European Journal of Taxonomy 3: 1-18

http://dx.doi.Org/10.5852/ejt.2011.3

www. europeanj ournaloftaxonomy. eu

2011 • Philippe J.R. Kok & Gilson A. Rivas

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

A new species of Anadia (Reptilia, Squamata) from the Venezuelan

'Lost World', northern South America

Kok P. J.R. & Rivas G.A. 2011. A new species of Anadia (Reptilia, Squamata) from the Venezuelan 'Lost World',

northern South America. European Journal of Taxonomy 3: 1-18. http://dx.doi.org/10.5852/ejt.201L3

Philippe J.R. KOK 1 2 4 & Gilson A. RIVAS 3

'Biology Department, Unit of Ecology and Systematics, Vrije Universiteit Brussel, 2 Pleinlaan,

B-1050 Brussels, Belgium

department of Vertebrates, Royal Belgian Institute of Natural Sciences, 29 rue Vautier, B-1000

Brussels, Belgium

3 Museo de Biologia, Lacultad Experimental de Ciencias, La Universidad del Zulia, Apartado Postal

526, Maracaibo 4011, Maracaibo, Venezuela

4 Email: [email protected], [email protected]

Abstract. A new gymnophthalmid lizard of the genus Anadia Gray, 1845 is described from the summit

of Abakapa-tepui, Bolivar State, Venezuela, between 2200-2242 m elevation. The new species, Anadia

mcdiarmidi sp. nov., is endemic to the Chimanta Massif and seemingly also occurs on Amuri-tepui

and Murei-tepui. The new taxon is ma inl y distinguished from all known congeners by the following

combination of characters: body fairly robust, dorsal scales small and quadrangular, middorsal scales

53-57, suboculars large, subequal in size, with sometimes one scale slightly protruding downward

between 4 th and 5 th supralabial, nasal entire, without sub-nostril groove, body uniform beige or greyish

to bluish brown in life, devoid of any conspicuous pattern in males, venter immaculate golden grey in

life, femoral pores 9-10 on each side in males, preanal pores absent, hemipenis globose, weakly bilobed,

bordered by numerous flounces (>20) bearing comblike rows of minute weakly mineralized spinules.

The presence of a species of Anadia , a primarily Andean genus, on the top of tepuis is of considerable

interest to the understanding of the Pantepui biogeography.

Key words. Gymnophthalmidae, Lizard, Hemipenes, Chimanta Massif, Pantepui.

Introduction

Gymnophthalmid lizards of the genus Anadia Gray, 1845 may be gracile, narrow-headed and semi-

arboreal or mostly terrestrial, more robust with a broader head (Oftedal 1974). This genus is primarily

Andean, with 17 currently recognized species distributed in the Ecuadorian Andes (A. peter si Oftedal,

1974), Colombian Andes [A. altaserrania Harris & Ayala, 1987;/l. bo go tens is (Peters, 1863 );A. pulchella

Ruthven, 1926], Venezuelan Andes [A. bitaeniata Boulenger, 1903; A. brevifrontalis (Boulenger, 1903);

A. hobarti La Marca & Garcia-Perez, 1990; A. pamplonensis Dumi, 1944], the central coastal range in

Venezuela [A. marmorata (Gray, 1846); A. steyeri Nieden, 1914], the eastern coastal range in Venezuela

(A. blakei Schmidt, 1932; A. pariaensis Rivas, La Marca & Oliveros, 1999), and at lower elevations

European Journal of Taxonomy 3: 1-18 (2011)

from Costa Rica to southwestern Ecuador and in southeastern Venezuela [A. ocellata Gray, 1845; A.

vittata Boulenger, 1913; A. bumanguesa Rueda-Almonacid & Caicedo, 2004; A. rhombifera (Gunther,

1859); A. escalerae Myers, Rivas & Jadin, 2009].

Oftedal (1974) provided a comprehensive review of the genus, which is still authoritative and included

11 species. Since Oftedal's work five additional species have been described in the genus: two from

Colombia (A. altaserrania and A. bumanguesa ), one from the Venezuelan Andes (A. hobarti ), one from

the peninsula of Paria in Venezuela (A. pariaensis), and one from the La Escalera region, Bolivar State in

southeastern Venezuela (A. escalerae ), which to date constitutes the easternmost locality for the genus.

Anadia pamplonensis was removed from the synonymy of A. bitaeniata by Harris & Ayala (1987). The

monophyly of the genus is still in question (Myers et al. 2009) and phylogenetic relationships are mostly

unknown.

In 1978 Roy McDiarmid (Smithsonian Institution) accompanied by the Venezuelan explorer Charles

Brewer-Carias collected an Anadia species on the summit of Murei-tepui in the Chimanta Massif

(Gorzula 1992; McDiannid & Donnelly 2005). Although the species was supposed to be described

by McDiannid and Harris (as stated by Gorzula 1992), Gorzula (1992) reported it from the Chimanta

Massif as Anadia breweri (a nomen nudum according to Gorzula & Senaris 1999; Myers & Donnelly

2008; and Myers et al. 2009) based on McDiannid's specimen and additional material collected by the

'Grupo Cientifico Chimanta'. The species was subsequently reported as ' Anadia species a' by Gorzula

& Senaris (1999), who mentioned it from three tepuis in the Chimanta Massif: Abakapa-tepui, Amuri-

tepui, and Murei-tepui, and as Anadia sp. A' by McDiannid & Donnelly (2005), who also reported it

from the three same mountains. The species is illustrated as Anadia breweri in a recent book by Brewer-

Carias & Audy (2010: unnumbered figure on page 210). Surprisingly, no description ever appeared and

the status of the new species (including its generic allocation) remains uncertain.

A recent expedition to Abakapa-tepui in the Chimanta Massif led in the framework of a study on the

evolution of the Pantepui herpetofauna resulted in the rediscovery of the species, which is described

below on the basis of four specimens.

Material and methods

Specimens were collected by hand and euthanized by intra-peritoneal injection of 2% Xylocaine. Whole

individuals were fixed in 10% formalin and later transferred to 70% ethanol for permanent storage.

Specimens have been deposited in the collections of the Royal Belgian Institute of Natural Sciences

(RBINS), Brussels, Belgium. Coordinates and elevations of localities were acquired using a Garmin

60CSx Global Positioning System unit and referenced to map datum WGS84.

Comparisons of external character states are based both on original descriptions and examination of

museum specimens (see Appendix for material examined). Terminology for morphological characters

and scale counts mostly follow Oftedal (1974) and Myers et al. (2009). All morphometric data were

taken from the preserved specimens to the nearest 0.01 mm and rounded to the nearest 0.1 mm, under

a Leica stereo dissecting microscope using an electronic digital calliper. For ease of comparison, the

description mainly follows the pattern of Myers et al. (2009). The following measurements and scale

counts were taken and abbreviated as follows: (1) snout-vent length, from tip of snout to posterior margin

of vent (SVL); (2) tail length, from posterior margin of vent to tip of tail (TL); (3) head length, from tip

of snout to anterior margin of ear opening (HL); (4) greatest width of head (HW); (5) greatest depth of

head (HD); (6) snout-axilla length (SAL); (7) neck length, from posterior edge of ear opening to forearm

held at right angle to body (NL); (8) axilla-groin distance (= trunk length), from base of forelimb to base

of hind limb (AXG); (9) forelimb length, from axilla to tip of claw of longest digit (FIL); (10) hind limb

KOK P.J.R. & RIVAS G.A., Anew Anadia from the Venezuelan 'Lost World'

length, from groin to tip of claw of longest digit (HIL); (11) number of subdigital lamellae under Finger

IV (SLF); (12) number of subdigital lamellae under Toe IV (SLT); (13) number of middorsal scales,

from postoccipitals to the level of posterior edge of thigh held at right angle to body (MD); (14) number

of ventrals in longitudinal rows, counted at midbody and including small ventrolateral rectangular plates

(VL); (15) number of ventrals in transverse rows, counted between collar and vent [thus including

preanals, which as pointed out by Oftedal (1974) are frequently difficult to distinguish from ventrals in

the genus] (VT); (16) number of scales around midbody, counted at mid-distance between collar and

vent (SAM); (17) number of supralabials (SL); (18) number of infralabials (IL); and (19) number of

femoral pores under each thigh (FP).

Males' genitalia were everted in the field by injection of 10% formalin at the base of the tail just prior

preservation. The left hemipenis of the holotype and both hemipenes of the adult male paratype (IRSNB

2674) were dissected in the laboratory, soaked over night in a solution of 2% potassium hydroxide,

washed with distilled water, and filled with petroleum jelly, mostly following the methods described

by Manzani & Abe (1988) and modified by Pesantes (1994) and Zaher (1999). To highlight the

calcareous spinules the organs were subsequently stained 24 hours in 1% potassium hydroxide saturated

with Alizarin Red [method adapted from Uzzell (1973)], and stored in 70% ethanol. Terminology of

hemipenis description mostly follows Dowling & Savage (1960), Savage (1997), Zaher (1999), and

Myers et al. (2009).

Colour pattern in life was taken from field notes and colour digital photographs. Sex was determined by

the presence/absence of hemipenes (base of tail dissected when hemipenes were not everted in the field).

Taxonomy follows Oftedal (1974), Pellegrino et al. (2001) and Rodrigues et al. (2005). Institutional

acronyms follow Frost (2011).

Results

Anadia mcdiarmidi sp. nov.

Figs 1-7, Table 1

Anadia breweri (nomen nudum ) — Gorzula 1992: 276, figs 168-171

Anadia species a - Gorzula & Senaris 1999: 114-115, fig. 88

Anadia sp. A - McDiarmid & Do nn elly 2005: 514

Anadia breweri (nomen nudum ) - Brewer-Carias & Audy 2010: 210, unnumbered figure

Holotype

IRSNB 2677 (field number PK 3565), 1 adult S collected by Philippe J.R. Kok, 03-05-2011 at llh39,

summit of Abakapa-tepui, Bolivar State, Venezuela, 05°11'09"N 062°17'36"W, 2201 m elevation.

Paratopotypes

(n=3) 1 adult S (IRSNB 2674, field number PK 3578) collected by Philippe J.R. Kok, 06-05-2011 at

9h40, summit of Abakapa-tepui, Bolivar State, Venezuela, 05°10'50"N 062°17'49"W, 2242 m elevation);

1 juvenile S (IRSNB 2675, field number PK 3577) collected by Philippe J.R. Kok, 06-05-2011 at 9hl0,

summit of Abakapa-tepui, Bolivar State, Venezuela, 05°11'00"N 062°17'34"W, 2200 m elevation; and

1 other juvenile <$ (IRSNB 2676, field number PK 3589) collected by Philippe J.R. Kok, 09-05-2011 at

9h00, summit of Abakapa-tepui, Bolivar State, Venezuela, 05°11'07"N 062°17'19"W, 2204 m elevation.

European Journal of Taxonomy 3: 1-18 (2011)

Fig. 1 . Anadia mcdiarmidi sp. nov. A. IRSNB 2677, 8 holotype in life. B. IRSNB 2675, juvenile male

paratype in life. C. Ventral view of the anaesthetized holotype (grid squares = 5 mm). D. Summit of

Abakapa-tepui, looking NW, showing the macrohabitat of the new species (03-05-2011). Photographs

by Philippe J.R. Kok.

KOK P.J.R. & RIVAS G.A., Anew Anadia from the Venezuelan 'Lost World'

Etymology

The specific epithet is a noun in the genitive case, honouring Roy McDiarmid, North Am erican

herpetologist (Smithsonian Institution), for his contribution to the knowledge of the Pantepui

herpetofauna.

Generic allocation

The new species is placed in the genus Anadia because it fits the 11 diagnostic traits for the genus

proposed by Oftedal (1974): dorsal scales smooth, essentially homogeneous; no distinct rows of much

smaller scales between lateral scales; ventral scales smooth, quadrangular; head scales smooth, without

striation; single frontonasal separating nasals; paired prefrontals; several large pigmented palpebrals;

gular scales smooth, essentially homogeneous, with no distinctly enlarged longitudinal rows; 7-10 large

preanal scales [maximum 9 in Oftedal (1974)]; femoral pores conspicuous in adult males; and limbs

well developed, pentadactyl with all toes clawed.

Definition and diagnosis

A fairly robust species currently not assigned to any species group. The new species is characterized

by the following combination of characters: (1) size small, body robust (maximum known SVL

62.2 mm); (2) tail longer than SVL; (3) dorsal scales small, quadrangular; (4) middorsal scales 53-57;

(5) suboculars large, unequal in size, with sometimes one scale slightly protruding downward between

4 th and 5 th supralabial, suboculars separated from palpebrals by 2-3 rows of mostly inconspicuous tiny

scales, a few of them distinctly enlarged with angular downward protrusion between suboculars; (6)

nasal entire, without sub-nostril groove; (7) 3-4 large pigmented palpebrals; (8) body uniform beige

Fig. 2. Anadia mcdiarmidi sp. nov. Ventral, dorsal, and lateral views of the head of the holotype (IRSNB

2677). Scale lines = 5 mm. Photographs by Philippe J.R. Kok.

European Journal of Taxonomy 3: 1-18 (2011)

or greyish to bluish brown in life, devoid of any conspicuous pattern in males; (9) absence of any dark

line on side of head in adult males; (10) venter immaculate golden grey in life; (11) femoral pores 9-10

on each side in males, unknown in females; (12) preanal pores absent; (13) hemipenis globose, weakly

bilobed, bordered by numerous flounces (>20, including about eight medioproximal asulcate flounces)

Fig. 3. Anadia mcdiarmidi sp. nov. Subpelvic region of the preserved $ holotype (IRSNB 2677),

showing preanal scales and femoral pores, and trunk of the preserved holotype in lateral view (head to

left), showing scalation at midbody. Scale lines = 5 mm. Photographs by Philippe J.R. Kok.

KOK P.J.R. & RIVAS G.A., Anew Anadia from the Venezuelan 'Lost World'

Fig. 4. Anadia mcdiarmidi sp. nov. Left hand and right foot of the holotype (IRSNB 2677) in life. Scale

line = 5 mm Photographs by Philippe J.R. Kok.

Fig. 5. Anadia mcdiarmidi sp. nov. The left fully everted—not maximally expanded—hemipenis of the

holotype (IRSNB 2677). Left: sulcate side. Centre: lateral view, turned with sulcate side facing left.

Right: asulcate side. BLS = fleshy bumplike structure; MPAF = medioproximal asulcate flounces; NSAF

= median nude space of the asulcate face; SB = sulcus branches; SF = spinulate flounces; SS = sulcus

spermaticus. Scale line = 2 mm. Although staining of the spinules by Alizarin Red is well visible under

the microscope, the structures are not clearly observable in the figure. Photographs by Philippe J.R. Kok.

European Journal of Taxonomy 3: 1-18 (2011)

bearing comblike rows of minute wealdy mineralized spinules, sulcus spermaticus divided in the crotch

by a small fleshy bumplike structure, sulcus branches barely detectable.

Thenewtaxonis immediately distinguished from Anadiaocellata, A. vittata,A. rhombifera, and A. petersi

[all members of the ocellata group of Oftedal (1974)] andH. bogotensis [single member of the bogotensis

group of Oftedal (1974)] in having quadrangular dorsal scales (subhexagonal in species of the ocellata

group, imbricate in A. bogotensis ), and in lacking a subnostril groove or divided nasal (nasal divided or

subnostril groove present in species of the ocellata group and in A. bogotensis ); from A. altaserrania

[a member of the bitaeniata group of Oftedal (1974) according to Harris & Ayala (1987)] in having

prefrontals (lacking in A. altaserrania ), in having more femoral pores in males (9-10 in A. mcdiarmidi

vs. 3-4 in A. altaserrania ), and in having a higher number of middorsal scales (53-57 in A. mcdiarmidi

vs. 45-47 in A. altaserrania ); from A. bitaeniata, A. brevifrontalis, A. hobarti, A. pamplonensis, and A.

pulchella [aft members of the bitaeniata group of Oftedal (1974) according to La Marca & Garcia-Perez

(1990)] in having a higher number of middorsal scales (53-57 in A. mcdiarmidi vs. maximum 50 in

species of the bitaeniata group); from A. blakei and A. marmorata [members of the marmorata group

of Oftedal (1974)] in having a higher number of middorsal scales (more than 50 in A. mcdiarmidi vs.

less than 35 in A. blakei and A. marmorata)', from A. bumanguesa [a member of the bitaeniata group

of Oftedal (1974) according to Rueda-Almonacid & Caicedo (2004), but a possible synonym of A.

steyeri and thus a member of the steyeri group of Oftedal (1974) according to Rivas et al. (in press)]

in having a lower number of scales around midbody (35-36 in A. mcdiarmidi vs. 41 in A. steyeri), and

in having pigmented palpebrals (unpigmented in A. bumanguesa)', from A. pariaensis (not assigned to

any species group in the original description) in having a much lower number of middorsal scales (54-

57 in A. mcdiarmidi vs. 70-72 in A. pariaensis)', from A. steyeri [single member of the steyeri group of

Oftedal (1974)] in having a lower number of scales around midbody (35-36 in A. mcdiarmidi vs. 40-45

in A. steyeri)', and from A. escalerae (not assigned to any species group in the original description),

the geographically closest relative, in having a robust body (slender in A. escalerae), in having smooth

dorsal scales (weakly keeled in A. escalerae), in lacking dorsolateral stripes (present in A. escalerae),

in having a lower number of middorsal scales (54-57 in A. mcdiarmidi vs. 63 in A. escalerae), and in

having a lower number of scales around midbody (35-36 in A. mcdiarmidi vs. 40 in A. escalerae).

Description of the Holotype

An adult male in good condition (Figs 1A, C, 2, 3, 4, 6), with 62.2 mm SVL and 110.0 mm TL (see

also Table 1). A rather robust Anadia species with moderately long limbs (digits overlap when limbs are

adpressed along body). Snout attenuate, flat in profile. HL 25% of SVL, 1.5 times longer than wide, 1.6

times wider than high; head conspicuously wider than neck. Neck long, 68% of HL, 35% of AXG. SAL

93% of AXG, 46% of SVL. Body wider than deep. Tail dorsoventrally flattened, about 1.8 times longer

than SVL. Limbs pentadactyl with aft digits clawed. Forelimb 27% of SVL, 52% of AXG; hind limb

36% of SVL, 73% of AXG.

Scutellation on the dorsal surface of head is typical for the genus [see fig. 1 in Oftedal (1974)], except

the condition of the parietals and interparietal (see below and Figs 2, 7). The frontal is damaged, the

frontoparietals are slightly damaged. Head scales are smooth with a few minute scattered pits, those

pits are especially concentrated on the upper and posterior margins of supralabials, on the lower and

posterior margins of infralabials, and on the posterior margin of rostral.

Rostral much wider than deep, laterally in contact with nasals and first supralabial, dorsally in contact

with large frontonasal. Frontonasal with nearly straight anterior margin, posteriorly pointed. Paired

prefrontals in contact, medial suture short. Frontal damaged, but obviously wider anteriorly and with

blunt anterior point. Paired frontoparietals slightly damaged, with long medial suture, in contact with

interparietal, parietals, and two posterior supraoculars. Three large supraoculars subequal in size.

KOK P.J.R. & RIVAS G.A., Anew Anadia from the Venezuelan 'Lost World'

Interparietal divided into about two scales unequal in size. Parietals not extending as far as interparietal

posteriorly, divided into two scales on the left side, into four scales on the right side. A series of six small

to large occipitals (postparietals) circling the common posterior margin of parietals and interparietal.

Fig. 6. Anadia mcdiarmidi sp. nov. Intrapopulation variation in dorsal (above) and ventral (below)

colour pattern in preserved specimens. From left to right: IRSNB 2677, IRSNB 2674, IRSNB 2676,

IRSNB 2675. Photographs by Philippe J.R. Kok.

European Journal of Taxonomy 3: 1-18 (2011)

Nasal scale entire, with a small indentation situated anterodorsally to nostril on the left side. Nasal scale

in contact with rostral, posteriorly in contact with prefrontal, and in broad contact with loreal. Loreal

large, pentagonal, in broad contact with first superciliary (= presuperciliary) and frenocular, in point

contact with preocular. One small preocular, three postoculars. First superciliary large, followed by

four shorter superciliaries. No small azygous scale between superciliaries and supraoculars. Frenocular

followed posteriorly by three suboculars. Third subocular slightly extending to lip between supralabials

4-5 on the left side only. Suboculars separated from palpebrals by 2-3 poorly defined rows of mostly

inconspicuous tiny scales, two of them distinctly enlarged with angular downward protrusion between

suboculars. Seven supralabials.

Eight ciliaries along upper eyelid. Lower eyelid scales translucent, slightly pigmented, with four higher-

than-wide palpebrals.

Temporal scales subimbricate, smooth, with flat surfaces, slightly larger above and smaller below. Ear

opening broad, ovoid, slightly inclined posterodorsally, edged with small, slightly pebblelike scales;

tympanum recessed, pigmented.

Fig. 7. Anadia mcdiarmidi sp. nov. Intrapopulation variation in the condition of the parietals and the

interparietal (frontoparietals in grey). Top, from left to right: IRSNB 2677, IRSNB 2676; bottom, from

left to right: IRSNB 2675, IRSNB 2674. Drawings not to scale.

KOK P.J.R. & RIVAS G.A., Anew Anadia from the Venezuelan 'Lost World'

Underside of head with six infralabials on each side. A large mental followed by a large postmental in

contact with first two infralabials. Two pairs of large genials in broad contact medially, in lateral contact

with infralabials 3-4; two pair of postgenials, the first pair large, each scale in contact with last genial

and infralabials 4-5, the second pair much smaller, each scale in contact with first postgenial and in point

contact with fifth infralabial. Gulars arranged in transverse rows, becoming slightly larger posteriad,

culminating in a well-defined collar row of eight subequal scales; no distinctly enlarged longitudinal

rows. Side of neck between ear and collar pebbled with subequal irregularly shaped juxtaposed scales.

Middorsal scales 54. Dorsal scales on neck subimbricate, irregularly shaped, with rounded surfaces.

Dorsal body scales smooth, juxtaposed, mainly quadrangular, a few irregularly shaped, but most are

longer than wide, in transverse rows only. Lateral scales smooth, similar to dorsals, but smaller.

Ventral scales much wider than dorsals, smooth, juxtaposed, quadrangular, slightly longer than wide

except for ca. four median longitudinal rows of square to wider than long scales on belly; in 14

longitudinal rows at midbody and 31 transverse rows between collar and vent.

Eight marginal preanal scales anterior to vent, the middle two narrower than the adjacent lateral ones,

most outer ones very small. Femoral pores well developed, in swollen pore scales mostly in linear

contact; 9 pores on left thigh, 10 on right. Pores do not extend onto preanal area (Fig. 3).

Caudal scales smooth, subimbricate, rectangular, in transverse rows around tail.

Scales on dorsal surfaces of arm large, smooth, imbricate; ventral surfaces of arm with similar, but

smaller and less imbricate scales. Hind limbs with large, smooth subimbricate to imbricate scales on

anterior face of thighs and on lower legs; scales similar but smaller on ventral side of thigh. Dorsal and

posterior sides of thigh with much smaller, irregularly shaped juxtaposed scales.

Moderate-size, imbricate scales atop hands and feet. Supradigital scales single; upper and lower ungual-

sheath scales covering base of claws, leaving tips well exposed. Palms and soles with small slightly

raised juxtaposed scales (Fig. 4). Two enlarged smooth thenar scales at base of pollex. Subdigital

lamellae mostly single. Famellae under first (I) through fifth (V) finger (right/left side): f. 7/6, II: 9/9,

III: 11/11, IV: 13/14, V: 10/9. Famellae under first (I) through fifth (V) toe (right/left side): I: 6/6, II:

10/10, III: 14/12, IV: 17/15, V: 15/6 (large part of Toe V missing on left side).

Tongue lanceolate; proximal two-third unpigmented, distal one-third heavily pigmented, including

fork. Upper surface behind fork covered with imbricate scalelike papillae that extend onto ventrolateral

sides of tongue. Raised medioventral side of tongue with numerous (>10) of thin, oblique, anteriorly

converging and pointed (chevronlike) infralingual plicae; anterior pair of plicae swollen, bluntly pointed

and larger than those following. Very shallow medioventral groove.

Anterior maxillary and dentary teeth conical, unicuspid, with no or very feeble recurvature, becoming

larger posteriorly.

The left hemipenis (Fig. 5) extends about four subcaudal rows when adpressed to the tail, and is about

7.0 mm in length and 4.0 mm across its widest point when fully everted (organ not maximally expanded

because one of the lobes was slightly damaged during the eversion process). Hemipenial body globose,

tapered near the base, weakly and symmetrically bilobed. Sulcus spermaticus broader near crotch,

relatively shallow, curving about halfway around the base of the organ, then running medially towards

the lobes. Sulcus flanked by a narrow nude area on each side, and divided in the crotch by a small fleshy

bumplike structure, from where two narrow, ill-defined and barely detectable sulcus branches seem

to extend laterally into the folded thickened 'ear-like' tissues of each lobe towards the apices of the

European Journal of Taxonomy 3: 1-18 (2011)

Atlantic Ocean

paKi

Venezuela

Suriname

Guiana

Colombia

Murei*tepui

Chi manta

Churi-tepui

mopo-iem

Fig. 8. Map of the Guiana Shield region (A), with enlarged black rectangle corresponding to the

Chimanta Massif (B). Black triangle indicates type locality, black dots indicate other know localities for

the species.

KOK P.J.R. & RIVAS G.A., Anew Anadia from the Venezuelan 'Lost World'

lobes. Lobes are complexly folded, not flattened terminally, and sulcus branches seem to end in small

depressions (from which residual seminal fluid was removed before hemipenis preparation).

Very base of hemipenis nude, pigmented in life. Lateral and asulcate faces of the organ with a series

of roughly equidistant oblique plicae or flounces with weakly calcified spinules. Twenty-three rows of

flounces extending along the organ. Basal flounces shorter, almost in straight transverse lines, the three

first rows almost continuous across the hemipenial base, except for narrow nude areas on each lateral

face and for nude areas edging each side of the sulcus spermaticus; five subsequent rows continuous

across hemipenial body (except for nude areas edging each side of the sulcus spermaticus), with the

seventh and eight rows distinctly pointing medially towards the apex on the asulcate face (those eight

flounces proximal to the median nude space of the asulcate face correspond to the 'medioproximal

asulcate flounces' sensu Myers et al. 2009); fifteen remaining rows chevron-shaped, also circling the

organ, but separated by a broad median nude space on the asulcate face and by the nude areas edging

each side of the sulcus spermaticus. The plicae bear comblike rows of minute weakly mineralized

spinules that project slightly from the edges of the plicae and usually increase in size towards the lateral

faces and the basal part of the organ (staining of the spinules by Alizarin Red is well visible under the

microscope although not observable in Fig. 5).

Colour of the Holotype in life

Dorsum greyish to bluish brown depending on light intensity, devoid of any conspicuous pattern, but

with a few scattered black scales on flanks. Arms and legs, including digits, greyish brown with a few

scattered black scales; rear of thighs with a few ill-defined dark markings. Ventral surfaces of head and

body golden grey, ventral surfaces of legs and tail bluish grey, palms and soles lemon yellow speckled

with brown and yellowish orange scales. Tympanum dark brown, iris pale yellow (Figs 1A, C, 4).

Colour of the Holotype in preservative

After approximately two months in preservative, the dorsal surfaces became dark brown (looking

slightly peppered when specimen is in liquid), the ventral surfaces are metallic blue, except the anterior

surface of head, which is light grey (Fig. 6). Palms and soles are grey. Tympanum is dark brown. It is

worthwhile to note that the ventral skin, and some parts that are light grey (like the anterior underside of

head), turn blue or bluer when pressure is applied to the skin.

Variation in paratypes

The paratypes consist in 1 adult S (SVL 60.4 mm), and 2 juvenile $ (SVL from 38.2 to 42.6 mm).

Table 1 provides measurements of the type series, and Fig. IB shows one of the juvenile paratypes in

life (IRSNB 2675).

One of the main differences among specimens of the type series is the occurrence of a presupraocular

scale in the three paratypes (absent in the holotype); that presupraocular scale is divided on both sides

in IRSNB 2675. IRSNB 2676 has four suboculars on both sides (vs. three in the holotype and other

paratypes). Supralabial counts are consistent, but infralabial counts vary from 5 (n=l) to 7 (n=l, one

side only). Pits on head scales are inconspicuous in the two juveniles (vs. well visible in the adult males).

Suboculars are always separated from palpebrals by 2-3 poorly defined rows of inconspicuous tiny

scales. In IRSNB 2674 three of these scales (vs. two in the holotype) are distinctly enlarged (larger than

in holotype) with angular downward protrusion between suboculars. In both juvenile paratypes, only

one of these scales is only slightly enlarged and slightly protrudes between suboculars. The condition of

the parietals and interparietal is highly variable among specimens and none of the specimens examined

has complete interparietal and parietals, which are always divided in several smaller scales (see Fig. 7

for comparison). Number of middorsal scales varies from 53 to 57. Ventrals are in 11 to 14 longitudinal

European Journal of Taxonomy 3: 1-18 (2011)

rows and in 29 to 30 transverse rows. The number of femoral pores varies from 8 to 10. Preanals vary

from 7 to 10.

The fully everted, maximally expanded, right hemipenis of the adult S paratype (IRSNB 2674) is identical

to the holotype's organ, except that maximal expansion makes the sulcus spermaticus bifurcation and

the sulcus branches even more difficult to appreciate. Maximal expansion does not highlight hemipenial

features that would have been overlooked in the not maximally expanded hemipenis of the holotype.

The hemipenis is 6.6 mm in length and 4.0 mm across its widest point when everted; there are twenty-

four rows of flounces extending along the organ, ca. eight of which are medioproximal asulcate flounces.

Colour in life ranges from beige to dark greyish, sometimes bluish, brown dorsally, depending on

light intensity. Rear of thighs usually has a few ill-defined dark markings. Some blue markings may be

visible around ear, or at the base of the tail, or on ventral parts. Ventral parts range from golden grey

to pale yellow. One of the juvenile paratypes (IRSNB 2675, see Fig. IB) has two poorly defined dark

Table 1. Sex, measurements (in mm) and scale counts of the type series of Anadia mcdiarmidi sp. nov.

1 = tail incomplete or obviously regenerated, 2 = data corresponding to right/left side, other abbreviations

are defined in the text.

Specimen

IRSNB 2677

IRSNB 2674

IRSNB 2675

IRSNB 2676

Sex

Juvenile $

SVL

62.2

60.4

38.2

42.6

110.0

57.0'

29.1'

74.0

15.8

13.6

9.0

10.2

10.8

8.7

5.8

6.6

6.7

5.7

3.9

4.4

SAL

28.4

23.8

14.6

18.9

10.8

6.7

5.2

6.0

AXG

30.7

29.9

18.4

19.8

F1L

16.6

10.2

11.6

H1L

22.4

18.5

14.3

15.6

SLF

13/14 2

14/13 2

12/13 2

13/14 2

SLT

17/17 2

17/15 2

17/17 2

SAM

7/7 2

in 2

nr-

6/6 2

7/6 2

5/5 2

10/9 2

9/9 2

9/8 2

10/10 2

KOK P.J.R. & RIVAS G.A., Anew Anadia from the Venezuelan 'Lost World'

brown stripes between eye and ear. In preservative all specimens became dark brown dorsally, looking

slightly peppered when in liquid. Ventral parts are always metallic blue (Fig. 6). Tympanum in preserved

specimens varies from dark brown to blue or golden blue.

Distribution and ecology

The new species is known only from the C hi m anta Massif in Venezuela (Fig. 8) where it has been

reported from Abakapa-tepui (type locality), Amuri-tepui, and Murei-tepui (sometimes named Eruoda-

tepui, see Discussion) between ca. 2100-2600 m elevation (McDiarmid & Donnelly 2005). The species

is probably widespread in the Chimanta Massif.

Anadia mcdiarmidi is diurnal and inhabits open, mostly flat sandstone areas on tepui summits (Fig.

ID). The holotype and two paratypes were collected during the day under rocks; IRSNB 2674 was

found during the day, crawling in a small crack between rocks. Two white fertile eggs (both measuring

12.3 mm) and several older eggshells attributable to the new species were found under a large rock,

suggesting that a same suitable nesting site may be reused [as already postulated by Gorzula (1992)],

and that females lay two eggs at a time. The only other lizard found syntopic with Anadia mcdiarmidi

was Neusticurus cf. rudis , which was also mainly found under rocks.

Discussion

As stated above, the new species has been mentioned previously in four publications: Gorzula (1992),

Gorzula & Senaris (1999), McDiarmid & Donnelly (2005), and Brewer-Carias & Audy (2010), but none

of these publications provides any comprehensive morphometric data, except a SVL of 86 mm and a

TL of 77 mm (suggesting a broken tail) in a large adult male (catalogued as MHNLS 10772, probably

the largest known specimen), and a SVL of 65 mm in a gravid female catalogued as MHNLS 10765

(Gorzula & Senaris 1999). The few published photographs [figs 168-171 in Gorzula (1992), fig. 88 in

Gorzula & Senaris (1999), and an unnumbered figure on page 210 in Brewer-Carias & Audy (2010)]

and data (Gorzula 1992; Gorzula & Senaris 1999) conform to the new species description and add the

following information: (1) large males obviously have a wider head than females, a character shared

by several Anadia species ( e.g. A. bogotensis , A. brevifrontalis, A. petersi, A. rhomb if era), (2) females

may have one or two faint darker stripes between orbit and ear (as also seen in one of our juvenile male

paratypes), (3) the known maximal size in the species is apparently 86 mm SVL.

Although Eruoda-tepui and Murei-tepui (occasionally spelled Murey-tepui) are sometimes considered

the same mountain (e.g. Gorzula & Senaris 1999), we consider them as separate tepuis (see Fig. 8).

McDiarmid & Donnelly (2005) provide 05°22'N, 062°05'W as the coordinates for Murei-tepui, and

clearly mention that it is the locality where Anadia sp. A (here described as Anadia mcdiarmidi) has

been collected. Eruoda-tepui is located north to Murei-tepui, and no report of the new species has been

recorded from that mountain yet.

Pellegrino et al. (2001) tentatively assigned the genus Anadia to the family Cercosaurinae (tribe

Cercosaurini) on the basis of morphology and our preliminary molecular results (Kok, unpublished)

corroborate this. Members of the genus Anadia have not been included in any modem phylogenetic

study and relationships between species, as well as their relationships with apparently close genera like

Euspondylus , remain unknown. The genus Euspondylus Tschudi, 1845 is ill defined (see Oftedal 1974;

Myers et al. 2009) and is mainly distinguished from Anadia by the presence of distinct rows of much

smaller scales between lateral scales (in Anadia lateral scales are subequal and mostly in unbroken

transverse rows with dorsal scales). A single study was conducted to test the phylogenetic position of

Euspondylus acutirostris (Peters, 1863) based on its cranial osteology and myology (Montero et al.

2002). Results from Montero et al. (2002) show E. acutirostris nested in a clade containing three species

European Journal of Taxonomy 3: 1-18 (2011)

of Anadia, thus suggesting that both genera are synonyms. However, Myers et al. (2009: 11) highlighted

that one of the E. acutirostris specimens used by Montero and colleagues could have been misidentified

and could actually be an Anadia marmorata , which is currently impossible to determine because that

specimen was dissected for the osteological portion of their study. Therefore, the phylogenetic position

of Anadia, intrageneric relationships within Anadia, and relationships to Euspondylus remain an enigma.

Based on morphology only it is challenging to infer relationships of the new taxon with other Anadia

species because many characters of the new species could be adaptations to high mountain habitat, and

thus shared by other high mountain species that may not be closely related. At this stage the lack of

adequate tissue sampling among Anadia species hampers any sound molecular analyses.

The origin of the Pantepui biodiversity still remains an evolutionary enigma (e.g. Rull 2009) and the

discovery of a species of Anadia (primarily an Andean genus) apparently endemic to tepui summits is

of considerable interest to the understanding of the processes underlying the origin and evolution of the

Pantepui herpetofauna. This matter is beyond the scope of this paper and is subject of ongoing research

that will be presented elsewhere.

Acknowledgements

The partial financial support from the King Leopold III Fund for Nature Exploration and Conservation

to PJRK is gratefully acknowledged. PJRK thanks B. Wilson for field companionship and C. Brewer-

Carias for clarification about some tepui names. Finally, we warmly thank R. Jadin and P. Sales Nunes

whose comments improved a previous version of this paper, and F. Bisbal, E. Camargo, D. Mora and D.

Sanchez (EBRG) for the loan of specimens under their care and various help.

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Appendix

Additional material examined

Anadia blakei (1) VENEZUELA: Sucre: Cerro Humo, Peninsula de Paria, EBRG 2746.

Anadia marmorata (2) VENEZUELA: Aragua: Estacion Biologica de Rancho Grande, 1100 m, EBRG

3423. Distrito Capital: sector Jerenba Parque Nacional Macarao, 2300 m (living specimen to be deposited

at EBRG).

Anadia escalerae (1) VENEZUELA: Bolivar: La Escalera, 132 km SE Rio Cuyuni (= 46 km SE km 88)

on the road to Santa Elena de Uairen, EBRG 1998 (holotype).

Euspondylus acutirostris (3) VENEZUELA: Aragua: Carretera El Junquito-Colonia Tovar, a 1.5 km

a la derecha del arco de la Colonia Tovar, 2350 m. MHNLS 17474. Falcon: Sierra de San Luis, cerro

Galicia, La Soledad, 1200 m. EBRG 3971. Yaracuy: Pico el Tigre, Parque Nacional Yurubi, 1700 m

(living specimen to be deposited at EBRG).

Manuscript received: 5 August 2011

Manuscript accepted on: 5 September 2011

Published on: 29 September 2011

Topic editor: Rudy Jocque

In compliance with Article 8.6 of the ICZN, printed versions of all papers are deposited in the libraries

of the institutes that are members of the EJT consortium.