were ex-

amined, we used data from Chavez et al. (2011). Twenty-

three qualitative and meristic morphological characters

Table 1 . Measurements (mm) of three specimens of Proctoporus machupicchu sp. nov. and the addition of Fig. 3 G, specimen of

subadult male not collected.

MHNC13373

MHNC13362

MHNC11815

Not collected (Fig. 3 G)

Characters (measurements mm)

Subadult male

Adult female

Subadult male

Snout- vent length

20.80

41.20

46.70

< 28,80

Tail length

32.30

60.82

61.40

—

Head length

8.50

10.70

11.00

—

Head width

4.80

5.80

5.70

—

Femoral pores

Supralabials

6-7 (left-right)

Loreal scale

PRESENT

Supraoculars

Genials

Postparietals (Occipitals)

Temporals

Scales around midbody

—

Transversal dorsal scale rows

—

Transversal ventral scale rows

Longitudinal dorsal scale rows

Longitudinal ventral scale rows

Lamellae under 4th finger

—

Lamellae under 4th toe

—

Postoculars

Superciliaries

Frontal

1.50

2.10

2.00

—

Frontonasal

1.50

1.95

2.10

—

Head length/Head width

1.77

1.84

1.93

—

Tail length/SVL

1.12

1.48

1.31

—

Frontal/frontonasal proportion

1.00

1.08

0.95

—

Amphib. Reptile Conserv.

April 2015 |

Volume 9 | Number 1 | e96

Results

Proctoporus machupicchu sp. nov.

urn:lsid:zoobank.org:act:216381E4-4C4B-4C3C-99AE-0DCEFEC45352

Figures 1-3.

Proposed standard English name:

Machu Picchu Andean Lizard

Proposed standard Spanish name:

Lagartija Andina de Machu Picchu

A new species of Andean lizard Proctoporus

Holotype: (Fig. 1; 2 A-C; 3 A-B), adult female, MHNC

13362 (field number LM 834), Peru, Department of Cus-

co, Province Urubamba, District Machu Picchu, from

Aobamba (13° 14' 17" S; 72° 33' 15" W), 2,760 m, col-

lected by Luis Mamani, Frank P. Condori, and Juan C.

Chaparro on 16 June 2013.

Paratypes: MHNC 13373, field number LM 845, (Fig.

3 E-F), immature male, same data as holotype; MHNC

13513, adult female (field number LM 637, Fig. 2 D-F;

3 C-D), Peru, Department of Cusco, Province Urubam-

ba, District Machu Picchu, from Winaywayna (13° IF

33.72” S; 72° 32’ 18.66” W), 2,800 m, collected by Luis

Mamani, Kateryne Pino, Alexander Pari, Andres Garcia,

and Gerardo Ceballos on 11 September 2012.

Diagnosis: (1) Frontonasal length equal to the frontal

length; (2) nasoloreal suture present in all specimens;

(3) Loreal scale present, not in contact with supralabi-

als; (4) supraoculars three; (5) superciliaries four, first

not expanded onto the dorsal surface of the head; (6)

postoculars two; (7) palpebral disc made up of a single,

undivided scale; (8) four supralabials anterior to the

posteroventral angle of the subocular; (9) three pairs of

genials in medial contact; (10) dorsal body scales quad-

rangular, keeled; (11) transverse rows of dorsals 38-39;

(12) transverse ventral rows 21; (13) a continuous se-

ries of small lateral scales separating dorsals from ven-

trals; (14) posterior cloacal plate made up of six scales

in both sexes; (15) anterior preanal plate scales paired;

(16) femoral pores present or not in males, when is pres-

ent six per hind limb (Fig. 3 G), absent in females; (17)

preanal pores absent; (18) subdigital lamellae on toe IV

16-17; subdigital lamellae on finger V 10-11; (19) limbs

overlapping when adpressed against body in adults; (20)

limbs pentadactyl, digits clawed; (21) dorsal and lateral

surfaces of head dark brown; lip irregularly yellow or or-

ange-cream stripes; ventral surface of head and pregular

region cream or orange, with or without irregular black

blotches; venter black or dark gray with cream or cream-

orange spots on the posterior margin of some scales, in

male juveniles is orange with black blotches.

All specimens of Proctoporus machupicchu have an

undivided palpebral eye disc, a putative synapomorphy

of the genus Proctoporus (Doan and Castoe 2005; Uzzell

1970). Proctoporus machupicchu can be distinguished

from all other species of the genus, except for P. iride-

scens, by the presence of three pairs of genials in me-

dial contact (two in all other species of Proctoporus ). It

can be distinguished from P. iridescens by having four

supralabials anterior to the posteroventral angle of the

subocular, by the presence of a loreal scale and a nasolo-

real suture (three supralabias, loreal scale, and nasoloreal

suture absent in P. iridescens ). It can further be differenti-

ated from P. pachyurus by having three supraoculars not

fused with superciliaries (four supraoculars in P pachy-

urus, first fused with first superciliary), and 38-39 trans-

verse dorsal scale rows (47-60 in P. pachyurus); from P.

sucullucu by having a frontonasal scale equal in length to

the frontal scale (frontonasal scale longer than the fron-

tal scale in P. sucullucu), and loreal scale not in contact

with the supralabials (in contact in P. sucullucu); from

Fig. 1. Holotype of Proctoporus machupicchu (MHNC 13362; SVL41.2 mm).

Amphib. Reptile Conserv. 3 April 2015 | Volume 9 | Number 1 | e96

Mamani et al.

Fig. 2. (A, B, C) Head of the holotype of Proctoporus machupicchu (MHNC 13362), lateral, dorsal, and ventral view; and (D, E,

F) Paratype (MHNC 13373) lateral, dorsal, and ventral view of the head. Scale bar 5 mm.

P. bolivianus by having frontonasal length equal to the

frontal length (frontonasal longer than frontal scale in P.

bolivianus ); first superciliary not fused with first supra-

ocular (fused in P. bolivianus ); from P unsaacae and P

guentheri by the absence of a series of continuous lateral

ocelli, loreal scale not in contact with supralabials, and

the absence of a pair of enlarged pregular scales in con-

tact (present in P. unsaacae and P. guentheri, loreal scale

in contact with supralabials in P. unsaacae and a pair of

enlarged pregular scales in medial contact in P. sucul-

lucu ); from P. carabaya and P. kiziriani by having a first

supraocular not fused with the first superciliary, (fused in

P carabaya and P. kiziriani) and limbs overlapping when

adpressed against body (not overlapping in P. carabaya

and P kiziriani)', from P lacertus by having first supra-

ocular not fused with the first superciliary (fused in P.

lacertus), and the presence of a loreal scale (absent in P.

lacertus); from P. xestus by the lack of prefrontal scales

(present in P. xestus) and the existence of keeled dorsal

scales (smooth in P xestus); and from P. chasqui by the

lack of prefrontal scales (present in P. chasqui), supraoc-

ulars three (four in P. chasqui ), and femoral pores absent

in females (present in females of P. chasqui).

Description of holotype: Adult female, snout-vent

length (SVL) 41.2 mm, tail length 60.8 mm; head scales

smooth, rounded in dorsal and lateral view, without stria-

tions or rugosities; rostral scale wider (1.9 mm) than tall

(0.9 mm), meeting the supralabials on either side at the

top of the supralabials, becoming higher medially, in

contact with frontonasal, nasals, and first supralabials;

frontonasal longer than wide, equal in length with frontal,

widest posteriorly, in contact with rostral, nasals, anterior

most supraocular, and frontal; prefrontals absent; frontal

longer than wide, roughly polygonal, not in contact with

superciliaries, in contact with frontonasal, first two su-

praoculars, and frontoparietals; frontoparietals polygonal

(right scale divided on the right anterior side), in contact

with frontal, second and third supraoculars, parietals, and

interparietal; supraoculars three, middle scale divided on

the posterior corner (in contact with frontoparietals on

both sides), all in contact with superciliaries, third in

contact with frontoparietal, parietal, and postocular; in-

terparietal longer than wide, polygonal, in contact with

frontoparietals anteriorly, with parietals laterally, and

with occipitals (or postparietals) posteriorly; parietals

polygonal, lateral suture in contact with temporals and

April 201 5 | Volume 9 | Number 1

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e96

A new species of Andean lizard Proctoporus

Figure 3. Dorsal and ventral views of living specimens of Proctoporus machupicchu. A-B adult female (MHNC 13362); C-D adult

female (MHNC 13513); and E-F inmature male (MHNC 13373); and G not collected of immature male showing femoral pores.

April 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

e96

V_jft

Mamani et al.

TBtDTCTW TD’QtrW

TS'CftTW 7VWW

Legend

Elevation

5700 - 6500

5000 - 5700

4300 - 5000

3600 - 4300

2800 - 3600

2100-2800

1400-2100

700-1400

0-700

Figure 4. Map showing the distribution of Proctoporus species known from southeast of Peru, based on species listed in Appendix I

and in Uzzell (1970), Doan and Castoe (2003), Doan et al. (2005), Chavez et al. (2011), and Goicoechea et al. (2013). Green circle,

Proctoporus machupicchu sp. nov.; blue triangle, P. bolivianus\ blue square, P. carabaya; red square, P. chasquv, blue circle, P. iri-

descens; blue pentagon, P. kizirianv, red circle, P. lacertus; red triangle, P. unsaacae; and red pentagon P. sucullucu.

postoculars, diagonally with temporals, posteriorly with

occipitals, anteriorly with third supraoculars and fronto-

parietals; three occipitals, smaller than parietals, medial

pentagonal, smaller than the laterals. Nasal divided, lon-

ger than high, in contact with first and second supralabi-

als; loreal present, not in contact with the supralabials, in

contact with nasal, first superciliary, and frenocular; four

superciliaries, first not fused with the first supraocular;

two preoculars, upper in contact with the first supercili-

ary and loreal scales, lower in contact with frenocular,

and first subocular; frenocular roughly pentagonal, in

contact with the second and third supralabials, lower

preoculars, first subocular, and loreal scales; palpebral

disc made up of a single transparent scale; three subocu-

lars; two postoculars; temporals smooth, polygonal; four

supralabials anterior to the posteroventral angle of the

suboculars. Mental wider (1.9 mm) than long (1.05 mm),

in contact with the first infralabial and postmental pos-

teriorly; postmental single, pentagonal, in contact with

the first infralabials and the first pair of genials; three

pairs of genials in medial contact, anterior pair in con-

tact with the first and second infralabials on the right side

and in contact with the second on the left side; second

pair of genials in contact with the second and third in-

fralabials; third pair of genials in contact with the third

and fourth infralabials laterally; one pair of chin shields,

separated by four smaller median pregulars; eight gular

scale rows; small lateral neck scales, round and smooth.

Dorsal scales rectangular, longer than wide, juxtaposed,

slightly keeled, in thirty-nine transverse rows; twenty-

three longitudinal dorsal scale rows at midbody; continu-

ous lateral scale series, smaller than dorsals, and partially

hidden in lateral fold; reduced scales at limb insertion re-

gions present; twenty-two transverse ventral scale rows;

ten longitudinal ventral scale rows at midbody; anterior

preanal plate scales paired; six posterior preanal plate

scales, lateralmost scales small; scales on the tail rectan-

gular (fewer square), juxtaposed; dorsal and dorsolateral

caudal scales slightly keeled anteriorly, smooth poste-

riorly; ventrolateral caudal scales smooth; midventral

subcaudal scales wider than the adjacent scales, almost

square, anteriormost midventral subcaudal scales subim-

bricate. Limbs pentadactyl; digits clawed; dorsal brachial

scales polygonal, subequal in size, subimbricate, smooth;

roundish ventral brachial scales, subimbricate, smooth;

dorsal antebrachial scales polygonal, subequal in size,

smooth; ventral antebrachial scales polygonal, smaller

than dorsals; dorsal manus scales polygonal, smooth,

subimbricate and arranged in three rows; palmar scales

small, rounded, and juxtaposed, domelike; dorsal scales

April 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

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A new species of Andean lizard Proctoporus

Figure 5. Type locality of Proctoporus machupicchu: (A, C) Montane forest, (B) Urubamba River, (D) Habitat of Proctoporus

machupicchu. Photo: (6A-C) Luis Mamani; 6 D ( Javier Farfan).

on fingers smooth, quadrangular, covering dorsal half of

digit, and overhanging subdigital scales, two on finger

I, four on II, six on III, six on IV, and four on V; scales

on anterodorsal surface of thigh polygonal, smooth,

subimbricate; scales on posterior surface of thigh small,

rounded, and juxtaposed; scales on ventral surface of

thigh small, enlarged, and smooth; femoral pores ab-

sent; preanal pores absent; scales on anterior surface of

crus polygonal, smooth, juxtaposed, decreasing in size

distally; scales on anterodorsal surface of crus rounded,

juxtaposed; scales on ventral surface of crus polygonal,

enlarged, smooth, flat, and subimbricate; scales on dor-

sal surface of toes polygonal, smooth; overhanging su-

pradigital lamellae, two on toe I, five on II, nine on III,

twelve on IV, seven on V; subdigital lamellae single, four

on toe I, eight on II, eleven on III, sixteen on IV, ten on

V; limbs overlapping when adpressed against the body.

Coloration in preservative: Dorsal and lateral surfaces

of head dark brown; ventral surface of head cream with

clusters of light and dark brown, and scales with black

spots inside. Gular region similar to the head, the mac-

ules in anterior side are light brown and diffuse, on pos-

terior side are thick. Lip irregularly barred with cream

coloring. Dorsal surface of the trunk same color as head.

Lateral surface of trunk of the same coloration as dor-

sum, fading to paler brown near venter. Ventral surface

of the trunk black with cream spots at posterior margin

of each scale. Color of limbs similar to body. Dorsal tail

coloration like that of body; ventral surface of tail dark

brown with cream spots.

Coloration in life: The coloration is similar to that in

preservative, but with orange spots along the ventral sur-

face of the body.

Variation: Scalation and morphometries of the paratypes

are similar to the holotype (Table 1). The coloration in

females is variable, the ventral surface of the head and

gular region are orange and pale yellow with brown and

black spots. In the sub-adult male the coloration in the

ventral surface of the head and gular region is an intense

orange and extends posteriorly to the ventral surface of

the trunk.

Etymology: The specific epithet is an indeclinable word

that refers to the distribution of the new species in the

Natural Protected Area of the Historic Sanctuary of Ma-

Amphib. Reptile Conserv.

April 201 5 | Volume 9 | Number 1 | e96

Mamani et al.

chu Picchu, in the Cordillera of Vilcanota, one of the

most important formations in the Andes of southern Peru.

Distribution: Proctoporus machupicchu is known only

from Aobamba (type locality), and Winaywayna, both

inside the Historic Sanctuary of Machu Picchu between

2,760-2,800 m (Fig. 4). With the addition of the new

species, the genus Proctoporus contains 12 species from

Peru; six of them ( Proctoporus guentheri, P. kiziriani , P.

lacertus, P. machupicchu sp. nov., P unsaacae, P. sucul-

lucu ) located in the Department of Cusco.

Habitat and ecology: Individuals were found during

the day under rocks in the montane forest, of the eastern

slope, of the Cordillera Oriental of the Andes (Fig. 5).

Conservation: The status of this species is unknown.

More herpetological surveys and population studies are

needed to adequately assess its status.

Discussion

Similarities in morphology and coloration would place P.

machupicchu closer to P. guentheri and P. unsaacae , but

further incorporation of DNA sequences and morpho-

logical data should provide a better resolution to the po-

sition of this new species within Proctoporus. Although

the description of P. machupicchu represents an increase

in the species richness within Proctoporus , the knowl-

edge of the actual species diversity of the genus is still

limited (Doan and Castoe 2003; Goicoechea et al. 2012)

and some taxonomic problems still remain to be solved.

Some authors (Goicoechea et al. 2012; Chavez et al.

2011; Kohler and Lehr 2004) have related Peruvian spe-

cies of Euspondylus (E. caideni Kohler, E. josyi Kohler,

E. nellycarrillae Kohler and Lehr, E. oreades Chavez,

Siu-Ting, Duran, and Venegas, E. rahmi (De Grijs), E.

simonsii Boulenger, and E. spinalis (Boulenger) with

Proctoporus. These species are found along central

and southern Peru, overlapping with the distribution of

Proctoporus and share with Proctoporus several derived

features including the presence of an undivided palpe-

bral eye disc. Recently, Goicoechea et al. (2012) found

molecular evidence to place a species of Euspondylus ,

E. chasqui within Proctoporus, nevertheless, the phy-

logenetic relationships of the remaining species of Pe-

ruvian Euspondilus and Proctoporus remains uncertain.

On the other hand, an additional species of Proctoporus,

P. cephalolineatus, presumably exist in Venezuela. This

species was previously described as belonging to the

Proctoporus luctuosus group (Garcfa-Perez and Yustiz,

1995). Nevertheless, because the holotype and unique

specimen of this species has limbs that do not overlap

when adpressed, Doan and Schargel (2003) removed P

cephalolineatus from the P. luctuosus group and related

Revised key to the genus Proctoporus

Key to the Species of Proctoporus from Peru, Bolivia, and Argentina (adapted from Goicoechea, Padial, Chaparro,

Castro viej o-Fisher, and De la Riva 2013)

la. Presence of prefrontals 2

lb. Absence of prefrontal scales 3

2a. Smooth dorsal scales, single large elongate subocular, presence of large spines at the base of the sulcus spermaticus

P. xestus

2b. Keeled dorsal scales, several small subocular scales P. chasqui

3a. Two pair of genial in contact 5

3b. Three pair of genial in contact 4

4a. Three supralabials anterior to the posteroventral angle of the subocular P. iridescens

4b. Four supralabials anterior to the posteroventral angle of the subocular P. machupicchu

5a. Two to three supraoculars 6

5b. Four supraoculars P. pachyurus

6a. Venter uniformly dark or with dark stippling or mottling near lateral scale rows 7

6b. Venter clear yellow or orange without dark mottling P. guentheri

7a. No continuous series of lateral ocelli 8

7b. Continuous series of lateral ocelli P. unsaacae

8a. Frontonasal scale longer than frontal scale 9

8b. Frontonasal scale equal in length to frontal scale 10

9a. Limbs overlapping when adpressed P. sucullucu

9b. Limbs not overlapping when adpressed P. bolivianus

10a. First supraocular not fused with first superciliary 11

10b. First supraocular fused with first superciliary P. carabaya

11a. Absence of loreal scale P. lacertus

lib. Presence of loreal scale P. kiziriani

April 201 5 | Volume 9 | Number 1

Amphib. Reptile Conserv.

e96

A new species of Andean lizard Proctoporus

this species with Eusponclylus and Pholidobolus based

on the presence of prefontals in P. cephalolineatus (a

character presumed not to be present in Proctoporus at

this time). This species shows the presence of palpe-

bral eye-disc divided vertically (J.E. Garcia-Perez, pers.

comm.). As the presence of an undivided palpebral eye

disc is a diagnostic character for the genus Proctoporus

(Doan and Castoe 2005; Goicoechea et al. 2012; 2013)

we believe that this species does not belong to this genus.

Further studies based on molecular and morphological

data are necessary to cast some light on these topics, as

well as on the relationships of P. machupicchu with other

species in the genus.

Acknowledgments. — For allowing access to collec-

tions under their care, we are grateful to Percy Yanque

and Rocio Orellana (MHNC), Carl J. Franklin (UTA),

Robert P. Reynolds (USNM), and Darrel Frost ( AMNH).

We are indebted to the Curso Fatinoamericano de Bi-

ologia de la Conservacion and the Asociacion para la

Conservacion de la Cuenca Amazonica (ACC A) for sup-

port fieldwork. We are grateful to the Center for Conser-

vation Education and Sustainability of the Smithsonian

Conservation Biology Institute for providing laboratory

equipment and to the rangers of the Historic Sanctuary of

Machu Picchu for their invaluable contribution to field

work. We would like to acknowledge Jose A. Ochoa for

providing camera lucida in composing the ink pen illus-

trations in Fig. 2 A-F. We thank David Kizirian, Claudia

Koch, Karen Wilke, and anonymous reviewers for their

valuable comments to the manuscript, Javier Farfan for

providing the pictures in Fig. 4, and Juan Elias Garcia-

Perez (MZ-UNEFFEZ) for sharing data of the holotype

(UFABG 3202) Proctoporus cephalolineatus. Noerrn

Goicoechea was supported by a FPI fellowship from the

Spain Ministry of Science and Innovation. Collecting

permits in Pern were issued by SERNANP-Machu Pic-

chu (054-20 1 2-SERNANP- JEF) .

Literature Cited

Boettger O. 1891. Reptilien und Batrachier aus Bolivien.

Zoologischer Anzeiger 14: 343-347.

Boulenger GA. 1885. Catalogue of the Lizards in the

British Museum ( Natural History ) /-///. Fondon,

United Kingdom.

Chavez G, Siu-Ting K, Duran V, Venegas PJ. 2011. Two

new species of Andean gymnophthalmid lizards of the

genus Euspondylus (Reptilia, Squamata) from central

and southern Peru. Zoo Keys 109: 1-17.

Doan TM, Castoe TA. 2003. Using morphological and

molecular evidence to infer species boundaries within

Amphib. Reptile Conserv. 9

Proctoporus bolivianus Werner (Squamata: Gymn-

ophthalmidae). Herpetologica 59: 433-450.

Doan TM, Schargel WE. 2003. Bridging the gap in Proc-

toporus distribution: A new species (Squamata: Gym-

nophthalmidae) from the Andes of Venezuela. Herpe-

tologica 59(1): 68-75.

Doan TM, Castoe TA. 2005. Phylogenetic taxonomy of

the Cercosaurini (Squamata: Gymnophthalmidae),

with new genera for species of Neusticurus and Proc-

toporus. Zoological Journal of the Linnean Society

145: 403-416.

Doan TM, Castoe TA, Arizabal- Arriaga W. 2005. Phylo-

genetic relationships of the genus Proctoporus sensu

stricto (Squamata: Gymnophthalmidae), with a new

species from Puno, southeastern Peru. Herpetologica

61:325-336.

Garcia-Perez JE, Yustiz EE. 1995. Una nueva especie de

Proctoporus (Sauria: Gymnophthalmidae) de los An-

des de Venezuela. Revista de Ecologi'a Latinoameri-

cana 4: 1-5

Goicoechea N, Padial JM, Chaparro JC, Castroviejo-

Fisher S, De la Riva I. 2012. Molecular phylogenetics,

species delimitation and biogeography of the Andean

lizards of the genus Proctoporus (Squamata: Gymn-

ophthalmidae). Molecular Phylogenetics and Evolu-

tion 65: 953-964.

Goicoechea N, Padial JM, Chaparro JC, Castroviejo-

Fisher S, De la Riva, I. 2013. A taxonomic revision

of Proctoporus bolivianus Werner (Squamata: Gymn-

ophthalmidae) with the description of three new spe-

cies and resurrection of Proctoporus lacertus Stejneg-

er. American Museum Novitates 3786: 1-32.

Kizirian DA. 1996. A review of Ecuadorian Proctoporus

(Squamata: Gymnophthalmidae) with descriptions of

nine new species. Herpetological Monographs 10:

85-155.

Kohler G. Felir E. 2004. Comments on Euspondylus and

Proctoporus (Squamata: Gymnophthalmidae) from

Peru, with the description of three new species and a

key to the Peruvian species. Herpetologica 60: 101—

118.

Stejneger F. 1913. Results of the Yale Peruvian Expedi-

tion of 1911. Batrachians and Reptiles. Proceedings

of the United States National Museum 45: 541-547.

Tschudi JJ. 1845. Reptilium conspectum quae in repub-

lica Peruana reperiuntur er pleraque observata vel col-

lecta sunt in itenere. Archiv fiir Naturgeschichte 11(1):

150-170.

Uzzell TM. 1969. A new genus and species of Teiid liz-

ard from Bolivia. Postilla 129: 1-15.

Uzzell TM. 1970. Teiid lizards of the genus Proctoporus

from Bolivia and Peru. Postilla 142: 1-39.

April 201 5 | Volume 9 | Number 1 | e96

Mamani et al.

Appendix 1

Specimens Examined. Museum acronyms refer to: AMNH, American Museum of Natural History, New York, USA; CBF, Colec-

cion Boliviana de Fauna, Fa Paz, Bolivia; MHNC, Museo de Historia Natural, Cusco, Peru; MNCN, Museo Nacional de Ciencias

Naturales, Madrid, Spain; USNM, Smithsonian Institution, National Museum of Natural History, Washington, USA; UTA, Univer-

sity of Texas, Arlington, USA. Numbers in brackets represents the original field numbers.

Proctoporus bolivianus

PERU: Puno: Sandia (UTA 52946-47 [TMD 01267, TMD 01271]); Cuyo-Cuyo (MHNC5333 [MNCN 4532]. MHNC 5348-49

[MNCN 4566, MNCN 4568], MNCN 43660-62 [MNCN 4531, MNCN 4534, MNCN 4567]); Patambuco (MHNC 5357 [MNCN

5357], MNCN 43663-64 [MNCN 4583, MNCN 4584]); BOFIVIA: Fa Paz: Pelechuco (MNCN 43655-56 [MNCN 4143, MNCN

4142]); Millipalla, 12 km S of Sorata (CBF 3437-39 [MNCN 4729, MNCN 4731, MNCN 4733], MNCN 43678-79 [MNCN 4730,

MNCN 4732]); Charazani (CBF 2329 [MNCN 4159]); Caalaya (CBF 2330 [MNCN 4162]).

Proctoporus chasqui

PERU: Ayacucho: Road between Abra Tapuna and San Francisco (MNCN 44407-08 [MNCN 4830, MNCN 4831]).

Proctoporus carabaya

Pent: Puno: Carabaya (MHNC 5428 [holotype: MNCN 4709], MHNC 5429-3 l[paratypes: MNC 4710, 4714, 4715]).

Proctoporus guentheri

PERU: Cusco: Urubamba (UTA 55366-67 [TMD 01322, TMD 01324]); Paucartambo (USNM 346179 [USNM 206266]); BO-

FIVIA: Fa Paz: Apolo (USNM 336148 [USNM 107286]).

Proctoporus iridescens

Peru: Puno: Sandia: Fimbani (MHNC 5359 [holotype: MNCN 4590]), MHNC 5361[paratype: MNCN 4593]); Puno: Carabaya

(MHNC 5699, MHNC 5701 [paratypes: MNCN 4793, 4795]).

Proctoporus kiziriani

Pern: Cusco: Quispicanchi (MHNC 5366 [holotype: MNCN 4602]), MHNC 5680, MHNC 5682-83, MHNC 5685 [paratypes:

MNCN 4750, 4751-4752, 4754]).

Proctoporus lacertus

PERU: Fa Convencion: Tincochaca (USNM 49551 [holotype], 49552 [paratype]); Calca (UTA 55315-23 [TMD 01301, TMD

01307, TMD 01309, TMD 01310, TMD 01311, TMD 01312, TMD 01313, TMD 01316, TMD 01317], USNM 298685-90 [JEC

6264, JEC 6265, JEC 6266, JEC 6267, JEC 6268, JEC 6269]); Ollantaytambo (USNM 49549 [paratype], USNM 107649); Nusta

Hispana (USNM 60699); Torontoy (USNM 60726); Paucartambo (AMNH 142921 [AMNH11568)].

Proctoporus pachyurus

PERU: Junrn: Cerro San Cristobal (MHNC 4693-94 [TA504, TA505], MHNC 4696 [TA507]); Tarma (UTA 55267-72 [TMD

01211, 01213, TMD 01214, TMD 01215, TMD 01216, TMD 01220], UTA 55314 [TMD 01195]); Palca (USNM 299581-82 [JEC

7092, JEC 7093]).

Proctoporus sucullucu

PERU: Apurfmac: Abancay (UTA 52950 [TMD 01146], UTA 55273-78 [TMD 01140, TMD 01141, TMD 01143, TMD 01144,

TMD 01157, TMD 01159]); Cusco: Quillabamba (USNM 298632-33 [JEC 6093, JEC 6094]); Puno: Ollachea (USNM 299125-27

[JEC 6591, JEC 6592, JEC 6593]); Ayacucho, Anco (MNCN 44474-82 [MNCN 5012, MNCN 5013, MNCN 5014, MNCN 5015,

MNCN 5016, MNCN 5017, MNCN 5018, MNCN 5019, MNCN 5020]).

Proctoporus unsaacae

PERU: Urubamba (UTA 55289-90 [TMD 01031, TMD 01032], UTA 55291-92 [TMD 01033, TMD 01035], UTA 55294-95 [TMD

01094, TMD 1037, TMD 01094]).

Proctoporus xestus

BOLIVIA: (AMNH 22740-41); Cochabamba (AMNH 38957-62).

Amphib. Reptile Conserv.

April 2015 | Volume 9 | Number 1 | e96

A new species of Andean lizard Proctoporus

Luis Mamani graduated in Biological Sciences from Universidad Nacional de San Antonio Abad del Cusco in

Pern. He received a bachelor degree in biology in 2013. He is currently an amphibian and reptile researcher at the

Herpetological Collection of the Museum of Natural History, Universidad Nacional de San Antonio Abad del Cusco,

Peru (MHNC). He has published a scientific paper in amphibian taxonomy and his interests include biodiversity,

taxonomy, systematics, and biogeography of gymnophthalmid lizards.

■ Noeim Goicoechea earned her B.S. in Biology from Universidad Autonoma de Madrid, Spain, in 2007 and her

M.S. in biodiversity and evolutionary biology from the same university in 2011. She is currently a Ph.D. student at

the Museo Nacional de Ciencias Naturales (Madrid, Spain), where she is studying species diversity and diversifica-

tion mechanisms in Andean lizards of the genera Proctoporus and Liolaemus within a phylogenetic and population

genetic framework.

Juan Carlos Chaparro graduated in Biological Sciences from Universidad Nacional Pedro Ruiz Gallo, Lam-

bay eque, Peru. He received a Master’s degree in Biodiversity in Tropical Areas and Conservation in 2013, from

a consortium between the International University of Menendez Pelayo (UIMP-Spain), Universidad Tecnologica

Indoamerica (UTI-Ecuador), and Consejo Superior de Investigaciones Cientificas (CSIC-Spain), under the supervi-

sion of Dr. Ignacio de la Riva. He is currently curator and researcher of the Herpetological Collection of the Museum

of Natural History, Universidad Nacional de San Antonio Abad del Cusco, Peru (MHNC). He has published more

than 20 scientific papers, and two book chapters on taxonomy, biodiversity, systematics, phylogeny, conservation,

and biogeography of South American amphibian and reptiles, with an emphasis on amphibians. He is interested in

biodiversity, conservation, taxonomy, systematics, phylogeny, and biogeography of the neotropical herpetofauna.

In accordance with the International Code of Zoological Nomenclature new rules and regulations (ICZN 2012), we have deposited this paper

in publicly accessible institutional libraries. The new species described herein has been registered in ZooBank (Polaszek 2005a, b), the of-

ficial online registration system for the ICZN. The ZooBank publication LSID (Life Science Identifier) for the new species described here

can be viewed through any standard web browser by appending the LSID to the prefix “http://zoobank.org/.” The LSID for this publication

is: um:lsid:zoobank.org:pub:ADEEE69A-964E-491F-93A8-EA7F4FE5303D.

Separate print-only edition of paper(s) (reprint) are available upon request as a print-on-demand service. Please inquire by sending a request

to: Amphibian & Reptile Conservation , amphibian-reptile-conservation.org, arc.publisher@gmail.com.

Amphibian & Reptile Conservation is a Content Partner with the Encyclopedia of Life (EOL), http:///www.eol.org/ and submits information

about new species to the EOL freely.

Digital archiving of this paper are found at the following institutions: ZenScientist, http://www.zenscientist.com/index.php/filedrawer; Ernst

Mayr Library, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (USA), http://library.mcz.harvard.edu/

emst_mayr/Ejournals/ARCns.

The most complete journal archiving and journal information is found at the official ARC journal website, amphibian-reptile-conservation,

org. In addition, complete journal paper archiving is found at: ZenScientist, http://www.zenscientist.com/index.php/filedrawer.

Citations

ICZN. 2012. Amendment of Articles 8,9,10,21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods

of publication. Zootaxa 3450: 1-7.

Polaszek A et al. 2005a. Commentary: A universal register for animal names. Nature 437: 477 .

Polaszek A et al. 2005b. ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological

Nomenclature 62(4): 210-220.

April 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

e96

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [Special Section]: 12-14 (e99).

SHORT COMMUNICATION

Noblella lynchi Duellman 1991 (Anura: Craugastoridae):

Geographic range extension, Peru

Daniel Rodriguez

Universidad National Mayor de San Marcos, Museo cle Historia Natural, Departamento cle Herpetoiogia. Av. Arenales 1256, Jesus Maria, Ap.

14-0434, Lima, PERU

Abstract . — Reported is a significant range extension of the Leaf Litter Frog, Noblella lynchi, an

endemic species to the Cordillera Central in northern Peru. The new record is in the Huiquilla Private

Conservation Area.

Key words. Leaf Litter Frog, Huiquilla Private Conservation Area, Cordillera Central, Abra Chanchillo, district of

Longuita, province of Luya, department of Amazonas, montane cloud forest

Citation: Rodriguez D. 2015. Noblella lynchi Duellman 1991 (Anura: Craugastoridae): Geographic range extension, Peru. Amphibian & Reptile Con-

servation 9(1) [Special Section]: 12-14 (e99).

NoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation', official journal website <amphibian-

reptile-conservation. org> .

Received: 17 March 2015; Accepted: 08 May 2015; Published: 30 June 2015

Noblella lynchi (Duellman 1991) is a leaf litter frog cat-

egorized as Data Deficient (DD) in the IUCN Red List,

because little is known about the extent of its occur-

rence and ecological requirements (Amphibian Special-

ist Group 2013). The holotype of Noblella lynchi (KU

212318, female, SVL 20.2 mm) and the paratype (KU

212319, subadult female, SVL 16.2 mm) were collected

on the slopes of Abra Chanchillo (06°49’ S, 77°54’ W,

elevation 2,870 m asl), 42 km (by road) ENE of Balsas,

province of Chachapoyas, department of Amazonas,

Peru, obtained on 22 January 1989 by John J. Wiens (Du-

ellman 1991).

During the course of fieldwork in the project “Es-

tado de Conservacion de anuros que habitan la Cuenca

del Rio Tingo en el Departamento de Amazonas” (Di-

rectorial Resolution N°033-2006-INRENA-IFFS-DCB),

three specimens of Noblella lynchi were collected in a

montane forest (06°23’10” S; 77°59’10.7” W, 2,700 m

asl) in the Huiquilla Private Conservation Area (ACP

Huiquilla), located in the district of Longuita, province

of Luya, department of Amazonas by Daniel Rodriguez

on 17 June 2006 (Fig. 1). The collected specimens were

the following: MUSM 24885 is a male, SVL 15.3 mm;

MUSM 24886 is a juvenile, SVL 7.8 mm, and MUSM

26448 is a female, SVL 19.5 mm) (Fig. 2).

Correspondence. Email: 1 daal911@yahoo.com

The holotype and paratype of Noblella lynchi (KU

212318-19) were found at 2,870 m asl. Both specimens

were under stones, during the day in cutover cloud for-

est. The type locality is along the road between Balsas

and Leimebamba (Duellman 1991). The new specimens

(MUSM 24885-86, 26448) were found at 2,700 m asl in

leaf litter during the night in montane forest of high and

low canopy (Young and Leon 1999). These specimens

represent a northward extension of 49.1 kilometers and

the first record for the province of Luya.

Currently, the species has some degree of protec-

tion, because it occurs in the ACP Huiquilla, which is

an important fragment of the montane cloud forest in the

northern part of the Cordillera Central.

Abbreviations: KU = Biodiversity Institute, University

of Kansas, Lawrence, Kansas, USA; MUSM = Museo

de Historia Natual, Universidad Nacional Mayor de San

Marcos, Lima, Peru; SVL = snout-vent length; m = me-

ters; asl = above sea level.

Acknowledgments — I thank Jesus H. Cordova

(MUSM) for access to specimens. For their helpful com-

ments on the manuscript, I thank William E. Duellman

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e99

Rodriguez

and Vanessa Correa. I also would like to thank Miguel

Chocce for helping with the map and to Jose G. la Torre

Montoya for access to ACP Huiquilla. Work was sup-

ported by the Asociacion Peruana para la Conservacion

de la Naturaleza (APECO) and their research fund “Ma-

ria Koepcke.”

Literature Cited

Amphibian Specialist Group 2013. Noblella lynchi. In:

IUCN 2014. IUCN Red List of Threatened Species.

Version 2014.3. Available: http://www.iucnredlist.

org/ [Accessed: 16 March 2015].

Duellman W. 1991. A new species of Leptodactylid frog,

genus Phyllonastes, from Peru. Herpetologica 47(1):

9-13.

Young KR, Leon B. 1999. Peru’s humid eastern montane

forests: An overview of their physical settings, bio-

logical diversity, human use and settlement, and con-

servation needs. DIVA, Technical Report no 5: 1-97.

Fig. 1 . Noblella lynchi (A and B: male, MUSM 28216, SVL 24.8 mm; C and D: young, MUSM 24886, SVL 7.8 mm; E and F:

female, MUSM 26448, SVL 19.5 mm). Photos by D. Rodriguez.

Amphib. Reptile Conserv.

June 201 5 I Volume 9 I Number 1 I e99

Range extension of Noblella lynchi

CHACHAPOYAS

LUYA

RODRIGUEZ DE MENDO

ENDI

MARISCAL CACERES

Fig. 2. Map showing distribution of Noblella lynchi in the Cordillera Central in northern Peru.

7B : 20'D n W

7fl’0 ™

50'0'W

77 , 4DB"W

77 , 30 , 0"W

Legend

Collection Localities

A New record

^ Type locality

ACP Huiquilla

| | Provincial Border

Regions \ Border

1:500,000

—

7B*20'0"W

/no'trw

5trC"W

77 : 30^w

Daniel Rodriguez is a Peruvian biologist involved in the conservation of natural areas, amphibians, and reptiles. He

is a researcher within the herpetology department at the Natural History Museum of San Marcos, Peru.

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e99

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [Special Section]: 15-25 (el 05).

Two new species of frogs of the genus Phrynopus (Anura:

Terrarana: Craugastoridae) from the Peruvian Andes

German Chavez, 2 Roy Santa-Cruz, 3 Daniel Rodriguez, and 4 Edgar Lehr

l Division de Herpetologla-Centro de Ornitologia y Biodiversidad (CORBIDI), Santa Rita N°105 Of. 202, Urb. Huertos de San Antonio, Surco,

Lima, PERU 2 Museo de Historia Natural Universidad Nacional San Agustin, Arequipa, PERU 3 Departamento de Herpetologia, Museo de Historia

Natural, Universidad Nacional Mayor de San Marcos. Av. Arenales 1256, Lince, Lima 14, PERU ^Department of Biology, Illinois Wesleyan

University, Bloomington, Illinois 61 701, USA

Abstract. — We describe two new species of Phrynopus from the western and eastern Andes of

northern and central Peru. One of them occurs in the Andean highlands of La Libertad region and is

described from 20 specimens. This species can be differentiated from other Phrynopus species that

lack a tympanum by the following combination of characters: skin of dorsum shagreen with scattered

low tubercles; skin of venter smooth; no tubercles on upper eyelids; dentigerous process of vomers

absent; vocal slits and nuptial pads absent; finger I slightly shorter than finger II; toe V slightly

longer than toe III; and maximum SVL of females is 31.2 mm. The other species is described from

three individuals found under moss-covered floors in the cloud forest of the Cordillera de Carpish,

Huanuco region. It can be easily distinguished from other Phrynopus species by its vermilion red

coloration on dorsum and venter.

Key words. Central Peru, Cordillera de Carpish, La Libertad, Huanuco, taxonomy

Resumen. — Describimos dos nuevas especies de Phrynopus de los Andes occidentales y de los

Andes orientales del centro de Peru. Una de ellases una especie de las zonas altoandinas de la

region La Libertad, que puede ser diferenciada de las otras especies de Phrynopus que no tienen

timpano por una combinacion unica de caracteres que consiste en: piel del dorso granulada con

presencia de tuberculos bajos desordenados; piel del vientre lisa; ausencia de tuberculos sobre los

parpados; procesos vomerianos de los dientes ausentes; machos sin sacos vocales ni excrescencias

nupciales; dedo I de la mano ligeramente mas corto que el dedo II; dedo V del pie ligeramente mas

largo que el dedo III; longitud hocico-cloaca de hembras alcanza los 31.2 mm. La otra especie es

descrita de tres individuos encontrados bajo el musgo del suelo en los bosques nublados de la

Cordillera de Carpish, en la region de Huanuco que puede ser facilmente diferenciada de las demas

especies de Phrynopus por su coloracion rojo bermellon, tanto en el dorso como en el vientre.

Palabras clave. Centro de Peru, Cordillera de Carpish La Libertad, Huanuco, taxononha

Citation: Chavez G, Santa-Cruz R, Rodriguez D, Lehr E. 201 5. Two new species of frogs of the genus Phrynopus (Anura: Terrarana: Craugastoridae)

from the Peruvian Andes. Amphibian & Reptile Conservation 9(1 ) [Special Section]: 1 5-25 (el 05).

NoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation, official journal website <amphibian-

reptile-conservation. org> .

Received: 30 May 2015; Accepted: 09 October 2015; Published: 18 October 2015

Introduction

Systematics of the frog genus Phrynopus were until re-

cently poorly known. Over the last seven years this taxa

was recognized as members of the family Craugastoridae

(Hedges et al. 2008; Pyron and Wiens 2011; Padial et al.

2014), and placed into the taxon Holoadeninae (genera

Bryophryne, Lynchius, Noblella, Psychophrynella , and

Oreobates; Padial et al. 2014).

The Andes of Peru hold a rich fauna of the genus

Phrynopus. For instance, from a total of 26 species, 22

have been described from 2000 to 2015 (AmphibiaWeb

2015). Remarkably, central Peru has the highest diversity

of this genus (Duellman and Lehr 2009; Lehr et al. 2000;

Correspondence. Emails: l vampflack@yahoo.com (Corresponding author); 2 chara53@hotmail.com\ 3 daal91 l@yahoo.com; 4 elehr@iwu.edu

Amphib. Reptile Conserv. 15 October 2015 | Volume 9 | Number 1 | el 05

Chavez et al.

Lehr 2002; Lehr et al. 2002; Lehr and Aguilar 2003; Lehr

et al. 2005; Lehr and Oroz 2012; Mamani and Malqui

2014) with 25 species ocurring in the Huanuco, Pasco,

and Jumn regions. Phrynopus thompsoni from La Liber-

tad region currently marks the northernmost distribution

of the genus. All species of Phrynopus are restricted to

cloud forests and puna regions from 2,200 to 4,400 me-

ters (m) of elevation (Duellman and Lehr 2009). During

2013 and 2014, fieldwork in the Andean highlands of La

Libertad region (northwestern Peru) and cloud forests of

the Huanuco region (Cordillera de Carpish, central Peru)

revealed two new species of Phrynopus, which are de-

scribed herein.

Material and Methods

Format of description follow Lynch and Duellman (1997)

and character definitions defined in Duellman and Lehr

(2009). We used preserved specimens (Appendix) and

original species descriptions for the comparative diagno-

ses. Specimens were preserved in 96% ethanol and per-

manently stored in 70%. The following variables were

measured to the nearest 0.1 mm with digital calipers un-

der a microscope: snout-vent length (SVL), tibia length

(TL), foot length (FL, distance from proximal margin of

inner metatarsal tubercle to tip of toe IV), head length

(HL, from angle of jaw to tip of snout), head width (HW,

at level of angle of jaw), eye diameter (ED), interorbital

distance (IOD), upper eyelid width (EW), intemarial dis-

tance (IND), eye-nostril distance, and (E-N, straight line

distance between anterior comer of orbit and posterior

margin of external nares). Fingers and toes numbered

preaxially to postaxially from I-IV and I-V respectively.

We determined comparative lengths of toes III and V by

adpressing both toes against toe IV; lengths of fingers I

and II were determined by adpressing the fingers against

each other. Specimens were sexed based on external

sexual characteristics (e.g., presence of vocal sacs) or

through dissections to evaluation of gonads. To reduce

reflections, preserved holotypes were photographed

submersed in ethanol. Photographs taken in the field by

the authors were used for descriptions of color in life.

Specimens were deposited in the herpetological collec-

tion of the Centro de Ornitologfa y Biodiversidad (COR-

BIDI), Lima, Museo de Historia Natural Universidad

Fig. 1 . Dorsal (left) and ventral (right) views of specimens of the type series of Phrynopus valquii sp. nov.: A-B) Holotype (COR-

BIDI 14005, adult female, SVL = 31.2 mm); C-D) Paratype (CORBIDI 14007, adult female, SVL = 28.07 mm); E-F (CORBIDI

13993, adult male, SVL = 25.2 mm). Photographs by German Chavez.

Amphib. Reptile Conserv.

October 201 5 I Volume 9 I Number 1 I el 05

Two new species of frogs of the genus Phrynopus

Nacional de San Marcos (MUSM), Lima, and Museo de

Historia Natural de la Universidad Nacional San Agustrn

(MUSA), Arequipa, all in Peru. For specimens examined

see Appendix I.

Results

Phrynopus valquii sp. nov.

um:lsid:zoobank.org:act:77C37E4F-4FC2-42DE-8105-ElA834169142

Holotype: CORBIDI 14005 (Figs. 1-2), an adult female

from “Cerro Alto Chucaro” (8° 3’0.60”S, 77°24’9.24”W)

4,025 m.a.s.l., Distrito de Parcoy, Provincia de Pataz, La

Libertad region, Pem, collected on 26 February 2014 by

German Chavez.

Paratypes: CORBIDI 13998-14001, 14003-04, 14006,

and 14008 adult males and CORBIDI 14007 adult female,

same data as holotype; CORBIDI 13989-94, 13996-97,

adult males and CORBIDI 13988 and, 13995, adult fe-

males, collected two km southern of “Cerro Mush Mush”

(8°4’12.08”S, 77°25’33.94”W) 4,123 m.a.s.l., Distrito

de Parcoy, Provincia de Pataz, La Libertad region, Pem,

on 27 February 2014 by German Chavez.

Diagnosis: A species of Phrynopus having the following

combination of characters: (1) skin on dorsum shagreen

with low scattered tubercles, skin on flanks areolate,

skin on throat, chest and belly smooth, ventral surface of

thighs coarsely areolate; discoidal and thoracic fold ab-

sent; dorsolateral folds absent; (2) tympanic membrane

and tympanic annulus absent, supratympanic fold absent;

(3) snout rounded in dorsal and lateral views; (4) upper

eyelid without conical tubercles; width of upper eyelid

narrower than IOD; cranial crests absent; (5) dentiger-

ous processes of vomers absent; (6) males lacking vocal

slits and nuptial pads; (7) finger I slightly shorter than

finger II; tips of digits rounded; (8) fingers without lat-

eral fringes; (9) ulnar and tarsal tubercles absent; (10)

heels lacking tubercles; inner tarsal fold absent; (11) in-

ner metatarsal tubercle ovoid, about 1.5 times larger as

rounded outer metatarsal tubercle; supernumerary plan-

tar tubercles absent; (12) toes without lateral fringes;

basal webbing present; toe V slightly longer than toe III;

toe tips rounded, about as large as those on fingers; (13)

in life, dorsum dark brown, reddish brown or olive green

with irregular darker blotches, dark brown post orbital

stripe present; throat cream, pearly white, pale salmon or

creamy yellow, chest and belly creamy white or bluish

white with or without brown irregular blotches; ventral

surfaces of forearms and thighs brown or salmon, ventral

surface of hands and feet cream or dark brown with irreg-

ular creamy white flecks or blotches; groin dark brown,

or reddish brown with pearly white flecks; (14) SVL in

females 26.4-31.2 mm (n = 4), in males 14.9-16.5 mm

(n = 16).

The assignment of the new species to Phrynopus is

based on the structure of the digital discs that lack cir-

cumferential groves as well as the overall morphological

similarity with other members of the genus. Phrynopus

valquii shares the absence of tympanic annulus and tym-

panic membrane with most species except P. auriculatus,

P montium (tympanic annulus visible beneath skin), and

P peruanus. The only other species of Phrynopus known

from La Libertad region is P. thompsoni , which inhabits

mountains of western Andes, approximately 123 km from

known localities of P valquii. The following characters

Table 1 . Range of measured characters (mm) and proportions of the type series of Phrynopus valquii and P. daemon. Range of

measured characters is followed by mean and standard deviation in parentheses.

Phrynopus valquii

Phrynopus daemon

Males (n = 16)

Females ( n = 4)

Male (n = 1)

Females (n = 2)

SVL

21.02-26.45 (x = 23.59 ± 1.34)

26.48-31.21 (x = 28.89 + 2.05)

21.7

21.42-24.35

6.35-7.64 (x = 7.03 ± 0.39)

7.31-8.22 (x = 7.85 + 0.40)

7.8

6.98-7.89

7.56-8.72 (x = 8.15 ± 0.39)

8.89-9.84 (x = 9.43 + 0.41)

7.6

8.02-8.08

7.63-8.93 (x = 8.33 ± 0.42)

8.91-9.36 (x = 9.17 + 0.19)

8.5

7.84-8.17

8.53-10.44 (x = 9.48 ± 0.63)

10.13-10.86 (x = 10.43 + 0.30)

7.98-9.61

2.02-2.81 (x = 2.22 + 0.21)

2.17-3.27 (x = 2.74 + 0.51)

2.3

1.64-1.81

E-N

1.74-2.19 (x= 1.93 ±0.12)

1.74-2.37 (x = 2.08 + 0.26)

1.8

1.63-1.82

IOD

2.14-2.93 (x = 2.44 ± 0.24)

2.68-2.91 (x = 2.78 + 0.10)

2.6

2.59-2.64

1.56-2.27 (x = 2.01 +0.16)

2.05-2.56 (x = 2.23 + 0.23)

1.7

1.96-1.98

IND

1.72-2.35 (x = 2.02 + 0.18)

2.05-2.61 (x = 2.36 + 0.23)

1.9

1.94-2.06

HL/SVL

0.27-0.32 (x = 0.29 + 0.01)

0.26-0.27 (x = 0.27 + 0.00)

0.35

0.32-0.32

HW/SVL

0.33-0.38 (x = 0.34 + 0.01)

0.31-0.33 (x = 0.32 + 0.00)

0.35

0.33-0.37

TL/SVL

0.32-0.38 (x = 0.35 + 0.01)

0.29-0.34 (x = 0.31 +0.01)

0.39

0.33-0.36

E-N/ED

0.61-1.08 (x = 0.87 + 0.10)

0.55-1.00 (x = 0.78 + 0.22)

0.78

0.99-1.00

EW/IOD

0.66-1.00 (x = 0.83 + 0.09)

0.76-0.90 (x = 0.80 + 0.07)

0.65

0.75-0.75

Amphib. Reptile Conserv.

October 2015

Volume 9 | Number 1 | el 05

Chavez et al.

Fig. 2. Ventral view of: A) right hand of the holotype of Phrynopus valquii; B) right foot of the holotype of Phrynopus valquii', C)

right hand of the holotype of Phrynopus daemon', D) right foot of the holotype of Phrynopus daemon. Photographs by German

Chavez.

distinguish the two species: P. valquii lack pustules on

skin of dorsum (P. thompsoni has skin on dorsum bear-

ing pustules arranged in longitudinal rows), has smooth

skin on venter (P. thompsoni has skin on venter coarsely

areolate), finger I slightly shorter than finger II (P. thomp-

soni has fingers I and II of equal lenght), body size with a

maximum SVL in females of 31.2 mm (P thompsoni has

a maximum SVL in females of 27.6 mm). Furthermore,

P. valquii can be distinguished from its congeners which

lack tympanic annulus and tympanic membrane by hav-

ing skin on dorsum shagreen with scattered low tubercles

(vs coarsely tuberculate in P. barthlenae, P. chaparroi,

and P. miroslawae; smooth in P. kauneorum, P. nicolae,

P. oblivius, and P. tautzorum; bearing conical or elon-

gated tubercles, wart or ridges in P. badius, P. bracki , P.

bufoides, P. curator , P. nicolae , P. paucari, P. vestigiatus,

and P. daemon sp. nov.); skin on throat, chest, and belly

smooth (vs weakly areolate in P. badius , P. curator, P.

interstinctus, P. lechriorynchus, and P. oblivius', areolate

in P. barthlenae, P. bufoides, P. chaparroi, P. dagmarae,

P. heimorum, P. horstpauli, P. kotosh, P. miroslawae, P.

nicolae, P. paucari, P. pesantesi, P. tautzorum, and P.

vestigiatus', coarsely areolate in P. daemon sp. nov.); dor-

solateral folds absent (vs present in P. daemon sp. nov.,

P. dagmarae, P. horstpauli, P. interstinctus, P. kotosh, P.

miroslawae, and P. vestigiatus)', supratympanic fold ab-

sent (vs present in P. kauneorum)', snout rounded in dor-

sal view (vs elongated in P. lechriorynchus', truncate in P.

vestigiatus)', tubercles on upper eyelid absent (vs present

in 72 barthlenae and P. curator)', dentigerous processes of

vomers absent (vs present in P. bracki, P. dagamarae, P.

kauneorum, P. kotosh, P. lechriorynchus, and P. nicolae',

and minute in P. interstinctus and P. vestigiatus)', nuptial

pads absent (vs present in P. barthlenae and P. chapar-

roi)', finger I slightly shorter than finger II (vs finger I

much shorter than finger II in P. dagmarae and P. horst-

pauli', equal length in P. juninensis; finger I longer than

finger II in P. daemon sp. nov.); lateral fringes on fingers

and toes absent (vs present in P. dagmarae, P. barthlenae,

and P. vestigiatus', and narrow lateral fringes present on

fingers in P. bracki)', ulnar tubercles absent (vs present in

P. kotosh)', tarsal fold absent (vs present in P. lechrioryn-

chus)', tubercles on heels absent (vs present in P. badius,

P. bracki, P. curator, P. dagmarae, P. horstpauli, and P.

tribulosus) and toe V slightly longer than toe III (vs toe

V elongated, much longer than toe III in P. bracki, and P.

horstpauli', equal length in P. miroslawae, P. kotosh, P.

tribulosus, and P. vestigiatus ; toe V slightly shorter tan

toe III in P. heimorum, P. juninensis, and P. daemon sp.

nov.).

Amphib. Reptile Conserv.

October 201 5 I Volume 9 I Number 1 I el 05

Two new species of frogs of the genus Phrynopus

Description of Holotype (Fig. 1): Head narrower than

body, wider than long, HW 119.7% of HL; HW 31.5%

of SVL; HL 26.3% of SVL; snout short, rounded in

dorsal and lateral views, (Fig. 1A), ED about as large

as E-N distance; nostrils slightly protuberant, directed

dorsolaterally; canthus rostralis short, straight in dorsal

view, rounded in profile; loreal region slightly concave;

lips rounded; upper eyelid without enlarged tubercles;

EW narrower than IOD (EW 77.31% of IOD); tym-

panic region bearing rounded tubercles distinguishable

in preservation; tympanic membrane and tympanic an-

nulus absent; postrictal tubercles present. Choanae small,

ovoid, not concealed by palatal shelf of maxilla; dentig-

erous processes of vomers absent; tongue broad, slightly

longer than wide, not notched posteriorly, posterior one

half free. Skin on dorsum shagreen with small scattered

tubercles, without dorsolateral folds; skin on flanks areo-

late; skin on throat, chest, and belly smooth; discoidal

and thoracic folds present; cloacal sheath not discernible;

large tubercles absent in cloacal region. Outer surface of

forearm without minute tubercles; outer and inner pal-

mar tubercles low, outer rounded, the same size of in-

ner, rounded palmar tubercle; supernumerary tubercles

absent, rounded, about half the size of subarticular tu-

bercles; subarticular tubercles prominent, ovoid in dorsal

view, rounded in lateral view, most prominent on base of

fingers; fingers without lateral fringes; finger I slightly

shorter than finger II; tips of digits rounded lacking mar-

ginal grooves (Fig. 2A). Hind limbs slender, short, TL

Fig. 3. Distribution of Phrynopus valquii sp. nov. (green cir-

cles) and Phrynopus daemon sp. nov. (red circles). Map by Ca-

terina H. Cosmopolis.

29.9% of SVL; FL 34.8% of SVL; upper surface of hind

limbs shagreen with small, scattered tubercles; posterior

and ventral surfaces of thighs coarsely areolate; heel

without conical tubercles; outer surface of tarsus without

tubercles; inner metatarsal tubercle ovoid, about one and

a third times larger as rounded outer metatarsal tubercle;

supernumerary plantar tubercles absent; subarticular tu-

bercles low, ovoid in dorsal view; toes without lateral

fringes; basal webbing present; toe tips rounded, lack-

ing marginal grooves, about as large as those on fingers;

relative lengths of toes: 1 < 2 < 3 < 5 < 4; toe V slightly

longer than toe III (Fig. 2B). Measurements (mm) of ho-

lotype: SVL 31.2; TL 9.3; FL 10.8; HL 8.2; HW 9.8; ED

2.4; IOD 2.9; EW 2.2; IND 2.4; E-N 2.3.

Coloration of holotype in life (Fig. 1): Dorsum reddish

brown with small creamy white irregular flecks; postric-

tal tubercles creamy white; dorsal surface of forearms

reddish brown; canthal and supratympanic regions dark

brown; flanks colored as dorsum, reddish brown with

small creamy white irregular flecks; axilla and groin red-

dish brown (Fig. IB); posterior surfaces of thighs and

concealed surfaces of shanks reddish brown with pearly

white blotches; throat pale salmon; chest, belly, and ex-

tremities (except ventral surfaces of hands and feet) blu-

ish white with pale salmon irregular blotches; fingers

and toes reddish brown, palmar and plantar surfaces and

subarticular tubercles pale salmon; iris dark brown with

creamy white reticulations.

Coloration of holotype in preservative: As described

above with reddish brown coloration being dark brown

and salmon coloration being creamy white; flanks dark

brown; iris gray.

Variation: The males are smaller than the females and

lack vocal slits and nuptial pads. The post-orbital stripe is

absent in female CORBIDI 13988 and males CORBIDI

13997 and CORBIDI 14001, which have dorsal cream

coloration with irregular weak brown blotches, and yel-

low blotches between the upper eyelids. Female COR-

BIDI 13988 bears tubercles on the upper eyelids and on

the dorsum, where they are higher in the occipital region.

Brown blotches on belly are present in most of individuals

except in males CORBIDI 13998 and CORBIDI 13993

that have dark brown venter; CORBIDI 13998 has little

white spots on the belly. Two males (CORBIDI 13989,

14008) have belly creamy white with weak brown irregu-

lar blotches. The color pattern of the ventral surfaces of

thighs and forearms in female CORBIDI 13988 and male

CORBIDI 14008 is salmon.

Etymology: The name is a patronym as tribute to Thom-

as Valqui, Ph.D. (Peru), ornithologist and founder of

CORBIDI, in recognition of his valuable contributions

and efforts for the conservation, research, and knowledge

of Peruvian birds.

Amphib. Reptile Conserv.

October 201 5 I Volume 9 I Number 1 I el 05

Chavez et al.

Fig. 4. Dorsal (left) and ventral (right) views of the type series of Phrynopus daemon sp. nov.: A-B)

Holotype (CORBIDI 15364, adult female, SVL = 24.3 mm); C-D) Female paratype (MUSA 4916, adult

female, SVL = 21.4 mm); E-F) Male paratype (MUSM 32747, SVL = 21.7). Photographs: A-D by Roy

Santa-Cruz and E-F by Daniel Rodriguez.

Distribution and Ecology: Phrynopus valquii is known

from eastern Andes of La Libertad Region, two km far

from southwestern limit of Rio Abiseo National Park

(Fig. 3). All individuals were collected during the dry

season, at daytime between 9:00 a.m. and 14:00 p.m in a

flat and very humid grassland area sorrounded by rocks

clusters and mountains, under stones which form the dry

bed of an old creek. The vegetation includes grasses of

the families Poaceae and Valerianaceae, lichens and fun-

gi growing on stone surfaces. The only sympatric anuran

species recorded is Gastrotheca phelloderma.

Phrynopus daemon sp. nov.

urn:lsid:zoobank.org:act:236 17A10-D 1 C 1-49B2-A1 84-2B42E9B20D 1 8

Holotype: CORBIDI 15364, (Figs. 2, 4A-B), an

adult female, from Achupampa, Cordillera de Carpish

(8°45’15.55”S, 76°34’39.03”W), 3,138 m.a.s.l., Distrito

de Chinchao, Provincia de Huanuco, Huanuco region,

Peru, collected on 20 October 2014 by Roy Santa Cruz,

Heidy Cardenas, Nelin Ramos, and Eduardo Crispin.

Paratypes: MUSA 4916 (Fig. 4 C-D), adult female,

same data as holotype; MUSM 32747 (Fig. 4 E-F), an

adult male from Unchog elfin forest, Cordillera de Car-

pish (09°43’1.93”S, 76°10’3.71”W), 3,341 m.a.s.l.,

Distrito de Churubamba, Provincia Huanuco, Huanuco

region, Peru, collected on 20 July 2013 by Daniel Ro-

driguez.

Diagnosis: A species of Phrynopus having the fol-

lowing combination of characters: (1) skin on dorsum

shagreen with small scattered tubercles, skin on flanks

areolate, skin on throat, chest, belly and ventral surface

of thighs areolate; thoracic fold evident; discoidal fold

absent; fragmented dorsolateral folds present; (2) tym-

Amphib. Reptile Conserv.

October 201 5 I Volume 9 I Number 1 I el 05

Two new species of frogs of the genus Phrynopus

panic membrane and tympanic annulus absent, short

supratympanic fold present; (3) snout rounded in dor-

sal view, subacuminate in lateral view; (4) upper eyelid

without enlarged tubercles; width of upper eyelid nar-

rower than IOD; cranial crests absent; (5) dentigerous

processes of vomers absent; (6) vocal slits and nuptial

pads absent, subgular vocal sac present; (7) finger I and

II of equal length or finger I shorter than finger II; tips

of digits rounded; (8) fingers with weak lateral fringes;

(9) ulnar tubercles absent, low tarsal tubercles present;

(10) heels lacking tubercles; inner tarsal fold absent; (11)

inner metatarsal tubercle ovoid, about 1.5 times larger

than rounded outer metatarsal tubercle; supernumerary

plantar tubercles absent; (12) toes with lateral fringes;

basal webbing absent; toe V slightly shorter than toe

III; toe tips rounded, about as large as those of fingers;

(13) in life, dorsum and flanks vermilion red or blackish

brown; throat orange-red, chest, and belly vermilion red

or blackish brown, ventral surface of hands and feet dark

grey, iris green with fine black reticulations; (14) SVL

in females 21.42-24.35 mm (n = 2), in single male 21.7

mm.

The assignment of the new species to Phrynopus is

based on the structure of the digital discs lacking circum-

ferential groves, as well as the overall morphological

similarity with the other members of the genus. Phryno-

pus daemon is readily distinguished from its congeners

by its uniform red or blackish-brown coloration with an

orange-red throat, and by its distinct fragmented dorso-

lateral folds. Phrynopus daemon shares with nine other

species of Phrynopus (P badius, P bracki, P. dagmarae,

P. heimorum, P. interstinctus , P. nicoleae, P. paucari, P

peruanus, P. vestigiatus ) coloration consisting of red in

the groin. However, none of these species has a color

combination consisting in the entire dorsum vermilion

red or blackish brown and throat orange-red. Likewise,

this species shares the absence of tympanic annulus and

tympanic membrane with most species except P. au-

riculatus , P. montium (tympanic annulus visible beneath

skin), and P. peruanus. Furthermore, P daemon can be

distinguished from the rest of its congeners by having

skin on dorsum shagreen with scattered small tubercles

(vs coarsely tuberculate in P barthlenae , P chaparroi,

and P miroslawae; smooth in P. kauneorum, P. nicolae,

P oblivius, and P. tautzorum ; bearing conical, elongated,

or low tubercles, warts or ridges in P. badius , P bracki , P.

bufoides, P curator , P nicolae , P paucari , P thompsoni ,

P vestigiatus, and P. valquii sp.no v.); skin on throat, chest

and belly areolate (vs weakly areolate in P badius, P. cu-

rator, P interstinctus, P. lechriorynchus, and P. oblivius

and smooth in P. bracki, P juninensis, P kauneorum, P

tribulosus, and P. valquii sp. nov.); fragmented dorsolat-

Fig. 5. A) Habitat of the type locality of Phrynopus valquii sp. nov.; B) Microhabitat of Phrynopus valquii sp. nov.; C) Habitat of the

type locality of Phrynopus daemon sp. nov.; D) Microhabitat of Phrynopus daemon sp. nov. Photographs: A-B by German Chavez

and C-D by Roy Santa-Cruz.

Amphib. Reptile Conserv. 21 October 2015 | Volume 9 | Number 1 | el 05

Chavez et al.

eral folds preent (vs continous dorsolateral folds present

in P. dagmame, P. horstpauli, P. interstinctus , P. kotosh,

P miroslawae , and P. vestigiatus ); snout rounded from

dorsal view (vs elongated in P. lechriorynchus; truncate

from dorsal view in P vestigiatus ); tubercles on upper

eyelid absent (vs present in P. barthlenae and P. cura-

tor); dentigerous processes of vomers absent (vs present

in P. bracki , P. dagmarae , P horstpauli , P kauneorum,

P. kotosh, P. lechriorynchus, and P. nicolae ; minute in

P interstinctus and P vestigiatus ); lateral fringes on fin-

gers absent (vs present in P dagmarae , P barthlenae,

P. thompsoni, and P vestigiatus ; narrow lateral fringes

present on fingers in P bracki ); ulnar tubercles absent (vs

present in P. kotosh ); tarsal fold absent (vs present in P.

lechriorynchus ); tubercles on heels absent (vs present in

P. badius, P. bracki, P curator, P dagmarae, P. horstpau-

li, and P. tribulosus ) and toe V slightly shorter than toe III

(vs toe V elongated, much longer than toe III in P bracki

and P horstpauli; of equal length in P. kotosh, P mirosla-

wae, P. thompsoni, P tribulosus, and P. vestigiatus; toe

V slightly longer than toe III in P. bufoides , P. chapcirroi,

P interstinctus, P kauneorum, P. oblivius, P paucari, P

pesantesi, and P valquii sp. nov.). Four other species of

Phrvnopus have been recorded from Cordillera de Car-

pish. Those are P. dagmarae (Lehr et al. 2002), P. interst-

inctus (Lehr and Oroz 2012, at San Marcos, 3,100-3,160

in), P kauneorum (Lehr et al. 2002), and P vestigiatus

(Lehr and Oroz 2012, at San Pedro de Carpish, 3,100 m),

but all of them can be differentiated from P. daemon sp.

nov. by their different color pattern on dorsum, throat,

and belly.

Description of the Holotype (Fig. 4 A-B): Head nar-

rower than body, wider than long, HW 102.4% of HL;

HW 33.2% of SVL; HL 32.4% of SVL; snout moderately

short, rounded in dorsal view, sub acuminate in lateral

view, (Fig. 4A), ED about as large as E-N distance; nos-

trils slightly protuberant, directed dorsolaterally; canthus

rostralis short, straight in dorsal view, sub acuminate in

profile; loreal region slightly concave; lips rounded; up-

per eyelid without enlarged tubercles; EW narrower than

IOD (EW 68.9% of IOD); tympanic region lacking tu-

bercles; tympanic membrane and tympanic annulus ab-

sent; postrictal tubercles absent. Choanae small, ovoid,

not concealed by palatal shelf of maxilla; dentigerous

processes of vomers absent; tongue broad, slightly lon-

ger than wide, not notched posteriorly, posterior one

half free. Skin on dorsum shagreen with a few conical

scattered tubercles, with ridges forming fragmented dor-

solateral folds; skin on flanks coarsely areolate; skin on

throat, chest, belly, and ventral surfaces of thighs coarse-

ly areolate; thoracic fold present, discoidal fold absent;

cloacal sheath not discernible; large tubercles absent in

cloacal region. Outer surface of forearms with low tuber-

cles; outer and inner palmar tubercles low, outer bilobate,

the same size as inner, rounded palmar tubercle; supernu-

merary tubercles indistinct in preservative; subarticular

Amphib. Reptile Conserv.

tubercles low, ovoid in dorsal view, most prominent on

base of fingers; fingers without lateral fringes; finger I

longer than finger II; tips of digits rounded lacking mar-

ginal grooves (Fig. 2C). Hind limbs slender, short, TL

33.5% of SVL; FL 39.4% of SVL; upper surface of hind

limbs shagreen with scattered rounded tubercles; pos-

terior and ventral surfaces of thighs coarsely areolate;

heel without conical tubercles; outer surface of tarsus

with low rounded tubercles; inner metatarsal tubercle

ovoid, about one and a half times larger than rounded

outer metatarsal tubercle; supernumerary plantar tuber-

cles absent; subarticular tubercles low, ovoid in dorsal

view; toes with lateral fringes; basal webbing absent; toe

tips rounded, lacking marginal grooves, about as large as

those of fingers; relative lengths of toes: 1<2<5<3<

4; toe V slightly shorter than toe III (Fig. 2D). Measure-

ments (mm) of holotype: SVL 24.35; TL 8.17; FL 9.61;

HL 7.89; HW 8.08; ED 1.81; IOD 2.64; EW 1.98; IND

2.06; E-N 1.82.

Coloration of holotype in life (Fig. 4 A-B): Dorsum,

dorsal surface of forearms, canthal and supratympanic

regions, flanks, axilla, and groin dark red (Fig. 4A); pos-

terior surfaces of thighs and concealed surfaces of shanks

of the same color as dorsum; throat, chest, belly, and ven-

tral surfaces of forelimbs and hindlimbs red (Fig. 4B);

fingers, toes, palmar, and plantar surfaces grayish black;

iris bronze with fine black reticulations.

Coloration of holotype in preservative: As described

above with dark red coloration being dark brown on dor-

sum and creamy white on venter and grayish black color-

ation being brown; iris gray.

Variation: Male MUSM 32747 has dorsal surfaces of

head and body blackish brown, with vermilion red spots

and black reticulations on dorsal surfaces of hind limbs,

forelimbs and lateral surfaces of head and body; throat

vermilion red, rest of ventral region is blackish brown.

Dorsolateral ridges in this individual form fragmented

dorsolateral folds (Fig. 3, E-F). Female paratype MUSA

4916 is smaller (SVL = 21.4) than the holotype, bearing

higher tubercles on the dorsum; its dorsal color pattern is

darker than the holotype, being the coloration of the belly

and similar in all specimens.

Etymology: The specific name daemon is a latin word

meaning “demon” in reference to the color pattern of the

new species (red) which reminds the authors of the color-

ation attributed to the devil’s servants, ancestrally called

demons.

Distribution and Ecology: Phiynopus daemon sp. nov.

is known from two localities, in the Huanuco region

(Fig. 3), central Peru. Both localities (Achupampa and

Unchog forest) are located in the cloud forests of the

Cordillera de Carpish, near the treeline on the eastern

October 2015 | Volume 9 | Number 1 | el 05

Two new species of frogs of the genus Phrynopus

side of the Andes. These cloud forests may be among

the ecosystems most affeced by new regimes of humid-

ity and temperature caused by climate change, and by

anthropogenic destruction (Gonzalez 2013). Individu-

als were collected during the dry season, at daytime, in

very humid microhabitats. Females CORBIDI 15364

and MUSA 4916 were found on the ground underneath

mosses and roots, at the transition between primary cloud

forest and grassland, whereas male MUSM 32747 was

found under a stone near a dry stream bed surrounded by

a few bushes and trees. Sympatric amphibians recorded

with the females are Phrynopus dagmarae , P. horstpauli,

Gastrotheca griswoldi, and G. stictopleura , all of them

more commonly observed than P. daemon sp. nov. Male

MUSM 32747 was sympatric with two undescribed spe-

cies of Plnynopus.

Discussion

The conservation status we propose for the two new spe-

cies is based on IUCN's Red List criteria (cite IUCN Red

List) and known distribution and threats. We consider

plausible the occurrence of Pbynopus valquii inside Rio

Abiseo National Park (RANP) because its type local-

ity is aproximately two km from the southwestern limit

of this protected area, which covers more than 274,000

km 2 ; part of this area contains grassland habitats and

there are no geographical barriers between this limit and

the type locality of P. valquii. Nevertheless, there are no

confirmed records of the species inside the RANP, so we

suggest to evaluate this species as Data Deficient based

on the limited information about its distribution range

and population status.

Phrynopus daemon inhabits cloud forests of the Cor-

dillera de Carpish, which is located in the Cordillera Ori-

ental (Atlantic drainage), Huanuco region. In this Cor-

dillera recent herpetological surveys (over the last five

years) have resulted in the discovery of five endemic

species of Plnynopus : P. dagmarae Lehr, Aguilar, and

Koehler 2001; P. inter stinctus Lehr and Oroz 2012; P

kauneorum Lehr, Aguilar, and Koehler 2001, P. vestigia-

tus Lehr and Oroz 2012, and P daemon sp. nov. Known

distribution ranges include, for most of them, only a few

localities. In fact, the Cordillera de Carpish is a chain of

mountains located between the Chinchao and Derrepente

rivers (both small tributaries of the Huallaga river, Hua-

nuco region), that might promote endemism because of

its varied topography and sharp altitudinal gradient pass-

ing from 600 m (at the bank of the Chinchao river) to

3,200 m of altitude over an airline distance of 28 km. Re-

markably, this important Cordillera is not protected un-

der Peruvian law, making it susceptible to deforestation

by agriculture and timber extraction. Reduction in cloud

forest cover is the main threath for Phrynopus daemon

sp. nov. Furthermore, an interstate road which crosses

Cordillera de Carpish (at both sides of Chinchao river)

Amphib. Reptile Conserv.

and the developing of mining concessions in the area,

add more threats to the conservation of the forests and

their fauna. On the basis of a known distribution range

smaller than 5,000 km 2 , fragmentation of habitats near

the type locality, and known threats from agriculture and

mining, we suggest categorizing this species as Endan-

gered (B 1 ab [ii,iii]) in the IUCN Red List.

Acknowledgments. — We are grateful to Ministerio

del Ambiente and SERFOR for collecting and research

permits. RSF fieldwork was possibe thanks to economi-

cal support of ODEBRECHT Peru, logistic support of

Rafael Tamashiro from the environmental staff of ODE-

BRECHT, and Daniel Cossios from the staff of BIOS-

FERA Consultores. Specimens were loaned to EL by

B. Milan and J. Cordova (MUSM). Likewise, we thank

Ignacio de la Riva and Alessandro Catenazzi for review-

ing the manuscript and providing critical suggestions and

corrections to improve this paper.

Literature Cited

Duellman W, Lehr E. 2009. Terrestrial-Breeding Frogs

(Strabomantidae) in Peru. Natur- und Tier-Verlag,

Naturwissenschaft, Munster. 382 p.

Hedges SB, Duellman WE, Heinicke MP. 2008. New

World direct-developing frogs (Anura, Terrarana),

molecular phylogeny, classification, biogeography,

and conservation. Zoo taxa 1737: 1-182.

Lehr E. 2002. Amphibien und Reptilien in Peru. Die Her-

petofauna entlang des 10. Breitengrades von Peru:

Arterfassung, Taxonomie, okologische Bemerkun-

gen und biogeographische Beziehungen. Dissertation

Natur-und Tier-verlag, Naturwissenschaft, Munster.

Germany. 208 p.

Lehr E, Aguilar C. 2003. A new species of Plnynopus

(Amphibia, Anura, Leptodactylidae) from the puna of

Maraypata (Departamento de Huanuco, Peru). Zoolo-

gische Abhandlungen (Dresden) 53: 87-92.

Lehr E, Aguilar C, Kohler G. 2002. Two sympatric new

species of Phrynopus (Anura: Leptodactylidae) from

a cloud forest in the Peruvian Andes. Journal of Her-

petology 36(2): 208-216.

Lehr E, Kohler G, Ponce E. 2000. A new species of

Pbynopus from Peru (Amphibia, Anura, Leptodac-

tylidae). Senckenbergiana biologica 80(1/2): 205-

212 .

Lehr E, Lundberg M, Aguilar C. 2005. Three new species

of Pbynopus from central Peru (Amphibia: Anura:

Leptodactylidae). Copeia 2005: 479—4-91.

Lehr E, Oroz A. 2012. Two new species of Pbynopus

(Anura: Strabomantidae) from the Cordillera de Car-

pish in central Peru (Departamento de Huanuco). Zoo-

taxa 3512: 53-63.

Lynch J, Duellman W. 1997. Frogs of the genus Eleu-

therodactylus (Anura: Leptodactylidae) in western

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Chavez et al.

Ecuador: Systematics, ecology, and biogeography.

Special Publication (University of Kansas. Museum

of Natural History) 23: 1-236.

Mamani L, Malqui S. 2014. A new species of Phrynopus

(Anura: Craugastoridae) from the central Peruvian

Andes. Zootaxa 3838(2): 207-214.

Padial JM, Grant T, Frost DR. 2014. Molecular systemat-

ic of Terraranas (Anura: Brachycephaloidea) with an

assessment of the effects of alignment and optimality

criteria. Zootaxa 3825(1): 1-132.

Pyron RA, Wiens JJ. 2011. A large-scale phylogeny of

Amphibia including over 2,800 species, and a revised

classification of extant frogs, salamander and caeci-

lians. Molecular Phylogenetics and Evolution 61:

543-583.

Appendix 1

Specimens examined:

Phrynopus barthlenae . — PERU: Huanuco: Maraypata (10°11’ 11.29”S, 76 o 05’58.4”W), 3,749 m: CORBIDI 14496-

98.

Phrynopus bracki . — PERU: Pasco: Abra Esperanza, Parque Nacional Yanachaga Chemillen (10°3r57.40”S,

75°20’58.20”W), 2,808 m: CORBIDI 10302-10304, 2750, 11616, 11621, 11628, 11631.

Phrynopus dagmarae . — PERU: Huanuco: Palma Pampa, 3,020 m: MUSM 20451 (holotype). Achupampa,

(09°43’48.87”S, 75°57 , 04.29”W), 3,122 m: CORBIDI 14552-59, 3,160 m: CORBIDI 14564-72, 14580-90.

Phrynopus heimorum . — PERU: Huanuco: ca. 10 lan E Conchamarca, 3,240 m: MUSM 20441 (holotype).

Phrynopus horstpauli . — PERU: Huanuco: Sacsahuanca (10°12’50.01”S, 76°07’ 10.18”W), 3,472 m: CORBIDI 14539.

Phrynopus interstinctus . — PERU: Huanuco: Cordillera de Caipish, San Marcos, 3,100 m: MUSM 29543 (holotype),

3,160 m: MUSM 29544-29545 (paratypes).

Phrynopus kauneorum . — PERU: Huanuco: Camino Pampa Alegre-Ruinas de Huanacaure (09°45’29.07”S,

75°53’21.37”W), 2,996 m: CORBIDI 14591, 14594-95, 14615, 14629; Cordillera de Carpish, 2,735 m: MUSM 18585.

Phrynopus peruanus . — PERU: Junin: Puna of Maraynioc (11 0 2U35.2”S, 75 0 28’52.6”W), 3,825 m: MUSM 19977-78.

Phrynopus tautzorum.— PERU: Huanuco: Maraypata (10°09’ 13.45”S, 76°04 , 39.56”W), 3,949 m: CORBIDI 14504-

05, 14507.

Phrynopus tribulosus . — PERU: Pasco: Abra Esperanza, Parque Nacional Yanachaga Chemillen (10°3r57.40”S,

75°20’58.20”W), 2,808 m: CORBIDI 10299-10301, 10°31’57.40”S, 75°20’57.02”W, 2,748 m, CORBIDI 11626,

11627, 11638.

Phrynopus vestigiatus . — PERU: Huanuco: Cordillera de Carpish, San Pedro de Carpish, 3,100 m: MUSM 29542

(holotype); Camino Pampa Alegre, Ruinas de Huanacaure (09°45’29.07”S, 75°53’21.37”W), 2,996 m: CORBIDI

14598-99, 14600-02, 14617-18.

Amphib. Reptile Conserv. 24 October 2015 | Volume 9 | Number 1 | el 05

Two new species of frogs of the genus Phrynopus

German Chavez is a peruvian scientist working on the diversity and conservation of amphibians and reptiles in

Peru since 2003. His research includes 16 scientific articles, describing four new species of reptiles, description

1 of two new species of amphibians, and rediscovering an endemic frog from central Peru. Currently, he is an as-

sociated researcher at the Center of Ornithology and Biodiversity (CORBIDI) in Lima, Peru and is working on

l ' ’ andean amphibians and reptiles.

Roy Santa-Cruz has the bachelor degree in biology (2007). He is currently an amphibian and reptile researcher

at the Herpetological Collection of the Museum of Natural History, Universidad Nacional de San Agustrn de

Arequipa, Pern (MUSA). His current research interests include taxonomy, biology, and ecology of amphibians

and reptiles of Pern.

Daniel Rodriguez is a peruvian biologist involved in the conservation of natural areas, amphibians, and reptiles.

He is a researcher at the Herpetology Department of the Natural History Museum of San Marcos University,

Lima, Pern.

Dr. Edgar Lehr is Associate Professor in the Department of Biology at Illinois Wesleyan University, USA. He

serves as Associate Editor for both the Journal of Herpetology (SSAR) and Salamandra (DGHT). Lehr’s research

focuses on the amphibian and reptile diversity of Peru and has resulted in the publication of over 100 articles, two

books, and the description of 84 new species (76 amphibians, eight reptiles) from Peru. Currently his research

involves the herpetofauna of protected areas in central Andean Peru.

In accordance with the International Code of Zoological Nomenclature new rules and regulations (ICZN 2012), we have deposited this paper in publicly acces-

sible institutional libraries. The new species described herein has been registered in ZooBank (Polaszek 2005a, b), the official online registration system for the

ICZN. The ZooBank publication LSID (Life Science Identifier) for the new species described here can be viewed through any standard web browser by append-

ing the LSID to the prefix “http://zoobank.org/.” The LSID for this publication is: um:lsid:zoobank.org:pub:833E0DC3-AFD3-4BA4-83EF-B738F386D415.

Separate print-only edition of paper(s) (reprint) are available upon request as a print-on-demand service. Please inquire by sending a request to: Amphibian &

Reptile Conservation , amphibian-reptile-conservation.org, arc.publisher@gmail.com.

Amphibian & Reptile Conservation is a Content Partner with the Encyclopedia of Life (EOL), http:///www.eol.org/ and submits information about new species

to the EOL freely.

Digital archiving of this paper are found at the following institutions: ZenScientist, http://www.zenscientist.com/index.php/filedrawer; Ernst Mayr Library, Mu-

seum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (USA), http://library.mcz.harvard.edu/ernst_mayr/Ejournals/ARCns.

The most complete journal archiving and journal information is found at the official ARC journal website, amphibian-reptile-conservation.org. In addition,

complete journal paper archiving is found at: ZenScientist, http://www.zenscientist.com/index.php/filedrawer.

Citations

ICZN. 2012. Amendment of Articles 8,9,10,21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods of publication.

Zootaxa 3450: 1-7.

Polaszek A et al. 2005a. Commentary: A universal register for animal names. Nature 437: All .

Polaszek A et al. 2005b. ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological Nomenclature 62(4):

210 - 220 .

Amphib. Reptile Conserv.

October 201 5 I Volume 9 I Number 1 I el 05

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [Special Section]: 26-33 (el 07).

A new elusive species of Petracola (Squamata:

Gymnophthalmidae) from the Utcubamba basin in

the Andes of northern Peru

Gourdes Y. Echevarria and 2 Pablo J. Venegas

1 1 Division de Herpetologfa-Centro de Ornitologia y Biodiversidad ( CORBIDI), Santa Rita N 105 36 Of. 202, Urb. Huertos de San Antonio, Surco,

Lima, PERU l2 Museo de Zoologia, Escuela de Ciencias Biologicas, Pontificia Universidad Catolica del Ecuador, Avenida 12 de Octubre 1076 y

Roca, Apartado 17-01-2184, Quito, ECUADOR

Abstract . — We described a new species of Petracola from the montane forests of the Utcubamba

basin in Amazonas department, Andes of northern Peru (elevation 1 ,889 m). The new species is easily

distinguishable from all other species of Petracola by having a gracile body, two supraoculars, one

loreal scale, nine femoral pores per limb in males, and white venter with black semicircular marks on

anterior margin of scales. An updated identification key for species of Petracola is provided.

Key words. Cordillera Central, Peruvian Yungas, Petracola labioocularis, precloacal pores, superciliaries, supraocu-

lars

Citation: Echevarria LY, Venegas PJ. 2015. A new elusive species of Petracola (Squamata: Gymnophthalmidae) from the Utcubamba basin in the

Andes of northern Peru. Amphibian & Reptile Conservation 9(1 ): 26-33 (el 07).

CommercialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium,

provided the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized

publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation', official journal website

< amphibian-reptile-conservation. org> .

Received: 05 September 2015; Accepted: 08 November 2015; Published: 23 November 2015

Introduction

Modern molecular phylogenetic approaches sometimes

combined with morphological data have revealed novel

phylogenetic relationship for many groups of reptiles,

viz.: South American pit viper genera Bothrops, Bothri-

opsis, and Bothrocophias by Fenwick et al. (2009); Te-

iidae lizards by Harvey et al. (2012); Jadin et al. (2013);

and skinks by Hedges and Conn (2012). As in the afore-

mentioned taxa, some important taxonomical changes

were proposed for gymnophthalmid lizards by Doan and

Castoe (2005), including the reallocation of Proctoporus

ventrimaculata Boulenger 1900 in the genus Petracola.

The genus Petracola Doan and Castoe 2005 is a clade

of small-sized Andean lizards with semifossorial habits

occurring in cloud forest and puna habitats (Kohler and

Lehr 2004; Kizirian et al. 2008). Three species endemic

to Peru are currently recognized: Petracola ventrimacu-

lata Boulenger 1900, known from multiple isolated lo-

calities between elevations of 3,050-3,600 m in the An-

des of northern Peru, on both sides of the Maranon river

(Kizirian et al. 2008); P. labioocularis Kohler and Lehr

2004, known only from the type locality at an elevation

of 2,980 m in Central Peru; and P. waka Kizirian, Bayef-

sky-Anand, Eriksson, Le, and Donnelly 2008, known

from four localities in Cajamarca department at eleva-

tions between 2,600 and 2,900 m.

In this paper, we describe a new species of Petracola

based on morphological data from a single specimen,

which was collected in 2007 in the Utcubamba basin of

northeastern Peru. Subsequent visits by one of us (PJV)

to the type locality of the taxon described herein, and

other localities along the Utcubamba basin between 2012

and 2014, revealed no additional specimens of the new

species. This discovery elevates the number of Petracola

species to four species.

Materials and Methods

Museum abbreviations are Centro de Ornitologia y Bio-

diversidad (CORBIDI) and Museo de Historia Natural de

la Universidad Nacional Mayor de San Marcos (MUSM),

both collections from Lima, Peru. The following mea-

surements were taken with a digital caliper and recorded

Correspondence. Email: Hourdese.20@ gmail.com (Corresponding author); 2 sancarranca@yahoo.es

Amphib. Reptile Conserv.

November 2015 I Volume 9 I Number 1 I el 07

Echevarria and Venegas

to the nearest 0.1 mm, except for tail length, which was

taken with a ruler and recorded within 1 mm: head length

(HL), head width (HW), shank length (ShL), axilla-groin

distance (AGD), and snout-vent length (SVL). For char-

acters recorded on both sides, condition on the right side

is presented first. Sex was determined by observation of

hemipenes from X-ray photographs. Scale counts and

color pattern data for P. labioocularis were taken from

Kohler and Lehr (2004) and some high quality photo-

graphs of an adult male paratype (SMF 80113) provided

by G. Kohler. Data for P. ventrimaculata and P. waka was

taken from Kizirian et al. (2008) and from examination

of specimens of P. ventrimaculata (see Appendix 1). For

comparisons with species of Euspondylus, Pholidobolus ,

Proctoporus, and Riama data was taken from Montanuc-

ci (1973), Kizirian (1996), and Doan and Castoe (2005).

We follow the terminology of Kizirian (1996) for scale

counts and morphological characters.

by lacking prefrontals. The morphological characters de-

fining Petracola are similar to those of the genera Eus-

pondylus, Pholidobolus, Proctoporus, and Riama. Thus,

placement of this new species in the genus Petracola is

tentative, based only on morphology and partly on distri-

bution (see discussion below).

New species description

Petracola angustisoma sp. nov.

urn:lsid:zoobank.org:act:ED95BB48-4C31-4161-B4A2-EE4A787EDBD4

Holotype: CORBIDI 00467 (Figs. 1-3A), adult

male from vicinity of Cocachimba (06°02’37.2”S,

77°53’15.8”W; Datum WGS 84) at 1,889 m, Bongara

province, Amazonas department, Peru, collected by P.J.

Venegas on 24 June 2007.

Results

Generic assignment

The new species is assigned to the genus Petracola as

defined by Doan and Castoe (2005) having, among other

characters, smooth head scales; transparent lower eyelid;

rectangular, smooth, and juxtaposed dorsal scales; and

Fig. 1 . Petracola angustisoma sp. nov., holotype CORBIDI

00467, male, 43.6 mm SVL. Dorsal (upper) and ventral (bot-

tom) views. Photographs by D. Quirola.

Diagnosis: Petracola angustisoma differs from P. la-

bioocularis (character states in parentheses) in having

two supraoculars (three), 37 transverse dorsal scale rows

(34-35), 22 transverse ventral scale rows (18-20), dor-

sum pale brown with seven discontinuous dark brown

longitudinal stripes extending to the base of tail (brown

or olive brown with a dorsolateral pale stripe on neck and

body), and venter white with black semicircular marks

on anterior margin of scales (grayish brown, with no pat-

tern); precloacal pores absent (present); and by lacking

supralabial-subocular fusion (present). The new species

can be distinguished from P. ventrimaculata by having

a gracile body (robust, Fig. 3B), three discontinuous su-

perciliaries (two), loreal present (usually absent), nine

femoral pores per hind limb in males (2-5), and venter

white with black semicircular marks on anterior margin

of scales (yellow with bold black bands); from P. waka it

differs in having three discontinuous superciliaries (four,

usually continuous), two genials (three), and nine femo-

ral pores per hind limb in males (5-7; Table 1).

Petracola angustisoma differs from all known species

of Euspondylus, except E. spinalis, by lacking prefrontal

scales and from E. spinalis by having two supraoculars

(three in E. spinalis)', from Pholidobolus by having dor-

sals smooth and juxtaposed (imbricate, striate, or keeled

in Pholidobolus ); from Proctoporus by having smooth

dorsal scales (keeled in Proctoporus ); and from Riama

by having smooth dorsals (smooth, striate, rugose, or

keeled in Riama).

Description of holotype: Rostral scale wider than

long, taller than adjacent supralabials, in contact with

frontonasal, nasals, and anteriormost supralabials. Fron-

tonasal longer than wide, widest at level of nares, dis-

tinctly larger than frontal. Frontal approximately as

long as wide, widest at anterior suture of anteriormost

supraocular, barely extending between frontoparietals.

Frontoparietals hexagonal, in contact medially and with

Amphib. Reptile Conserv.

November 201 5 I Volume 9 I Number 1 I el 07

A new species of Petracola from northern Peru

Fig. 2. Dorsal (A), lateral (B), and ventral (C) views of the head

of Petracola angustisoma sp. nov. Holotype, CORBIDI 00467,

male. Photographs by D. Quirola.

supraoculars. Interparietal hexagonal. Parietals polygo-

nal, longer than wide. Postparietals two, with posterior

sutures forming a nearly straight line. Supraoculars two.

Anterior supraocular larger than posterior supraocular.

Superciliary series discontinuous, 2— 1/2— 1, the ante-

riormost extending onto dorsal surface of head. Nape

scales immediately posterior to head scales larger than

adjacent dorsals. Nasal subtriangular, pierced in center

by nostril, with shallow groove extending dorsally to

loreal. One loreal on each side, not in contact with su-

pralabials. Palpebral disc transparent with minute brown

flecks. Supralabials eight. Suboculars three. Postoculars

two. Supratympanic temporals 3/3. Tympanum recessed,

transparent. Infralabials six. Genials two, meeting at

broad midventral sutures. Pregulars in somewhat regu-

lar transverse rows, anteriormost two rows larger than

posterior rows. Gulars in eight rows. Gular fold distinct,

concealing three rows of small scales.

Dorsals smooth, juxtaposed, rectangular, in 37 trans-

verse rows and 19 longitudinal rows (at tenth transverse

ventral scale row). Ventrals smooth, in 22 transverse and

10 longitudinal rows. Dorsals and ventrals separated by

Fig. 3. Two species of Petracola in life. Holotype of P. angus-

tisoma sp. nov. (A), and adult male of P. ventrimaculata (COR-

BIDI 03630) from Laguna Norte, Cajamarca (B). Photographs

by P.J. Venegas.

approximately three longitudinal rows of small to granu-

lar scales, which are continuous with small to granular

scales at insertion of limbs. Cloacal plate with two large

anterior and four large posterior scales. Tail complete.

Caudal scales rectangular, smooth, glossy, in 98 trans-

verse rows.

Forelimbs pentadactyl, with claws. Anterodorsal

scales smooth, glossy, polygonal, and larger than those

on posterior side. Dorsal scales on brachium much larger

than ventrals. Ventral scales on brachium subgranular.

Anterior scales on antebrachium polygonal, smooth, and

glossy. Posteroventral scales on antebrachium roundish,

smooth, and glossy. Palmar scales domed, some with

central pits. Dorsal scales on finger I 2/3, II 5/5, III 8/8,

IV 10/9, and V 6/6. Subdigital lamellae undivided; on

finger I 5/5, II 10/9, III 13/12, IV 13/13, and V 7/8.

Scales on anterior surface of thigh polygonal, smooth,

glossy, and distinctly larger than adjacent scales. Scales

on anteroventral surface of thigh polygonal smooth and

glossy. Femoral pores 9/9. Scales on posterior surface of

thigh small, round, smooth, and glossy. Scales on dorsal

surface of crus polygonal to round, subimbricate, smooth,

and glossy. Scales on ventral surface of crus significantly

larger than adjacent scales. Scales on dorsal surface of

foot polygonal, irregularly arranged, subimbricate, and

of varying sizes. Dorsal scales on toe I 3/3, II 6/6, III

9/9, IV 12/12, V 7/7, and single. Plantar scales ovoid and

domed. Subdigital lamellae divided from a point halfway

Amphib. Reptile Conserv.

November 201 5 I Volume 9 I Number 1 I el 07

Echevarria and Venegas

Table 1 . Scutellational characters in species of Petracola. Range followed by mean + standard deviation is given for quantitative

characters if applicable.

Characters

Supraoculars

Petracola angustisoma

(n= 1)

Petracola labioocularis

Superciliaries

Postoculars

Genials

Transverse dorsal

scale rows

Longitudinal dorsal

scale rows

Transverse ventral

scale rows

Longitudinal ventral

scale rows

Femoral pores per

hind limb in males

Femoral pores per

hind limb in females

2-3

(2.7 ± 0.45)

34-35

34.7 ± 0.45

Petracola ventrimaculata

(n = 107)

2/2 (n = 106)

3/3 in = 1)

1- l/l-l {n = 77)

2 - 1 / 2-1 in = 8 )

4/4 ( n = 3)

Other in = 19)

2/2 in = 103)

Other in = 4)

0/0 in = 97)

1/1 in = 6)

Other in =16)

6/6 in = 39)

7/7 in = 34)

6/7 in = 10)

Other in = 23)

6/6 in = 60)

7/6 in = 11)

6/5 in = 11)

Other in = 24)

2/2 in = 103)

3/3 in = 3)

3/2 in = 1)

30-41

34.24 ± 2.25

in = 107)

15-23

18.71 ± 1.52

in = 107)

21.88 ± 1.43

in = 106)

8-13

10.82 + 1.02

in= 106)

Petracola waka

2/2 in = 35)

3/2 in = 2)

Other in = 2)

4/4 (>i = 32)

4/3 in = 2)

5/4 in = 1)

3/3 in = 1)

2 - 1 / 2-1 in = 1 )

l-l/l-l in = 1)

Other (/? = 1)

2/2 {n = 17)

3/3 {n = 14)

2/3 in = 3)

3/2 in = 4)

Other (/? = 1)

2/2 in = 5)

1/1 in = 7)

2/1 in = 1)

1/2 in = 3)

0/0 in = 1)

8/8 in = 14)

7/7 in = 13)

8/7 in = 6)

7/8 in = 2)

7/6 in = 2)

7/7 in = 14)

6/6 in = 15)

5/6 (« = 2)

6/5 (« = 1)

8/7 in = 1)

6/7 (« = 1)

5/5 in = 1)

3/3 in = 35)

36-49

in = 39;

42.2 ±4.16)

23-28

(n = 37;

25.0+ 1.23)

between base and tip to the base of toes in 7/6 rows on

toe I, 12/12 on II, 17/16onIII, 20/19onIV, 11/13 onV.

Measurements (mm) and proportions of the holotype:

SVL 43.6 mm; TL 100 mm; HL 9.6; HW 6.7; ShL 5;

AGD 23.4; TL/SVL 2.3; HL/SVL 0.2; HW/SVL 0.2;

ShL/SVL 0. 1 ; and AGD/SVL 0.5.

Holotype color in life: Dorsal background pale brown

with seven discontinuous black longitudinal stripes on

dorsum and two on flanks (continuous on neck) extend-

ing onto base of tail; dorsal and dorsolateral aspects with

six longitudinal rows of dark yellow flecks extending

onto tail; limbs covered by dark brown reticulations;

Amphib. Reptile Conserv.

November 201 5 I Volume 9 I Number 1 I el 07

A new species of Petracola from northern Peru

Altitude

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I I 1000-1.500

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Fig. 4. Distribution map of species of Petracola. Petracola

angustisoma sp. nov. (blue square), P ventrimaculata (black

triangles), P. labioocularis (red pentagon), P. waka (green cir-

cles).

tail with scattered dark brown marks; sides of head with

longitudinal black stripes; labials and ventrolateral re-

gion of head white with black marks; sides of neck cov-

ered with black reticulations; and four faint ocelli with

creamy brown center above the forelimb insertion (Fig.

3A). Ventral surface of body including limbs and tail

white; ventral surface of head with irregular black marks

on the center of each scale; ventral surface of neck with

two transverse black stripes; ventral surface of trunk and

tail with black semicircular marks on anterior margin

of scales, arranged in transverse rows; ventral surface

of forelimbs with black reticulations; ventral surface of

hind limbs with roundish black marks.

Color in preservative: Si mil ar to color in life but

the dark yellow flecks on dorsum, dorsolateral region

and tail are cream instead of dark yellow, and the center

of the four ocelli above forelimb are cream instead of

creamy brown (Fig. 1).

Distribution and natural history: The new species

is known only from its type locality in a montane forest

of the Utcubamba basin, northern portion of Cordillera

Central, Peru, at 1,889 m (Fig. 4). Following the ter-

restrial ecoregions of the world by Olson et al. (2001),

this locality lies in the Peruvian Yungas ecoregion. The

holotype of P. angustisoma was found under a pile of

decaying vegetation inside an abandoned house, along

with a juvenile specimen of Mastigodryas boddaerti. The

type locality lies in the vicinity of Cocachimba and is

composed of croplands with scattered large boulders and

bushes, embedded in a matrix of humid montane forest.

However, the forest has been almost completely removed

and only some small patches of secondary forest remain

close to ravines. Sympatric species of squamate reptiles

are Atractus sp., Chironius exoletus, M. boddaerti, and

Stenocercus huancabambae.

Etymology: The specific epithet angustisoma is a noun

(in apposition) in the nominative singular and derives

from the Latin words angusti (= narrow) and soma (=

body). It refers to the slender body of this species.

Discussion

Although the practice of describing species with a single

specimen is not recommended, we feel confident about

the identity of Petracola angustisoma as a different

taxon, due to its strikingly distinctive diagnostic charac-

ters. Unfortunately we were not able to collect additional

specimens, despite repeated surveys to the type locality

and other adjacent localities along the Utcubamba basin.

The genus Petracola was erected by Doan and Castoe

(2005) based on the results of a phylogenetic study of

gymnophthalmid lizards using molecular evidence (Cas-

toe et al. 2004), where samples of Proctoporus ventri-

maculata (KU 219838) and Proctoporus cf. ventrimacu-

lata (KU 212687) resulted paraphyletic to Proctoporus

(see Fig. 6 in Castoe et al. 2004). Based on morphological

evidence, the authors assigned Proctoporus labioocularis

(Kohler and Lehr 2004), a species from central Peru (Fig.

4), to the genus Petracola. Three years later Kizirian et

al. (2008) described Petracola waka, remarking that the

new species allocation to the genera was based on con-

venience rather than confidence regarding phylogenetic

relationships; however, they overlooked the fact that one

of the paratypes of P. waka (KU 212687) was included

in Castoe et al. (2004) phylogeny. Therefore, the assigna-

tion of this species to the genus Petracola is supported by

molecular evidence as well.

The identification of superciliaries and supraoculars in

gymnophthalmid lizards has not been consistent. Herein

we follow the terminology of Kizirian (1996), who de-

fines the superciliaries as those scales between the supra-

oculars and the ciliaries, in a continuous or discontinuous

series, and in Petracola ventrimaculata the anteriormost

supraocular is fused with the first superciliary. Doan and

Castoe (2005) consider that P. ventrimaculata has three

supraoculars and Kohler and Lehr (2004) consider that

P labiocularis has four supraoculars. Following Kizirian

(1996), the anteriormost “supraocular” in both P. ven-

trimaculata and P. labiocularis is the first superciliary.

Thus, P. ventrimaculata has two supraoculars and two

discontinuous superciliaries, of which the anteriormost

extends onto the dorsal surface of head (Uzzell 1970;

Amphib. Reptile Conserv.

November 201 5 I Volume 9 I Number 1 I el 07

Echevarria and Venegas

Fig. 5. Close-up of the heads of Petracola angustisoma sp. nov.

(A), P. labioocularis (SMF 80113) (B), and P ventrimaculata

(CORBIDI 03630) (C), showing the anteriormost superciliary

(red arrows) and supraoculars (blue arrows). Scale bar = 1 mm.

Photographs by D. Quirola, G. Kohler, and J. C. Chavez, re-

spectively.

Kizirian et al. 2008), a condition shared with P. labioocu-

laris , P. waka, and the species described herein (Fig. 5).

We found conspicuous differences between Petracola

labioocularis and all other species of Petracola , such as

the presence of three supraoculars, supralabial- subocular

fusion, and precloacal pores in males and females (or no

scales between femoral pores; Fig. 6), a rare condition

in Gymnophthalmidae, also present in some Riama spe-

cies (Kizirian 1996). It also has a disjunct distribution

being the only species of the genus present in central

Peru which makes its allocation in the genus doubtful.

We tentatively assigned the new species to the genus Pe-

tracola due to overall similarities with other species of

the genus, especially the presence of two supraoculars,

anteriormost superciliary extended onto dorsal surface

of head (or fused with anteriormost supraocular), and

its distribution in the Andes of northern Peru (Fig. 4).

Increased taxon sampling in a molecular phylogeny is

needed to resolve the phylogenetic relationships among

Petracola angustisoma, P labioocularis, and other spe-

cies currently recognized as Petracola.

Acknowledgments. — We are grateful to the staff of

Asociacion Ecosistemas Andinos (ECOAN) and Consul-

tores Asociados en Naturaleza y Desarrollo (CANDES)

for their valuable help and logistic support in two dif-

ferent field trips. The authors are very grateful to Dr. G.

Kohler for his kind willingness to cooperate with high

quality photographs and information of Petracola la-

bioocularis. We are also thankful to D. Quirola and J.C.

Chavez for taking photographs of preserved specimens.

Collecting permits were issued by the Instituto Nacional

de Recursos Naturales (110-2007-INRENA-IFFS-DCB),

Lima, Peru.

Literature Cited

Castoe TA, Doan TM, Parkinson CL. 2004. Data Parti-

tions and Complex Models in Bayesian Analysis: The

Phylogeny of Gymnophthalmid Lizards. Systematic

Biology 53(3): 448-469.

Doan TM, Castoe TA. 2005. Phylogenetic taxonomy of

the Cercosaurini (Squamata: Gymnophthalmidae),

with new genera for species of Neusticurus and Proc-

toporus. Zoological Journal of the Linnean Society

143: 405-416.

Fenwick AM, Gutberlet Jr. RL, Evans JA, Parkinson CL.

2009. Morphological and molecular evidence for phy-

logeny and classification of South American pitvipers,

genera Bothrops, Bothriopsis, and Bothrocophias

(Serpentes: Viperidae). Zoological Journal of the Lin-

nean Society 156(3): 617-640.

Harvey MB, Ugueto GN, Gutberlet Jr. RL. 2012. Review

of Teiid Morphology with a Revised Taxonomy and

Phylogeny of the Teiidae (Lepidosauria: Squamata).

Zootaxa 3459: 1-156.

Hedges SB, Conn CE. 2012. A new skink fauna from Ca-

ribbean islands (Squamata, Mabuyidae, Mabuyinae).

Zootaxa 3288: 1-244.

Jadin RC, Burbrink FT, Rivas GA, Vitt LJ, Barrio-

Key to species of Petracola

l.a. Supralabial-subocular fusion and precloacal pores present P. labioocularis

1. b. Supralabial-subocular fusion and precloacal pores absent 2

2. a. Superciliary series complete, genials three, and venter pale with scattered dark brown spots P. waka

2. b. Superciliary series incomplete, genials two, and venter pale with black bands or spots 3

3. a. Loreal absent, 2-5 femoral pores per hind limb in males and venter pale with bold black transverse band P. ventrimaculata

3.b. Loreal present, nine femoral pores per hind limb in males and venter pale with transverse rows of black spots. ..P. angustisoma

Amphib. Reptile Conserv.

November 201 5 I Volume 9 I Number 1 I el 07

A new species of Petracola from northern Peru

Fig. 6. Ventral view of waist of Petracola labioocularis (paratype, SMF 80113) showing the

precloacal and femoral pores. Scale bar = 1 mm. Photograph by G. Kohler.

Amoros CL, Guralnick RP. 2013. Finding arboreal

snakes in an evolutionary tree: phylogenetic place-

ment and systematic revision of the Neotropical bird-

snakes. Journal of Zoological Systematics and Evolu-

tionary Research 52(3): 257-264.

Kizirian D. 1996. A review of Ecuadorian Proctoporus

(Squamata: Gymnophthalmidae) with descriptions of

nine new species. Herpetological Monographs 10:

88-155.

Kizirian D, Bayefsky-Anand S, Eriksson A, Le M, Don-

nelly M. 2008. A new Petracola and a re-description

of P. ventrimaculata (Squamata: Gymnophthalmi-

dae). Zootaxa 1700: 53-62.

Kohler G, Lehr E. 2004. Comments on Euspondylus and

Proctoporus (Squamata: Gymnophthalmidae) from

Peru, with the description of three new species and

a key to the Peruvian species Herpetologica 60(4):

501-518.

Montanucci RR. 1973. Systematics and evolution of the

Andean lizard genus Pholidobolus (Sauria: Teiidae).

Miscellaneous Publications of the Museum of Natural

History University of Kansas 59: 1-52.

Olson DM, Dinerstein E, Wikramanayake ED, Burgess

ND, Powell GVN, Underwood EC, D’amico JA, It-

oua I, Strand HE, Morrison JC, Loucks CJ, Allnutt TF,

Ricketts TH, Kura Y, Lamoreux JF, Wettengel WW,

Hedao P, Kassem KR. 2001. Terrestrial Ecoregions of

the World: A New Map of Life on Earth. BioScience

51(11): 933-938.

Uzzell TM. 1970. Teiid lizards of the genus Proctoporus

from Bolivia and Peru. Postilla 142: 1-39.

APPENDIX I

Petracola ventrimaculata. — PERU: C A JAMARC A DEPARTMENT: Celendin province: La Victoria 6° 51’ 20. 11”S,

78° 18’ 25.40”W, 3,449 m, MUSM 26352, 26354-56, 26359; Conicorgue 6° 50’ 0.29”S, 78° 19’ 13.40”W, 3,573

m, MUSM 26238-43, 26245, 26353, 26259; Tingo (Sorochuco district) 6° 55’ 33.16”S, 78° 18’ 59.16”W, 3,410 m,

26357, 27297, 27300, 6° 56’ 1.611” S, 78° 20’ 34.4862”W, 3,550 m, MUSM 26230; La Chorrera (Sorochuco district)

6° 55’ 52.30”S, 78° 18’ 55.02”W, 3,267-3,835 m, MUSM 26229, 27295, 27299, 27301; La Encanada 6° 56’ 17.02”S,

78° 19’ 18.71”W, 3,267 m, MUSM 27298, 27302, 27303; Cerro Las Gordas (Hualgayoc district), MUSM 29879-80;

Chugunnayo 6° 53’ 35.15”S, 78° 18’ 44.17” W, 3,420 m, MUSM 26231-32; Puente Hierba Buena 6° 58’ 48.73”S, 78°

22’ 44.62”W, 3,360 m, MUSM 26233-35; Jadibamba 6° 50’ 17.20”S, 78° 22’ 58.54”W, 3,560 m, MUSM 26236; Bajo

Quengorio 6° 50’ 28.59”S, 78° 25’ 52.17” W, 3,410 m, MUSM 26237; Sorochuco y Huasmin 6° 58’ 38.98”S, 78° 22’

46.0 1”W, 3,576 m, MUSM 29504-07, 7° V 7.93”S, 78° 18’ 46.90”W, 3,466 m, MUSM 29508; NO DATA, MUSM

26260, 26261, 26262; Pampa Grande 7° V 42.30”S, 78° 17’ 52.69”W, 3,892m, CORBIDI 09239; Michiquillay 7° V

41.69”S, 78° 18’ 56.09”W, 3,817 m, CORBIDI 9247; Cajamarca province: Laguna Norte 6° 43’ 42.8874 S, 78° 42’

57.311” W, 3,593 m; CORBIDI 03630; PIURA DEPARTMENT: Huancabamba province: El Tambo 5° 21’ 40.39”S,

79° 33’ 9.72”W, 2,981 m, CORBIDI 10482-91, 10498, 10503, 11268-73; 2,700 m, MUSM 15452-56; AMAZONAS

DEPARTMENT: Chachapoyas province: Barro Negro 6°42’58.2”S, 77°51’53.8”W, 3,290 m, CORBIDI 11067-75,

11104-26; Leimebamba 6°44’48”S, 77°47’11.6”W, 2,799 m, CORBIDI 11010.

Amphib. Reptile Conserv. 32 November 201 5 | Volume 9 | Number 1 | el 07

Echevarria and Venegas

Lourdes Y. Echevarria graduated in biological sciences from Universidad Nacional Agraria La Molina, Lima,

Peru, in 2014. As a student, she collaborated constantly in the order and management of the herpetological

collections of Centro de Omitologia y Biodiversidad, Lima, developing a great interest in reptiles, especially

lizards. For her undergraduate thesis, Lourdes worked on the “Review of the current taxonomic status of Pe-

tracola ventrimaculata (Cercosaurini: Gymnophthalmidae) using morphological and ecological evidence.” She

worked as a researcher of the Museo de Zoologia (QCAZ), Pontificia Universidad Catolica del Ecuador in

Quito during 2015. This manuscript is her first lizard species description. Lourdes is preparing a monograph on

the systematics of the Petracola ventrimaculata complex based on the results of her undergraduate thesis, as

well as other papers about taxonomy of lizards and snakes.

Pablo J. Venegas graduated in Veterinary Medicine from Universidad Nacional Pedro Ruiz Gallo, Lambayeque,

Peru, in 2005. He is currently curator of the herpetological collection of Centro de Ornitologia y Biodiversidad

(CORBIDI). He worked as a researcher of the Museo de Zoologia QCAZ, Pontificia Universidad Catolica del

Ecuador in Quito during 2015. His current research interest is focused on the diversity and conservation of the

Neotropical herpetofauna with an emphasis on Peru and Ecuador. He has published more than 40 scientific

papers on taxonomy and systematics of Peruvian and Ecuadorian amphibians and reptiles.

In accordance with the International Code of Zoological Nomenclature new rules and regulations (ICZN 2012), we have deposited this paper in publicly acces-

sible institutional libraries. The new species described herein has been registered in ZooBank (Polaszek 2005a, b), the official online registration system for the

ICZN. The ZooBank publication LSID (Life Science Identifier) for the new species described here can be viewed through any standard web browser by append-

ing the LSID to the prefix “http://zoobank.org/.” The LSID for this publication is: um:lsid:zoobank.org:pub:FlC772A4-E580-4EAA-9BD8-E8A83D276743.

Separate print-only edition of paper(s) (reprint) are available upon request as a print-on-demand service. Please inquire by sending a request to: Amphibian &

Reptile Conservation, amphibian-reptile-conservation.org, arc.publisher@gmail.com.

Amphibian & Reptile Conservation is a Content Partner with the Encyclopedia of Life (EOL), http:///www.eol.org/ and submits information about new species

to the EOL freely.

Digital archiving of this paper are found at the following institutions: ZenScientist, http://www.zenscientist.com/index.php/filedrawer; Ernst Mayr Library, Mu-

seum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (USA), http://library.mcz.harvard.edu/emst_mayr/Ejoumals/ARCns.

The most complete journal archiving and journal information is found at the official ARC journal website, amphibian-reptile-conservation.org. In addition,

complete journal paper archiving is found at: ZenScientist, http://www.zenscientist.com/index.php/filedrawer.

Citations

ICZN. 2012. Amendment of Articles 8,9,10,21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods of publication.

Zootaxa 3450: 1—7.

Polaszek A et al. 2005a. Commentary: A universal register for animal names. Nature 437: All .

Polaszek A et al. 2005b. ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological Nomenclature 62(4):

210 - 220 .

Amphib. Reptile Conserv.

November 201 5 I Volume 9 I Number 1 I el 07

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [Special Section]: 34-44 (el 09).

Publication LSID:

urn:lsid:zoobank.org:pub:EBC93EDE-B09B-407C-92DE-966BBD2CC2A2

A new species of montane gymnophthalmid lizard, genus

Cercosaura (Squamata: Gymnophthalmidae), from the

Amazon slope of northern Peru

Gourdes Y. Echevarria, 2 Andy C. Barboza, and 3 Pablo J. Venegas

1 " ' Division de Herpetologia-Centro de Ornitologia y Biodiversidad (CORBIDI), Santa Rita N°105 36 Of. 202, Urb. Huertos de San Antonio, Surco,

Lima, PERU l i Museo de Zoologia, Escuela de Ciencias Biologicas, Pontificia Universidad Catolica del Ecuador, Avenida 12 de Octubre 1076 y

Roca, Apartado 17-01-2184, Quito, ECUADOR

Abstract . — Based on morphological and previously published molecular and phylogenetic evidence,

we report the discovery of a new species of Cercosaura from the northern portion of Cordillera

Central, northern Peru. The new species inhabits the montane forests of northeastern Peru at

elevations between 1,788-1,888 m. It differs from other species of Cercosaura by having the dorsum

lighter than flanks, a white labial stripe that continues along the ventrolateral region until the hind

limb insertion, subdigital lamellae on toes not tuberculate, 6-8 longitudinal rows of ventral scales,

32-36 transverse rows of dorsal scales, and dorsal surface of forelimbs and fingers dark brown.

Key words. Central Andes, collar scales, Peruvian Yungas, San Martin, systematics, new species, lizard, South Amer-

ica

Citation: Echevarria LY, Barboza AC, Venegas PJ. 2015. A new species of montane gymnophthalmid lizard, genus Cercosaura (Squamata: Gymn-

ophthalmidae), from the Amazon slope of northern Peru. Amphibian & Reptile Conservation 9(1 ): 34^4 (el 09).

Copyright: © 201 5 Echevarria et al. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial-

NoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation', official journal website <amphibian-

reptile-conservation. org> .

Received: 03 September 2015; Accepted: 01 December 2015; Published: 16 December 2015

Introduction

The New World lizard clade Gymnophthalmidae Merrem,

1820 comprises 248 extant species belonging to 47 taxa

ranked as genera (Colli et al. 2015; Uetz 2015). Gymn-

ophthalmid lizards are small, often secretive and many

species have fossorial or semi-fossorial habits (Colli et al.

1998; Pianka and Vitt 2003; Mesquita et al. 2006). They

are primarily limited to tropical latitudes, present in the

lowland Amazonian forest and foothills, valleys and hill-

sides of the Andes (Presch 1980). There are also species

that inhabit the Quaternary sand dunes in the Sao Fran-

cisco Basin in Brazil (Rodrigues 1996) to high elevations

in the Andes, such as Proctoporus bolivianus that can be

found at 4,080 m elevation in the Peruvian Andes (Du-

ellman 1979).

The genus Cercosaura Wagler, 1830 was reviewed by

Doan (2003) using morphological data in a phylogenetic

analysis, as a result the genus was redefined to include

the genera Pantodactylus and Prionodactylus, a view that

was corroborated by subsequent molecular studies (Cas-

toe et al. 2004; Doan and Castoe 2005). With the taxo-

nomic changes by Doan (2003), the genus Cercosaura

contained 1 1 species distributed throughout the Amazo-

nian, Guianan, and savannah regions of South America,

occurring in all its countries, except for Chile, and ex-

tending into Panama. Some years later, Pantodactylus

steyeri was assigned to Cercosaura (Bernils et al. 2007),

and C. hypnoides Doan and Lamar 2012 was described,

increasing the number of Cercosaura species to 13 (Uetz

2015). Recently, Torres-Carvajal et al. (2015) presented

a molecular phylogeny of Cercosaura and related taxa,

“ Cercosaura ” vertebralis and “ Cercosaura ” dicra were

found to be nested within Pholidobolus , and therefore

referred to that genus. Furthermore, their phylogenetic

hypothesis supported the recognition of C. argula and

C. oshaughnessyi as different species (Avila Pires 1995),

and of C. ocellata bassleri, as separate species (i.e., C.

bassleri and C. ocellata). Therefore 13 species of Cer-

cosaura are currently recognized: C. argulus Peters,

Correspondence. Email: 1 lourdese.20@gmail.com (Corresponding author); 2 andy_barboza@yahoo.es\ ~sancarranca@yahoo.es

Amphib. Reptile Conserv.

December 2015 I Volume 9 I Number 1 I el 09

Echevarria et al.

1863; C. bassleri (Ruibal, 1952); C. eigenmanni (Griffin,

1917); C. hypnoides Doan and Lamar, 2012; C. manicata

O’shaughnessy, 1881; C. nigroventris (Gorzula and Se-

naris, 1999); C. ocellata Wagler, 1830; C. oshaughnessyi

(Boulenger, 1885); C.parkeri (Ruibal, 1952); C. phelpso-

rum (Lancini, 1968); C. quadrilineata (Boettger, 1876);

C. schreibersii Wiegmann, 1834; and C. steyeri (Tedesco,

1998). It should be highlighted that the inclusion of spe-

cies such as C. hypnoides, C. nigroventris, C. parkeri, C.

phelpsorum, and C. steyeri need to be confirmed by a ro-

bust phylogeny that include the mentioned taxa.

Recent fieldwork in the montane forests of San Martin

department, northeastern Peru, resulted in the discovery

of a potentially new species of Cercosaura, which was

later confirmed after examination of Cercosaura manica-

ta manicata and C. manicata boliviana specimens and its

position in Torres-Carvajal et al. (2015) phylogeny.

Materials and Methods

Specimens were fixed in 10% formalin for 24 hours, and

later stored in 70% ethanol. All type specimens were de-

posited in the herpetological collection of the Centro de

Omitologia y Biodiversidad (CORBIDI), Lima, Peru.

Other species of Cercosaura examined in this study are

deposited at CORBIDI and the Museo de Zoologia Pon-

tificia Universidad Catolica del Ecuador, Quito (QCAZ),

and are listed in Appendix I. Scale counts and color pat-

tern information for Cercosaura argulus, C. eigenman-

ni, C. oshaughnessyi, and C. ocellata were taken from

Avila-Pires (1995); C. quadrilineata, C. schreibersii, and

C. phelpsorum from Doan (2003); and C. parkeri from

Soares-Barreto et al. (2012) and C. steyeri from Tedesco

(1998). Snout- vent length (SVL) and tail length (TL) mea-

surements were taken to 1 mm with a ruler. For characters

recorded on both sides, condition on the right side is pre-

sented first. Egg volume was calculated using the fonnula

for a prolate spheroid V= 4/3 tc (length/2) (width/2) 2 . Sex

was determined by dissection or by noting the presence

of hemipenes. We follow the terminology and general de-

scription format of Doan and Lamar (2012).

Results

Cercosaura doanae sp. nov.

urn:lsid:zoobank.org:act:3810814C-BA99-418B-9612-0AllB4C72450

Cercosaura sp. Torres-Carvajal et al., 2015: 282 (see dis-

cussion).

Figs. 1-3, 4 (upper), 5A, 6C.

Holotype: CORBIDI 00651, adult male from Laguna

Negra (06°53’29.3”S, 77°23’18.3”W; WGS 84), 1,788

m, Mariscal Caceres Province, San Martin Department,

Peru, collected by P.J. Venegas and D. Vasquez on 3 Feb-

ruary 2008.

Fig. 1. Holotype (CORBIDI 00651; SVL = 52 mm) of Cerco-

saura doanae sp. nov. in dorsal (upper) and ventral (bottom)

views. Photographs by G. Chavez.

Fig. 2. Head of the holotype (CORBIDI 00651) of Cercosaura

doanae sp. nov. in lateral (A), dorsal (B), and ventral (C) views.

Photographs by G. Chavez.

Amphib. Reptile Conserv.

December 201 5 I Volume 9 I Number 1 I el 09

A new species of montane gymnophthalmid lizard, genus Cercosaura

Fig. 3. Holotype of Cercosaura doanae sp. nov. in life: dorso-

lateral (upper) and ventral (bottom) views. Photographs by P.J.

Venegas.

Paratypes (19): PERU: San Martin Department: Mar-

iscal Caceres Province: from type locality CORBIDI

00649, 00656, 00658, 00659 adult females, CORBIDI

00650, 00652, 00660, 00662 juveniles, CORBIDI 00663,

00654, 00655, 00657, 00661 adult males, CORBIDI

00653 subadult female, collected between 2-4 February

2008, by P.J. Venegas and D. Vasquez; Anasco Pueblo

(06°50’11.6”S, 77°29’09.7”W), 1,888 m, CORBIDI

00648 a juvenile collected on 2 February 2008 by P.J

Venegas and D. Vasquez; Fajasbamba (06°44’48.4” S,

77°38’25.6” W), 1,814 m altitude, CORBIDI 15074 adult

female, CORBIDI 15075 juvenile female, CORBIDI

15076 juvenile male, CORBIDI 15088 adult male col-

lected between 25-28 October 2014 by F.Y. Echevarria

and A.C. Barboza.

Diagnosis: Cercosaura hypnoides from the Amazon

slope of Colombia (Doan and Famar 2012), C. manicata

manicata from the Amazon slope of Ecuador and cen-

tral Peru, and C. manicata boliviano Werner, 1899 from

southern Peru and Bolivia (Uzzel 1973) are the most sim-

ilar species to C. doanae by having the dorsum lighter

than flanks and a light labial stripe. Nevertheless, Cerco-

saura doanae can be distinguished from C. hypnoides by

having 6-7 supralabials (5 in C. hypnoides ), dorsal scales

in transverse rows (transverse and oblique rows in C. hyp-

noides ■), and 0-3 lateral scale rows (4-7 in C. hypnoides ).

The new species can be distinguished from C. manicata

boliviano Werner 1899 (character state of C. manicata

boliviano in parenthesis) by having a cream labial stripe

beginning before the eye, on first or second supralabial,

continuing along the ventrolateral region up to hind limb

insertion (light labial stripe beginning under eye and end-

ing before collar fold; Fig. 4, middle), two conspicuous

widened collar scales at midline (three or four enlarged

collar scales at midline; Fig. 5B), and three posterior cloa-

cal plates in males and five in females (four in males and

females). Furthermore, C. doanae differs from C. mani-

cata manicata (character state of C. manicata manicata in

parenthesis) by having subdigital lamellae on toes not tu-

berculate (tuberculate for entire length of toes); and dor-

sal surface of forelimbs dark brown (brown with a white

broad line on brachium, antebrachium, and fingers I, II,

III; Fig. 5C).

Furthermore, Cercosaura doanae is easily distin-

guished from C. argula, C. bassleri, C. eigenmanni, C.

ocellata, C. oshaughnessvi, C. parkeri , C. quadrilineata,

and C. schreibersii (Fig. 6) in having brown labials with

a white stripe extending from the first or second supral-

abial towards forelimb insertion. Cercosaura argula and

C. oshaughnessyi have labials white or light cream, C.

eigenmanni has brown labials with white broad vertical

bars, C. bassleri and C. ocellata have creamy gray or

gray labials with thin black vertical bars, C. parkeri has

creamy gray or white labials with dark or faint brown ver-

tical bars, C. quadrilineata and C. schreibersii have labi-

als varying from creamy gray or dirty cream to white with

dark flecks, spots or mottling. Additionally, C. doanae can

be distinguished from C. argula by having an undivided

frontonasal (divided in C. argula ), two genials (three),

single lamellae on fingers and toes (mostly divided),

32-36 transverse dorsal scale rows (38-45), 34-42 scales

around midbody (27-35), and venter pale orange (white);

from C. eigenmanni by having 34-42 scales around mid-

body (26-32 in C. eigenmanni ), and 9-12 femoral pores

in males (6-7); from C. bassleri and C. ocellata by having

hexagonal dorsal scales (quadrangular in C. bassleri and

C. ocellata ), scales on flanks slightly smaller than dorsals,

Fig. 4. Lateral views of male specimens of (upper) Cercosaura

doanae (holotype), (middle) C. manicata boliviano (CORBIDI

16500), and (bottom) C. manicata manicata (CORBIDI 08797).

Photographs by J. C. Chdvez-Arribasplata.

Amphib. Reptile Conserv. 36 December 201 5 | Volume 9 | Number 1 | el 09

Echevarria et al.

Fig. 5. Ventral views of heads and dorsal surface of the fore-

limbs of (A) Cercosaura doanae sp. nov. (holotype), (B) C.

manicata boliviana (CORBIDI 14272), and (C) C. manicata

manicata (CORBIDI 08797); showing the collar scales and the

striking white line along the brachium, antebrachium, and fin-

gers I, II, III of C. manicata manicata. Red arrows indicate the

collar scales at midline. Photographs by D. Quirola and J.C.

Chavez-Arribasplata.

keeled (scales on flanks distinctly smaller than dorsals,

smooth or slightly keeled), lamellae on toes single (most-

ly divided), 10-13 lamellae under fourth finger (14-18),

and 15-18 lamellae under fourth toe (16-24); from C. os-

haughnessyi by having a single frontonasal (divided in C.

oshaughnessyi ), 32-36 transverse dorsal scale rows (37-

52), scales on flanks slightly smaller than dorsals (scales

on flanks distinctly smaller and sharply delimited from

dorsals and ventrals), and venter orange (white); from C.

parkeri by having 34-42 scales around midbody (24-30

in C. parkeri), 9-12 femoral pores in males (2-6), and

lateral scales slightly smaller than dorsals (lateral scales

similar in size to dorsals); from C. quadrilineata by hav-

ing 6-8 longitudinal rows of ventral scales (four in C.

quadrilineata), 16-19 transverse rows of ventral scales

(21-23), and 9-12 femoral pores in males (eight); from

C. schreibersii by having 16-19 transverse ventral scale

rows (17-24 in C. schreibersii), and 9-12 femoral pores

in males (3-5).

Cercosaura doanae can be distinguished from both

C. nigroventris and C. phelpsorum by having subdigital

lamellae on toes not tuberculate (tuberculate in C. nigro-

ventris and C. phelpsorum) and by ventral coloration in

preservative, having creamy tail (beige and dark brown in

C. nigroventris and C. phelpsorum, respectively) (Doan

2003).

The new species differs from the poorly known Cerco-

saura steveri in having 6-8 longitudinal rows of ventral

scales (four in C. steveri), dorsal scales not mucronate

(strongly mucronate), 34-42 scales around midbody (17),

and 15-18 lamellae on Toe IV (14).

Pholidobolus hillisi and the former Cercosaura spe-

cies, P. dicra and P. vertebralis (see Torres-Carvajal et

al. 2015), are also very similar to C. doanae in having

dorsum lighter than flanks, brown labials with a white or

light cream labial stripe that extends towards the fore-

limb insertion, and hexagonal and strongly keeled dorsal

scales. However, the new species can be readily distin-

guished from all these Pholidobolus species by lacking a

light vertebral stripe, which in P dicra bifurcates anteri-

orly at midbody, and by having the loreal scale in contact

with supralabials (in the aforementioned species of Pholi-

dobolus the loreal scale is not in contact with supralabi-

als). Additionally, it can be distinguished from P. hillisi

(in parenthesis) by lacking a distinct diagonal white stripe

on each side of the chin, extending from the fourth genial

to the forelimb (present); from P dicra (in parenthesis)

by having three supraoculars (four); and from P. verte-

bralis (in parenthesis) by having palpebral disc single or

divided, usually into 2-3 scales (divided, into 5-8 scales).

Characterization: (1) supraoculars three; (2) supercili-

aries 3—4, first expanded onto dorsal surface of head; (3)

palpebral eye-disc made up of a single or divided trans-

parent scale; (4) supralabials 6-7; (5) infralabials 5-7; (6)

dorsal body scales hexagonal, strongly keeled; (7) dorsal

scales only in transverse rows; (8) transverse rows of dor-

sals 32-36; (9) a continuous series of small lateral scales

(usually two) separating dorsals from ventrals; (10) two

conspicuous widened collar scales at midline; (11) trans-

verse rows of ventrals 16-19; (12) longitudinal rows of

ventrals 6-8; (13) femoral pores per hind limb 9-12 in

males, 0-9 in females; (14) precloacal pores absent; (15)

posterior cloacal plates three in males, five in females;

(16) subdigital lamellae on toe IV 15-18; (17) limbs

overlapping when adpressed against body; (18) dorsum

brown; light brown dorsolateral stripe present, extending

from loreal onto the tail; flanks chocolate brown, darker

than dorsum; cream labial stripe, starting on second or

third supralabial and running continuously as a ventrolat-

eral creamy brown stripe along the body; black ocelli with

white or creamy brown center present along the flanks;

(19) ventrally throat and neck pinkish gray; chest yellow-

ish orange; venter yellowish orange in males and pink-

ish brown in females, ventral surface of hind limbs and

ventral surface of tail orange, becoming grayish cream

towards the tip; and ventral surface of forelimbs yellow.

Description of holotype: Adult male (CORBIDI

00651), hemipenes not everted, SVL = 52 mm, tail com-

plete, TL = 108 mm; head scales rugose and porous; ros-

tral scale wider than tall, meeting supralabials on either

Amphib. Reptile Conserv.

December 201 5 I Volume 9 I Number 1 I el 09

A new species of montane gymnophthalmid lizard, genus Cercosaura

Fig. 6. Some species of Cercosaura : (A) C. argula (CORBIDI 12634) from Ere river, Loreto, Peru; (B) C. bassleri (CORBIDI

13208) from Bahuaja Sonene National Park, Puno, Peru; (C) C. doanae sp. nov. (CORBIDI 661) from Laguna Negra, San Martin,

Peru; (D) C. eigenmanni from Porto Yelho, Rondonia, Brazil; (E) C. manicata manicata (CORBIDI 9217) from Cordillera Azul

National Park, San Martin, Peru; (F) C. manicata boliviana (CORBIDI 16500) from San Pedro, Cusco, Peru; (G) C. ocellata from

Para, Brazil; (H) C. oshaughnessyi (CORBIDI 12637) from Ere river, Loreto, Peru; and (I) C. schreibersii from Ipero, Sao Paulo,

Brazil. Photographs by: A-C, E, and H by P.J. Venegas; F by A. Catenazzi; G by P Melo-Sampaio; D and I by M. Teixeira-Junior.

side at above the height of supralabials, in contact with

frontonasal, nasals, and first supralabials; frontonasal

wider than long, hexagonal, in contact with nasals and

prefrontals, shorter than frontal; prefrontals present, not

in contact; frontal longer than wide, hexagonal, in con-

tact with anteriormost supraocular, prefrontals, and fron-

toparietals; frontoparietals pentagonal, in contact with

all three supraoculars, parietals and interparietal; supra-

oculars three, first supraocular in contact with first three

superciliaries, second supraocular in contact with third

superciliary, third supraocular in contact with fourth su-

praciliary, one postocular, and parietal; interparietal lon-

ger than wide, heptagonal, in contact with parietals and

occipitals; parietals hexagonal, in contact with one post-

ocular, a subequally large supratemporal, and one occipi-

tal; occipitals three, smaller than parietals, median one

smallest, extending further posteriorly than two lateral

occipitals. Nasal longer than high, nostril situated anteri-

orly, in contact with first and second supralabials and lo-

real; loreal irregularly pentagonal, in contact with second

supralabial, frenocular, and first superciliary; frenocular

subtriangular, dorsal most comer in contact with first su-

perciliary, in contact with second and third supralabials,

preocular, and first subocular; four superciliaries, first ex-

panded onto dorsal surface of head; two preoculars (right)

and one (left), in contact with first superciliary in both

sides; palpebral eye-disc made up of a single (right) and

divided into two transparent scales (left); suboculars five;

postoculars two; temporal rugose polygonal; supratempo-

rals two, first largest; ear opening oblong, tympanum re-

cessed; supralabials seven; infralabials six. Mental wider

than long, in contact with first infralabial and postmental

posteriorly; postmental single, pentagonal, posterior su-

ture angular with point directed posteriorly, in contact

with first and second infralabials and first pair of genials;

two pairs of genials, anterior pair in contact with second

and third infralabials, second genials in contact with third

and fourth infralabials; two pairs of chin shields, sepa-

Amphib. Reptile Conserv.

December 201 5 I Volume 9 I Number 1 I el 09

Echevarria et al.

rated by irregular pregulars; four rows of pregular scales;

gular scale rows three; collar fold distinct; lateral neck

scales granular.

Dorsals hexagonal, longer than wide, with posterior

margins slightly curved, imbricate, with a single high,

rounded keel, in 34 transverse rows, oblique rows absent;

longitudinal dorsal scale rows 25 at fifth transverse ven-

tral scale row, 27 at tenth transverse ventral scale row, 17

at fifteenth transverse ventral scale row; small, slightly

keeled lateral scale series, two scales wide, approximately

half the size of dorsals, ovoid, smaller and more numer-

ous rounded lateral scales at limb insertion regions; lat-

eral fold present. Ventral scales smooth, squarish with

rounded posterior margins, imbricate, in 17 transverse

rows; longitudinal ventral scale rows at midbody six;

anterior precloacal plate paired, three scales on posterior

precloacal plate. Scales on tail like those on body; dorsal

and dorsolateral caudal scales hexagonal and keeled, ven-

tral and ventrolateral caudal scales square, smooth.

Limbs pentadactyl; digits clawed; dorsal brachial

scales polygonal, subequal in size, imbricate, keeled; ven-

tral brachial scales much smaller than dorsal scales, round,

juxtaposed, smooth; dorsal antebrachial scales polygonal,

subequal in size, multicarinate; ventral antebrachial scales

polygonal, subequal in size, smooth. Scales on dorsal sur-

face of manus polygonal, smooth, subimbricate; scales

on palmar surface of manus small, rounded, subimbri-

cate, domelike; thenar scales two, smooth; finger length

formula IV > III > II > V > I; scales on dorsal surfaces

of fingers smooth, quadrangular, covering dorsal half of

digit, overhanging supradigital lamellae 4 on I, 6 on II,

8 on III, 9 on IV, 5 on V; subdigital lamellae 6 on I, 1 0/9

on II, 13/13 on HI, 14/14 on IV, and 9/9 on V. Scales on

anterodorsal surface of thigh large, polygonal, keeled, im-

bricate; scales on posterior surface of thigh small, round-

ed, subimbricate; scales on ventral surface of thigh large,

rounded, flat, smooth; femoral pores nine (right) and 10

(left); precloacal pores absent; scales on anterior surface

of crus polygonal, keeled, subimbricate, decreasing in

size distally; scales on anterodorsal surface of crus po-

lygonal, subimbricate, keeled; scales on ventral surface of

crus large, polygonal, smooth, flat, and imbricate. Scales

on dorsal surface of pes polygonal, keeled, subimbricate;

scales on palmar surface of pes small, rounded, subim-

bricate, domelike; toe length formula IV > III > V > II >

I; scales on dorsal surface of digits single, quadrangular,

smooth, of varying sizes, overhanging supradigital lamel-

lae 3 on I, 6 on II, 9 on III, 10 on IV, and 7 on V; subdigital

lamellae single and tuberculate along the toes, 6/6 on I,

10/11 on II, 15/16 on III, 18/18 on IV, and 15 on V; limbs

overlapping when adpressed against the body.

Coloration of holotype in life: Dorsal and lateral sur-

faces of head brown; a distinct cream stripe initiates on

the ventral most portion of second supralabial scale and

continues through all supralabials onto body, after second

supralabial the stripe bends dorsally across third supra-

Amphib. Reptile Conserv.

labial and to the top of supralabials 4, 5, 6, and 7, then

the stripe bends ventrally in a straight line to the bottom

of the auricular opening, onto the body as a ventrolateral

stripe. Dorsal surface of body brown. Faint light brown

dorsolateral stripes from first superciliary onto tail. Lat-

eral surface of body chocolate brown; lateral stripe from

head continues over forelimb insertion, ending at hind

limb insertion. Nine black ocelli with cream center, from

neck to hind limb insertion, and six ocelli on the base of

tail. Dorsal surface of forelimbs same color of dorsum,

with a faint ocellus of creamy brown center on dorsal sur-

face of ante brachium; dorsal surfaces of manus brown

with cream pigmentation on some scales. Dorsal surfaces

of hind limbs brown with an ocellus near hind limb inser-

tion and few creamy brown spots; dorsal surfaces of pes

brown with coppery brown pigmentation on some scales.

Dorsal tail coloration brown with scattered light brown

and black marks. Lateral surfaces of tail brown.

Ventral surface of head and neck pinkish gray. Venter

yellowish orange. Ventral surface of forelimbs yellow,

ventral surface of hind limbs pale orange with dark gray

flecks. Ventral surface of tail orange becoming cream to-

wards the end. Iris brown.

Coloration of holotype in preservative (ethanol

70 %) : Dorsum darker brown; faint light brown dorsolat-

eral stripes light gray and barely distinct from dorsum col-

oration; flanks lighter brown. Ventrally head, neck, chest,

and venter are dark gray and ventral surface of limbs and

tail dirty cream.

Variation: Measurements and scale counts of Cerco-

saura doanae are presented in Table 1. Supraoculars

usually three; one specimen (CORBIDI 00659) has two

supraoculars on left side and three on right side. Usu-

ally four superciliaries; only specimen CORBIDI 00662

presents three superciliaries on each side. Seven supra-

labials in most specimens; 6/6 in specimens CORBIDI

00662, 00648, and 15088 (15%), and 7/6 in specimens

CORBIDI 15075, 15076 (10%). Usually six infralabials;

5/4 in CORBIDI 15075, 5/5 in CORBIDI 00652, 00654,

15074, and 15076 (20%), 5/6 in CORBIDI 00656, 15088

(10%), 6/5 in CORBIDI 00648, 00662 (10%), and 6/7

in CORBIDI 00658. Specimen CORBIDI 00659 (5%)

has 3/2 postocular scales. Usually 3/3 supratympanic

temporals (50%); specimens CORBIDI 00649, 00652,

00655, 00658, 00661 have 4/4 (25%), specimens COR-

BIDI 15074, 16076, 15088 have 4/3 (15%), and COR-

BIDI 00653 has 3/4. Specimen CORBIDI 15075 has only

one pair of genials. Sexual dimorphism present in num-

ber of cloacal plates; male specimens have two anterior

and three posterior cloacal plates (88%), only CORBIDI

00654 has four anterior plates. Female specimens have

usually two anterior and five posterior cloacal plates

(67%); CORBIDI 00659 has four anterior plates and

CORBIDI 00649 has four posterior plates. Palpebral disc

transparent, undivided in specimens CORBIDI 00648,

December 2015 | Volume 9 | Number 1 | el 09

A new species of montane gymnophthalmid lizard, genus Cercosaura

Table 1 . Variation in scutellation, sexual dimorphism in SVL (mm), and color pattern of Cercosaura doanae sp. nov., C. manicata

manicata, and C. manicata boliviano . Range followed by mean ± standard deviation is given for quantitative characters if appli-

cable.

Character

Cercosaura doanae

(n = 20)

Cercosaura manicata

manicata (n = 15)

Cercosaura manicata

boliviana {n = 3)

Supraoculars

Superciliaries

3-4

3.95 ± 0.22

3-4

3.87 + 0.35

Genials

1-2

1.95 + 0.22

2-3

2.67 + 0.58

Supralabials

6-7

6.85 + 0.37

5-7

5.53 + 0.83

5-7

6.33 + 1.15

Infralabials

4-7

5.65 + 0.67

4-5

4.4 + 0.51

Transverse dorsal scale rows

32-36

33.1 + 1.07

34-41

37.33 + 2.41

35-40'

Longitudinal dorsal scale rows

22-30

24.8 + 1.88

29-35

32.67 + 1.91

26-27

26.33 + 0.58

Transverse ventral scale rows

16-19

17.4 + 0.82

17-21

19.13 + 1.06

19(18)'— 23

21 + 1.15

Longitudinal ventral scale

rows

6-8

7.9 + 0.45

6 '-8

Scales around midbody (at

10 th transverse ventral scale

row)

34-42

37.45 + 1.93

41-50

45.4 + 2.29

33-41'

Lateral scale rows

0-3

0-4

Femoral pores per hind limb

in males

9-12

10.5 + 1.19

10-14

11.83 + 1.33

Femoral pores per hind limb

in females

0-9

6 + 3.11

8-13

10.33 + 2.52

Posterior cloacal plates in

males

2-3

Posterior cloacal plates in

females

4-5

4.86 + 0.38

4-5

Lamellae on 4 th toe

15-18

16.7 + 0.86

15-17

15.73 + 0.8

19-23

Lamellae on 4 th finger

10-13

11.55+0.76

10-13

11.33 + 0.82

13-15

14 + 1

Maximum SVL in males (mm)

52.06

61.62

56'

Maximum SVL in females

(mm)

55.59

59.35

58 1

Collar scales at midline

Two conspicuous and

widened

Two conspicuous and

widened

Three or four, enlarged

Beginning and extent of labial

stripe

Before the eye, on first

or second supralabial,

continuing along the

ventrolateral region up

to hind limb insertion

First supralabial, continu-

ing along the ventrolateral

region up to hind limb inser-

tion

Under eye and ending be-

fore collar fold

Color on dorsal surface of

forelimbs

Dark brown

Brown with a white broad

line on brachium, antebra-

chium and fingers I, II, III

Brown

'Data from Uzzel (1973).

Amphib. Reptile Conserv.

December 201 5 I Volume 9 I Number 1 I el 09

Echevarria et al.

00652, 00653, 00660, 00662, 15074-76, 15088 (45%),

and divided in two or three sections in CORBIDI 00649,

00650, 00654-59, 00661, 00663(55%).

Dorsal coloration is identical in all specimens, includ-

ing juveniles. Faint ocelli, with white or creamy brown

center, on dorsal surface of antebrachium and few or

several creamy brown spots are present in adults and ju-

veniles. Ventral coloration of head and venter in males

vary from grayish pink and pale orange, respectively,

like in the holotype, to complete white throat and ven-

ter (CORBIDI 15088). Adult females differ from adult

males by having the ventral surface of head, throat, and

venter pinkish brown, and the ventral surface of tail yel-

low. Lateral ocelli present in male specimens CORBIDI

00651, 00654, 0657, 00661, 15088 (25%), and female

CORBIDI 00659 (5%). Ocelli on hind limb present in

two specimens (CORBIDI 15088 and holotype).

Distribution and natural history: Cercosaura doa-

nae is known only from three localities in the head wa-

ters of the Huayabamba basin, San Martin department, at

elevations of 1,788-1,888 m, along the Amazon slope of

the extreme northern portion of Central Andes in north-

ern Peru (Fig. 7). It inhabits the Amazonian premontane

forest in the Yungas ecoregion (500-2,300 m) accord-

ing to Brack (1986) and Penaherrera del Aguila (1989),

and Peruvian Yungas ecoregion according to Olson et al.

(2001). The new species was found active on sunny days

in pasturelands for cattle surrounded by forest in Anasco

Pueblo and Lajasbamba (Fig. 8A). All observed indi-

viduals were moving through the herbaceous vegetation,

and hiding in it when disturbed. At Laguna Negra, the

new species was very abundant, moving at day through

the leaf litter in primary forest (Fig. 8B). When disturbed,

individuals run and hid within leaf litter, fallen trees, and

in roots at the base of trees. Female specimen CORBIDI

00659, collected on 4 February 2008, contained two ovi-

ductal eggs, right egg was 12.1 mm x 6.6 mm and left

egg 12.3 mm x 6.7 mm, and their respective volumes as

279.86 mm 3 and 291.38 mm 3 .

Etymology: The specific epithet is a noun in the geni-

tive case and patronym for Tiffany Doan, in recognition

of her contribution to the systematics of gymnophthalmid

lizards (e.g., Doan 2003; Doan and Castoe 2005), and to

the knowledge of the herpetofauna from southern Peru.

Discussion

The Neotropical genus Cercosaura is a poorly sampled

taxa that surely has several undescribed species. In a

recent molecular phylogeny of Cercosaura and related

taxa, Torres-Carvajal et al. (2015) showed distinction be-

tween C. doanae sp. nov. (cited as Cercosaura sp.) and

C. manicata manicata (cited as C. manicata ) as sister

species with strong support (PP = 1.00), and separated

5'C‘vJ‘N -

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-a r-T

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75-Q nyw

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| 0- 250

| 2M-SW

|socl saoo

~! 1000 ■ 1500

_j ISM -2000

□ ZWH- 3 $W

31 2600. MOO

Q 3000-3600

I] 3600-^000

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Fig. 7. Distribution of Cercosaura doanae sp. nov. (circles), C.

manicata boliviano (green pentagons), C. manicata manicata

(blue triangles), and C. hypnoides (sky blue square). Red circle

indicates the type locality of the new species. Locality data

from the literature (Doan and Lamar 2012; Uzzel 1973) and

specimens deposited at Centro de Ornitologia y Biodiversidad

(CORBIDI) and Museo de Zoologia of Pontificia Universidad

Catolica del Ecuador.

by branches that are similar in length to other branches

separating sister species among Cercosaura. Addition-

ally, the position of both species within the Cercosaura

clade is strongly supported (PP = 1.00) as a basal sub-

clade. However, the genetic distance values between the

new species and C. manicata manicata are lower (12S =

0.015, 16S = 0.016, ND4 = 0.032, and c-mos = 0.004)

than the interspecific ranges reported by the authors. For

example, the genetic distance values between two largely

recognized species as C. eigenmanni and C. ocellata are

0.031 for 12S, 0.019 for 16S, 0.060 for ND4, and 0.007

c-mos. We are confident in the separation of C. doanae

sp. nov. and C. manicata manicata , since the morpholog-

ical differences between both species are clear (i.e., the

absence versus presence of tuberculate lamellae and the

coloration of forearms; see diagnosis above and Figure

5C), and support the phylogenetic distinction. Although,

samples of C. manicata boliviana are not included in the

phylogenetic tree of Torres-Carvajal et al. (2015), clear

differences can be recognized between the new species

and C. manicata boliviana (e.g., beginning of labial

stripe, size, and number of collar scales at midline and

the number of posterior cloacal plates; see diagnosis

above and Figures 4-5).

Amphib. Reptile Conserv.

December 201 5 I Volume 9 I Number 1 I el 09

A new species of montane gymnophthalmid lizard, genus Cercosaura

Uzzel (1974) gave clear differences between C. mani-

cata boliviana and C. manicata manicata, all of them

confirmed in the specimens examined by us. Both sub-

species can be easily distinguished from each other mor-

phologically (see Figures 4-6) and occur in allopatry (see

Figure 7). In fact, we consider that there is enough mor-

phological evidence to consider C. manicata boliviana as

a distinct taxa, but it needs to be confirmed with robust

molecular data. We believe that Cercosaura doanae sp.

nov. and both subspecies of C. manicata represent a spe-

cies complex.

Acknowledgments. — Echevarria, Barboza, and Ven-

egas are indebted to M. Salas and R. Wagter for logistic

support and companionship in the field. The expedition

in 2008 where PJV discovered the new species was fund-

ed by the simultaneous efforts of the non-governmental

organization UCUMARI and the Gobierno Regional de

San Martin (GORESAM). The expedition in 2014 where

LYE and ACB collected the rest of type specimens was

financed by UCUMARI with funds provided by Apen-

heul Primate Conservation Trust (Apenheul Primate

Park), Otterfonds IUCN-NL, and US Fish and Wildlife

Services (Critically Endangered Species Fund). This re-

search was funded by the Secretaria de Educacion Su-

perior, Ciencia, Tecnologia e Innovation del Ecuador

and Pontificia Universidad Catolica del Ecuador. Speci-

mens of Cercosaura doanae were collected with the

following permits: 110-2007-INRENA-IFFS-DCB and

1 1 8-2007-INRENA-IFFS-DCB .

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Fig. 8. Habitat of Cercosaura doanae sp. nov.: (A) landscape

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Appendix I

Specimens examined

Cercosaura manicata manicata . — ECUADOR: Provincia Pastaza: Campo Oglan (AgipOil), QCAZ 5793, 5821; Pablo

Lopez de Oglan Alto, QCAZ 11818; Campamento K10, Campo Villano (AgipOil), 1°28’32.12”S, 77°32’5.53” W,

QCAZ 11831. PERU: Departamento Loreto: Provincia Datem del Maranon: Pongo Chinim, 3° 6’ 46.8”S, 77° 46’

34.4” W, 365 m, CORBIDI 09406. Departamento San Martin: Provincia Picota: Puesto de Control 16 Chambirillo (PN

Cordillera Azul), 7°4’8.9”S, 76°’0’55.2”W, 1,122 m, CORBIDI 08796, 08797, 08836, 08837, 09217, 10419, 10421,

10422; rio Chambira, CORBIDI 03659; Shamboyacu, CORBIDI 06774.

Cercosaura manicata boliviano . — PERU: Departamento de Cusco: Capire 13° 25’ 22.27 “S, 70°54’ 12.16” W, 1,237

m, CORBIDI 14272; Pitumarca, 13° 55’ 5.64 “S, 71°0’43.81” W, 4,889 m, CORBIDI 14704; San Pedro, 13° 3’ 51.012

“S, 71°33’37.44” W, 1,560 m, CORBIDI 16500.

Pholidobolus dicra . — ECUADOR: Provincia Morona Santiago: Guarumales, 2° 34’ 0.0006” S, 78° 30’ 0” W, 1,700 m,

QCAZ 5292, 5304. Provincia Tungurahua: Rio Blanco, Via Banos-Puyo, 1° 23’ 55.6434” S, 78° 20’ 24” W, 1,600 m,

QCAZ 6936, no locality data QCAZ 8015.

Pholidobolus hillisi. — ECUADOR: Provincia Zamora-Chinchipe: near San Francisco Research Station on Loja-

Zamora road, 3°57’57”S, 79°4’45”W, WGS84, 1,840 m, QCAZ 4998-99, 5000; San Francisco Research Station,

3°58’ 14”S, 79°4’41”W, 1,840 m, QCAZ 6840, 6842, 6844.

Pholidobolus vertebralis . — ECUADOR: Provincia Carchi: Chilma Bajo, 0°51’53.83”N, 78°2’59.26”, W, 2,071 m,

QCAZ 5057, 8671-8673, 8678, 8679,8717, 8724, 0°51’50.31”N, 78°2’50.05” W, 2022, QCAZ 8684-8689. Provincia

Pichincha: Mindo, 0°3’2.41”S, 78°46’ 18.77” W, 1,700 m, QCAZ 2911, 2912, 2915, 0°4’40.98”S, 78°43’55.02” W,

l, 601 m, QCAZ 7528; Cooperativa El Porvenir, El Cedral 0°6’50.40”N, 78°34’ 11.75” W, 2297 m, QCAZ 5081, 5082;

Santa Lucia de Nanegal, 0°6’48.70”N, 78°36’48.60” W, 1,742 m, QCAZ 10667, 0°7’8.51”N, 78°35’58.70” W, 1900

m, QCAZ 10750.

Amphib. Reptile Conserv.

December 201 5 I Volume 9 I Number 1 I el 09

A new species of montane gymnophthalmid lizard, genus Cercosaura

Lourdes Y. Echevarria graduated in Biological Sciences from Universidad Nacional Agraria La Molina,

Lima, Peru, in 2014. As a student, she collaborated constantly in the order and management of the herpeto-

logical collection of Centro de Omitologia y Biodiversidad (CORBIDI) developing a great interest in reptiles,

especially lizards. For her undergraduate thesis, Lourdes worked on the “Review of the current taxonomic

status of Petracola ventrimaculatus (Cercosaurini: Gymnophthalmidae) using morphological and ecological

evidence.” She has continued to work as a researcher at the Museo de Zoologia (QCAZ), Pontificia Universidad

Catolica del Ecuador in Quito in 2015. Lourdes is preparing a monograph on the systematics of the Petracola

ventrimaculatus complex based on the results of her undergraduate thesis, as well as other papers about the

taxonomy of lizards and snakes.

Andy C. Barboza graduated in Biological Sciences from Universidad Nacional de Trujillo, La Libertad, Peru

in 2012. She works as collection manager of the amphibian collection of Centro de Omitologia y Biodivers-

idad (CORBIDI). For her undergraduate thesis she worked on the “Composition and altitudinal distribution of

amphibians from Otishi National Park,” in collaboration with Missouri Botanical Garden (GMB). Her current

research interest is focused on the systematics, diversity, and conservation of Neotropical herpetofauna, particu-

larly from Peru, and the evolutionary history and behavior of amphibians facing climate change.

Pablo J. Venegas graduated in Veterinary Medicine from Universidad Nacional Pedro Ruiz Gallo, Lam-

bay eque, Peru, in 2005. He is currently curator of the herpetological collection of Centro de Omitologia y

Biodiversidad (CORBIDI) and worked as a researcher at the Museo de Zoologia QCAZ, Pontificia Universidad

Catolica del Ecuador in Quito in 2015. His current research interest is focused on the diversity and conserva-

tion of the Neotropical herpetofauna with emphasis in Pern and Ecuador. So far he has published more than 40

scientific papers on taxonomy and systematics of Pemvian and Ecuadorian amphibians and reptiles.

Amphib. Reptile Conserv.

December 201 5 I Volume 9 I Number 1 I el 09

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [General Section]: 1-13 (e94).

A new species of Anolis (Squamata: Iguanidae) from Panama

1 3 Steven Poe, 2 Simon Scarpetta, and 1 Eric W. Schaad

department Of Biology, Castetter Hall, University of New Mexico, Albuquerque, New Mexico 87131, USA 2 Field Conservation Facility, 477 Oak

Road, Stanford University, Stanford, California 94305, USA

Abstract. — We describe Anolis elcopeensis, a new species of anole lizard from low to moderate

elevations of the Pacific slope of the Cordillera Central of central Panama. Anolis elcopeensis is

a close relative of and resembles the Amazonian species A. fuscoauratus but differs from it and

similar species mainly in body size, male dewlap color, and mitochondrial DNA. We estimate the

phylogenetic position of the new species relative to all species of Anolis, and analyze variation

in the mitochondrial COI gene among some populations of the new species. We also discuss the

mythical presence of Anolis fuscoauratus in Panama, document the possible occurrence of A.

maculiventris in Panama, and present preliminary evidence for multiple cryptic fuscoauratus- like

species in eastern Panama.

Key words. Central America, cryptic species, Darien, lizard, Panama, Reptilia

Citation: Poe S, Scarpetta S, Schaad EW. 2015. A new species of Anolis (Squamata: Iguanidae) from Panama. Amphibian & Reptile Conservation

9(1) [General Section]: 1-13 (e94).

NoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official journal website <amphibian-

reptile-conservation. org> .

Received: 16 Feburary 2014; Accepted: 20 April 2015; Published: 27 April 2015

Introduction

Panama continues to yield new species of lizards and

frogs annually despite already displaying one of the high-

est herpetofaunal diversities in Central America (Kohler

2008, 2011). Thirteen new species of Anolis have been

described from Panama since 2007 to bring the number

of Anolis species known from Panama to 44. The dy-

namic biogeographic history of Panama as a land bridge

between North and South America has been cited as an

explanation for the extraordinary faunal diversity of this

country (Savage 1983).

Herpetologists working in Panama have long known

of an undescribed species of Anolis similar to A. fuscoau-

ratus from Barro Colorado Island (BCI) and other areas

(Myers and Rand 1969; Ibanez et al. 1994; Stan Rand,

pers. comm. 2003). This abundant species has remained

undescribed probably due to its lack of distinctive char-

acteristics and resemblance to other nondescript anoles

in Central and South America. Anolis species similar to

A. fuscoauratus frequently are straightforwardly differ-

entiable only by male dewlap color. We have collected

numerous examples of this undescribed species from its

known localities and several new localities and have con-

firmed its uniqueness using molecular and morphologi-

cal data. We describe this form from material collected

west of the Panama Canal Zone and provisionally assign

the well-known Canal Zone populations to this species.

We also present evidence that this form is part of a com-

plex of central and eastern Panamanian species similar to

Anolis fuscoauratus.

Materials and Methods

We adopt the evolutionary species concept (Simpson

1961; Wiley 1978) and operationalize this concept by

identifying species based on consistent differences be-

tween populations. That is, we hypothesize that popula-

tions or sets of populations that are diagnosable by major

differences in the frequencies of traits are distinct evolu-

tionary lineages or species.

Measurements were made with digital calipers on

preserved specimens and are given in millimeters (mm),

usually to the nearest 0. 1 mm. Snout-vent length (SVL)

was measured from tip of snout to anterior margin of clo-

aca. Head length was measured from tip of snout to ante-

rior margin of ear opening. Femoral length was measured

Correspondence. Email: ~anolis@unm.edu (Corresponding author)

April 2015 | Volume 9 | Number 1 | e94

Amphib. Reptile Conserv.

Poe et al.

from midline of venter to knee, with limb bent at a 90°

angle. Head width was measured at the broadest part of

the head, between the posterolateral corners of the orbits.

Scale terminology and characters used mainly follow

standards established by Ernest Williams for species de-

scriptions of anoline lizards (e.g., Williams et al. 1995).

Museum abbreviations follow Sabaj Perez (2014). Type

specimens were deposited in the Museum of Southwest-

ern Biology (MSB) of the University of New Mexico.

Phylogenetic analyses

We first identified a hypothesized undescribed species

of Anolis based on a discovered population’s all-orange

male dewlap, small body size, and morphological and

genetic distinctness (see below). In order to determine

the phylogenetic position of the new species and iden-

tify appropriate species for comparison, we included the

putative new species in phylogenetic analyses of all rec-

ognized species in the genus Anolis as of 01 June 2014

(results not shown). We collected morphological and mi-

tochondrial COI data for the putative species and com-

bined these with existing data for multiple genes (Alfoldi

et al. 2012; the informal name “sunni” in supplementary

appendices refers to this form but is not listed in the pub-

lished paper) and additional new collected data. Prelimi-

nary phylogenetic analyses of this hypothesized species

and all known species of Anolis suggested this form to

be a member of a strongly supported clade of 14 Central

and South American species similar to A. fuscoauratus

(we henceforth refer to these species as the “fuscoaura-

tus group”). We analyzed this sample of 15 species with

two outgroups (A. carpenteri, A. polylepis ) using a par-

titioned Bayesian Analysis (Ronquist et al. 2012) with

one “mixed” GTR model with rate heterogeneity for the

DNA sequence data (24879 sites) and another “standard”

model for morphological data (46 characters). We ran the

analysis for 2,000,000 generations, sampling every 1,000

trees, and discarded the first 50% of samples as bumin.

100

fuscoauratus

tolimensis

elcopeensis

bocourti

gruuo

pseudokemptoni

fortunensis

kemptoni

100

maculiventris

medemi

100

antonii

mariarum

94 I

monteverde

tenorioensis

aitae

I carpenteri

I polylepis

Fig. 1 . Phylogenetic estimate of placement of A. elcopeensis sp. nov. based on Bayesian analysis of morphological and molecular

data. Numbers are clade credibility values.

April 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

e94

A new species of Anolis from Panama

We also performed phylogenetic analysis of the mito-

chondrial COI gene for multiple individuals of the puta-

tive new species and single individuals of close relatives

according to the above analysis (Appendix 1). Data were

from the Barcode of Life initiative (see www.barcode-

oflife.org for data collection techniques) and published

data. We used Partitionfinder (Lanfear et al. 2012) to

identify a best model for this gene and assigned “mixed”

GTR models with rate heterogeneity to each partition.

We ran the analysis for 2,000,000 generations, sampling

every 1,000 trees, and discarded the first 50% of samples

as bumin.

Statistical analyses

Among geographically proximal forms, the new species

is most similar to Anolis gruuo in external morphology

(see below). In order to test the distinctiveness of the

hypothesized new species relative to this form, we per-

formed a discriminant function analysis of 14 individuals

of the new species and three individuals of A. gruuo us-

ing 10 characters of scalation, with a-priori grouping of

individuals as either A. gruuo (individuals from Santa Fe,

Veraguas, Panama; Lotzkat et al. 2012) or the putative

new species (individuals from near El Cope, Penonome,

and El Valle; Code; see below; Table 1). Characters in-

corporated were number of scales across the snout at the

second canthals (snsc), number of scales between supra-

orbital semicircles (sosc), number of scales between in-

terparietal and supraorbital semicircles (ipsosc), number

of postrostrals (pr), number of postmentals (pm), number

of loreal rows (lorr), number of supralabials from rostral

to center of eye (sle), number of expanded lamellae on

fourth toe (lm), number of ventral scales counted longi-

tudinally in 5% of snout to vent length (v5), and number

of dorsal scales counted longitudinally in 5% of snout to

vent length (d5). We observed no consistent differences

in these traits between males and females and therefore

pooled sexes in the analysis.

Results

The phylogenetic analysis of fuscoauratus group Ano-

lis placed the putative new species with South Ameri-

can species A. fuscoauratus and A. tolimensis with weak

support (Fig. 1). There was strong support (probability =

85%) for the new species clustering with a clade separate

from species similar to A. altae (i.e., A. altae, A. mon-

teverde, A. tenorioensis ) within the fuscoauratus group

clade.

The Partitionfinder analysis of the COI dataset sug-

gested two partitions, one for amino acid positions one

100

kemptoni

elcopeensis Caimito

elcopeensis NW Penonome

elcopeensis Ell Valle

elcopeensis El Cope

cf elcopeensis Lake Bayano

fuscoauratus

— cf elcopeensis Pipeline Road

- gruuo

altae

— fortunensis

0.05

maculiventris

Fig. 2. Phylogenetic estimate of the mitochondrial COI gene for samples of A. elcopeensis and close relatives based on Bayesian

analysis. Numbers are clade credibility values.

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Amphib. Reptile Conserv.

e94

Poe et al.

Table 1 . Specimens and data used in Discriminant Function Analysis (svl listed for reference; not used in dfa). See text for abbre-

viations.

Species

Museum number

snsc

sosc

ipsosc

lorr

sle

svl

elcopeensis

MSB 95543

8.5

14.5

41.9

elcopeensis

MSB 95544

9.5

15.5

38.8

elcopeensis

MSB 95545

7.5

36.0

elcopeensis

MSB 95546

13.5

37.0

elcopeensis

MSB 95547

5.5

9.5

35.5

elcopeensis

MSB 95548

7.5

33.5

elcopeensis

MSB 95549

41.8

elcopeensis

MSB 95550

4.5

41.9

elcopeensis

MSB 95551

8.5

42.1

elcopeensis

MSB 95552

5.5

14.5

39.2

elcopeensis

MSB 95554

38.0

elcopeensis

MSB 95555

13.5

42.3

elcopeensis

MSB 95556

41.4

elcopeensis

MSB 95557

13.5

44.5

A. elcopeensis mean

12.1

1.9

3.6

6.4

4.9

8.3

5.5

11.4

8.9

14.2

39.6

gruuo

POE 1626

4.5

9.5

13.5

45.0

gruuo

POE 1627

8.5

43.1

gruuo

POE 1628

4.5

43.1

A. gruuo mean

9.3

2.0

3.7

6.3

4.7

8.7

6.7

8.0

9.0

14.5

43.7

and two and a second partition for position three. The

resulting tree (Fig. 2) shows substantial divergence of the

putative new species. A mitochondrial clade composed

of individuals from Parque Omar Torrijos, El Valle, and

two localities near Penonome is minimally divergent

among populations (0.2-1. 4% p distances) and at least

7.3% divergent from other included Anolis species. We

consider these populations, which are uniform in male

dewlap color, to inhabitat the holotype and paratype lo-

calities of our putative new species and we describe this

form below. We also discuss the other two samples (from

Pipeline Road near the Panama Canal Zone and Lake

Bayano) that show substantial divergence from included

Anolis.

The discriminant function analysis accurately clas-

sified 100% of A. gruuo and 100% of the putative new

species.

Systematics

Anolis elcopeensis sp. Nov

urn:Isid:zoobank.org:act:9D828BDE-E151-48FE-91E7-E176E693B382

Figs. 3 A, 4A-B, 5 A, 6.

Holotype

MSB 95571, adult male from Parque Nacional G.D.

Omar Torrijos H., Code Province, Panama (8.66815,

-80.59267, 801 m), collected by Eric Schaad on 13 De-

cember 2008 from the trails near the visitor center.

Paratypes scored for traits

MSB 95570 bears the same data as the holotype. MSB

95550-2 bear the same locality data as the holotype, col-

lected by Steven Poe and Caleb Hickman in December

2003. MSB 95543-9, west of El Valle de Anton, road

ending at Chorro Las Mozas, Code Province, Panama

(8.859476, -80.14686, 570 m), collected by Poe and

Hickman in December 2003. MSB 95554-7, 95560-1,

same west of El Valle de Anton locality, collected by Poe,

Erik Hulebak, and Heather Maclnnes during 30 July-4

August 2004. MSB 95559, Hotel Campestre, El Valle de

Anton, Code Province, Panama (8.6129, -80.1251, 617

m), collected by Poe, Hulebak, and Maclnnes on 3 1 July

2004.

Additional topotypical paratypes

MSB 95569, 95572-9 bear the same data as the holotype,

collected by Schaad during December 2008.

Diagnosis

Anolis elcopeensis is a small grayish-brown anole with

smooth ventral scales and short limbs. We diagnose this

species relative to its 11 closest relatives (Fig. 1). Anolis

elcopeensis is unlikely to be confused with A. fuscoau-

ratus (Amazon basin; solid pink male dewlap), A. bo-

Amphib. Reptile Conserv.

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A new species of Anolis from Panama

Figure 3. A) Anolis elcopeensis (female, El Cope, Code, Panama). B) A. cf elcopeensis (male, south of Gamboa, Panama, Panama).

C) A. cf maculiventris (female, Yaviza, Darien, Panama).

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Figure 3 (continued). D) A. maculiventris (male, near Buenaventura, Colombia).

courti (Amazon basin; white male dewlap), A. tolimensis

(northeastern Andes; pink and orangish-red male dew-

lap), A. medemi (Gorgona Island, Colombia; pink and

orange male dewlap), A. antonii (northwestern Andes;

pink and reddish-orange dewlap), or A. mariarum (ex-

treme northwestern Andes; orange-red and yellow dew-

lap) based on geography. Nevertheless, A. elcopeensis

differs from each of these species in its solid orange male

dewlap color pattern.

Anolis elcopeensis is most easily distinguished from

the Central American members of its clade by male

dewlap color (Fig. 4; except for A. gruuo ) and smaller

body size (maximum SVL = 45 mm, n = 35): A. gruuo

(solid orange male dewlap; maximum SVL = 52 mm);

A. pseudokemptoni (red-orange anterior, pink posterior

male dewlap; maximum SVL = 55 mm); A. kemptoni

(red-orange anterior, pink posterior male dewlap; maxi-

mum SVL = 53 mm); A. fortunensis (red anterior, orange

posterior male dewlap; maximum SVL = 49 mm).

In the field A. elcopeensis is most likely to be confused

with A. gruuo and A. carpenteri, which have similar solid

orange male dewlaps (A. altae, which also is similar, does

not occur in Panama); and A. limifrons and A. gaigei,

with which it is frequently sympatric {Anolis elcopeensis

is amply genetically distinct from each of these species;

Figs 1, 2; A. gaigei is phylogenetically very distant from

the fuscoauratus group, data not shown). In addition to

being larger, male A. gruuo display an externally bulg-

ing tailbase in our photos, presumably indicating larger

hemipenes, which we did not observe in male A. elco-

peensis (Fig. 5). Anolis gruuo is found at mid to high

elevations (860-1,530 m) of the Serrania Tabasara from

Santa Fe west 80 km to just past Hato Chami (Lotzkat

et al. 2012). We found Anolis elcopeensis at mid to low

elevations (245-801 m) from El Cope east to Altos de

Campana and possibly further (see below). Anolis car-

penteri has a dorsal greenish tint and we have observed

it to become patterned only when stressed. Anolis elco-

peensis is never green, and usually displays banding on

the tail and a dark interorbital bar regardless of mood.

Anolis carpenteri is found on the Caribbean slope at mid

to low elevations. All of our collections of A. elcopeensis

are on the Pacific slope. Anolis elcopeensis and A. limi-

frons differ in male dewlap color (solid orange in A. elco-

peensis, dirty white with or without basal yellow spot in

A. limifrons). Females of these frequently sympatric spe-

cies may be distinguished by limb length. The adpressed

hindlimb of A. elcopeensis usually reaches to the ear,

whereas in A. limifrons the hindlimb is longer, reaching

anterior to the eye. Anolis elcopeensis and A. gaigei dif-

fer in the condition of the ventral scales (strongly keeled

in A. gaigei ; smooth in A. elcopeensis ).

Etymology

The name honors the type locality, Parque Nacional G.

D. Omar Torrijos H., and the people who have worked

to maintain this wonderful forest. The park previously

was named P. N. El Cope. Several new species of Anolis

recently have been described from the park (i.e., A. ku-

nayalae, A. ibanezi, A. elcopeensis ).

External description of holotype (paratype

variation in parentheses, measurements in

mm)

Snout to vent length 43.5 (males to 44.6; females to 44.4);

head length 0.23 SVL (0.22-0.26 SVL); head width 0.14

SVL (0.13-0.15 SVL); femoral length 0.25 SVL (0.25-

0.28 SVL); ear height 0.03 SVL (0.02-0.03 SVL); tail

length 1.7 SVL (1.5-1. 7 SVL); fourth toe length 0.17

SVL (0.13-0.17 SVL).

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A new species of Anolis from Panama

Figure 4. A) Anolis elcopeensis (male dewlap, El Valle de Anton, Code, Panama). B) Anolis elcopeensis (female dewlap, El Cope,

Code, Panama). C) A. cf. elcopeensis (male dewlap, south of Gamboa, Panama, Panama). D) A. cf. elcopeensis (male dewlap, Cerro

Azul, Panama, Panama). E) A. cf. elcopeensis (male dewlap, Lake Bayano, Panama). F) A. cf. elcopeensis (male dewlap, Meteti,

Darien, Panama). G) A. cf. maculiventris (male dewlap, Yaviza, Panama). H) A. maculiventris (male dewlap, near Buenaventura,

Colombia).

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Figure 5. A) Anolis elcopeensis (male, El Cope, Code, Panama). B) A. gruuo (male, Hato Chami, Chiriquf, Panama). Note bulging

tail base in A. gruuo.

Dorsal head scales multicarinate on frontal area and uni-

to multicarinate on nasal area, mostly unicarinate in su-

praocular disc, some smooth scales between supraorbital

semicircles and posterior to orbits (or frontal and supra-

ocular areas nearly smooth); frontal depression pres-

ent; dorsal surface of rostral scale smooth, not notched;

thirteen (10-14) scales across the snout between second

canthals; supraorbital semicircles distinct, separated by

three (1-3) scales; three (2-5) scales separate interpari-

etal and supraorbital semicircles; supraocular disk with

some enlarged scales, bordered by a complete row of

small scales; one elongate supraciliary scale followed by

a series of small scales; seven (4-7) loreal rows; greater

than 35 total loreals; elongate anterior nasal scale con-

tacts sulcus between rostral and first supralabial (or nasal

not greatly elongate); preoccipital absent; seven (6-9)

supralabials to center of eye; six (4-7) postrostrals; 6 (4-

8) postmentals; gradually enlarged scales in supraocular

disc; mental completely (or partially) divided posteriorly,

extends posterolaterally beyond rostral (or nearly even

with sulcus), with posterior border in concave arc (or

straight line transverse to head); sublabials weakly en-

larged, not much larger than medial scales; dewlap large,

reaching posterior to axillae (proportionately smaller

in small males; variable in females: absent or small, to

axillae); six-seven rows of single scales on male dew-

lap; tubelike axillary pocket absent; enlarged postcloa-

cal scales present (or absent in all females and some

males); nuchal, dorsal, and caudal crests absent; dorsal

scales keeled; approximately two (0-3) enlarged middor-

sal rows; twelve (9-15) longitudinal scale rows in 5% of

SVL; ventral scales smooth, in transverse rows; eleven

(8-11) longitudinal scale rows in 5% of SVL; anterior

thigh scales large, keeled, becoming smaller and smooth

posteriorly; supradigitals multicarinate; toepads expand-

ed and overlap first phalanx; fifteen (13-16) expanded

lamellae under third and fourth phalanges of fourth toe

(counted using the approach of Williams et al., [1995]);

tail with single row of keeled middorsal scales.

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A new species of Anolis from Panama

Figure 6. A) Ventral and B) dorsal head scales of holotype of A. elcopeensis.

Skeletal description (based on dry skele-

tons MSB 95560-1)

Parietal roof slightly convex, with Y-shaped parietal

crests with a short stem, with no casquing, lacking crenu-

lation on edges, with anterolateral comers flush with

posterolateral edges of frontal; posterior roof of parietal

slopes inferiorly; supraoccipital crests completely vis-

ible dorsally (no “half-funnel”); pineal foramen extends

posteriorly into parietal, forming a U that opens at the

parietal-frontal suture; dorsal skull bones smooth; post-

frontal present, appears partially fused; prefrontal sepa-

rated from nasal by anterior extension of frontal; fron-

tal sutures anteriorly with nasals; no parallel crests on

nasals; external nares bordered posteriorly by nasals;

premaxilla dorsally nonoverlapping, laterally flush with

nasals; dorsal aspect of jugal terminates on lateral or

posterior surface of postorbital; posterior aspect of jugal

slightly convex; epipterygoid contacts parietal dorsally;

pterygoid and palatine teeth absent; lateral edge of vomer

is smooth, without posteriorly directed lateral processes;

maxilla extends posteriorly beyond ectopterygoid on

ventral surface of skull; crest between basipterygoid pro-

cesses of basisphenoid absent; lateral shelf of quadrate

absent; black pigment is present on parietal only; pos-

teriormost mandibular tooth is posterior to anterior my-

lohyoid foramen; splenial absent; ventral aspect of an-

teromedial process of coronoid juts posteriorly or slopes

smoothly anteriorly; external opening of surangular fora-

men bordered by both dentary and surangular; posterior

suture of dentary blunt or pronged; anteriormost aspect

of posterior border of dentary is well within mandibular

fossa; labial process of coronoid present; coronoid does

not extend posterolaterally beyond surangular foramen;

jaw sculpturing absent; angular absent; teeth unicuspid

anteriorly, tricuspid posteriorly; angular process of ar-

ticular present; ten premaxillary tooth positions.

Color in life (adapted from field notes and

color photos by SP)

Anolis elcopeensis is a brown or brownish-gray lizard.

Its dorsum may be marked with faint lateral banding,

rows of spotting, or striations (Fig. 3A) or appear lightly

speckled (Fig. 5A), or may be nearly patternless pale

gray or dark brown. The tail usually is banded, and there

usually is a dark dorsal interorbital bar (Fig. 1A). The

ventral body usually displays dark brown markings on

white background. The iris is brown. The throat is pale.

Distribution and natural history

Anolis elcopeensis is known from Code Province in

Panama and potentially from Panama and Darien Prov-

inces (Fig. 7; see below). These provinces have been

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Poe et al.

Figure 7. Map of eastern Panama showing type locality of Anolis elcopeensis (star; sampled for both morphology and COI), locali-

ties sampled for COI (white circles), and localities for additional specimens referenced in text (black circles). Closed circles and

star indicate localities for confirmed A. elcopeensis according to morphological and molecular comparisons. Open circles indicate

localities for specimens referred to as Anolis cf. elcopeensis that may represent A. elcopeensis or undescribed species. Square marks

locality for A. maculiventris or a similar species (see text). Appendix 2 lists voucher specimens for localities.

well surveyed for herpetofauna, however much of Code

is difficult to access and remains unexplored. Habitat in

this province ranges from lowland rainforest in the Canal

Zone up to cool tropical premontane rainforest in Parque

Nacional Omar Torrijos. This range spans an elevation

gradient from sea level up to over 1,000 meters. We have

collected topotypical and paratopotypical A. copeensis

from 245 to 801 meters. We have found A. elcopeensis in

near-pristine primary and old selectively logged forests

(i.e., at the type locality) and in heavily disturbed road-

side vegetation.

All of our collections of A. elcopeensis occurred at

night when anoles sleep. Among 35 recorded observa-

tions at El Cope, mean sleeping perch height was 4.15

m (standard deviation 2.35). Among 30 individuals for

which sleeping perch type was recorded, 24 were on

twigs, three were on leaves, and three were on vines.

Thus, among perches easily survey able by humans (i.e.,

excluding high canopy, burrows, etc.), narrow perches

that are relatively high seem to be preferred by A. el-

copeensis. Diumally, the species has been observed to

be active on the ground and on vegetation at heights up

to 1.5 m (Mason Ryan, pers. comm.). On 23 July 2002,

Ryan observed a Cocoa Woodcreeper ( Xiphorhynchus

susurrans) catching and consuming an adult male A. el-

copeensis that was displaying from a tree buttress.

Anolis fuscoauratus and A. maculiventris in

Panama?

Anolis elcopeensis is more similar to the South American

species A. fuscoauratus than to any species in Central

America. Anolis fuscoauratus is a common forest anole

in Amazonian South America (Avila-Pires 1995). This

nondescript species is difficult to distinguish morpho-

logically from the species described here and from its

Andean and Pacific Colombian lowland congeners (A.

antonii, A. mariarum, A. tolimensis , A. maculiventris, A.

medemi ), and there are doubtless multiple cryptic species

among supposed A. fuscoauratus in Amazonia and the

eastern Andes (Poe, unpublished). We have collected A.

fuscoauratus from its type locality in Bolivia, and anoles

nearly or completely indistinguishable from topotypical

A. fuscoauratus in Peru, Colombia, Panama, and Ecua-

dor. Some authors (e.g., Kohler 2008) list A. fuscoaura-

tus from Panama, but this occurrence seems unlikely as

true A. fuscoauratus is replaced in the western lowlands

of Colombia by A. maculiventris.

We have collected fuscoauratus-like Anolis in eastern

Panama from the Panama Canal to Pirre Station, Darien.

The fuscoauratus -like anole we have collected along the

Pan American highway out to Metetf usually possesses

a bicolor orange/yellow dewlap (Fig. 4), although we

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A new species of Anolis from Panama

have collected specimens near Lake Bayano with solid

orange dewlaps (Fig. 4E), as in A. elcopeensis, Although

our current assignment for these eastern populations is A.

elcopeensis , we suspect this form may represent one or

multiple species distinct from A. elcopeensis. The vari-

ability in dewlap color (Fig. 4) and mitochondrial DNA

(Fig. 2; note positions of samples from Pipeline Road

and Lake Bayano) suggests the presences of a species

complex of fuscoauratus-hke anoles in central and east-

ern Panama. Given the local variation we have observed

in limb length and body color pattern, it also is possible

that we have failed to recognize multiple sympatric small

grayish-brown anole species with orange dewlaps at our

study sites. We currently are investigating these issues.

Near Yaviza in Darien, the fuscoauratus/ elcopeensis-

like anoles we have collected possesses a bicolor dew-

lap with pink posteriorly (Fig. 5G), similar to the Pacific

Colombian lowland form A. maculiventris (Fig. 5H). In

addition, some of our collections of this Darien popula-

tion appeared strongly dorsally patterned (Fig. 3C), as

also is common in South American A. maculiventris (Fig.

3D; but also occasionally evident in A. elcopeensis east

of the canal; pers. obs.). This population may represent

A. elcopeensis or an additional undescribed species, but

for now we tentatively assign these to A. maculiventris.

If this species inference is accurate, the number of recog-

nized anole species in Panama is increased to 46.

Acknowledgments. — Thanks to Julie Ray and Rober-

to Ibanez for facilitating our field work in Panama. Caleb

Hickman, Mason Ryan, Ian Latella, Julian Davis, Erik

Hulebak, and Heather Maclnnes helped collect individu-

als of the new species. Thanks to Sierra Llorona Lodge

for allowing us to collect lizards on their property. Mason

Ryan, Ian Latella, and three anonymous reviewers made

helpful comments on the manuscript. Thanks to Gunther

Kohler (SL tissue) and Omar Torres-Carvajal (QCAZ tis-

sue) for providing tissues. Collecting and export permits

were provided by ANAM. Funding was provided by the

National Science Foundation.

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Appendix 1

Samples used in phylogenetic analysis of COI.

POE = field numbers of SP; MSB = Museum of Southwestern Biology; QCAZ = Museo de Zoologia at Pontificia

Universidad Catolica del Ecuador; SL = field series of Sebastian Lotzkat. Voucher, Anolis species, Locality: POE 1457,

fortunensis, Panama: Chiriquf: near Fortuna dam. POE 1474, kemptoni, Panama: Chiriquf: North of Boquete. SL 355,

gruuo, Panama: Ngobe Bugle: North of Escopeta Camp. POE 2761, fuscoauratus, Bolivia: Marban: Camiaco. QCAZ

4724, maculiventris. POE 3294, altae, Costa Rica: San Jose: Volcan Barva. MSB 95565, elcopeensis , Panama: Code:

Northeast of Caimito, “La Cascada.” MSB 95566, elcopeensis, Panama: Code: 25 km NW of Penonome. POE 1894,

elcopeensis, Panama: Code: Parque Nacional Omar Torrijos. MSB 95547, elcopeensis, Panama: Code: Chorro Las

Mozas at El Valle. POE 1644, cf. elcopeensis, Panama: Panama: Pipeline Road. POE 1665, cf. elcopeensis, Panama:

Panama: East of Lake Bayano.

Appendix 2

Voucher specimens for referenced localities (all in Panama; see Fig. 7).

MVUP = Museo de Vertebrados, University of Panama. See Appendix 1 for other abbreviations. MVUP 2142 Anolis

cf. maculiventris, Darien, 4-6 km W of Yaviza. POE 4528 Anolis cf. elcopeensis, Darien, 8 km S of Metetf. MSB

95563 Anolis cf. elcopeensis, Panama, between Rio Maje and Rio Urti E of Lake Bayano (9.0622222, -78.986111).

MSB 95568 Anolis cf. elcopeensis, Panama, 11-15 km N of El Llano on El Llano-Carti Road (9.4866667, -79.026389;

463 m). MSB 95553 Anolis cf. elcopeensis, Panama, Cerro Azul (9.22, -79.38 885 m). MSB 95562 Anolis cf. elco-

peensis, Panama, Pipeline Road north of Gamboa (9.12006, -79.71468, 44 m). MVUP 2140 Anolis cf. elcopeensis,

Panama, Radisson Summit Hotel on Gailard (9.04957, -79.63373, 130 m). MSB 95567 Anolis cf. elcopeensis, Panama,

Sierra Llorona Lodge near Santa Rita Arriba (9.342, -79.776, 200 m). MSB 95558 Anolis elcopeensis, Panama, Altos

de Campana (8.68, -79.94, 800 m). POE 1626-1628 Anolis gruuo, north of Santa Fe, Veraguas, Panama.

■ Steven Poe is Associate Professor in the Department of Biology and Associate Curator in the Division of Amphib-

ians and Reptiles of the Museum of Southwestern Biology at the University of New Mexico, USA. His research

' M focuses on taxonomy, phylogenetics, and comparative evolution, especially of Anolis lizards. He has collected over

m 250 species of Anolis in 15 countries.

Simon G. Scarpetta is a field conservation biologist at Stanford University and a soon to be graduate student of

paleontology at The University of Texas at Austin. He received his B.S. in Biology with an emphasis in ecology

and evolution from Stanford University, where he studied phylogenetics and biogeography of Ameiva lizards. He

has also been involved with lab and field work at the University of New Mexico studying Anolis lizards. Simon is

currently interested in osteology and evolutionary patterns of lizards in western Australia.

« Eric W. Schaad is a senior associate biologist for Quad Knopf Inc. in Central California. He received his M.S. in

biology at the University of New Mexico where he studied the evolution of ecomorphs and community structure in

Anolis lizards. Eric currently works in conservation biology with a focus on reptile and amphibian conservation and

the impacts of development on threatened and endangered species in Central California.

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A new species of Anolis from Panama

In accordance with the International Code of Zoological Nomenclature new rules and regulations (ICZN 2012), we have deposited this paper

in publicly accessible institutional libraries. The new species described herein has been registered in ZooBank (Polaszek 2005a, b), the of-

ficial online registration system for the ICZN. The ZooBank publication LSID (Life Science Identifier) for the new species described here

can be viewed through any standard web browser by appending the LSID to the prefix “http://zoobank.org/.” The LSID for this publication

is: um:lsid:zoobank.org:pub:22ED2728-2093-46D6-AE9B-A77AC56A7412.

Separate print-only edition of paper(s) (reprint) are available upon request as a print-on-demand service. Please inquire by sending a request

to: Amphibian & Reptile Conservation, amphibian-reptile-conservation.org, arc.publisher@gmail.com.

Amphibian & Reptile Conservation is a Content Partner with the Encyclopedia of Life (EOL), http:///www.eol.org/ and submits information

about new species to the EOL freely.

Digital archiving of this paper are found at the following institutions: ZenScientist, http://www.zenscientist.com/index.php/filedrawer; Ernst

Mayr Library, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (USA), http://library.mcz.harvard.edu/

emst_mayr/Ejournals/ARCns.

The most complete journal archiving and journal information is found at the official ARC journal website, amphibian-reptile-conservation,

org. In addition, complete journal paper archiving is found at: ZenScientist, http://www.zenscientist.com/index.php/filedrawer.

Citations

ICZN. 2012. Amendment of Articles 8,9,10,21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods

of publication. Zootaxa 3450: 1-7.

Polaszek A et al. 2005a. Commentary: A universal register for animal names. Nature 437: 477.

Polaszek A et al. 2005b. ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological

Nomenclature 62(4): 210-220.

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Amphib. Reptile Conserv.

e95

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [General Section]: 14-23 (e95).

A single species of mangrove monitor (Varanus) occupies

Ambon, Seram, Buru and Saparua, Moluccas, Indonesia

1 3 Valter Weijola and 2 4 Samuel S. Sweet

Zoological Museum, University of Turku, 20520 Turku, FINLAND 2 Department of Ecology, Evolution and Marine Biology, University of California,

Santa Barbara, California 93106, USA

Abstract . — According to current literature the islands of the central Moluccan region harbor at

least three species of monitor lizards. This suggests similar patterns of species richness to the

northern Moluccas and could imply significant taxonomic and ecological complexity throughout the

Moluccan region. Field investigations in habitats from sea level up to 300 m elevation failed to locate

more than one widespread species, by definition referable to Varanus indicus (type locality Ambon).

Reassessments of records for other species of mangrove monitors show that these can either be

attributed to taxonomic mis-identifications or to colonial-era specimens lacking reliable collection

data. We test Principal Components Analysis of scalation characters as a diagnostic tool for some

of the island populations and species within the Varanus indicus group.

Key words. Monitor, Euprepiosaurus, Varanus indicus, Varanus cerambonensis, Varanus rainerguentheri, Moluccas,

habitat use

Citation: Weijola V, Sweet SS. 201 5. A single species of mangrove monitor ( Varanus) occupies Ambon, Seram, Buru and Saparua, Moluccas, Indo-

nesia. Amphibian & Reptile Conservation 9(1) [General Section]: 14-23 (e95).

Copyright: © 201 5 Weijola and Sweet. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommer-

cialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation', official journal website <amphibian-

reptile-conservation. org> .

Received: 16 Feburary 2014; Accepted: 20 April 2015; Published: 25 April 2015

Introduction

The island of Ambon has had a history of commercial

and strategic importance and has been connected to the

European economy for five centuries. As a consequence,

many early faunal collections came from there, and it is

the type locality for a considerable number of Indonesian

species, some of which have proved to be native to the

island whereas others were merely shipped from Am-

bon (e.g., Daan and Hillenius 1966; Hoek Ostende et al.

1997). Despite this long European presence, little first-

hand information has been published on the biology of

the local Varanus species. The first recorded observations

and possible specimen collection of monitors on Ambon

were made in 1792 by Claude Riche, one of the natural-

ists of the d’Entrecasteaux Expedition, and reported by

F.M. Daudin in the description of Tupinambis indicus a

decade later (Daudin 1802). For the next two hundred

years this was the only species reported from the cen-

tral Moluccas (to which we refer to the islands of Seram,

Buru, Ambon, and the other islands in the Lease group).

This changed when Philipp et al. (1999) revised the

identity of V. indicus and described a second species

from Ambon, Seram, and Buru, which they named V. ce-

rambonensis (Fig. 1 A-D), distinguishable from V. indi-

cus through the presence of a yellow temporal stripe, a

banded dorsum, and a bi-colored tongue. In 2012 Somma

and Koch reported that a third species, V. rainerguentheri

(Fig. 1 E-F), also occurs on Buru in sympatry with V.

cerambonensis (and possibly V. indicus). Varanus Salva-

tor has also been reported to occur on Seram on the basis

of a single voucher specimen (Koch et al. 2007). These

records are discussed here and Principal Components

Analysis (PCA) is tested as a tool to detect differences

between island populations of species in the V. indicus

group (Fig. 2).

With a surface area of 17,400 km 2 Seram is the sec-

ond-largest island in the Moluccas (after Halmahera)

(Monk et al. 1997). It is estimated to have emerged as a

land mass around 5-6 MYA along the Outer Banda Arc

and rotated westward (Hall 2002), thus always having

been isolated from New Guinea (Audley-Charles 1993;

Fortuin and de Smet 1991). For animal groups with good

dispersal abilities, such as Lepidopterans, this appears to

have had little impact on current diversity and commu-

nity composition when compared to the slightly larger

Correspondence. Email: 3 vweijola@ gmailcom (Corresponding author); 4 sweet@lifesci.ucsb.edu

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Weijola and Sweet

Fig. 1 . Mangrove monitors and their habitats: V. cerambonensis on Ambon (A), Seram (B), and Burn (C, D). Varanus rainerguen-

theri on Halmahera (E) and Obi (F). Coastal vegetation on Ambon (G) and Nipa swamp (H). Photographs by Valter Weijola.

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e95

Single species of mangrove monitor in central Moluccas

island of Halmahera (de Jong 1998). For monitors specif-

ically, the lack of a land connection with New Guinea ap-

pears to have restricted the number of successful coloniz-

ers to just one (this study) whereas the composite island

of Halmahera has a larger set of species (Weijola 2010).

There are strong patterns in the distribution of the

members of the subgenus Euprepiosaurus , the Indoaus-

tralian radiation of gracile terrestrial and arboreal species

containing mangrove and blue-tailed monitors and the

slender tree monitors of the V. pr as inns group. Members

of the latter clade are largely restricted to landmasses on

the Sahul shelf. The blue-tailed monitors in the V. indi-

cus group ( V. caerulivirens, V. doreanus, V. finschi, V. jo-

biensis , and V. yuwonoi ) show a similar pattern, with the

exceptions of also inhabiting Halmahera (and adjacent

islands) as well as the island of New Britain (Ziegler et

al. 2007). The only lineage with a demonstrated ability

for significant oversea dispersal is that mainly inhabit-

ing coastal areas, traditionally known as a variable and

widespread mangrove monitor, V. indicus. This lineage,

with one representative in the central Moluccas, has late-

ly been split up into several closely related species that

all appear to exhibit allopatric distributions: V. cerambo-

nensis (Burn, Ambon, Lease Islands, Seram); V. indicus

(Australia, New Guinea, and satellite islands, as well as

many Pacific islands); V. melinus (Mangole and possi-

bly Taliabu); V. juxtindicus (Rennert); V. rainerguentheri

(northern Moluccas); V. lirungensis (Talaud); and V. obor

(Sanana) (Fig. 3). Additional populations of uncertain

status occur in the Ara, Kei, and Tanimbar island groups.

In 2008 to 2009 fieldwork was conducted to study the

niche partitioning among monitor species on several Mo-

luccan islands (Weijola 2010; Weijola and Sweet 2010).

On Ambon, Seram, Saparua, and Burn the species com-

munities were initially presumed to be composed of V.

indicus utilizing coastal habitats and V. cerambonensis

occupying habitats farther inland as suggested by Philipp

et al. (1999) and mirroring the ecological roles of V. rain-

erguentheri and V. caerulivirens on Halmahera (Weijo-

la 2010) or V. indicus and V. jobiensis on New Guinea

(Philipp 1999). This hypothesis was rejected during field-

work as it became evident that only one of the species, V.

cerambonensis (, sensu Philipp et al. 1999), functioned as

a habitat generalist and occurred throughout each island,

and that V. indicus (, sensu Philipp et al. 1999) was absent

from these islands altogether.

The absence of V. indicus {sensu Philipp et al. 1999)

is problematic inasmuch as Ambon is the type locality

for this species. The only two specimens, ZMA 11146c

and ZFMK 70650 (formerly ZMA 11146d), indicating a

sympatry between V. indicus and V. cerambonensis on

Ambon (and in the central Moluccas) turned out to have

belonged to a colonial-era collector stationed on Ambon,

but there is no evidence to suggest that they were actually

collected there. The identity of V. indicus has been re-

viewed in detail by Weijola and a Case to synonymize V.

cerambonensis with V. indicus has been submitted to the

International Commission on Zoological Nomenclature

(ICZN) (Weijola, In press). As this nomenclatural issue

is yet to be resolved we follow the current name uses and

diagnoses here but note that future changes are possible.

Methods

Fieldwork was conducted during March and December

2009 near the following settlements: Ambon - Liang

(VW); Hitu (VW, SS); Soya di Atas (VW, SS); upper

Ambon Bay (VW, SS); Waitami (VW, SS); Latuhalat

(VW, SS); Seram (VW) - Besi; Burn (VW) - Namlea,

Wamlana, Samleko; and Saparua (VW, SS) - Kulur. Spe-

cies identification in the field followed the diagnostic

characters provided by Philipp et al. (1999). Accordingly

Varanus cerambonensis can be identified by its distinct

yellow temporal band and yellow markings (dots and /

or ocelli) arrayed in a pattern of transverse bands on the

dorsum. These characters can effectively be used in the

field even at a distance with a pair of binoculars.

Observations on habitat use were obtained by quietly

traversing all major habitat types from coastal (man-

groves, natural coastal scrub, coconut plantations, park-

lands) to lowland rainforests and hill forests up to 300

m elevation. Searches were made on foot or by canoe.

For each observation date, time, location, habitat, and

vegetation type, altitude, and activity were recorded. Ac-

tive animals could often be heard running through dry

litter in the undergrowth before fleeing up a tree where

their identity could be confirmed. Basking animals often

remained still unless approached within flight distance

(normally 10-30 m).

The examination of museum vouchers allowed for a

larger set of characters including scale counts to be as-

sessed. According to Philipp et al. (1999) V. cerambo-

nensis has on average smaller scales and higher scale

counts than does V. indicus : e.g., scales around midbody

(131-150 vs. 106-144), or transverse rows of dorsal

scales (126-163 vs. 105-137).

Fig. 2. Principal Components Analysis of scalation characters

for several island populations of V. cerambonensis, V. indicus,

and V. rainerguentheri. The two Xs represent ZFMK 70650 and

ZMA 11146c.

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Weijola and Sweet

Museum specimens at Naturalis (RMNH) and the

Zoological Museum Amsterdam (ZMA) were identified

(VW) and scale counts for a Principal Components Anal-

ysis were extracted from Brandenburg (1983). Counts

employed were: midbody scale rows (S), dorsal scale

rows from dorsal margin of tympanic recess to anterior

margin of hind limbs (XY), transverse rows of ventral

scales from gular fold to anterior margin of hind limbs

(T), transverse rows of dorsal scales from posterior mar-

gin of tympanic recess to gular fold (X), scales around

neck at anterior margin of gular fold (m), scales from

rictus to rictus across dorsum of head (P), scales around

tail base (Q), scales around the tail 1/3 from the base (R),

and number of ventral scales from the tip of snout to gu-

lar fold (N).

Principal Components Analysis was performed in

PAST (Hammer et al. 2001) using all the above-men-

tioned scale characters for specimens from Ambon, Se-

ram, Burn, Halmahera, Temate, Bacan, New Guinea,

Waigeo, and Biak (Appendix 1).

Results

Morphology

The Principal Components Analysis of scalation char-

acters (Fig. 2) worked well to differentiate the included

island populations with partial overlap found only be-

tween V. rainerguentheri and V. indicus. PCI and PC2 ac-

counted for almost 90% of the total variance. The factor

loadings for PCI were all positive with highest values on

factors XY (0.78) and S (0.42). On PC2 all loadings were

positive except for XY and R, with highest values on T

(0.71) and m (0.61). PC3 gave more overlap between the

population clusters. Eigenvalues and factor loadings for

PC1-PC3 are presented in Table 1.

Habitat use

All field observations, involving a total of 8 1 sightings

(Ambon, 31, Burn, 21, Seram, 9, and Saparua, 20) were

identified as V. cerambonensis. Monitors were most nu-

merous on Ambon, Burn, and Saparua whereas fewer

observations were made on Seram. A majority of obser-

vations ( n = 70) was made in coastal areas where moni-

tor population densities appear to peak. Encounter rates

were high both in littoral forest ( n = 38) in sandy and

karst (n = 9) areas, as well as in mangroves (n = 14) and

Nipa swamps (n = 9). The preferred areas usually had a

bushy undergrowth used for hiding and larger trees for

basking and hiding in tree cavities. Seven observations

were made in coconut or mixed-crop plantations in low-

land areas.

Far fewer monitors were observed in lowland rainfor-

ests (n = 1), swamp/sago forest (n = 1), and hill forests ( n

= 2), with the highest altitude observation at around 300

m near a small stream at Soya di Atas on Ambon. There

Sulawesi

6 D

Talaud

Halmahera

Bacan

Mangole

Morotai

Waigeo

Varan us cerambonensis

• Varanus rainerguentheri

Varanus indicus

• Varanus melinus

• Varanus obor

• Varanus Urungensis

• Taxonomically undetermined

X Study sites

Biak

Taliabu

fsanana

r - : x-x

Burn

Obi

Seram

-K -----

*7 \

'X i Xg a p artia

Ambon

New Guinea

Banda ^ei i

Aru

Tanimbar

Timor

134 s

Fig. 3. Distribution map of mangrove monitors in the Moluccas and western New Guinea, the blue tailed monitors not included.

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Single species of mangrove monitor in central Moluccas

Table 1. Loading values, proportion of variance, and eigenval-

ues for PCA. The two highest loading factors on the three first

components highlighted.

Factor

Comp 1

Comp 2

Comp 3

0.111204

0.099216

-0.054745

0.19027

0.24899

0.3686

0.77924

-0.57179

-0.10626

0.36133

0.60913

-0.23433

0.41622

0.45941

0.036529

0.27334

0.71204

0.1145

0.16643

0.10618

0.15529

0.0088444

-0.052658

0.8697

Proportion of

variance

81.245

8.594

4.1006

Eigenvalue

711.843

75.2985

35.9281

is however an almost fully melanistic specimen (ZMA

15416g) at the Naturalis Museum collected at Lake Rana

on Bum (at 770 m elevation) which shows that the spe-

cies also occupy higher altitudes.

Activity and foraging

Fifty-six of the monitors were first observed while active-

ly moving or foraging on the ground whereas the other

25 were first seen while basking on tree trunks/branches

(n = 21) or on the ground (/? = 4). Monitors usually be-

came active and emerged to bask at around 0815-0845

and returned to their retreats late in the afternoon, the

latest observation of an active individual was made on

Ambon at 1600. The only specific foraging events ob-

served during this study were several specimens on Bum

actively digging out sea turtle nests in search of eggs, one

individual on Seram digging for sago gmbs in a rotten

Metroxylon trunk, and another individual (also on Se-

ram) digging through a pile of garbage at the edge of a

mangrove swamp.

Discussion

Natural history

Observations of habitat use of V. cerambonensis cor-

respond well with that reported from field studies of

mangrove monitors (V. indicns senso lato ) in other re-

gions (Iyai and Pattiselanno 2006; Philipp 1999; Smith

and Griffiths 2009; Weijola 2010). Densities appear to

peak in coastal and saltwater influenced areas with suit-

able vegetation cover and decrease with increasing alti-

tude where animals also become more restricted to areas

near bodies of freshwater. Dietary studies show Varanus

cerambonensis to be an opportunistic predator with the

single largest component being crustaceans which makes

up almost half of the diet (Philipp et al. 2007). As is usual

throughout the Moluccas monitors were more frequently

Amphib. Reptile Conserv.

encountered near Muslim than Christian settlements,

presumably reflecting dietary restrictions and the scarcity

of hunting dogs.

In more species rich-communities such as that of

Halmahera, mangrove monitors (on Halmahera V. rain-

erguentheri ) are rarely observed at higher altitudes where

instead V. caerulivirens is common (Weijola 2010). Vara-

nus indicus on New Guinea may similarly be restricted

in upland areas by competition from V. jobiensis and V.

doreanns. On some single species islands the mangrove

monitors appear to persist higher up and can occasionally

be found up to at least 700-900 m elevation (as demon-

strated by their presence at Lake Rana on Bum). On New

Ireland, Papua New Guinea, the senior author has col-

lected mangrove monitors as high up as the Lelet Plateau

at 900 m elevation (Weijola, unpub. data).

Biogeography

Whereas many of the larger islands in the northern Mo-

luccas (e.g., Halmahera, Obi, Bacan, and Morotai), and

island arcs moving along the northern coast of New

Guinea, have several monitor species with evident eco-

logical specialization (Weijola 2010), the other Moluc-

can islands, including Ambon, Seram, Bum, Tanimbar,

and Kei, have only single members of ecological gen-

eralists of the V. indicus group present (this study; Wei-

jola, unpub. data). These are joined by members of the

V. salvator group in the Sula islands and on Obi (Weijola

2010; Weijola and Sweet 2010), but the presence of V.

salvator on Seram (Koch et al. 2007) has not gained sup-

port from recent fieldwork (Edgar and Lilley 1993; this

study) and they were unknown to several experienced

hunters contacted by VW. This is usually a conspicuous

animal wherever it occurs; for example, the new records

for Taliabu and Sanana were established on the first and

second days of fieldwork (VW and SS), on the first day

on Mangole (VW) and on the second day on Obi (VW).

Varanus rainerguentheri

Somma and Koch’s (2012) distribution record of V. rain-

erguentheri, and their claim of its co-existence with V.

cerambonensis, on Bum is based on a preserved speci-

men (Senckenberg Museum, Frankfurt [SMF 56469])

and a photo taken in the field (Somma and Koch 2012,

Fig. 6). Both were identified as V. rainerguentheri from

the occurrence of rows of dorsal ocelli. However, there

are eleven vouchered Varanus from Burn at the Natu-

ralis Museum (ZMA 15416a-j, RMNH 7223), which are

similar in color pattern to those presented as V. rainer-

guentheri by Somma and Koch, and which were exam-

ined and identified as V. cerambonensis by Philipp et al.

(1999) (forming the record of V. cerambonensis for that

island). All above-mentioned specimens conform in col-

or pattern to those observed in the field during this study.

As is indicated by Weijola (2010) there are typically no

April 201 5 | Volume 9 | Number 1 | e95

Weijola and Sweet

distinct bands of dorsal ocelli on adult V. rainerguentheri

but these are instead characteristic of the V. cerambonen-

sis populations on Ambon and especially Bum (ZMA

15416, Weijola field observations). For these reasons we

regard Somma and Koch’s records of V. rainerguentheri

from Bum to be mis-identifications of V. cerambonensis.

Principal Components Analysis

The results of the PCA illustrates its potential to recover

geographic clusters among the sampled islands. As the

increasing number of island endemics and cryptic spe-

cies has made identifications more problematic, and the

use of single color pattern characteristics can be mislead-

ing, we acknowledge its usefulness as an additional di-

agnostic tool.

Although considered conspecific the distance between

the Ambon/Seram and Bum populations detected by the

PCA indicate morphological separation between the two

populations. In addition to scalation differences the pop-

ulation of Bum also differ in color pattern from those of

Ambon and Seram, notably by the brown/orange throat

and abdomen color (seen in live specimens), as well as

having more evident dorsal rosettes.

Conclusions

Recent research on Indonesian monitors has relied heav-

ily on colonial-era museum voucher specimens and re-

cent animals obtained from the pet-trade (Bohme and

Ziegler 1997; Philipp et al. 1999; Somma and Koch

2012; Ziegler et al. 2007a, b). This has obscured the fact

that some of the newly described island endemics such

as V. melinus, V. cerambonensis, and V. rainerguentheri,

are not previously unknown animals co-occurring with a

widespread V. indicus, but are instead local forms previ-

ously assigned to a variable V. indicus that have now been

recognized as separate taxa. This has inflated the number

of species thought to be present, with several islands al-

legedly harboring multispecies communities. Although

we cannot rule out the possible existence of additional

secretive species we conclude that there currently is no

evidence for more than one species of mangrove monitor

in the central Moluccas.

TO avoid future confusion in the taxonomic and bio-

geographic interpretations of this group we call for more

critical scrutiny of unique colonial-era museum speci-

mens with single-word localities and no further support-

ing information.

Acknowledgments. — Travel support was provided by

the National Geographic Society, Svenska studiefonden,

Nordenskiold-Samfundet, and Svensk-Osterbottniska

samfundet. The Jenny and Antti Wihuri Foundation pro-

vided Weijola with a graduate- student scholarship. Of all

Amphib. Reptile Conserv.

the generous and hospitable local people we are particu-

larly grateful to Pak Aiuti of Seram and Pak Assis of Sa-

parua. We thank Daniel Bennett, Fred Kraus, and Harold

de Lisle for constructive comments on the manuscript,

and Mikael von Numers for technical advice.

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view of the subgenus Euprepiosaurus of Varanus

(Squamata: Varanidae): Morphological and molecular

phylogeny, distribution and zoogeography, with an

identification key for the members of the V. indicus

and the V. prasinus species groups. Zootaxa 1472:

1-28.

Valter Weijola has a B.S and M.S in biology from Abo Akademi University and is now a Ph.D. student at

the University of Turku. He is currently working on the biogeography and phylogenetics of Pacific moni-

tors and has traveled extensively in Indonesia, Papua New Guinea, and the Solomon islands in pursuit of

his interests.

Sam Sweet received a B.S. at Cornell University, and M.S. and Ph.D. at the University of California,

Berkeley. He is Professor of Ecology and Evolution at the University of California Santa Barbara, and di-

vides his time between recovery efforts for endangered herpetofauna in southwestern North America, and

investigations of the spatial ecologies of Indoaustralian monitor lizards.

April 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

e95

Weijola and Sweet

Appendix 1

Naturalis Museum specimens and scalation data included in the PCA.

Catalog number

Island

ZMA 10202

Alkmaar Island

134

131

RMNH 7297a

Ambon

150

105

140

RMNH 7297b

Ambon

146

100

140

RMNH 7297d

Ambon

152

104

142

RMNH 7297e

Ambon

139

104

138

RMNH 7297f

Ambon

148

101

131

RMNH 7297g

Ambon

153

105

146

100

RMNH 7196

Ambon

149

141

RMNH 3152

Ambon

154

110

143

100

RMNH 3150

Ambon

138

110

142

RMNH 3800

Bacan

134

128

RMNH 2 103 lg

Biak

125

131

RMNH 2 103 lh

Biak

124

128

RMNH 21033

Biak

122

135

RMNH 21026a

Biak

127

128

RMNH 21026b

Biak

121

126

RMNH 21024

Biak

115

122

RMNH 21021

Biak

123

122

RMNH 7223

Buru

147

145

ZMA 15416a

Buru

192

104

151

104

ZMA 15416b

Buru

187

141

104

ZMA 15416c

Buru

178

138

103

ZMA 15416d

Buru

187

150

100

ZMA 15416e

Buru

179

142

100

ZMA 15416f

Buru

183

143

101

ZMA 15416g

Buru

164

143

ZMA 15416h

Buru

174

142

103

ZMA 154161

Buru

179

135

105

ZMA 154 16j

Buru

159

142

ZMA 15414a

Halmahera

163

128

ZMA 15414b

Halmahera

155

139

ZMA 15414c

Halmahera

147

132

RMNH 7197

Haruku

144

101

142

RMNH 21041

Insoemarr Island

120

121

RMNH 21045

Japen

107

106

RMNH 21052

Japen

105

120

RMNH 21053

New Guinea

113

107

RMNH 21054

New Guinea

105

108

RMNH 21055a

New Guinea

111

109

RMNH 21047

New Guinea

115

113

RMNH 21042

New Guinea

115

113

RMNH 21046

New Guinea

110

RMNH 21036a

New Guinea

116

110

RMNH 21036b

New Guinea

116

110

RMNH 21036e

New Guinea

114

112

ZMA 10201

New Guinea

131

125

Amphib. Reptile Conserv.

April 2015 | Volume 9 | Number 1 | e95

Single species of mangrove monitor in central Moluccas

Appendix 1 (continued)

Naturalis Museum specimens and scalation data included in the PCA.

Catalog number

Island

ZMA 10208

New Guinea

125

114

RMNH 21037

New Guinea

128

RMNH 21038

New Guinea

118

RMNH 21034

New Guinea

128

123

RMNH 21046

New Guinea

129

120

RMNH 5359

New Guinea

115

113

RMNH 21018

New Guinea

115

112

RMNH 21050

New Guinea

118

116

RMNH 6726

New Guinea

121

122

RMNH 21040

New Guinea

128

122

RMNH 21020

New Guinea

123

125

ZMA 10194b

New Guinea

143

136

ZMA 10194c

New Guinea

124

122

ZMA 10200

New Guinea

121

115

RMNH 21035

New Guinea

119

111

RMNH 5260

New Guinea

116

110

RMNH 21048

New Guinea

113

115

RMNH 3151

Ravak

139

115

RMNH 3189

Seram

152

100

150

RMNH 3190a

Temate

136

120

RMNH 3190b

Temate

133

128

ZMA 15417

Temate

140

127

ZMA 11146c

Unknown

140

123

ZMA 1 1 146d

Unknown

131

121

ZMA 10192a

Waigeo

146

122

ZMA 10192b

Waigeo

151

123

ZMA 10192c

Waigeo

138

120

ZMA 10192d

Waigeo

147

119

ZMA 10192f

Waigeo

139

117

ZMA 10192g

Waigeo

141

125

ZMA 10192h

Waigeo

140

123

Amphib. Reptile Conserv.

April 201 5 | Volume 9 | Number 1 | e95

A dark morph of a male Fowler’s Toad (Anaxyrus fowleri) in a cranberry ( Vaccinium macrocarpon ) bog. The toad was vocalizing

just before this picture was taken. The uninflated and darkly pigmented vocal sac is partially visible. Photo by Brad Timm.

May 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

e97

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [General Section]: 24-33 (e97).

Fowler’s Toad ( Anaxyrus fowleri) occupancy in the

southern mid-Atlantic, USA

^ara S. Jones and 23 Todd A. Tupper

1 Department of Environmental Science and Policy George Mason University, Fairfax, Virginia 22030, USA 2 Department of Science, Technology and

Business Northern Virginia Community College Alexandria, Virginia 22311, USA

Abstract .— We assessed the effects of landscape structure on Anaxyrus fowleri ( Fowler’s Toad) site

occupancy using 14 years of call survey data collected from 250 sites in Virginia and Maryland, and

landscape variables derived from the National Wetlands Inventory, U.S. Census Bureau, National

Land Cover Databases, and U.S. Department of Agriculture. We also conducted a time series

analysis on A. fowleri occupancy rates using call survey data collected throughout Virginia and

Maryland. We found A. fowleri site occupancy to be negatively affected by deciduous forest, hay

crops, development and agricultural pesticides, and we identified a negative interannual trend in

occupancy rates between 1999 and 2012.

Key words. Fowler’s toad, amphibian declines, calling anuran surveys, North American Amphibian Monitoring Pro-

gram, landscape ecology, species-habitat modeling, anurans

Citation: Jones KS, Tupper TA. 201 5. Fowler’s Toad ( Anaxyrus fowleri) occupancy in the southern mid-Atlantic, USA. Amphibian & Reptile Conserva-

tion 9(1) [General Section]: 24-33 (e97).

cialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official journal website <amphibian-

reptile-conservation. org> .

Received: 27 January 2015; Accepted: 30 April 2015; Published: 02 May 2015

Introduction

Amphibian populations are declining globally, with an-

thropogenic degradation of landscapes near wetlands

having a major impact on many species of pond-breeding

amphibians (Findlay and Houlahan 1997; Blaustein and

Kiesecker 2002). Upland habitats surrounding wetlands

are vital for successful dispersal, foraging, and non-

breeding activities, making upland habitat quality critical

to the life history of pond-breeding amphibians (Wind-

miller 1996; Semlitsch 2000; Gibbons 2003; Bartlet et

al. 2004). The negative effects of landscape degradation

on many amphibian species, including hydroperiod al-

teration, pollution of wetlands from roadway runoff and

agricultural chemicals, and mechanical disturbance of

foraging, retreat and burrowing sites (Luo et al. 1999;

Turtle 2001; Gray et al. 2004b) are fairly well under-

stood. These anthropogenic disturbances can ultimately

impact mobility and survival of larval, juvenile, and adult

amphibians, and can lead to population declines and ex-

tirpations (Blaustein and Kiesecker 2002; Gray et al.

2004a). Thus, understanding how natural and anthropo-

Correspondence. Email: i ttupper@nvcc.edu

genic landscape-level processes effect amphibian popu-

lations is critical to amphibian conservation.

The Fowler’s Toad, Anaxyrus (Bufo) fowleri, is wide-

ly but irregularly distributed throughout the eastern Unit-

ed States, occurring from southern New England to the

Florida Panhandle and as far west as Missouri, Arkansas,

and Louisiana (Netting and Goin 1945; Green 1992; Kl-

emens 1993; Conant and Collins 1998). Though typically

associated with coastal dune systems and scrub-pine for-

ests, A. fowleri also occurs in rocky and sparsely veg-

etated areas in dry, sandy, deciduous woodlands, and in

agricultural and developed areas (Schlaugh 1976, 1978;

Klemens 1993; Zampella and Bunnell 2000; Rubbo and

Kiesecker 2005; Gooch et al. 2006). Some biologists and

naturalists have thus described A. fowleri as being toler-

ant of urbanization, and scarification in agricultural ar-

eas (Ferguson 1960; Martof et al. 1980; Klemens 1993;

Rubbo and Kiesecker 2005; Gooch et al. 2006). How-

ever, other studies suggest that A. fowleri are habitat spe-

cialists sensitive to environmental perturbations (Breden

1988; Green 2005; Tupper and Cook 2008).

May 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

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Jones and Tupper

In Canada, A. fowled are federally protected (Oldham

2003), but they are not considered a species of concern in

the United States. Anaxyrus fowled populations are be-

lieved to be relatively stable and abundant in the eastern

United States (Conant and Collins 1998). However, A.

fowled extirpations have been documented in the north-

eastern and southeastern United States (Breden 1988;

Klemens 1993; Mierzwa et al. 1998; Tupper and Cook

2008; Walls et al., 2011; Milko 2012). These extirpations

were largely attributed to anthropogenic disturbances,

such as habitat degradation, pesticide application, road

mortality (National Park Service, unpubl. data), hydro-

period alteration, competition from invasive species, and

probable increased predation pressures from urban toler-

ant predators such as skunks {Mephitis mephitis) and rac-

coons ( Procyon lotor ) (Schaff and Garton 1970; Lazell

1976; Groves 1980; Klemens 1993; Tupper and Cook

2008; Milko 2012).

In the southern mid- Atlantic region, A. fowled occur

throughout Virginia and Maryland, but are less common

outside of the Coastal Plain (Mitchell and Reay 1999).

Coastal regions are thought to contain more favorable

upland habitats for this species (see Mart of et al. 1980;

Mitchell and Reay 1999; Cook et al. In prep), but much

of the Coastal Plain in Virginia and Maryland is more

densely populated and intensely developed than west-

ern regions. For instance, the mid-Atlantic Coastal Plain

has the highest population density and second-highest

growth rate of all ecoregions in Virginia (VGDIF 2005).

If A. fowled are sensitive to landscape perturbations, hu-

man population growth and development may lead to A.

fowled population declines in the southern mid- Atlantic.

To the best of our knowledge, quantitative data de-

scribing the effects of landscape-level variables on A.

fowled are non-existent for the mid- Atlantic and are lim-

ited elsewhere (see Gooch et al. 2006; Tupper and Cook

2008; Birx-Raybuck 2010; Eskew et al. 2011). Occupan-

cy trend analyses of A. fowled populations indicate that

they are stable in most mid- Atlantic states (except Mary-

land; see Weir et al. 2014), but these analyses are tempo-

rally limited (Weir et al. 2009). Thus, critical thresholds in

landscape-level variables essential to A. fowled occupan-

cy are unknown and it is unclear if southern mid- Atlantic

populations are stable over the long term. Therefore our

objectives were to identify and describe landscape-level

variables that influence A. fowled site occupancy and to

complete a more comprehensive time-series analysis for

this species in the southern mid- Atlantic.

Materials and Methods

Site selection and data collection

We randomly selected 250 sites in Virginia and Mary-

land for landscape-level analyses (Fig. 1). Selected sites

were North American Amphibian Monitoring Program

(NAAMP) calling anuran survey points (adjacent to

wetlands) that were surveyed with anuran call counts be-

tween 1999 and 2012 (Weir and Mossman 2005). Move-

ment data for A. fowled are limited, but available studies

indicate that a 1 km buffer surrounding breeding wetlands

is a biologically meaningful distance for analyzing the

effects of landscape features on anuran (including Anaxy-

rus spp.) occurrence (Clarke 1974; Miaud et al. 2000;

Muths 2003; Bartlet et al. 2004; Smith and Green 2005;

Forester et al. 2006). Therefore, our landscape variables

were derived from 1 km buffers surrounding calling sur-

vey points. Any calling survey points found to have over-

lapping buffers were removed from analysis. Anuran call

data (ranked ordinal values based on chorus intensity [0-

3]) and sampling covariates (ambient temperatures, sky

and wind conditions, and noise disturbance levels) were

collected in accordance with NAAMP guidelines by

trained NAAMP volunteers (Weir and Mossman, 2005).

Fig. 1 . Map of calling anuran surveys conducted in Maryland and Virginia. Closed circles indicate sites occupied by A. fowled and

open circles indicate unoccupied sites.

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Amphib. Reptile Conserv.

e97

Fowler’s Toad occupancy in the southern mid-Atlantic

Calculation of landscape variables

We quantified landscape variables using data from four

publicly available sources: (1) the National Wetlands

Inventory (NWI) from the U.S. Fish and Wildlife Ser-

vice; (2) 2012 TIGER/Line road data from the U.S.

Census Bureau; (3) the National Land Cover Database

(NLCD2006) from the Multi-Resolution Land Charac-

teristics Consortium; and (4) the National Pesticide Use

Database from the U.S. Department of Agriculture. Ini-

tial data manipulation was done in QuantumGIS (QGIS;

QGIS Development Team 2011). For NWI data, we ex-

tracted distance from calling survey sites to nearest wet-

land and determined the number and types of wetlands

within a 1 km buffer zone of calling survey sites. Using

TIGER/Line road files, we calculated road length and

type within 1 km buffers. We prepared land cover data

by clipping NLCD2006 data for each buffer into an indi-

vidual raster file. These files were then imported into R

(R Core Team 2013) and analyzed using the SDMTools

package (VanDerWal 2013). Total pesticide application

rate (kg/km 2 ) at each site was determined by calculating

the sum of application levels within each buffer for all

pesticides listed in the National Pesticide Use Database.

Data Analyses

We used the R package Unmarked (Fiske and Chandler

2011) to identify landscape-level variables associated

with A. fowled occupancy. Landscape level habitat data

were only available for a single year, so we used MacK-

enzie et al.’s (2002) occupancy model to account for

imperfect detectability, particularly false-negative detec-

tions, when evaluating habitat variables. Lalse-positive

detections can also result in high site occupancy biases,

but false-positive detection rates vary between species,

and previous tests of NAAMP volunteers resulted in no

false-positive detections of A. fowled , even amongst

inexperienced volunteers (Genet and Sargent 2003; Ro-

yle and Link 2006; McClintock et al. 2010). Therefore,

false-positive detections of A. fowled were unlikely to be

at levels high enough to bias site occupancy and were not

considered in the modeling process.

We assessed models using a multimodel inference

approach (see Burnham et al. 2011). We used 19 non-

correlated site covariates (Table 1) considered to be bio-

logically meaningful in anuran breeding site selection

when creating a priori models (Cushman 2006). Julian

date (date) and temperature (temp) were found to af-

fect detection probability; therefore these two sampling

covariates were used in all models. We ranked compet-

ing models with Akaike Information Criterion (for data

sets with high independent to dependent variable ratio

[AICc]) by calculating differences between candidate

models and the lowest AICc (A AICc) model. We used

Akaike weight (u\) for each model to guide selection.

To determine change in occupancy between years,

we fit a colonization-extinction model (MacKenzie et al.

2003) using date and temp as covariates to account for

differences in repeated sampling periods. We then used

a smoothed trajectory to determine mean occupancy for

each year (Weir et al. 2009). Serial autocorrelation in

Table 1 . Landscape variables used in analyses of A. fowled calling anuran survey data for in Virginia and Maryland.

Variable Description

Crops

Dec

Dev

Ever

For

Grass

Hay

Mix

Patch

Pesticides

Road

Shrub

Wavg

Wdis

Wet

Wnear

Wnum

Wtype

Proportion of area used for annual crops or perennial woody crops

Proportion of forest with >75% canopy cover of deciduous trees

Proportion of area that has been developed, including suburban and urban areas

Proportion of forest with >75% or more evergreen trees

Proportion of all forest habitats (Dec + Ever + Mix)

Proportion of area with graminoids or herbaceous vegetation covering over 80% of land which might be grazed but not tilled

Habitat Diversity, calculated as Shannon’s diversity index using habitat proportions

Proportion of area planted with grass/legume mixtures used for grazing or hay crops

Proportion of mixed forest with neither deciduous nor evergreen dominant

Number of terrestrial habitat patches divided by total number of possible habitat patches (i.e., if each raster square represented a

different type of habitat)

Total kg/km 2 of agricultural pesticides applied within 1 km radius of buffer from CSS

Total length of all roads in a 1 km radius buffer from CSS

Proportion of area with canopy less than 5 m tall (e.g., shrubs and early successional forest)

Average size of wetlands in a 1 km radius buffer from CSS ± in m 2

Distance (m) of nearest wetland from Calling Survey Site (CSS)

Proportion of total buffer area covered by wetlands

Size (ha) of the wetland nearest to the CSS

Number of wetlands in a 1 km radius buffer from CSS

Number of different types of wetlands in a 1 km radius buffer from CSS

May 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

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Jones and Tupper

the residuals violated assumptions of a parametric lin-

ear regression analysis; therefore we used a non-seasonal

Autoregressive Integrated Moving Average (ARIMA)

analysis to better understand changes in A. fowleri oc-

cupancy rates over time. We determined significance of

parameters using a conditional least squares estimation.

We assessed model fit with a Ljung-Box Q-test whereby

a high P-value indicates that autocorrelation functions

are not significantly different than white noise (Ljung

and Box 1978).

We used an empirical Bayesian approach to deter-

mine conditional distribution of occurrence from the

colonization-extinction model and then extrapolated best

unbiased predictions of occupancy probability at each

site. All sites with an occupancy probability > 0.75 were

considered occupied. To determine distributions of spe-

cific variables at occupied and unoccupied sites, we cre-

ated site occupancy accumulation curves for each habitat

variable. Habitat recommendations are based on maxi-

mum values found at 90% of occupied sites.

Trend analysis was completed in Minitab v.16 (Minit-

ab Inc., Pennsylvania, USA) and all other analyses were

completed in R v.3.0.2. Maps and figures were created

using QGIS 2.0 and Excel 2010 (Microsoft Corp., Wash-

ington, USA).

Results

Landscape analyses

Two hundred fifty sites were sampled between 1999 and

2012 throughout Maryland and Virginia, 108 (42.8%) of

which had at least one detection of A. fowleri. Approxi-

mately eight-percent (300/3841) of sampling events re-

sulted in detections. Not all sites were sampled every

year (ranging from 1-14 years, x = 6.22; + 0.197), but

most sites were surveyed > 10 times (i = 23.1; + 1.09).

Table 2. Top five models and full model from unmarked analy-

sis of A. fowleri occupancy in Virginia and Maryland. Mod-

els are ranked from lowest to highest with AICc values. Julian

date and temperature were used as sampling covariates in all

models. The full model was constructed using the maximum

number of site covariates which would create a model that con-

verged: Wnum + Wavg + Road + Ever + Dec + Mix + Crops +

Hay + Dev + Wet + Core + Patch + Pesticides.

Model

AICc

A. AICc

Dec + Hay + Dev + Pesticides

1768.65

0.62

Dec + Hay + Dev

1769.55

0.91

0.38

Dec + Hay + Pesticides

1780.54

11.90

0.00

Dec + Dev + Pesticides

1791.16

22.52

0.00

Pesticides

1964.45

195.80

0.00

Full model

2958.68

1190.03

0.00

Table 3. Transformed and untransformed beta coefficients from

top occupancy models.

Variable

Untransformed

Transformed

Estimated SE

(Intercept)

3.887

0.980

0.017

Dec

-6.148

0.002

Hay

-5.373

0.005

Dev

-4.814

0.008

0.009

Pesticides

-0.005

0.499

0.013

Bayesian analysis indicated that only two sites where no

detections occurred had an occupancy probability >0.75.

We found strong support for a model indicating that

deciduous forest, hay, development and pesticides nega-

tively influenced A. fowleri occupancy (AICc = 1768.65,

w. = 0.62; Tables 2 and 3). The buffers ranged from

0-97% deciduous forest (. x = 0.32; + 0.015), 0-74%

hay (v = 0.18; + 0.011), 0-83% development (x = 0.12;

+ 0.009), and 0-535 kg pesticides applied (v = 55.7; +

5.231). Ninety-percent of occupied sites were covered by

less than 25% development, 35% hay, and 50% decidu-

II.H

U.6

♦ ft*

♦ •

♦ ♦

* •

I Mi

U.fi

0.4

&• o.:

0.2 0.4 0.6

Proportion of habitat variable in buffer

IIkMuoiis forcst

* Duvdopfflcnl

O.ft

§ 0.4

0.2

hill 200 300 41 ID

Pesticides applied (kg/km-)

500

(410

Fig. 2. Accumulation curve showing the maximum proportions

of habitat variables found at each occupied site. Ninety -percent

of occupied sites were covered by less than 0.25 development,

0.35 hay, and 0.50 deciduous forest cover.

Fig. 3. Accumulation curve showing the maximum agricultural

pesticide application rates at occupied sites. The maximum

amount of pesticides applied at 90% of occupied sites was 165

kg/km 2 .

May 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

e97

Fowler’s Toad occupancy in the southern mid-Atlantic

ous forest (Fig. 2). The maximum amount of pesticides

applied at 90% of occupied sites was 165 kg/km 2 (Fig. 3).

Time-series analyses

Using ARIMA analysis, we found that a single autore-

gressive term was contributing to interannual changes

in A. fowleri occupancy rates ( t = 4.32, P < 0.001). We

confirmed that the model was valid, with uncorrelated

residuals (Q = 13.2, df = 9, P = 0.152). Trend analysis in-

dicated a downward trend in occupancy rates over time,

with occupancy decreasing from 55.3% in 1999 to 29.5%

in 2012 (Fig. 4).

'e.

£ 0,2

2000 2002 2006 20flX 2010 2012

Year

Discussion

Forest cover

Fig. 4. Mean site occupancy and confidence intervals of A.

fowleri in Virginia and Maryland from 1999-2012. Time se-

ries analysis indicates a 53% decrease in site occupancy, from

55.3% in 1999 to 29.5% in 2012.

Many contemporary landscape-level studies indicate

a positive relationship between amphibian species and

forested habitat within buffers of varying sizes around

breeding ponds (see Cushman 2006). We were able to

distinguish between forest types on a fairly large scale

and identified a negative relationship between A. fowleri

occupancy and deciduous forest. Although both species

can be sympatric, A. fowleri is largely replaced by Amer-

ican Toads ( Anaxyrus americanus) in later successional

forests that are dominated by moister, more nutrient rich

soils, and hardwood trees (Wright and Wright 1967;

Lazell 1976; Klemens 1993). Our results confirm long-

standing observations made across A. fowleri ' s range

that suggest they are more common in early successional

habitats that are either relatively open or dominated by

mixed or coniferous forest (Hubbs 1918; Hoopes 1930;

Netting and Goin 1945; Littleford 1946; Cory and Man-

ion 1955; Wright and Wright 1967; Clarke 1974; Green

1989; Lazell 1976; Klemens 1993; Zampella and Bun-

nell 2000; Tupper and Cook 2008).

Hay

The proportion of area covered by grass/legume mixtures

used for grazing or hay crops within the 1 km buffer was

The Provincelands of Cape Cod National Seashore, Barnstable County, Massachusetts, USA. The reddish vegetation in the center

of the photo is a cranberry ( Vaccinium macrocarpon) bog, a wetland used for breeding by the Fowler’s toad. The surrounding land-

scape is ideal for the Fowler’s toad and supports one of the largest populations of this species in the United States. The landscape

contains a patchwork of sand, pitch pine ( Pinus rigida ), scrub oak ( Quercus ilicifolia ), and dune grass ( Ammophila breviligulata).

Photo by Rebecca Flaherty.

May 2015 | Volume 9 | Number 1

Amphib. Reptile Conserv.

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Jones and Tupper

found to have a significantly negative impact on A. fowl-

ed occurrence. Agricultural development can negatively

affect anuran dispersal abilities, and soil compaction as-

sociated with agricultural landscape alterations may pro-

hibit anuran burrowing (Whalley et al. 1995; Jansen et

al. 2001; Gray et al. 2004b). Wetlands within agricultural

landscapes may be altered physically and biologically

such that postmetamorphic anurans emerge smaller and

presumably less fit (Beja and Alcazar 2003; Gray et al.

2004a, b).

Development

We found development to be another significant variable

negatively affecting A. fowled occurrence. Development

contributes to reduced genetic diversity in pond breeding

amphibians, increased pollution of upland and wetland

habitats, increased road mortality, and microclimate al-

teration of remaining habitat patches (Soule 1987; Reh

and Seitz 1989; Fahrig 1995; deMayndier and Hunter

1998; deMaynadier and Hunter 2000; Turtle 2001; Tinnn

and McGarigal 2014). Various studies indicate that de-

velopment and fragmentation is detrimental to amphib-

ian persistence (see review in Cushman 2006), including

A. fowled and congener A. amedcanus (Schlauch 1976,

1978; Gibbs et al. 2005; Walls et al. 2011). However,

studies conducted throughout the United States (e.g.

New Jersey [Zampella and Bunnell 2000], Pennsylvania

[Rubbo and Kiesecker 2005], North Carolina [Gooch

et al. 2006], and Louisiana [Milko 2012]) suggest that

A. fowled are urban tolerant. While these studies have

shown that A. fowled can occur in developed habitats,

they do not include pre-development population sizes,

are temporally and spatially limited, and are likely refer-

ring to suburbanization rather than large-scale urbaniza-

tion (see Schlauch 1978).

Pesticides

While the differences in AICc values between the top

two models were small, four out of the top five mod-

els included agricultural pesticide application levels as

a negative covariate, indicating that pesticide exposure

may play an important role in A. fowled site occupancy.

Data suggest that chemical pesticides associated with ag-

riculture have wide-ranging direct effects on amphibians,

including endocrine disruption, immunosuppression, de-

velopmental delays, and increased mortality (Mann et al.

2009). Exposure to insecticides was found to be highly

toxic to larval A. fowled in laboratory studies (see review

in Green 2005) and had sub-lethal effects its congener, A.

amedcanus , causing eye and limb deformities, increased

time to metamorphosis, and reduced post-metamorphic

body size (Harris et al. 2000; Boone and James 2003;

Howe et al. 2004). Agricultural runoff containing pes-

ticides may be contaminating certain wetlands in this

study, thus potentially accounting for reduced A. fowled

occupancy rates in agricultural landscapes.

Trends

By pooling data from Maryland and Virginia, we esti-

mate that A. fowled occupancy has decreased by approx-

imately 53% over the last 14 years. Weir et al. (2009)

found a significant, but negligible, negative occupancy

trend for A. fowled in Delaware, and indicated unchang-

ing occupancy rates in Virginia, Maryland, West Virgin-

ia, and New Jersey between 2001 and 2007. However, a

more recent study (conducted as the same time as ours,

with a similar data set, see Weir et al. 2014) also indicat-

ed A. fowled declines in Maryland. Differences in trend

estimates between our study and Weir et al. (2009, 2014)

may be due to differing sample sizes. Since we nearly

doubled the scope of analysis of Weir et al. (2009) and

have three more years than Weir et al. (2014), we be-

lieve our results more accurately describe trends in A.

fowled occupancy in Maryland and Virginia. Although

a more comprehensive analysis is needed to identify the

proximate causes of decline in A. fowled in Maryland

and Virginia, we suspect that its declines are in part due

to the recent loss of subclimax communities. Virginia

has lost 51.6% of its softwood forest since 1940 (VDOF

2014) and early successional habitats have been steadily

juxtaposing to later successional seres throughout Mary-

land: As of 2008 less than 10% of existing Maryland for-

ests were occupied by early successional regimes (Lister

2011).

Anaxyrus fowled may be able to persist longer in mod-

erately developed coastal environments (Schlauch 1978)

than other pond-breeding amphibians due to their high

fecundity rates, salt tolerance, desiccation resistance,

and ability to breed in wetlands with varying hydrope-

riod regimes (Wright and Wright 1967; Claussen 1974;

Markow 1997; Tupper and Cook 2008; Birx-Raybuck

2010; Eskew et al. 2012). The ability of A. fowled to oc-

cupy these types of habitats is advantageous because they

harbor fewer interspecific amphibian competitors (see

Martof et al. 1980; Klemens 1993; Mitchell and Reay

1999). If early successional habitats continue to become

less widespread in the mid-Atlantic, coastal regions may

become more important to the long-term persistence of

A. fowled. However, much of the southern mid- Atlantic

coastal plain is densely populated (VGDIF 2005) and in-

tensely developed. Thus, successional changes occurring

further inland coupled with increased urbanization of the

southern mid- Atlantic coastal plain could potentially ex-

acerbate declines.

Conclusions

Amphibian populations are more vulnerable to habitat

loss and fragmentation when located on the margins of

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Amphib. Reptile Conserv.

e97

Fowler’s Toad occupancy in the southern mid-Atlantic

their geographic range (Swihart et al. 2003). Our data in-

dicate that even in the middle of their range, A. fowleri

occupancy rates are declining. Landscapes most appro-

priate for this species appear to contain only moderate

amounts of deciduous forest (< 50%), few hay crops (<

35%), relatively little development (< 25%), and low

pesticide application rates.

Acknowledgments. — This project was made possible

by D. Marsh (Washington and Lee University), coordina-

tor of the National Science Foundation supported project,

“Toads, Roads, and Nodes: Collaborative Course-Based

Research on the Landscape Ecology of Amphibian Popu-

lations.” We would like to thank biologists and volun-

teers of the North American Amphibian Monitoring Pro-

gram for their efforts in data collection. C. Knights, B.

Bowman, and P. Maphumulo of Northern Virginia Com-

munity College provided logistical and financial sup-

port. We thank P. Paton (University of Rhode Island), B.

Timm (University of Massachusetts), A. Royle (Patuxent

Wildlife Research Center), G. Casper, D. Bradford, Mac

Given (Newman University), and Robert Cook (Cape

Cod National Seashore) for comments on drafts of this

manuscript.

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Kara Jones is a naturalist and graduate student in the Department of Environmental Science and Policy at

George Mason Univeristy. Her area of specilization is in worm snake ecology and species-habitat modeling.

Photographed by Kara Jones.

Todd Tupper is an associate professor of biology at Northern Virginia Community College. He teaches biol-

ogy, zoology, and biostatistics. His area of interest is amphibian monitoring and conservation. Photographed

by Natasha Abner.

May 201 5 | Volume 9 | Number 1

Amphib. Reptile Conserv.

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Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(1) [General Section]: 34-51 (e98).

Two new species of the genus Cylindrophis Wagler, 1828

(Squamata: Cylindrophiidae) from Southeast Asia

1 6 A.A. Thasun Amarasinghe, 2 Patrick D. Campbell, 3 Jakob Hallermann,

4 lrvan Sidik, Matna Supriatna, and 5 lvan Ineich

l Research Center for Climate Change, University of Indonesia, Gd. PAU Lt. 8.5, Kampus UI, Depok 16424, INDONESIA 2 Department of Life

Sciences, Darwin Centre, Natural History Museum, Cromwell Road, South Kensington, London SW7 5BD, ENGLAND 3 Zoologisches Museum

Hamburg, Universitdt Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, GERMANY ^Museum Zoologicum Bogoriense, The Indonesian

Institute of Science (LIPI), Widyasalwaloka Building, Jl. Raya Jakarta Bogor, Cibinong 16911, INDONESIA 5 Museum national d’Histoire naturelle,

Sorbonne Universites, ISYEB (Institut de Systematique, Evolution et Biodiversite), UMR 7205 (CNRS, MNHN, UPMC, EPHE), 57 rue Cuvier, CP

30 (Reptiles), 75005 Paris, FRANCE

Abstract. — The original description of Anguis ruffa (now Cylindrophis ruffus) given by Laurenti in

1768 is not sufficiently comprehensive for the morphological identification of the species, and the

type locality, given as “Surinami,” is in error. However, Schlegel in 1844 corrected the type locality

as “Java in Indonesia.” There is also, currently, no proof of the existence of a type specimen of

Anguis ruffa. Therefore, we accept Schlegel’s correction of the type locality being Java. Anguis ruffa

is here redescribed based on museum specimens collected from Java only. Because the original

description of C. r. burmanus is insufficiently comprehensive we here redescribe this species using

the presumed type series collected from Myanmar, and we also designate a lectotype. We examined

a large number of Cylindrophis specimens deposited in European and Indonesian museums,

using morphological and meristic characters, plus coloration. We identified four groups based

on the number of scale rows around the midbody (17, 19, 21, and 23). Among the Cylindrophis

collections at Natural History Museum, London and Museum national d’Histoire naturelle, Paris, we

have discovered several specimens which do not fit any recognized species descriptions. We here

describe two new species chosen from among them: C. jodiae sp. nov. from Vietnam and C. mirzae

sp. nov. from Singapore. Finally, we provide color plates showing the different body colorations for

all the recognized species in the genus Cylindrophis.

Key words. Biogeography, Indonesia, pipe snake, Singapore, taxonomy, Vietnam

Citation: Amarasinghe AAT, Campbell PD, Hallermann J, Sidik I, Supriatna J, Ineich I. 2015. Two new species of the genus Cylindrophis\Nag\er, 1828

(Squamata: Cylindrophiidae) from Southeast Asia. Amphibian & Reptile Conservation 9(1) [General Section]: 34-51 (e98).

Copyright: © 201 5 Amarasinghe et al. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommer-

cialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation', official journal website <amphibian-

reptile-conservation. org> .

Received: 23 May 201 5; Accepted: 03 June 201 5; Published: 1 0 June 201 5

Introduction

The first species of pipe snake was described by Linnae-

us (1758) as Anguis maculata from Sri Lanka (America

in error fide Deraniyagala 1955), followed by Anguis

ruffa described by Laurenti (1768). The genus Cylindro-

phis was established by Wagler (1828) with a type spe-

cies from Java, Cylindrophis resplendens Wagler 1828, a

binomen later synonymized with Cylindrophis ruffus by

Schlegel (1844). After Wagler (1828), several additional

species (e.g., Cylindrophis melanotus Wagler 1830, Cyl-

indrophis lineatus Blanford 1881, Cylindrophis isolepis

Correspondence. Email: fhasun@rccc.ui.ac.id

Boulenger 1896, Cylindrophis opisthorhodus Boulenger

1897, Cylindrophis boulengeri Roux 1911, Cylindro-

phis aruensis Boulenger 1920, Cylindrophis celebensis

Smith 1927, Cylindrophis heinrichi Ahl 1933, Cylindro-

phis engkariensis Stuebing 1 994, Cylindrophis yamdena

Smith and Sidik 1998) and one subspecies, Cylindrophis

rufus burmanus. Smith 1943) were added to the genus.

Most of the taxa are endemic to one island or small island

group. The Asian genus Cylindrophis was formerly in-

cluded in the family Uropeltidae, later McDowell (1975)

included the genus in the family Cylindrophiidae, along

with the genus Anomochilus Berg 1901. Furthermore,

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e98

Amarasinghe et al.

McDowell (1975) synonymized C. celebensis and C.

heinrichi with the Sulawesi endemic C. melanotus. Re-

cently, Wallach et al. (2014) synonymized the trinomen,

C. r. burmcinus with C. ruffus. Therefore, the genus pres-

ently consists often valid monotypic species (Wallach et

al. 2014) of which nine are distributed in Southeast Asia

(Stuebing 1994), and one (C. maculatus) is endemic to

Sri Lanka (Somaweera 2006). There are no Cylindrophis

in the Indian peninsular (Smith 1943). Later, Cundall et

al. (1993) allocated the genus Anomochilus to its own

family, Anomochilidae, thus rendering the family Cylin-

drophiidae monotypic. The family Cylindrophiidae can

be distinguished from its sister family Anomochilidae by

having a mental grove, nasals in contact, and no preocu-

lar (Das et al. 2008).

The original description of Anguis rujfa is limited to

only a few words: “Corpore aequali, ruffo, lineis trans-

versalibus albis interruptis; abdomine vario,” and the type

locality was given as “Surinam!, ” in error. An English

translation of the original description was given by Adler

et al. (1992) as “Body uniform, red, broken white trans-

verse bands; abdomen various. Lives in Surinam; housed

in Gronovius’s Museum.” The holotype was deposited

first at “Mufeo Gronoviano” / Museo Laurentii Theodori

Gronovii [may be Museum Gronovianum], Lugdunum

Batavorum (= Leiden) and later believed to have been

transferred to the Naturalis Biodiversity Center, Nation-

aal Natuurhistorische Museum (Rijksmuseum), Leiden,

Netherlands (RMNH.RENA). Even though Iskandar and

Colijn (2002) regarded the type materials of Anguis rujfa

as lost from Naturalis Biodiversity Center, there is no ev-

idence to support that the type was ever deposited there

in the first place. Wagler’s (1828) species Cylindrophis

resplendens , which was described from Java, was syn-

onymized with Cylindrophis ruffus by Schlegel (1844).

The type locality of Anguis rujfa was later corrected from

Suriname to Java in Indonesia {fide Schlegel 1844). Al-

though Gray (1849) considered that the Javanese popula-

tion is a variation ( javanica ) of Anguis rujfa , subsequent

authors have accepted that the type locality is indeed

Java (e.g.. Smith 1943, Taylor 1965). Furthermore, Tay-

lor (1965) invalidated [id est nomen oblitum ] Laurenti’s

(1768) name and choose the next available name, Tortrix

rufa Schlegel 1844 [id est nomen protectum ].

Although the species name was usually spelled erro-

neously as “ rufus ” in older publications, a justification is

given for this by Adler et al. (1992) as the original spell-

ing given by Laurenti (1768) is “rujfa.” Smith (1943)

described a subspecies, Cylindrophis rufus burmanus ,

from Burma (now Myanmar). In the original descrip-

tion of C. r. burmanus, he failed to mention how many

specimens he examined, but it is clear from the descrip-

tion that he had several specimens at his disposal at the

time. According to Smith (1943), ventrals varied from

201-225, and subcaudals from 5-7, but he only provid-

ed the measurements for the largest specimen as “Total

length: 330, tail 10 mm” [i.e., SVL 320 mm]. The dis-

Amphib. Reptile Conserv. 35

tribution was given as “Range. Tenasserim and Burma

as far North as Myitkyina” but a precise type locality

was not given. Furthermore, Smith (1943) extended the

distribution of “Cylindrophis rufus rufus ” (forma typica)

from Java to Siam (now Thailand), French Indo-China,

Malay Peninsula and Archipelago. Although subsequent

authors (e.g., Taylor 1965, Iskandar and Colijn 2002)

have accepted the above trinomen from Myanmar, re-

cently Wallach et al. (2014) included the subspecies un-

der Cylindrophis ruffus because subspecies are not rec-

ognized in their catalogue. To date Cylindrophis ruffus

has been widely recorded from Thailand, Laos, Vietnam,

Myanmar, Cambodia, China, Malaysia, Singapore, and

several Indonesian islands including Sumatra, Borneo,

Java, and Sulawesi (Gray 1849; Boulenger 1888, 1893;

Smith 1943; Deuve 1970; Dowling and Jenner 1988;

Adler et al. 1992; Geissler et al. 2011; de Lang 2011,

201 3). Several of these authors refer to undescribed spe-

cies and it has also already been argued that C. ruffus

being a complex of several species, is in need of a critical

examination (Stuebing 1994; Smith and Sidik 1998). Our

results below will be a preliminary step to the recognition

of several unidentified species within this complex. An

additional number of undescribed species are included

in our examined material but pending larger samples, we

prefer not to describe them at this time.

Materials and Methods

We have examined more than 150 Cylindrophis speci-

mens deposited in various museum collections (Appen-

dix 1): Museum Zoologicum Bogoriense, Bogor, Indo-

nesia (MZB); Natural History Museum [formerly British

Museum (Natural History)], London, United Kingdom

(UK) (BMNH); Museum national d’Histoire naturelle,

Paris, France (MNHN-RA); Senckenberg Forschung-

sinstitut und Naturmuseum, Frankfurt, Germany (SMF);

Western Australian Museum, Perth, Western Australia,

Australia (WAM); and Museum fur Naturkunde, Berlin,

Germany (ZMB). We compared all our examined speci-

mens with past descriptions and other published data of

all known congeners (Appendix 1). Museum acronyms

follow Sabaj Perez (2014).

We obtained distribution data from examined speci-

mens, published literature as well as personal observa-

tions. The following characters were measured with a

Mitutoyo digitmatic caliper to the nearest 0.1 mm and

only along the left side of the body for symmetrical char-

acters: snout-vent length (SVL), measured from tip of

snout to anterior margin of vent; tail length (TL), mea-

sured from anterior margin of vent to tail tip. We counted

supralabial and infralabial scales from the gape to the

rostral and mental scales, respectively. We counted mid-

body dorsal scale rows around the body, in three posi-

tions, on the neck (at the point of the 10 th scale starting

from the first scale after the mental groove on the ventral

June 2015 I Volume 9 I Number 1 I e98

Two new species of the genus Cylindrophis from Southeast Asia

side), midbody (at the point of half of the ventral count),

and at one scale anterior to precloacal, always excluding

the ventral scale from counts. When counting the number

of ventral scales, we scored specimens according to the

method described by Dowling (1951), but started from

the first scale after the mental groove. We counted sub-

caudal scales from the first postcloacal scale to the scale

before the tip of the tail.

All color descriptions and other associated color char-

acters are based on preserved specimens. The presence

and absence of white bands on the nape and the back,

plus the shape of the band on the nape (narrow, when

the band is wider than one scale-width; or wide, when

the band is wider than one scale-width), the shape of the

bands at back (complete, when the band is a complete

dorsal ring; or interrupted, when the bands do not meet

mid-dorsally), and the arrangements of the bands at back

(constant, when the bands are regularly arranged and

each part of the band arranged confronting each other;

or alternating, when the bands are irregularly arranged

and each part of the band arranged avoiding each other)

are considered as morphological characters. We have not

recorded the sex of the specimens other than where the

hemipenis was everted because most of the examined

specimens are old, having huge historical value, we de-

cided to keeping them intact.

The distribution of each species (in Fig. 8) does not

show precise localities (due to the general lack of precise

localities in historical collections). Therefore, the whole

biogeographical area or country is shaded for each spe-

cies.

Results

The original description of Anguis ruffa given by Lau-

renti (1768) is not comprehensive enough for a morpho-

logical identification of the species. Our recent attempt

to locate the type material of C. ruffus at RMNH was

unsuccessful (Marinus Hoogmoed pers. comm, to Ivan

Ineich on 23 October 2014). We believe that there are

several species masquerading today within the current-

ly accepted name Cylindrophis ruffus. We also believe

that the type of C. ruffus first arrived in the Netherlands

from Jakarta, Indonesia because of the following: (1) the

Dutch East Indies (now Indonesia) was a Dutch colony

under the administration of the Dutch Government since

the early 17 th century; (2) most of the specimens arriv-

ing at the Netherlands natural history collections prior

to 1850 originated from Java, Indonesia, especially West

Java which was where the administration capital was

based, Batavia (now Jakarta); (3) there was a town called

“Batavia” in the former Dutch colony of Suriname which

could be misidentified with Batavia in Indonesia. There-

fore, we accept Schlegel’s (1844) correction for the type

locality of Anguis ruffa as Java. Furthermore, our attempt

to locate the type material of Cylindrophis resplendens

Wagler, 1828 (type locality: Java) which was believed to

be deposited at MNHN-RA was again unsuccessful.

Our species examination and comparison also shows

that Cylindrophis ruffus burmanus has morphological

and meristic character differences large enough to el-

evate it to species level. Although Iskandar and Colijn

(2002) raised it previously to the species level, they gave

no justification for this. There are six specimens (see Ta-

ble 1) collected from Burma in the BMNH today, among

them two specimens (BMNH 1940.3.3.1-2) are labelled

“ Cylindrophis ruffus burmensis ” and all the others as

“ Cylindrophis rufus .” As these specimens were probably

present at the time of Smith when his manuscript was

completed in 1938 [ fide the preface of Smith (1943)] but

delayed because of the second world war, they may be

considered syntypes of C. r. burmanus. Although, there

Table 1. Details of the possible syntype series of Cylindrophis ruffus burmanus Smith, 1943 compared to the data provided in the

original description; “ — ” = not applicable, “?” = not given, = damaged.

Character

Smith (1943)

Catalogue Number (BMNH)

1940.3.3.1

1940.3.3.2

1891.11.26.28

1908.6.23.3

1925.12.22.4

1925.4.2.2

Species name

on the label

—

“ Cylindrophis ruffus

burmensis ”

“ Cylindrophis rufus ”

Location

Tenasserim

and Burma as

far North as

Myitkyina

Rangoon,

Burma

Rangoon,

Burma

Pyimnana, Up-

per Burma

Burma

Sahmaw, Myit-

kyina District,

Burma

Thandoun,

Burma

Presenter (col-

lector unknown)

F.J. Meggitt

E.W. Oates

F. Wall

Total length in

330

217.2

286

299.5

264

288

SVL in mm

[320]

320

212

280

293

256

280

Tail length in

5.2

6.5

Ventrals

201-225

213

209

221

-225*

201

Subcaudals

5-7

-6-7*

-6-1*

-5*

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Amarasinghe et al.

Table 2. Comparison of some morphometric, meristic, and morphological characters of Cylindrophis species which have 23, 19,

and 17 midbody scale rows, based on examined materials; “ — ” = Not applicable.

Character

23 scale rows at midbody

19 scale rows at midbody

17 scale

rows at

midbody

C. aruensis

(« = 3)

C. opistho-

rhodus in = 5)

C. bouleng-

eri {n = 3)

C. burma-

nus {n = 6)

C. melano-

tus in = 14)

C. ruffus

in =14)

C. eng-

kariensis

in = 1)

Location

Dammar

Flores

Wetar

Myanmar

Sulawesi

Java

Borneo

SVL (in mm)

155-305

270-470

240-310

212-320

292-575

257-715

473

Scale rows around neck

25-26

21-23

19-21

17-19

20-23

Scale rows around midbody

Scale rows around precloacal

18-20

18-21

16 or 17

17 or 18

Midventral scales

173-182

185-210

193-200

201-225

233-275

186-197

234

Subcaudals

6-7

5-6

5-7

6-8

5-8

5+1

Frontal > prefrontal ( 1) or < (0)

Pale band/ring on the nape pres-

ent (1) or absent (0)

Pale band/ring wide (1) or narrow

(0)

—

Pale band/ring complete (1) or

dorsally interrupted (0)

—

Crossbands on the back present

(1) or absent (0)

Crossbands complete (1) or inter-

rupted (0)

—

Crossbands constant (1) or alter-

nating (0)

—

Crossbands wide (1) or narrow

(0)

—

is no indication of a holotype designation in the descrip-

tion, the specimen BMNH 1940.3.3.1 is exactly match-

ing with the morphometric, meristic, and morphological

characters given in the original description. We note also

that the original description of C. r. burmanus was not

comprehensive enough for identification. Therefore, we

here designate the closely matching specimen (BMNH

1940.3.3.1), for which measurements were given in the

original description, as the lectotype of Cylindrophis ruf-

fus burmanus in order to stabilize the name with a recog-

nized type specimen. Furthermore, we provide a compre-

hensive redescription on the basis of that lectotype, and

its five paralectotypes located at the BMNH.

Among our examined sample at the BMNH and

MNHN-RA, we found several specimens representing

two morphospecies that do not fit the diagnoses of any

known species. These specimens are morphologically

distinct, geographically isolated, and well outside of the

corrected distribution range of C. ruffus. The differences

of those two morphospecies are large enough to consider

them as “undescribed species.” Therefore, we formally

describe them as new species in this paper. They differ

from all other known species of the genus Cylindro-

phis (see Tables 2-3; Figs. 1-16) with respect to their

coloration and body scalation, especially their midbody

dorsal scale counts and ventrals. We assign the two new

species to the genus Cylindrophis based on the follow-

ing character combination: a medium-sized snake with

a cylindrical body, of nearly equal diameter throughout

its length; a small head, not really distinct from the thick

neck; a depressed snout; small eyes, with rounded pupils;

the nostril pierced in the middle of a single nasal shield,

slightly directed forwards; the upper head scales large

and symmetrical; no intemasals, loreals, or preoculars; a

mental groove present; 2 1 rows of smooth and iridescent

dorsal scales; the ventral scales barely enlarged, and the

tail very short and blunt.

Furthermore, based on the number of scale rows

around the midbody we have identified four morphologi-

cal groups within the Cylindrophis genus. Note however

that the phylogenetic validity of those groups has not yet

been tested:

(1) 17 midbody scale rows (one species): C. eng-

kariensis — see Table 2.

(2) 19 midbody scale rows (five species): C. ruffus , C.

melanotus, C. boulengeri , C. burmanus — see Table 2.

(3) 21 midbody scale rows (seven species): C. macu-

latus , C. lineatus , C. isolepis , C. yamdena, C. jodiae

sp. nov. (see below), C. mirzae sp. nov. (see below) —

see Table 3.

(4) 23 midbody scale rows (two species): C. opistho-

rhodus, C. aruensis — see Table 2.

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June 201 5 I Volume 9 I Number 1 I e98

Two new species of the genus Cylindrophis from Southeast Asia

Systematics: We redescribe Cylindrophis rujfus and C.

burmanus and describe two new species from Vietnam

and Singapore, respectively, as follows:

Cylindrophis ruff us (Laurenti 1768)

Anguis ruffa Laurenti 1768: 71.

(Figs. 1, 8; Table 2)

Synonyms:

Cylindrophis resplendens Wagler 1828: pi. 5, fig. 1. Type

locality, Java.

Cylindrophis rufa javanica Gray 1849: 112. Type local-

ity, Java.

Proposed standard English name: Red-Tailed Pipe-

Snake

Proposed standard Indonesian name: Ular Pipa Ekor

Merah

Remarks: Here we accept the correction of the type lo-

cality made by Schlegel (1844). We have failed to find

another species of Cylindrophis sympatric with C. ruf-

fus in Java among the specimens examined. However the

biogeographical range of C. rujfus could extend beyond

Java, e.g., Borneo and Peninsular Malaysia — see Stue-

bing (1994: Table 1).

Examined materials (14): MZB 1418, (SVL 715 mm),

Burial, Bogor, West Java, Indonesia; MZB 3816, 1433,

(SVL 325 mm, 350 mm), Banten, Indonesia; MZB

300, 301, 304, 309, 1049, 2000, (SVL 580 mm, 550

mm, 520 mm, 560 mm, 540 mm, 650 mm); MNHN-

RA 1975.0073-74, 3280, 2007.2452 (formerly 3280A),

7182, (SVL 258 mm, 300 mm, 517 mm, 466 mm, 257

mm), Java, Indonesia.

Diagnosis: Cylindrophis rujfus is distinguished from

all congeners by having the following characters: 19

midbody scale rows (vs. 17 in C. engkariensis; 21 in C.

isolepis, C. lineatus, C. maculatus, C. yamdena; 23 in C.

aruensis, C. opisthorhodus ), 186-197 ventrals (vs. 233-

275 in C. melanotus; 201-225 in C. burmanus), wide and

constant bands encircling dark body (vs. dorsum uniform

black with no cross bands in C. boulengeri:, narrow and

alternating bands on paler body in C. burmanus), an in-

terrupted and wide band on the nape (vs. no ring on the

nape in C. boulengeri:, a complete and narrow ring encir-

cling the nape in C. burmanus).

Description of examined materials: SVL 257-715 mm;

body elongate, rounded in cross section; head not distinct

from neck, broadened and dorsoventrally flattened in the

orbital and sagittal regions; snout rounded in dorsal and

lateral view.

Rostral shield large, somewhat visible from a dorsal

perspective with a conical apex; a single nasal, widely in

contact behind the rostral, no internasals; nasals in con-

tact with rostral anteriorly, with prefrontal dorsally, and

the first and second supralabials ventrally; nostrils large;

canthus rostralis weakly defined; prefrontal somewhat

larger than the frontal and quadrangular; frontal large, tri-

angular, and length same as its width; supraocular wide,

triangular, posteriorly wider; parietal small, triangular,

Table 3. Comparison of some morphometric, meristic, and morphological characters of Cylindrophis species which have 2 1 mid-

body scale rows, based on examined materials; “ — ” = Not applicable.

Character

21 scale rows at midbody

C. isolepis

in = 7)

C. lineatus

(n = 2)

C. maculatus

(n = 33)

C. yamdena

(n = 5)

C. jodiae sp.

nov. (« = 11)

C. mirzae sp.

nov. (n = 4)

Location

Jampea

Borneo

Sri Lanka

Yamdena

Vietnam

Singapore

SVL (in mm)

320-500

540-713

262-378

500-610

146-656

220-693

Scale rows around neck

21-23

20-21

19 or 20

Scale rows around midbody

Scale rows around precloacal

16 or 17

15-18

17 or 18

Mid ventral scales

217-225

215-218

195-208

179-193

182-196

196-217

Subcaudals

5-6

6-8

4-6

5-7

4-7

Frontal > prefrontal (1) or < (0)

Pale band/ring on the nape present (1) or

absent (0)

Pale band/ring wide (1) or narrow (0)

—

Pale band/ring complete (1) or dorsally

interrupted (0)

—

Crossbands on the back present (1) or

absent (0)

Crossbands complete (1) or interrupted

—

Crossbands constant (1) or alternating (0)

—

Crossbands wide (1) or narrow (0)

—

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June 201 5 I Volume 9 I Number 1 I e98

Amarasinghe et al.

Fig. 1. Coloration of Cylindrophis ruffus MZB 1418 (A) head

in dorsal view, (B) head in ventral view, (C) head in lateral

view, (D) midbody in dorsal view, (E) midbody in ventral view,

and (F) tail in ventral view.

Fig. 2. Coloration of Cylindrophis burmanus lectotype, BMNH

1940.3.3.1 (A) head in dorsal view, (B) head in ventral view,

body in ventral view, and (F) tail in ventral view.

Fig. 3. Scalation of Cylindrophis burmanus lectotype, BMNH 1940.3.3.1 (A) head in dorsal

view, (B) head in ventral view, (C) head in lateral view, (D) tail in ventral view, (E) midbody in

dorsal view, (F) midbody in ventral view.

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June 2015 I Volume 9 I Number 1 I e98

Two new species of the genus Cylindrophis from Southeast Asia

Fig. 4. Coloration of Cylindrophis jodiae sp. nov. holotype,

MNHN-RA 1911.0196 (A) head in dorsal view, (B) head in

ventral view, (C) head in lateral view, (D) midbody in dorsal

view, (E) midbody in ventral view, and (F) tail in ventral view.

Fig. 6. Coloration of Cylindrophis mirzae sp. nov. holotype,

MNHN-RA 3279 (A) head in dorsal view, (B) head in ventral

view, (C) head in lateral view, (D) midbody in dorsal view, (E)

midbody in ventral view, and (F) tail in ventral view.

Fig. 5. Scalation of Cylindrophis jodiae sp. nov. holotype, MNHN-RA 1911.0196 (A) head in

dorsal view, (B) head in ventral view, (C) tail in ventral view, (D) head in lateral view, (E) tail in

lateral view, (F) body in dorsal and ventrolateral view.

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e98

Amarasinghe et al.

its rear border pointed, bordered by supraocular, frontal

shield, upper posterior temporal shield, occipital shield,

and two dorso-nuchal shields posteriorly on each side,

the occipital shield is of equal size as other dorso-nuchal

scales; loreal and preocular absent; eye small, pupil

rounded; eye in broad contact with supraocular dorsally,

prefrontal and third supralabial anteriorly, fourth supral-

abial ventrally, and postocular posteriorly; a single small

postocular, quadrangular, posteriorly wider, in broad

contact with supraocular, anterior temporal, and fourth

supralabial; temporals 1+2, triangular; anterior temporal

larger than posteriors; anterior temporal in contact with

supraocular and both posterior temporal with parietal

dorsally, 4 ,h -6 th supralabials ventrally; anterior temporal

does not meet parietals.

Six supralabials, 3 rd -5 th larger in size; first supralabi-

al in contact with rostral anteriorly and nasal dorsally;

second supralabial in contact with nasal and prefrontal

dorsally, third supralabial in contact with prefrontal and

eye dorsally, fourth supralabial in contact with the eye,

postocular, and anterior temporal dorsally; fifth supra-

labial in contact with anterior and posterior temporals;

sixth supralabial in contact with lower posterior temporal

dorsally and body scales posteriorly.

Mental small, triangular; first infralabial pair larger

than mental plate and in broad contact with each other,

in contact with anterior chin shield posteriorly; six in-

fralabials in total, l st -3 rd in contact with first chin shield,

4 th -6 th in contact with gular scales, and not touching the

chin shields; anterior chin shields larger than posterior

ones; a mental groove continues from the posterior tip of

the mental until the posterior chin shields.

Body slender; transverse dorsal scale rows (20-23)-

1 9 — (17 — 18), all smooth, subcycloid, and weakly imbri-

cate; vertebrals and midventrals undifferentiated from

adjacent scales; 186-197 ventrals; cloacal plate divided,

precloacal undivided and triangular, tail extremely short,

relative TL (TL/total length) 2. 1-2.9%, with a conical ro-

bust and thick tip; 5-8 entire subcaudals.

Coloration: All the examined specimens have a reddish

brown back with wide and incomplete lighter bands en-

circling the body along dorsal surface from behind nape

to tail, each band covering about two scales; head entirely

darker, an incomplete, wide ring encircling the nape; the

venter is dark brown with regular, cream colored stripes,

divided at midline. See Fig. 1 for details of coloration in

preservative.

Distribution: Cylindrophis rujfus is recorded from Java,

Indonesia (Fig. 8). Possible type locality is Batavia (now

Jakarta) in Indonesia (not Batavia in Suriname).

Cylindrophis burmanus Smith 1943

Cylindrophis rufus burmanus Smith 1943: 97

(Figs. 2, 3, 8; Tables 1, 2)

Amphib. Reptile Conserv. 41

Proposed standard English name: Burmese Pipe-Snake

Lectotype (designated herein): BMNH 1940.3.3.1,

(SVL 320 mm), collected from Rangoon, Burma (now

Myanmar) by an unknown collector, collection date

unknown. This specimen was presented to BMNJi by

Professor F.J. Meggitt, University College Rangoon (ac-

cording to the museum registry). Although Smith (1943)

had several specimens at his disposal at the time, he pro-

vided the measurement for only the largest specimen in

the series. Because the original description is not com-

prehensive enough, and because of the fact that the Cyl-

indrophis population in Myanmar may represent more

than one species, in order to stabilize the name with a

recognized type specimen, we here designate BMNH

1940.3.3.1 as the lectotype.

Paralectotypes (6): BMNH 1940.3.3.2, (SVL 212 mm),

collected from Rangoon, Burma by an unknown collec-

tor, presented by F.J. Meggitt; BMNH 1908.6.23.3, (SVL

293 mm), Burma, collector and date unknown, presented

by Major F. Wall; BMNH 1891.11 .26.28, (SVL 280 nun),

Pyinmana, Upper Burma, collector and date unknown,

presented by E.W. Oates; BMNH 1925.4.2.2, (SVL 280

nun), Thandoung, Burma, collector and date unknown,

presented by F. Wall; BMNH 1925.12.22.4, (SVL 256

nun), Sahmaw, Myitkyina District, Burma, collector and

date unknown, presented by F. Wall; and probably ZMB

3094 (fide Iskandar and Colijn 2002; indicated no justi-

fication). All these paralectotypes share the same charac-

ters as the lectotype and belong to the same species.

Diagnosis: Cylindrophis burmanus is distinguished from

all congeners by having the following characters: 19

midbody scale rows (vs. 17 in C. engkariensis ; 23 hi C.

aruensis , C. opisthorhodus; 21 in C. isolepis, C. linea-

tus , C. maculatus , and C. yamdena ), 201-225 ventrals

(vs. 233-275 in C. melanotus ; 193-200 in C. boulengeri;

186-197 in C. rujfus), narrow and alternating bands on

paler body (vs. dorsum uniform black with no crossbands

in C. boulengeri; wide, constant, dorsally interrupted

bands encircling the dark body in C. rujfus), a complete

and narrow ring encircling the nape (vs. no ring on the

nape in C. boulengeri; a wide, dorsally interrupted band

encircling the nape hi C. rujfus).

Description of lectotype: SVL 320 mm, tail length 10

mm; body elongate, rounded in cross-section; head not

distinct from neck, broadened and dorsoventrally flat-

tened in the orbital and sagittal regions; snout rounded in

dorsal and lateral view.

Rostral shield large, visible from above with a conical

apex; a single nasal, widely hi contact behind the rostral,

no intemasals; nasals in contact with rostral anteriorly,

with prefrontal dorsally, and the first and second supra-

labials ventrally; nostrils large; canthus rostralis weakly

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Two new species of the genus Cylindrophis from Southeast Asia

defined; prefrontal hexagonal, larger than frontal; fron-

tal large, triangular, and longer than width; supraocular

wide, triangular, wider posteriorly; parietal small, trian-

gular, its rear border rounded, bordered by supraocular,

frontal shield, upper posterior temporal shield, occipital

shield, and two dorso-nuchal shields posteriorly on each

side, the occipital shield smaller than other dorso-nuchal

scales; loreal and preocular absent; eye small, pupil

rounded; eye in broad contact with supraocular dorsally,

prefrontal and third supralabial anteriorly, fourth supral-

abial ventrally, and postocular posteriorly; a single large

postocular, subtriangular, posteriorly narrow, in broad

contact with supraocular, anterior temporal, upper pos-

terior temporal, and fourth supralabial; temporals 1+2,

all triangular; anterior temporal smaller than upper pos-

terior; anterior temporal in contact with both posterior

temporals, 4 th and 5 th supralabials ventrally; anterior tem-

poral does not meet parietals.

Five supralabials, 3 rd and 4 th largest in size; first supral-

abial in contact with rostral anteriorly and nasal dorsally;

second supralabial in contact with nasal and prefrontal

dorsally; third supralabial in contact with prefrontal and

eye dorsally; fourth supralabial in contact with eye, post-

ocular, and anterior temporal dorsally; fifth supralabial in

contact with anterior and posterior temporals.

Mental small, triangular; first infralabial pair larger

than mental plate and in broad contact with each other, in

contact with anterior chin shield posteriorly; five infral-

abials in total, l st -3 rd in contact with first chin shield, 4 th

and 5 th in contact with gular scales, and not touching the

chin shields; anterior chin shields larger than posterior

ones; a mental groove continues from the posterior tip of

the mental until the posterior chin shields.

Body slender; transverse dorsal scale rows 19-19-17,

all smooth, subcycloid, and weakly imbricate; vertebral

and midventral scales undifferentiated from adjacent

scales; 213 ventrals; cloacal plate divided, precloacal un-

divided and triangular, tail extremely short, relative TL

(TL/total length) 3.0%, with a conical thick and robust

tip; 6 or 7 (damaged) entire subcaudals.

Coloration: The lectotype (the largest specimen of the

original syntypes) has a brown back with narrow and al-

ternating white stripes along dorsal surface from behind

nape to tail, each stripe covering about half of one scale;

head entirely dark, a complete, narrow ring encircling

the nape; the venter is brown with regular, mottled cream

colored bars. See Fig. 2 for details of coloration in pre-

servative.

Variation of paralectotypes: SVL range from 256-293

mm; body scale rows at neck ranges from 17-19; ven-

trals 201-225; relative TL 2. 1-2.9%.

Distribution: Cylindrophis burmanus is only reported

from Myanmar (Fig. 8).

Cylindrophis jodiae sp. nov. Amarasinghe, In-

eich, Campbell & Hallermann

(Figs. 4, 5, 8; Table 3)

urn:lskl:zoobank.org:act:2D375EF3-B484-49F6-8F19-C9D5B7CD0CCC

Proposed standard English name: Jodi’s Pipe-Snake

Holotype: MNHN-RA 1911.0196, SVL 415 mm, col-

lected from Annam, Central Vietnam, by the French bot-

anist Philippe Eberhardt, without precise date, but before

1911.

Paratypes (10): MNHN-RA 1974.1251, (SVL 391 mm),

collected in the area of Saigon, southern Vietnam, by Ser-

gent Poilane before 1974; MNHN-RA 1885.0100-103,

(SVL 265, 264, 146, 177 mm), collected in Cochinchina,

southern Vietnam, by Girard before 1885; MNHN-RA

1885.0098-99, (SVL 375, 656 mm), collected in Co-

chinchina, southern Vietnam, by Girard before 1885;

MNHN-RA 1935.0001, (SVL 271 mm), collected in

Cochinchina, southern Vietnam, by Rene Bourret before

1935; MNHN-RA 1974.1253, (SVL 192 mm), collect-

ed in the area of Saigon, southern Vietnam, by Sergent

Poilane before 1974; BMNH 1920.1.20.2649, (SVL 345

mm), collected from Long-Xuyen, Vietnam by F. Lataste,

collection date unknown.

Diagnosis: Cylindrophis jodiae sp. nov. is distinguished

from all congeners by having the following characters:

21 midbody scale rows (vs. 17 in C. engkariensis; 19 in

C. boulengeri , C. burmanus , C. melanotus, C. ruff us; 23

in C. aruensis , C. opisthorhodus), 182-196 ventrals (vs.

217-225 in C. isolepis ), wide and interrupted bands on

the back (vs. lateral and middorsal stripes along the body

in C. lineatus; two series of large reddish-brown spots

along the back, which are enclosed by a black network in

C. maculatus ; no bands and paler back in C. yamdena).

Description of holotype: An adult, SVL 420 mm, tail

length 10.1 mm; body elongate (largest body diameter at

midbody is 23.8 mm), flattened laterally in cross section;

head not distinct from neck, broadened and dorsoven-

trally flattened in the orbital and sagittal regions; snout

rounded in dorsal and lateral view.

Rostral shield large, visible from above with a conical

apex; a single nasal, widely in contact behind the ros-

tral, no intemasals; nasals in contact with rostral anteri-

orly and prefrontal posteriorly, and the first and second

supralabials ventrally; the holotype has its right nasal

in contact with the left prefrontal by a point, which is

an anomaly; nostrils large; canthus rostralis weakly de-

fined; prefrontals slightly larger than the frontal, and

pentagonal; frontal small, triangular, and same length as

its width (length 3.8 mm, width 3.7 mm), equal or some-

what smaller than supraocular; supraocular wide, subtri-

angular, wider posteriorly; parietals smaller than frontal

June 201 5 I Volume 9 I Number 1 I e98

Amphib. Reptile Conserv.

Amarasinghe et al.

Fig. 7. Scalation of Cylindrophis mirzae sp. nov. holotype, MNHN-RA 3279 (A) head in dorsal

view, (B) head in ventral view, (C) tail in ventral view, (D) head in lateral view, (E) tail in lateral

view, (F) body in dorsal view, (G) body in ventral view.

which are in large median oblique contact oriented from

right to left antero-posteriorly, subtriangular, their rear

border bluntly pointed, bordered by supraoculars, frontal

shield, upper posterior temporal shields, occipital shield,

and two dorso-nuchal shields posteriorly on each side,

the occipital shield of the same size as other dorso-nuchal

scales; left parietal in larger contact than the right (just a

point) with the frontal; loreal and preocular absent; eye

small (diameter 1.8 mm), pupil rounded; eye in broad

contact with supraocular dorsally, prefrontal and third

supralabial antero-ventrally, fourth supralabial ventrally,

and postocular posteriorly; a single postocular, quadran-

gular, posteriorly roundish and wider, in broad contact

with supraocular, anterior temporal, and narrow contact

with fourth supralabial; temporals 1+2, triangular; ante-

rior temporal larger than posteriors; anterior temporal in

contact with supraocular and posterior temporal dorsally,

4 th and 5 th supralabials ventrally, anterior temporal does

not meet parietal on both sides; upper posterior temporal

slightly larger than lower posterior temporal.

Five supralabials, 3 rd -5 th larger in size; first supral-

abial in contact with rostral anteriorly and nasal dorsally;

second supralabial in contact with nasal and prefrontal

dorsally; third supralabial in contact with prefrontal and

eye dorsally; fourth supralabial in contact with the eye,

postocular, and anterior temporal dorsally; fifth supral-

abial in contact with anterior and posterior temporals

dorsally and body scales posteriorly.

Mental small, triangular; first infralabial pair larger

than mental plate and in broad contact with each other;

1 st infralabials in contact with anterior chin shield poste-

riorly; five infralabials in total, l st -3 rd in contact with first

chin shield, 4 th and 5 th in contact with gular scales and

not touching the chin shields; anterior chin shields larger

than posterior ones; a mental groove continues from the

posterior tip of the mental until the posterior chin shields.

Body slender; transverse body scale rows 21-21-17,

all smooth, subcycloid, and weakly imbricate; vertebral

and midventral scales undifferentiated from adjacent

scales; 188 ventrals; cloacal plate divided, precloacal un-

divided and triangular; tail extremely short, relative TL

(TL/total length) 2.5%, with a conical robust and thick

tip, and six paired subcaudals.

Coloration: The holotype has a dark brown back with

wide and interrupted white bands along dorsal surface

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e98

Two new species of the genus Cylindrophis from Southeast Asia

from behind nape to tail, each band covering about two

scales; head entirely dark, an incomplete, wide band en-

circling the nape; the venter is dark brown with regular,

cream colored bars, divided at midline. See Fig. 4 for

details of coloration in preservative.

Variation of paratypes: SVL range from 146-656 nun,

but MNHN-RA 1885.0102-3, 1974.1253 are juveniles;

body scale rows at one scale prior to precloacal ranges

from 16-18; ventrals 182-196; subcaudals 4-6; all the

subcaudals entire except MNHN-RA 1885.0100 (2 nd di-

vided), MNHN-RA 1885.0103 (3 rd divided); relative TL

2. 0-3. 3%.

Etymology: The species epithet is an eponym latinized

as a noun in the genitive singular, honoring Dr. Jodi

Rowley for her generous friendship, and remarkable

contributions and expeditions assessing amphibian de-

cline due to various diseases, conservation status, and in

documenting amphibian biodiversity. Jodi Rowley is an

Australian herpetologist. She has conducted amphibian

research in Southeast Asia, mainly in Vietnam. Currently

she is a co-ordinator of Australian Museum Research

Institute, a member of the IUCN Amphibian Red List

Authority and the co-chair for Mainland Southeast Asia

of the IUCN Species Survival Commission Amphibian

Specialist Group.

Distribution: The new species is only reported from

Vietnam (Fig. 8). The specimens from Cambodia and

Thailand are much closely related to this new species,

however for the moment we exclude these specimens as

it seems now, after having examined these specimens,

that there may be many more species in existence in

Cambodia and Thailand.

Cylindrophis mirzae sp. nov. Amarasinghe, In-

eich, Campbell & Hallermann

(Figs. 6, 7, 8; Table 3)

urn:lsid:zoobank.org:act:D0BBDECC-22AE-4D9A-AEF4-2BAlF9C115A0

Proposed standard English name: Mirza’s Pipe-Snake

Proposed standard Indonesian name: Ular Pipa Mirza

Holotype: MNHN-RA 3279, (SVL 419 mm), collect-

ed at Singapore, by Joseph Fortune Theodore Eydoux

(1802-1841), certainly during the expedition on the ves-

sel La Favorite (1829-1832).

Paratypes (3): BMNH 1847.2.9.23, (SVL 693 mm), col-

lected from Singapore, by A.F. Gardiner, collection date

unknown; BMNH 1938.9.8.1, (SVL 580 mm), collected

from Singapore, by Dr. A.G.H. Smart (Assistant Medical

Advisor, Colonial Office S.W.I.), presented by Dr. H.B.

Newham (London School of Hygiene and Tropical Med-

icine), collection date unknown; BMNH 1880.9.10.23,

(SVL 298 mm), collected from Singapore, collector and

the date unknown, presented by Dr. Dennis.

INDIA

LAOS

C. h nr mantis

Myanmar (Burma)

THAILAND

U, PHILIPPINES

CAMBODIA

C. maculattis

Sri Lanka

s Malays) a

C. lineattis

Sarawak .

C. metanotus

Sulawesi

C. rmrzae sp. no;

-^Singapore /

C. engkariansis

Sarawak I

BORNEO

SUMATRA } 1

C. aruensis

Da mar Is

C. I&olapis^ c. btmlongarl

Jampea Is. V Wetar Is. A

C. aruensis

Aru Is. ?

C. ruttus

Java

C. ymmdvnm

Yamtiena is.

C. cpisfftorftoch*

Lsw*r Sunda v

Fig. 8. Current distribution pattern of the genus Cylindrophis.

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e98

Amarasinghe et al.

Diagnosis: Cylindrophis mirzae sp. nov. is distinguished

trom all congeners by having the following characters:

21 midbody scale rows (vs. 17 in C. engkariensis ; 19 in

C. boulengeri, C. burmcmus, C. melanotus, C. ruffiis; 23

in C. aruensis, C. opisthorhodus), narrow and completed

lighter rings encircling the dark body at anterior and pos-

terior parts of the body (vs. no bands on the paler back in

C. isolepis and C. yamdena; lateral and middorsal stripes

along the body in C. lineatus ; wide and interrupted bands

on the back in C. jodiae sp. nov.; two series of large red-

dish-brown spots along the back, which are enclosed by

a black network in C. maculatus).

Description of holotype: An adult, SVL 419 mm, tail

length 10.0 mm; body elongate (largest body diameter at

midbody is 14.6 mm), rounded in cross section; head not

distinct from neck, broadened and dorsoventrally flat-

tened in the orbital and sagittal regions; snout blunt in

dorsal and lateral view.

Rostral shield small, slightly visible from above with

a conical apex; a single nasal, widely in contact behind

the rostral, no intemasals; nasals in contact with rostral

anteriorly, with prefrontal posteriorly, and the first and

second supralabials ventrally; nostrils large; canthus ros-

tralis weakly defined; prefrontals larger than the frontal,

and quadrangular; frontal large (length 2.7 mm and width

3.1 mm), triangular, and with the same length as width,

equal or somewhat larger than supraocular; supraocular

wide (length 2.6 mm and width 2.3 mm), triangular, pos-

teriorly wider; parietals equal in size to frontal, subtrian-

gular, their rear border rounded, bordered by supraocular,

frontal shield, upper posterior temporal shield, occipital

shield, and two dorso-nuchal shields posteriorly on each

side, the occipital shield is of same size as other dorso-

nuchal scales; loreal absent; no preocular; eye small

(diameter 1 .0 mm), pupil rounded; eye in broad contact

with supraocular dorsally, prefrontal and third supralabi-

al anteriorly, fourth supralabial ventrally, and postocular

posteriorly; a single postocular, trapezoidal, posteriorly

wider, in broad contact with supraocular, anterior tem-

poral, and wide contact with fourth supralabial ventrally;

temporals 1+2, subtriangular; anterior temporal much

larger than posteriors; anterior temporal in contact with

supraocular and upper posterior temporal dorsally, lower

posterior temporal posteriorly, 4 th and 5 th supralabials

ventrally; anterior temporal well separated from the pari-

etal by the supraoculars and the upper posterior temporal.

Six supralabials, 3 rd and 4 th larger in size and touch-

ing the eye; first supralabial in contact with rostral ante-

riorly and nasal dorsally; second supralabial in contact

with nasal and prefrontal dorsally, third supralabial in

contact with prefrontal and eye postero-dorsally, fourth

supralabial in contact with the eye, postocular, and an-

terior temporal dorsally; fifth supralabial in contact with

anterior and lower posterior temporal dorsally; sixth su-

pralabial in contact with posterior temporals dorsally and

body scales posteriorly.

Mental large, triangular; first infralabial pair slightly

smaller than mental plate and in narrow contact with each

other, and with anterior chin shield posteriorly; six infral-

abials in total, l st -3 rd in contact with first chin shield, 4 th -

6 th in contact with gular scales but not touching the chin

shields; anterior chin shields larger than posterior ones;

a mental groove continues from the posterior tip of the

mental until the posterior chin shields.

Body slender; transverse body scale rows 19-21-18,

all smooth, subcycloid, and weakly imbricate; vertebrals

and midventrals undifferentiated from adjacent scales;

213 ventrals; cloacal plate divided, precloacal undivided

and triangular; tail extremely short, relative TL (TL/total

length) 2.3%, with a conical robust and thick tip; five

subcaudals, the first three entire, the following divided

and the last one entire again.

Coloration: The holotype has a brown back with narrow

and completed lighter rings encircling the body along

dorsal surface from behind nape to tail, each band cover-

ing about one scale; head lighter, an incomplete, narrow

ring encircling the nape; the venter is dark brown with

regular, cream colored stripes, some divided at midline.

See Fig. 6 for details of coloration in preservative.

Variation of paratypes: SVL range from 298-693 mm;

ventrals 196-217; six subcaudals in all paratypes; rela-

tive TL 2. 0-3.3%.

Etymology: The species epithet is an eponym latinized

as a noun in the genitive singular, honouring Dr. Mirza

Kusrini for her generous friendship and support, for her

dedication and important contributions to herpetological

conservation and ecology in Indonesia. Mirza Kusrini is

an Indonesian herpetologist and currently she is a lec-

turer at Bogor Agricultural University, Indonesia and a

steering committee member of IUCN Species Survival

Commission Amphibian Specialist Group.

Distribution: The new species is evidently recorded

from Singapore (Fig. 8).

Discussion

Although, it has been confirmed that the types of Anguis

ruffa and Cylindrophis resplendens are lost, it is now

clear that the type locality of Cylindrophis rujfns is Java

(fide Schlegel 1844). The International Code of Zoo-

logical Nomenclature (ICZN) supports the designation

of a neotype in order to stabilize the taxonomic status.

However, we have decided not to undertake such action

due to the following reasons: (1) our available samples

from Java were too small (n = 14), (2) we have not yet

compared the C. cf. ruffiis populations (which also have

19 midbody scale rows) from other Sundaic Islands and

Peninsular Malaysia (including specimens mentioned by

June 2015 I Volume 9 I Number 1 I e98

Amphib. Reptile Conserv.

Two new species of the genus Cylindrophis from Southeast Asia

Fig. 9. Coloration of Cylindrophis aruensis syntype BMNH

1946.1.16.72 (A) head in dorsal view, (B) head in ventral view,

body in ventral view, and (F) tail in ventral view.

Fig. 11. Coloration of Cylindrophis engkariensis holotype ZRC

8821 (A) head in dorsal view, (B) head in ventral view, (C)

head in lateral view, (D) midbody in dorsal view, (E) midbody

in ventral view, and (F) tail in ventral view.

Stuebing 1994), and (3) we have not found a preserved

adult specimen from Jakarta with associated DNA sam-

ples to support its designation as a neotype. We believe it

would be better to designate a specimen of which DNA

samples are available to solve the taxonomic issues men-

tioned in number (2) above.

Based on morphological and meristic characters, par-

ticularly the number of dorsal scale rows at midbody,

Fig. 10. Coloration of Cylindrophis boulengeri MZB 5284 (A)

head in dorsal view, (B) head in ventral view, (C) head in lateral

view, (D) midbody in dorsal view, (E) midbody in ventral view,

and (F) tail in ventral view.

Fig. 12. Coloration of Cylindrophis isolepis MZB 1926 (A)

head in dorsal view, (B) head in ventral view, (C) head in lateral

view, (D) midbody in dorsal view, (E) midbody in ventral view,

and (F) tail in ventral view.

number of ventrals, and coloration, we include C. ruf-

fus and C. burmanus in the morphological group which

has 19 middorsal scale rows. However Taylor’s (1965)

specimen of “ Cylindrophis rujfus rujfus ” from Thailand

and the populations from Thailand and Cambodia might

either represent an undescribed species or represent C.

jodiae sp. nov. which is distributed in Vietnam. How-

ever, we defer from attempting to answer such questions

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e98

Amarasinghe et al.

Fig. 13. Coloration of Cylindrophis lineatus holotype BMNH

1946.1.16.5 (A) head in dorsal view, (B) head in ventral view,

body in ventral view, and (F) tail in ventral view.

Fig. 15. Coloration of Cylindrophis melanotus MZB 2999 (A)

head in dorsal view, (B) head in ventral view, (C) head in lateral

view, (D) midbody in dorsal view, (E) midbody in ventral view,

and (F) tail in ventral view.

because we believe such questions should be addressed

with the support of molecular evidence and with the

comparison involving large samples from each of the

representative countries. We did not compare C. mirzae

sp. nov. from Singapore with the populations in Sumatra,

Peninsular Malaysia, and Borneo because the available

samples from those locations were too small, thus we

have voluntarily excluded those areas from our study. It

Fig. 14. Coloration of Cylindrophis maculatus BMNH

1962.861 (A) head in dorsal view, (B) head in ventral view, (C)

head in lateral view, (D) midbody in dorsal view, (E) midbody

in ventral view, and (F) tail in ventral view.

Fig. 16. Coloration of Cylindrophis opisthorhodus MZB 1515

(A) head in dorsal view, (B) head in ventral view, (C) head in

lateral view, (D) midbody in dorsal view, (E) midbody in ven-

tral view, and (F) tail in ventral view.

is probable that C. mirzae sp. nov. may be distributed

in some parts of Sumatra (e.g., C. cf. mirzae specimen

listed in Appendix 1 below).

It seems also that Cylindrophis melanotus might be

a species complex or at least consisting of two cryptic

species (note the wide range of ventrals: 233-275). Al-

though, the taxonomy of the genus Cylindrophis should

be examined critically with larger samples and with the

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e98

Two new species of the genus Cylindrophis from Southeast Asia

Fig. 17. Coloration of Cylindrophis yamdena holotype WAM

R1 12252 (A) head in dorsal view, (B) head in ventral view, (C)

head in lateral view, (D) midbody in dorsal view, (E) midbody

in ventral view, and (F) tail in ventral view.

support of molecular analyses (especially for the species

which have 19 and 21 midbody scale rows), we have

described the above two new species due to their clear

morphological differences and because of their biogeo-

graphically isolation from all other known taxa.

Acknowledgments. — We thank the Ministry of Re-

search and Technology of the Republic of Indonesia

(RISTEK), S. Wahyono and L. Shalahuddin for coor-

dinating and granting research permissions to AATA;

the staff members of LIPI-MZB including A. Hamidy,

Syaripudin, and W. Trilaksana for facilitating in-house

study of specimens; Robert Stuebing and Kelvin Lim

(Lee Kong Chian Natural History Museum) for kindly

sending the photos of Cylindrophis engkariensis type;

Ruchira Somaweera and Paul Doughty (Western Austra-

lian Museum) for kindly examining and sending photos

of Cylindrophis yamdena ; and Gemot Vogel for kind

support, valuable comments, and data issued from his

specimen examination. We wish to thank M. Hoogmoed

and E. Dondorp (RMNH, Leiden) for data about the col-

lections under their care. We also thank N.K. Amaras-

inghe and the staff of RCCC-UI for their kind support,

and Howard O. Clark, Jr. for excellent graphic design of

the manuscript. Finally, we thank Van Wallach, Olivier

Pauwels, and Gernot Vogel for reviewing the manuscript

and their valuable comments.

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Stuttgart, and Tubingen, Germany. 29 p.

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mit vorangehender Classification der Sdugetiere und

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Wallach V, Williams KL, Boundy J. 2014. Snakes of the

World: A Catalogue of Living and Extinct Species.

CRC Press, Taylor and Francis Group, Florida, USA.

1,237 p.

Appendix 1

Comparative materials examined

Cylindrophis aruensis Boulenger, 1920 (Fig. 9) -Aru Island, Indonesia: BMNH 1946.1.16.72-73 (syntypes); Dainmer

Island, Indonesia: MZB 305.

Cylindrophis boulengeri Roux, 1911 (Fig. 10) - Ilwaki, Wetar Island, Barat Daya, Maluku, Indonesia: SMF 16996

(holotype), MZB 5243, 5284; Madura Island, East Java, Indonesia (doubtful location): MZB 314.

Cylindrophis engkariensis Stuebing, 1994 (Fig. 11) -Nanga Segerak, Sarawak, Malaysia: ZRC 8821 (holotype).

Cylindrophis isolepis Boulenger, 1896 (Fig. 12) - Jampea Island, Selayar, South Sulawesi, Indonesia: BMNH

1946.1.1.47 (holotype); MZB 299A-B, 1926, 3149, 3365-66.

Amphib. Reptile Conserv. 49 June 2015 | Volume 9 | Number 1 | e98

Two new species of the genus Cylindrophis from Southeast Asia

Appendix 1 (continued)

Cylindrophis cf. jodiae - Trapeang-Chan, Cambodia: MNHN-RA 1970.0411-13; Snoc Trou, Cambodia: MNHN-

RA 1963.0713; Trabeang Thum lake, Choam Khsant, Cambodia: MNHN-RA 2010.0909; Ban Chao Samran, Muang

District, Thailand: MNHN-RA 1998.0576; Ban Pong, Thailand: MNHN-RA 1999.7634; Ban Bang Ba, Muang Dis-

trict, Phang Nga Province, Thailand: MNHN-RA 1997.6582; Bangkok, Thailand: MNHN-RA 3281; Thailand: BMNH

1865.4.28.17, 1897.10.8.18, 1947.1.1.8, 1969.324, 1969.819, 1969.1693, 1987.1723-24.

Cylindrophis lineatus Blanford, 1881 (Fig. 13) - Singapore: BMNH 1946.1.16.5 (holotype); Borneo: BMNH

1901.5.17.1.

Cylindrophis maculatus Linnaeus, 1758 (Fig. 14) - Sri Lanka: BMNH 1962.861, 1892.11.3.3, 1969.2755, 1968.77,

1905.3.25.76-81, 1894.9.11.5-7, 1845.8.7.5, 1897.10.20.18, 1931.5.13.1-5, 1915.5.3.1, 1930.5.8.48, 1930.5.8.51,

1930.5.8.50, 1930.5.8.49, 1930.5.8.52, 1962.254, 1964.1632-1633, 1964.1687; MNHN-RA 3282-83.

Cylindrophis melanotus Wagler, 1830 (Fig. 15) - Dumoga West, North Sulawesi, Indonesia: MZB 3246; Manado,

North Sulawesi, Indonesia: MNHN-RA 5779, 1999.8281; Rantepao, North Toraja, South Sulawesi, Indonesia: MZB

3826; Majene, West Sulawesi, Indonesia: MZB 310; Lindu Lake, Tornado, Central Sulawesi, Indonesia: MZB 1553,

3621; Butung Island, South-East Sulawesi, Indonesia: MZB 2834, 2999; Tinanggea, South Konawe, South East Su-

lawesi, Indonesia: MZB 4567; Tinukari, Wawo, North Kolaka, South East Sulawesi, Indonesia: MZB 4568; Halma-

heira (=Halmahera), Indonesia: ZMB 34313 (holotype of Cylindrophis heinrichi ); Sulawesi, Indonesia: MNHN-RA

3278, 7180, 7180A.

Cylindrophis cf. mirzae - Sumatra: MNHN-RA 1884.0115.

Cylindrophis opisthorhodus Boulenger, 1897 (Fig. 16) - Lombok, Indonesia: BMNH 1946.1.16.148-149 (syntypes);

Ruteng, Watu, Manggarai, East Nusa Tenggara, Indonesia: MZB 1286; Flores, East Nusa Tenggara, Indonesia: MZB

1515; Ndao Nuse, West Rote, Rote Ndao, East Nusa Tenggara, Indonesia: MZB 1532.

Cylindrophis yamdena Smith & Sidik, 1998 (Fig. 17) - Yamdena Island, Indonesia: WAM R1 12252 (holotype),

109947, 109971-72, 109980.

A. A. Thasun Amarasinghe is an Indonesian resident and herpetologist with a special interest in the patterns

and processes of speciation. He has been carrying out reptile taxonomy work in South and Southeast Asia

since 2005, mainly focussing on the following geographic regions: Sri Lanka, India, Andaman/Nicobar, and

Indonesia. He is the editor-in-chief of TAPROBANICA the Journal of Asian Biodiversity (ISSN: 1800-427X)

which is now regarded as a leading journal for biodiversity and conservation in the tropical Asian region. As a

conservationist, he has a strong commitment to furthering the IUCN’s vision and mission. He is a commission

member of the following IUCN committees: CEM South East Asia including thematic groups and the IUCN

Species Survival Commission (Amphibian Specialist Group and Crocodile Specialist Group). He is currently

a research scientist at the Research Center for Climate Change, University of Indonesia, Depok, Indonesia.

R Patrick D. Campbell is a British citizen and Senior Curator of Reptiles, managing over an estimated 174,000

herpetology specimens in the Natural History Museum (London) at which he has been employed for almost 30

years. He is interested in collection management, reptile taxonomy, and the processes of speciation. He has car-

ried out herpetological fieldwork recently in Kenya and the French Guiana and is also interested in the reptilian

fauna of South and Southeast Asia, mainly focussing on the countries of Sri Lanka, India, Andaman/Nicobar,

and Indonesia. As a BSAC (British Sub Aqua Club) advanced trained professional diver and dive instructor, he

has dived on various expeditions including the RAF lead Benthic Orchid III examining the effect of the 2004

Tsunami in the Similan Islands Thailand, conducting underwater surveys for the burrowing starfish Astropecten

in Spain, conducting marine biological surveys in the Cliffe Lagoons/Portland Harbour UK sifting and identify-

ing macro invertebrate samples, and investigating shallow subtitle hydrothermal vents off the Greek island of

Milos. He has lead various collection improvement projects at the museum including the recent CSIP (Collec-

tion Storage Improvement Project) Wildebeest project involving over 20,000 specimens.

Amphib. Reptile Conserv. 50 June 2015 | Volume 9 | Number 1 | e98

Amarasinghe et al.

Jakob Hallermann studied biology at the Universities of Tubingen and Hamburg, and then completed his

Ph.D. in 1994 at the University of Tubingen focussing on the morphology of the Iguanian nose including a

phylogenetic analysis. After two years as a volunteer at the Museum of Natural History Stuttgart, he became

the curator of the herpetological collection of the Zoological Museum Hamburg (now Centre of Natural His-

tory, Cenak). His scientific interests are systematics, biogeography, speciation, and the comparative anatomy of

herpetofauna. Since 1997 his research focus has shifted to the South and Southeast Asian herpetofauna. He has

described many new species of the agamid lizard genera Calotes, Pseudocalotes, and Bronchocela. Currently,

he is conducting research on the phylogeny of the African house snake genus Boaedon.

H Irvan Sidik was born in Bandung, West Java Province, Indonesia. Irvan obtained an M.Sc. in the field of phy-

logenetics at the Institute Technology of Bandung. Since 1992 Irvan has been working as a staff researcher in

the laboratory of herpetology at the Museum Zoologicum Bogoriense, Indonesian Institute of Sciences (LIPI)

in the Cibinong Science Center. Beginning as an auxiliary field survey researcher, and then as a local CITES of-

ficer, Irvan became interested and developed a great interest in the snakes of the region of Sundaland. Irvan has

continued with more scholarly work on the mountainous areas of the western part of Indonesia. Irvan’s research

is based on museum collections of specimens and field research in Indonesia’s regions mentioned above. Irvan

has been involved in several international research collaborations, and is currently working with the University

of Texas at Arlington, USA on research of amphibians and reptiles in the mountains of Java and Sumatra. Irvan

has published on the herpetofauna of Kalimantan and his first book was about snakes that are traded in Indone-

sia (CITES appendices I, II, and III) written in Indonesian. Currently, Irvan is studying the phylogeography of

the reed snake genera Calamaria for his Ph.D. at the University of Brawijaya, Malang.

Jatna Supriatna is the chairman of Research Centre for Climate Change, University of Indonesia. Jatna is

, v one of the leading research biologists, primatologists, and conservationists in Indonesia. He serves as a senior

K lecturer of the Biology Department, and a coordinator for the Graduate Program on Conservation Biology of

the University of Indonesia. He has practiced as an editor for many international journals. In 2007 he was as-

K tv signed as the chairman of the IUCN/SSC PSG South East Asia. He served on several assignments including

those for the government of Indonesia : the National Research Council, the Steering Committee on Biodiversity

Action Plan (Ministry of Planning), and the Biodiversity Taskforce (Ministry of Research & Technology). For

his dedication to conservation, he received the Golden Ark Award (1999) from his royal highness Prince of

Berhard of the Netherlands. He also has received “the Habibie Award” (1999) from the Indonesian president.

He has published ten books, mostly on Indonesia’s biodiversity conservation, as well as over 100 research ar-

ticles in journals such as Science , Nature, Conservation Biology, Primates, Evolution, Primate Conservation,

Herpetologica, and others.

Ivan Ineich is a French herpetologist especially interested in systematics, biogeography, and processes of spe-

ciation. He has mostly been carrying out reptile taxonomy work in Africa and on the Pacific Islands, but also

South and Southeast Asia. He was curator of lizards and snakes at Paris Natural History Museum (NNHN) from

1988 to 2014. He was editor-in-chief of the French Bulletin cle la Societe Herpetologique cle France for many

years. He is currently researcher at the Institute for Systeamtic, Evolution and Biodiversity at the Museum

national d’Histoire naturelle of Paris, a position that he occupied since 1988.

In accordance with the International Code of Zoological Nomenclature new rules and regulations (ICZN 2012), we have deposited this paper in publicly acces-

sible institutional libraries. The new species described herein has been registered in ZooBank (Polaszek 2005a, b), the official online registration system for the

ICZN. The ZooBank publication LSID (Life Science Identifier) for the new species described here can be viewed through any standard web browser by append-

ing the LSID to the prefix “http://zoobank.org/.” The LSID for this publication is: um:lsid:zoobank.org:pub:A4C569A0-36DB-4E6D-B3CE-3533 1FE535F2.

Separate print-only edition of paper(s) (reprint) are available upon request as a print-on-demand service. Please inquire by sending a request to: Amphibian &

Reptile Conservation, amphibian-reptile-conservation.org, arc.publisher@gmail.com.

Amphibian & Reptile Conservation is a Content Partner with the Encyclopedia of Life (EOL), http://Avww.eol.org/ and submits information about new species

to the EOL freely.

Digital archiving of this paper are found at the following institutions: ZenScientist, http://www.zenscientist.com/index.php/filedrawer; Ernst Mayr Library, Mu-

seum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (USA), http://library.mcz.harvard.edu/emst_mayr/Ejoumals/ARCns.

The most complete journal archiving and journal infomiation is found at the official ARC journal website, amphibian-reptile-conservation.org. In addition,

complete journal paper archiving is found at: ZenScientist, http://www.zenscientist.com/index.php/filedrawer.

Citations

ICZN. 2012. Amendment of Articles 8,9,10,21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods of publication.

Zootaxa 3450: 1—7.

Polaszek A et al. 2005a. Commentary: A universal register for animal names. Nature 437: 477.

Polaszek A et al. 2005b. ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological Nomenclature 62(4):

210 - 220 .

Amphib. Reptile Conserv.

June 2015 I Volume 9 I Number 1 I e98

CONTENTS

Special Section

Luis Mamani, Noemi Goicoechea, and Juan C. Chaparro — A new species of Andean lizard Proctoporus (Squa-

mata: Gymnophthalmidae) from montane forest of the Historic Sanctuary of Machu Picchu, Peru 1

Daniel Rodriguez — Noblella lynchi Duellman 1991 (Anura: Craugastoridae): Geographic range extension, Peru. ... 12

German Chavez, Roy Santa-Cruz, Daniel Rodriguez, Edgar Lehr — Two new species of frogs of the genus

Phrynopus (Anura: Terrarana: Craugastoridae) from the Peruvian Andes 15

Lourdes Y. Echevarria and Pablo J. Venegas — A new elusive species of Petracola (Squamata: Gymnoph-

thalmidae) from the Utcubamba basin in the Andes of northern Peru 26

Lourdes Y. Echevarria, Andy C. Barboza, and Pablo J. Venegas — A new species of montane gymnophthalmid

lizard, genus Cercosaura (Squamata: Gymnophthalmidae), from the Amazon slope of northern Peru 34

General Section

Steven Poe, Simon Scarpetta, and Eric W. Schaad — A new species of Anolis (Squamata: Iguanidae) from

Panama 1

Valter Weijola and Samuel S. Sweet — A single species of mangrove monitor ( Varanus ) occupies Ambon,

Seram, Buru and Saparua, Moluccas, Indonesia 14

Kara S. Jones and Todd A. Tupper — Fowler’s Toad (Anaxyrus fowleri) occupancy in the southern mid-Atlantic,

USA 24

A. A. Thasun Amarasinghe, Patrick D. Campbell, Jakob Hallermann, Irvan Sidik, Jatna Supriat-

na, Ivan Ineich — Two new species of the genus Cylindrophis Wagler, 1828 (Squamata: Cylindrophi-

idae) from Southeast Asia 34

Table of Contents Back cover

Cover: An adult individual of Canelos Treefrog ( Ecnomiohyla tuberculosa ) collected in a tree hole during a herpe-

tological survey, carried out by the Field Museum, in the Putumayo basin at the Peruvian Amazonia. The Canelos

Treefrog is one of the most enigmatic Neotropical frogs and its occurence has been documented at scattered locali-

ties along the upper Amazon basin of Brazil, Colombia, Ecuador, and Peru. Although the phylogenetic position and

natural history of this species remains a mystery, it is currently under inventigation by a team of herpetologists from

Ecuador and Peru led by Dr. Santiago Ron. Photograph: Pablo J. Venegas.

Instructions for Authors: Located at the Amphibian & Reptile Conservation website:

http://amphibian-reptile-conservation.org/submissions.html

VOLUME 9

2015

NUMBER 1