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Research article

um:lsid:zoobank.org:pub:67D17AEB-B622-4184-9265-D3D2D6DA50QA

Two new charismatic Pristimantis species (Anura: Craugastoridae)

from the tepuis of “The Lost World” (Pantepui region, South America)

Philippe J. R. KOK'-"

^Department of Vertebrates, Royal Belgian Institute of Natural Seienees, rue Vautier 29, B-1000

Brussels, Belgium

E-mail: [email protected]

^Amphibian Evolution Eab, Department of Biology, Vrije Universiteit Brussel, Pleinlaan 2, B-1050

Brussels, Belgium

E-mail: [email protected]

um:lsid:zoobank.org:author:62BA0CB3-FB94-4Q7F-9QAl-06AB6401B911

Abstract. Two new eolourfiil speeies of direet-developing frogs of the genus Pristimantis are deseribed

from the summit of two isolated tepuis (sandstone table mountains) in the Eastern Pantepui Distriet

of the Guiana Shield highlands. Pristimantis jamescameroni sp. nov. is deseribed from the summit of

Aprada-tepui from 2557-2571 m elevation, andP imthurni sp. nov. is deseribed from the summit ofPtari-

tepui at 2471 m elevation. Both speeies share the absenee of a differentiated tympanie membrane and

external tympanie annulus (but presenee of tiny pharyngeal ostia), the presenee of nuptial pads in males,

and the presenee of lateral fringes on fingers and toes, a eombination of eharaeters that immediately

distinguishes them from all other known Pantepui eongeners. The two new speeies are morphologieally

similar to eaeh other and are phylogenetieally elosely related, but they ean be distinguished based

on eolour pattern and morphologieal eharaeters sueh as head proportions, dorsal skin texture, and

eondition of the supratympanie fold. The lUCN eonservation status of the new speeies is eonsidered as

Endangered (EN) owing to their apparent very restrieted ranges. The number of deseribed Pristimantis

speeies oeeurring exelusively on tepui (and faunistieally related granitie mountains) summits and upper

slopes now reaehes eleven.

Keywords. Anura, Guiana Shield, Systematies, Taxonomy, Terrarana.

Kok PJ.R. 2013. Two new charismatic Pristimantis species (Anura: Craugastoridae) from the tepuis of “The Lost

World” (Pantepui region. South America). European Journal of Taxonomy 60: 1-24. http://dx.doi.org/10.5852/

eit.2013.60

Introduction

The outstanding biogeographie region where the tepuis (a word used to deseribe South Ameriean table

mountains eonsisting of Preeambrian sandstone) and the faunistieally related granitie mountains oeeur,

was named Pantepui by Mayr & Phelps (1967) in their seminal study on the origin of the bird fauna of the

south Venezuelan highlands. The Pantepui region (Fig. lA) lies in the northwestern part of the Guiana

Shield and is traditionally seen as one of the foremost eentres of endemism in the Neotropies (Berry et al.

1995; Davis et al. 1997). Historieally, Pantepui has often been limited to the upper slopes and summits

European Journal of Taxonomy 60 : 1-24 ( 2013 )

Fig. 1. A. Map of the Pantepui region, loeated in the western part of the Guiana Shield. The white

reetangle eorresponds to the area enlarged in (B). B. Distribution of known tepui summit Pristimantis

speeies oeeurring east of the Rio Caroni in Venezuela and in western Guyana: R aureoventris (Wei-

Assipu-tepui and upper slopes of Mount Roraima), R yuruaniensis (Yuruani-tepui), R muchimuk

(Churi-tepui), P. abakapa (Abakapa-tepui), P. auricarens mdP pulvinatus (Auyan-tepui); (1) Aprada-

tepui, type loeality of Pristimantis jamescameroni sp. nov. (area enlarged in Fig. 6A); (2) Ptari-tepui,

type loeality of Pristimantis imthurni sp. nov. (area enlarged in Fig. 6B).

KOK P. J.R., Two new Pristimantis from Pantepui

of the Guiana Shield highlands eovering a total surfaee of about 5000 km^, with an altitudinal range of

1,500-3,014 m above sea level (Mayr & Phelps 1967; Rull & Nogue 2007). Nonetheless, Steyermark

(1982), followed by Kok et al. (2012) and Kok (2013), expanded the original definition of Pantepui to

inelude the intervening Pantepui lowlands (200-400 m asl) and uplands (400 - ca. 1200 m asl), whieh

better refieets the biogeography and the past and eurrent biotie interaetions in the area (Kok 2013).

The Seottish writer Sir Arthur Conan Doyle was the first to eoin the expression “The Lost World” in the

title of his famous fietion novel published in 1912, in whieh he depiets an extraordinary expedition to

an isolated table mountain in the tropieal jungle of northern South Ameriea in seareh of dinosaur-like

ereatures and a forgotten eivilisation (Doyle 1912). Doyle was largely inspired by the aeeount of an

expedition of the Royal Geographieal Soeiety of London to the summit of Mount Roraima in 1884, led

by Sir Everard im Thum (MeDiarmid & Donnelly 2005; Dalziell 2007).

The vertieal walls of tepuis ean reaeh 1000 m, whieh makes most of these mountains isolated horizontally

from one another, but also vertieally from their surroundings. In addition to physiographie isolation,

eurrent edaphie, elimatie and eeologieal faetors eontribute to further isolate most tepui tops from the

surrounding savannah and tropieal forest.

Kok et al. (2012) demonstrated that, despite their eurrent unique geomorphology, the tepuis were

insuffieient barriers to loeal {i.e., within Pantepui) gene fiow, leading to a low genetie diversity among

most tepui summit amphibians and reptiles. The same authors suggested that single-tepui endemism

might not be as eommon as previously stated, sinee some tepui summit speeies, deseribed as being single-

tepui endemies, turned out to be synonyms of more widespread taxa {e.g. , in the gQnQraAnomaloglossus

and Tepuihyla, see for instanee Kok et al. 2013 and Jungfer et al. 2013). Evolutionary proeesses in the

area remain puzzling given that on the one hand, speeies previously reported as highly restrieted often

also oeeur on nearby tepuis, or in some eases in the surrounding uplands {Tepuihyla), and on the other

hand, some populations oeeurring on different tepui summits and identified as a single speeies turned

out to be eomplexes of distinet taxa phenotypieally hard to distinguish from one another {e.g, in the

genus Stefania, Kok pers. obs.). Kok et aPs (2012) diseovery has a potentially strong implieation for the

taxonomie status of several deseribed tepui speeies and highlights the diffieulty of taxonomie deeisions

in the region, whieh until now was often strongly infiueneed by the unique topography of the area, and

probably also by the legaey of Doyle’s novel. However, from a eonservation point of view it is erueial to

delimitate speeies and speeies’ distributions as aeeurately as possible, espeeially in these mountainous

areas reported as partieularly sensitive to global warming (Nogue et al. 2009).

The frog genus Pristimantis is the most speeiose vertebrate genus, with more than 450 nominal speeies

eurrently reeognized (Frost 2013). Twenty of these deseribed speeies are distributed in Pantepui (Kok

& Barrio-Amoros 2013; Rojas-Runjaie et al. 2013), nine of them stated as being restrieted to tepui

summits and tepui upper slopes {sensu Kok 2013; Rojas-Runjaie et al. 2013). These numbers do not

inelude the many putative undeseribed and highly restrieted speeies (at least 15), reported from the

region by MeDiarmid & Donnelly (2005) and still waiting to be named.

New material was reeently obtained from several poorly explored tepuis, whieh eonfirms the oeeurrenee

of two distinet, ummmQd Pristimantis speeies on Aprada-tepui and Ptari-tepui, respeetively, two isolated

tepuis loeated east of the Rio Caroni, in the Eastern Pantepui Distriet, Bolivar state, Venezuela. These

new speeies are deseribed below and their lUCN Red Eist Category is diseussed.

European Journal of Taxonomy 60: 1-24 (2013)

Material and methods

Fieldwork and deposition of specimens

Tepui summits harbour fragile ecosystems and extreme care was taken to ensure a minimum impact on

the environment (e.g., no waste was left, vegetation was not damaged and turned rocks were replaced as

they were found whenever possible). Specimens were collected by hand and euthanized by immersion in

a 2% lidocaine solution (Linisol), fixed in 10% formalin for a few days and transferred to 70% ethanol

for permanent storage. A piece of liver and/or thigh muscle was taken from most specimens prior to

fixation and preserved in 95% ethanol for later molecular analyses. Specimens were deposited in the

collections of the Institut Royal des Sciences Naturelles de Belgique (IRSNB). Tissue samples were

deposited in the Amphibian Evolution Lab, Biology Department at the Vrije Universiteit Brussel (VUB).

Coordinates and elevations of the collection localities were acquired using a Garmin Global Positioning

System unit and referenced to map datum WGS84.

Morphology

Terminology for morphological characters mostly follows Lynch & Duellman (1997), Savage et al.

(2004), Kok & Kalamandeen (2008), Means & Savage (2007), Duellman & Lehr (2009), and Kok et

al. (2011). 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, descriptions mainly follow the pattern of recent descriptions in the

genus (e.g, Myers & Donnelly 2008; Barrio-Amoros et al. 2010; Kok et al. 2011; Rojas-Runjaic et al.

2013).

Classical measurements for the genus were taken and are abbreviated as follows:

HANDl

HAND 11

HAND 111

HAND IV

SVL

TIL

WFD

WTD

eye length, horizontal distance from the posterior margin to the anterior margin of the eye

eye to naris distance from anterior comer of eye to posterior margin of naris

foot length from proximal edge of outer metatarsal tubercle to tip of Toe IV

length of Finger 1 from proximal edge of palmar tubercle to tip of finger

length of Finger 11 from proximal edge of palmar tubercle to tip of finger

length of Finger 111 from proximal edge of palmar tubercle to tip of finger, also equi¬

valent to hand length

length of Finger IV from proximal edge of palmar tubercle to tip of finger

head length from angle of jaw to tip of snout

greatest width of the head

intemarial distance, taken between the median margins of the nares

interorbital distance, taken between the inner margins of the orbits

snout length from anterior comer of eye to tip of snout

snout-vent length, from tip of snout to posterior margin of vent

tibia length from outer edge of fiexed knee to heel

width of disc on Finger 111

width of disc on Toe IV

Colour pattern in life was taken from field notes and was later refined based on high definition colour

digital photographs. Sex and maturity were determined by dissection and direct examination of gonads,

and by examination of secondary sexual characters (such as nuptial pads in males). Internal soft anatomy

was examined by dissection of preserved specimens.

Taxonomy follows Pyron & Wiens (2012). Institutional acronyms follow Frost (2013).

KOK P.J.R., Two new Pristimantis from Pantepui

Results

Generic allocation of the new species

No identifiable morphologieal synapomorphy supporting the genus Pristimantis (Hedges et al. 2008) has

been proposed yet. The new taxa are assigned to the genus Pristimantis based on moleeular phylogenetie

relationships (Kok et al. 2012, Kok pers. obs., August 2012) as well as on their morphologieal

eharaeteristies, whieh fall within the range of oihQx Pristimantis speeies (see diagnoses).

Class Amphibia Gray, 1825

Order Anura Fiseher von Waldheim, 1813

Family Craugastoridae Hedges, Duellman & Heinieke, 2008

Subfamily Pristimantinae Pyron & Wiens, 2011

Pristimantis jamescameroni sp. nov.

Figs 2-5, Table 1

um:lsid:zoobank.org:aet:67EB6C36-0F31-4587-9805-3136BEF4E010

Eleutherodaetylinae series b (in part) - Gorzula & Senaris 1999: 55 + plate 46.

Eleutherodactylus sp. B -MeDiarmid & Donnell, 2005: 514 [table 18A.1], 524.

Pristimantis sp. - Hedges et al. 2008: 179.

Pristimantis “Aprada” - Kok a/. 2012: Suppl. Information: 13.

Definition and diagnosis

A small speeies of the genus Pristimantis eurrently not assigned to any speeies group, but morphologi-

eally most similar to speeies of the polyphyletie unistrigatus speeies group {sensu Hedges et al. 2008,

see also diseussion in Kok et al. 2011) mainly eharaeterized in having Finger I shorter than II, Toe V

longer than III, extending to the distal edge of the distal subartieular tuberele of Toe IV when toes are

adpressed, and by the absenee of eranial erests and the presenee of vomerine teeth. The new speeies

is eharaeterized by the following unique eombination of eharaeters: (1) body small, male 22.8 mm

SVE, females 26.3-27.5 mm (n=3); (2) dorsal skin shagreen, belly skin eoarsely areolate; (3) tympanum

absent (tympanie membrane not differentiated and tympanie annulus not visible externally); (4) tiny

pharyngeal ostia present; (5) snout rounded to subovoid in dorsal view, rounded to slightly sloping in

profile, canthus rostralis nearly straight in dorsal view, rounded in eross seetion, loreal region slightly

eoneave, fiaring slightly at upper lip; (6) upper eyelid shagreen with 1-2 distinetly enlarged tubereles on

eaeh eyelid; (7) ehoanae small, oval, dentigerous proeesses of vomers very small, barely visible, slightly

oblique, ovoid to triangular, posterior and medial to ehoanae, eaeh bearing 1-5 teeth; (8) absenee of voeal

slits in male; (9) tongue eordiform; (10) two large, mostly unpigmented nuptial pads on eaeh thumb in

male; (11) Finger I shorter than II; (12) fingers with lateral fringes, best developed preaxially on Fingers

II-III; (13) ulnar tubereles absent or ineonspieuous, when present low, not forming a distinet line; (14)

tarsal tubereles absent, one ineonspieuous ealear tuberele present; (15) inner metatarsal tuberele oval,

about four times the size of the round, projeeting outer metatarsal tuberele; (16) Toe V longer than III,

extending to the distal edge of the distal tuberele on Toe IV when toes are adpressed; (17) toes with

lateral fringes, best developed preaxially on Toes III-IV, webbing basal between Toes IV-V; (18) in life

main dorsal baekground eolouration is orange to orangish dark brown, either with irregular dark brown

ehevrons or a darker middorsal band, and/or a light middorsal stripe, ventral eolouration is white with

brown retieulation and an orange pateh posteriorly; (19) in preservative dorsal baekground eolouration

is light to dark brown, either with irregular brown ehevrons or a darker middorsal band, and/or a light

middorsal stripe, ventral eolouration is eream with brown retieulation.

European Journal of Taxonomy 60: 1-24 (2013)

Etymology

The specific epithet is a noun in the genitive case, honouring the Canadian film director, producer,

enviro nm entalist and explorer Mr. James F. Cameron in recognition of his efforts to alert the general

public to environmental problems through pioneering high quality “blockbuster” movies and adventu¬

rous documentaries. James Cameron also encourages people to go vegan (a diet excluding animal

products), one of the effective ways to reduce human environmental impacts such as global climate

change, identified as a serious threat to tepuis ecosystems (see Nogue et al. 2009).

Material examined

Holotype

VENEZUELA: an adult male collected by Philippe J. R. Kok, 15 Jun. 2012 at 08hl8, summit of Aprada-

tepui, Bolivar state, 5°24’42”N, 62°27’00” W, 2570 m elevation, IRSNB 4160 (field number PK 3636).

Paratopotypes (n=4)

VENEZUELA: three adult females (IRSNB 4161-63, field numbers PK 3632, PK 3637, PK 3635) and

one juvenile (IRSNB 4164, field number PK 3639), collected by Philippe J. R. Kok (except IRSNB

4161, collected by Brad Wilson) 14-15 Jun. 2012, all from the summit of Aprada-tepui between 2557-

2571 m elevation.

Fig. 2. Pristimantis jamescameroni sp. nov. Intraspecific variation in dorsal and ventral colour pattern

in living specimens. A. Dorsolateral view of the S holotype (IRSNB 4160). B. Ventral view of the same

specimen. C. Dorsolateral view of a $ paratype (IRSNB 4163). D. Ventral view of the same specimen.

E. Dorsolateral view of a $ paratype (IRSNB 4162). F. Ventral view of the same specimen. Photographs

by the author.

KOK P.J.R., Two new Pristimantis from Pantepui

Description of the holotype

An adult male 22.8 mm SVL (Figs 2A-B, 3,4D, H, see Table 1 for measurements), in very good eondition,

exeept a large sear under the right thigh where a pieee of musele was removed prior to preservation. Head

slightly longer than wide (HW 94.7% of HL), wider than body; HW 39.0% of SVL; HL 41.2% of SVL;

eranial erests absent. Snout longer than eye length (SL 117.1% of EL), rounded to subovoid in dorsal

view, rounded to slightly sloping in profile; canthus rostralis nearly straight in dorsal view, rounded in

eross seetion, loreal region slightly eoneave, fiaring slightly at upper lip; eye-naris distanee shorter than

eye length (EN 71.4% of EE). Nares slightly protuberant, direeted posterolaterally, visible in frontal and

dorsal views. Widest upper eyelid width narrower than interorbital distanee (upper eyelid width 82.7%

of 10), upper eyelid shagreen with 1-2 enlarged tubereles on eaeh eyelid. Tympanum absent (tympanie

membrane not differentiated and tympanie annulus not visible externally); tiny pharyngeal ostia present.

Supratympanie fold eonspieuous in life, slightly arehed, originating at posterior eomer of eye, failing

to reaeh shoulder; post-rietal tubereles evident. Choanae small, oval, not eoneealed by palatal shelf of

maxillary areh; dentigerous proeesses of vomers very small, barely visible, slightly oblique, ovoid to

triangular, posterior and medial to ehoanae, eaeh bearing 3-4 teeth. Tongue eordiform, slightly longer

than wide, rounded posteriorly, posterior half free. Voeal slits and voeal sae absent.

Fig. 3. Pristimantis jamescameroni sp. nov. Ventral view of left hand and left foot of the S holotype

(IRSNB 4160). Photographs by the author.

European Journal of Taxonomy 60: 1-24 (2013)

Fig. 4. Pristimantis jamescameroni sp. nov. Intraspecific variation in dorsal (above) and ventral (below)

colour pattern in preserved specimens. A, E. $ paratype (IRSNB 4161). B, F. $ paratype (IRSNB

4162). C, G. $ paratype (IRSNB 4163). D, H. S holotype (IRSNB 4160). Photographs by the author.

KOK P. J.R., Two new Pristimantis from Pantepui

Dorsal skin shagreen, ineluding on head; middorsal raphe deteetable; no dermal folds or ridges visible

on dorsal surfaee; flanks granular. Throat surfaee shagreen; upper ehest shagreen, slightly “wrinkled”;

weak thoraeie fold; belly skin eoarsely areolate; weak diseoidal fold anterior to groin; posteroventral

thigh and eloaeal region eoarsely areolate; eloaeal sheath absent.

Hand length 27.2% of SVL. Finger I 80.0% of II. Relative length of Angers III > IV > II > I; adpressed

Finger I fails to reaeh proximal edge of digital pad of Finger II; adpressed Finger IV reaehes the

interealary eartilage of Finger III on the left side, the base of the dise of Finger III on the right side. Two

large, not eonneeted, non-spinous, mostly unpigmented (translueent when wet, white when dry) nuptial

pads on eaeh thumb, one extending along the preaxial surfaee of the thenar tuberele and invading most

of it, the other one extending along the dorsal and the preaxial surfaee of the thumb. Lateral fringes on

all Angers, best developed preaxially on Fingers II-III (Fig. 3). Finger dises broadly expanded, elliptieal,

broader than long, eireumferential groove eonspieuous, distal edge of dise rounded; dise of Finger III

2.1 times wider that the distal end of the adjaeent phalanx. Palmar tuberele large, poorly defined, not

fully pigmented, deeply bifid; thenar tuberele large, protuberant, ovoid; supernumerary tubereles few,

large (almost equal in size to the subartieular tubereles), slightly protuberant; subartieular tubereles

large, round and protuberant, one eaeh on FI and FII, two eaeh on Fill and FIV. Ulnar tubereles few,

ineonspieuous, forming an ill-defined line; presenee of a small antebraehial tuberele.

Hind limbs moderate in length, heels slightly overlap when held at right angles to sagittal plane; TIL

50.4% of SVL; FL 41.7% of SVL. Relative length of Toes IV > V > III > II > I; tip of Toe V extends

to the distal edge of the distal subartieular tuberele on Toe IV; tip of Toe III extends to the distal

edge of the penultimate subartieular tuberele on Toe IV on the left side, to the proximal half of the

penultimate subartieular tuberele on Toe IV on the right side. Lateral fringes on aft toes, best developed

preaxially on Toes III-IV; webbing basal between Toe IV-V (Fig. 3). Toe dises mostly equal in size to

Anger dises, WTDAVFD = 1; toe dises broadly expanded, elliptieal, broader than long, eireumferential

groove eonspieuous, distal edge of dise rounded. Inner metatarsal tuberele elongate, oval, about four

times the size of the projeeting, round outer metatarsal tuberele; subartieular tubereles round, large and

protuberant; supernumerary plantar tubereles small, low and round, inereasing in size distally. Single,

ineonspieuous and very small round ealear tuberele; no outer tarsal tubereles deteetable; inner tarsal

fold not deteetable.

Colour of holotype in life (see Fig. 2A-B)

Dorsal baekground eolour orangish brown eovered with numerous tiny white fleeks, top of head slightly

darker; ineonspieuous darker (than adjaeent dorsum) middorsal band outlined by ill-defined dark brown

dorsolateral lines; irregular white band on the canthus rostralis\ bluish white line on outer edge of upper

eyelids; upper lips white; most post-rietal tubereles white; enlarged tubereles on eyelids light brown.

Flanks orangish brown, with numerous tiny white fleeks and some irregular white spots of variable

sizes; groins, anterior thighs, and armpits bright orange. Arms and legs eoloured as dorsal surfaees,

with a few small irregular white spots on upper arms and on the distal part of tarsi. Throat, ehest, and

lower flanks white with brown anastomosed retieulation; belly and most of the undersurfaee of thighs

bright orange; undersurfaee of distal thighs, shanks, and tarsi brown with a few small irregular white

spots; posterior surfaee of thighs and eloaeal area brown. Upper faee of hands and feet orangish brown,

exeept tip of Fingers I-II, whieh is bright orange, and tip of toes, whieh is whitish. Palms and soles

brown; undersurfaee of Fingers I-III bright orange, ineluding dises; undersurfaee of tip of Toes I-III

bright orange ineluding dises. Iris silver with dark brown venation and an ill-defined, broad horizontal

brownish stripe; greenish silver spot on the posterior upper part of the iris.

European Journal of Taxonomy 60: 1-24 (2013)

Table 1. Sex, measurements (in mm), and HL/SVL ratio of type speeimens ofPristimantisjamescameroni

sp. nov. and Pristimantis imthurni sp. nov. Abbreviations are defined in the text.

Pristimantis jamescameroni sp. nov.

Aprada-tepui

Pristimantis imthurni sp. nov.

Ptari-tepui

Charaeter

IRSNB

4160

IRSNB

4161

IRSNB

4162

IRSNB

4163

IRSNB

4164

IRSNB 4165

Sex

Male

Female

Juvenile

Male

SVL

22.8

27.0

27.5

26.3

18.3

22.9

9.4

10.9

11.2

10.9

8.0

8.7

HL/SVL

41.2%

40.3%

40.7%

41.4%

43.7%

38.0%

8.9

10.1

10.3

9.9

7.4

8.5

4.1

4.5

4.7

4.3

3.3

3.7

2.5

2.9

2.7

2.0

2.4

2.1

2.5

2.4

2.5

2.1

2.3

3.5

3.3

3.4

3.2

2.1

3.3

2.6

2.9

3.0

2.4

2.7

HANDl

3.6

4.2

4.0

3.1

3.9

HAND 11

4.5

4.7

4.5

5.1

3.4

4.4

HAND 111

6.2

6.7

6.6

4.8

6.2

HAND IV

4.8

5.3

4.9

5.6

3.7

4.9

WFD

1.3

1.4

1.0

1.3

9.5

10.5

10.4

10.6

7.4

9.3

WTD

1.3

1.5

1.0

1.4

TIE

11.5

13.2

12.7

12.8

9.9

11.5

Colour of holotype in preservative (see Fig. 4D, H)

After 13 months in 70% ethanol (July 2013), orangish brown faded to light brown. Orange and white

faded to eream. Dorsal pattern turned generally more eonspieuous. Brown retieulation on throat,

ehest, and lower fianks faded to light brown. Granules in the eloaeal area, as well as subartieular and

supernumerary tubereles beeame dark brown. Dises are greyish brown.

Sexual dimorphism and variation among paratypes

See Table 1 for measurements of the type series, and Figs 2, 4, 5 for intraspeeifie variation. Sexual

dimorphism evident in size, with adult females being larger than the adult male (26.3-27.5 v^. 22.8 mm

SVL), and by the presenee of two nuptial pads in the male. A sexual diehromatism in belly eolouration

seems to oeeur, with the presenee of an orange spot restrieted to the middle area of the belly in living

females, whereas the orange area eovers the belly, the posterior part of the ventral surfaee, and the

undersurfaee of thighs in a more eontinuous way in the living male (see Fig. 2B, D, F). Hands and feet

slightly longer in the male than in the females (HAND 111 27.2% v^. 24.0-25.1% of SVL; FL 41.7% v^.

37.8-40.3% of SVL). No additional signifieant differenee is deteeted in other size ratios. No signifieant

variation in skin texture oeeurs among the preserved speeimens, but in life IRSNB 4163 (adult female.

Fig. 2C) had a eonspieuous dorsolateral fold extending from behind the eye to the end of the body;

that fold totally disappeared in preservative. Eyelid tubereles are more difheult to deteet in preserved

speeimens and range from 1 to 2 on eaeh eyelid. Number of teeth on dentigerous proeesses of vomers

KOK P.J.R., Two new Pristimantis from Pantepui

varies from 2 to 6. Weak thoraeie and diseoidal folds deteetable in IRSNB 4163 (female) and IRSNB

4164 (juvenile), not seen in the other paratypes. Tip of Toe V fails to reaeh the distal edge of the distal

subartieular tuberele on Toe IV on the left side in IRSNB 4161 (female) only.

Colour pattern variable. In life, dorsal baekground eolouration varies from bright orange to orangish

dark brown, usually eovered with tiny white fleeks; dorsal pattern varies from irregular brown ehevrons

(e.g., in IRSNB 4161 and IRSNB 4162) to a slightly darker (than adjaeent dorsum) middorsal band

outlined by ill-defined dark brown dorsolateral lines (e.g, in IRSNB 4160) and/or a pale middorsal

stripe (e.g, in IRSNB 4163); pattern on fianks varies from a few irregular oblique white (e.g, IRSNB

4163) or brown/white (e.g., IRSNB 4162) stripes to white spots (holotype); eolour of ventral retieulation

varies from brown to dark brown; ineonspieuous transverse bands on arms and legs oeeur in two female

paratypes (IRSNB 4162 and IRSNB 4163). Enlarged tubereles on eyelids are usually white (light brown

in the holotype). The juvenile (IRSNB 4164, Fig. 5) has a eomplex and more marked dorsal pattern

eonsisting of a greyish brown interorbital line preeeded by a greyish brown eirele and followed by

a wide W-shaped marking outlined by greyish brown in the seapular region; the W-shaped marking

is followed by a greyish brown areh, itself followed by two baek to baek C-shaped light grey lines.

It also differs from adults in having an irregular greyish band on the canthus rostralis, the upper lip

not eompletely white, granules in the loreal region, dark brown labial stripes, the supratympanie fold

underlined by a dark brown band, brown transverse bands on the forearm, and reddish transverse bands

on the legs. Iris varies from silver or greenish silver to bronze, with dark brown venation and an ill-

defined, broad horizontal brownish stripe.

Fig. 5. Pristimantisjamescameroni sp. nov. Juvenile speeimen (IRSNB 4164,18.3 mm SVL). Photograph

by the author.

European Journal of Taxonomy 60: 1-24 (2013)

Araopan-tepul

Aprada-tepui

Posaipa-tepui

Perediaca-tepui

^■4 ■

ft . /rti- ■ ^

- * Chirca-tepui

‘■'l.AWJ

'ievlyv-..

Ptari-tepui

Carrao-tepui

Sororopan-tepui

Fig. 6. A. Satellite map of the Aprada massif, showing the loeation of Aprada-tepui and neighbouring

smaller table mountains. Yellow star indieates the type loeality of Pristimantis jamescameroni.

B. Satellite map of the Ptari massif, showing the loeation of Ptari-tepui and neighbouring mountains.

Yellow star indieates the type loeality of Pristimantis imthurni. Satellite maps © Google Maps.

KOK P. J.R., Two new Pristimantis from Pantepui

Morphological comparisons with congeneric species

Available data suggest that tepui summit Pristimantis speeies have relatively restrieted distributions in

the Guinea Shield highlands; therefore, eomparisons with eongeners only foeus on Pristimantis speeies

known to oeeur in the Pantepui region (as defined by Kok 2013). Comparisons of external eharaeter

states are based both on original deseriptions and examination of museum speeimens (see Appendix for

material examined).

Twenty speeies of Pristimantis are eurrently reported from the Pantepui region: P. aureoventris Kok,

Means & Bossuyt, 2011, only known from the summit of Wei-Assipu-tepui and the upper slopes of

Mount Roraima, Cuyuni-Mazaruni Distriet, Guyana; P abakapa Rojas-Runjaie, Salerno, Senaris &

Pauly, 2013, deseribed from the summit of Abakapa-tepui, Bolivar state, Venezuela; P. auricarens

(Myers & Donnelly, 2008), known only from Auyan-tepui, Bolivar state, Venezuela; P avius (Myers

& Donnelly, 1997), reported from Pieo Tamaeuari, Amazonas state, Venezuela; P cantitans (Myers &

Donnelly, 1996), known from Cerro Yavi and Cerro Yutaje, Amazonas state, Venezuela; P. dendrobatoides

Means & Savage, 2007, only known from the Wokomung massif, Potaro-Siparuni Distriet, Guyana;

P. guaiquinimensis (Sehliiter & Rodder, 2007), deseribed from Guaiquinima-tepui, Bolivar state,

Venezuela; P jester Means & Savage, 2007, known from the Wokomung massif, Potaro-Siparuni Distriet,

Guyana, and the slopes of Maringma-tepui, Cuyuni-Mazaruni Distriet, Guyana (Kok, pers. obs. 2007);

P. marahuaka (Fuentes-Ramos & Barrio-Amoros, 2004), endemie to Cerro Marahuaka, Amazonas state,

Venezuela; P. marmoratus (Boulenger, 1900), reported as widespread in the Guiana Shield from eastern

Venezuela to Freneh Guiana and northern Brazil (Frost 2013), but several speeies may hide under this

name; P. memorans (Myers & Donnelly, 1997), known only from the Sierra Tapirapeeo, Amazonas

state, Venezuela; P. Barrio-Amoros, Mesa, Brewer-Carias & MeDiarmid 2010, apparently

endemie to the summit of Churi-tepui, Bolivar state, Venezuela; P. pruinatus (Myers & Donnelly, 1996),

known only from Cerro Yavi, Amazonas state, Venezuela; P. pulvinatus (Rivero, 1968), reported from

the Gran Sabana region and Auyan-tepui, Bolivar state, Venezuela to western Guyana (Frost 2013),

but more than one speeies may hide under this name (see Kok et al. 2012); P. saltissimus Means &

Savage, 2007, known from the Wokomung massif, Potaro-Siparuni Distriet, Guyana, and the slopes

of Maringma-tepui, Cuyuni-Mazaruni Distriet, Guyana (Kok pers. obs. 2007); P. sarisarinama Barrio-

Amoros & Brewer-Carias, 2008, endemie to Sarisarinama-tepui, Bolivar state, Venezuela; P. vilarsi

(Melin, 1941), known from upper Amazonian Brazil, Colombia, Venezuela and Peru (Frost 2013);

P yaviensis (Myers & Donnelly, 1996), reported from Cerro Yavi and Cerro Yutaje, Amazonas state,

Venezuela; P yuruaniensis Rodder & Jungfer, 2008, known with eertainty only from the summit of

Yuruani-tepui, Bolivar state, Venezuela (possible oeeurrenee on the neighbouring Kukenan-tepui, see

Magdefrau & Magdefrau 1994; Rodder & Jungfer 2008; Kok et al. 2011); and P. zeuctotylus (Lyneh &

Hoogmoed, 1977), widespread in northeastern South Ameriea (Frost 2013).

Pristimantis jamescameroni sp. nov. is most easily distinguished from P. abakapa, P aureoventris, P

avius, P cantitans, P dendrobatoides, P guaiquinimensis, P marmoratus, P memorans, P pruinatus,

P pulvinatus, P saltissimus, P sarisarinama, P vilarsi, P yuruaniensis and P. zeuctotylus by laeking a

differentiated tympanie membrane and an external tympanie annulus (at least one of these struetures is

deteetable in the latter 15 speeies). It further differs from P. vilarsi and P. zeuctotylus in having Finger

I < II (Finger I > II in P vilarsi and P. zeuctotylus).

Pristimantis jamescameroni sp. nov. is immediately distinguished from P. auricarens, P. jester, P.

marahuaka, P. muchimuk, and P. yaviensis (the only known Pantepui “earless” Pristimantis) in having

eonspieuous lateral fringes on fingers and toes (absent or limited to a weak keel in the latter 5 speeies).

European Journal of Taxonomy 60: 1-24 (2013)

Fig. 7. A The summit of Aprada-tepui lying above the elouds, photographed from a helieopter looking

to the SW (2 May 2011). B. Summit of Aprada-tepui (15 Jun. 2012). Photographs by the author.

KOK P. J.R., Two new Pristimantis from Pantepui

Distribution and ecology

Pristimantis jamescameroni sp. nov. is only known from the summit of Aprada-tepui (Figs IB, 6A,

7A-B), Bolivar state, Venezuela, where it oeeurs from 2557 to 2571 m elevation. Aprada-tepui lies in

the Aprada massif (Fig. 6A), in the Chimanta subdistriet (MeDiarmid & Donnelly 2005). It is loeated

ca. 22 km NW of the Chimanta massif and ca. 30 km S of Auyan-tepui (airline). Aeeording to the

GPS, Aprada-tepui reaehes a maximal elevation of about 2575 m above sea level. The summit area of

Aprada-tepui is ca. 4.3 km^, and is mainly eovered by open roek vegetation and small islands of tepui

forests (Huber 1995, Fig. 7B); it is eharaeterized by a high number of small lakes (Fig. 7B) and some

deep eanyons.

Aside from the male holotype, whieh was eolleeted by day on the ground while ealling in a small

shallow roek ereviee, hidden by the vegetation, all speeimens were eolleeted under roeks. Other males

were heard ealling very sporadieally from shallow roek ereviees by day, but eould not be loeated. As

in most other tepui summit Pristimantis, the new speeies is not abundant and individuals are diffieult

to eolleet. The only other anuran reported from the summit of Aprada-tepui is Stefania satelles Senaris,

Ayarzagtiena & Gorzula, 1997, whieh oeeurs in higher density and might partly feed on P. jamescame¬

roni sp. nov. (pers. obs.).

Pristimantis imthurni sp. nov.

Figs 8-10, Table 1

um:lsid:zoobank.org:aet:9AFC18C4-23AQ-4C2Q-96EB-480F0F512791

Eleutherodactylus sp. H - MeDiarmid & Donnelly 2005: 514 [table 18A.1], 521.

Definition and diagnosis

A small speeies of the genus Pristimantis eurrently not assigned to any speeies group, but morphologi-

eally most similar to speeies of the polyphyletie unistrigatus speeies group {sensu Hedges et al. 2008,

see also diseussion in Kok et al 2011) mainly eharaeterized in having Finger I shorter than II, Toe V

longer than III, extending to the distal edge of the distal subartieular tuberele of Toe IV when toes are

adpressed, and by the absenee of eranial erests and the presenee of vomerine teeth. The new speeies

is eharaeterized by the following unique eombination of eharaeters: (1) body small, male 22.9 mm

SVL; (2) dorsal skin smooth, belly skin eoarsely areolate; (3) tympanum absent (tympanie membrane

not differentiated and tympanie annulus not visible externally); (4) tiny pharyngeal ostia present; (5)

snout broadly rounded in dorsal view and in profile, canthus rostralis nearly straight in dorsal view,

rounded in eross seetion, loreal region eoneave, flaring slightly at upper lip; (6) upper eyelid smooth

with one barely distinet enlarged tuberele; (7) ehoanae very small, round, dentigerous proeesses of

vomers ineonspieuous, ovoid, posterior and medial to ehoanae, eaeh bearing 3-4 teeth; (8) absenee

of voeal slits in male; (9) tongue eordiform; (10) two large, broadly eonneeted, mostly unpigmented

nuptial pads on eaeh thumb in male; (11) Finger I shorter than II; (12) Angers with lateral fringes, best

developed preaxially on Fingers II-III; (13) ulnar tubereles absent; (14) tarsal tubereles absent, one

ineonspieuous ealear tuberele present; (15) inner metatarsal tuberele oval, about four times the size of

the round, projeeting outer metatarsal tuberele; (16) Toe V longer than III, extending to the distal edge

of the distal tuberele on Toe IV when toes are adpressed; (17) toes with lateral fringes, best developed

preaxially on Toes III-IV, webbing basal between Toes IV-V; (18) in life dorsal baekground eolouration is

vivid yellow with a ehoeolate brown middorsal stripe and ehoeolate brown flanks, ventral eolouration is

translueent brown with a few white spots and fleeks; (19) in preservative dorsal baekground eolouration

is pale yellow with a light brown middorsal stripe and light brown flanks, ventral eolouration is light

grey with a few white spots and fleeks.

European Journal of Taxonomy 60: 1-24 (2013)

Etymology

The specific epithet is a noun in the genitive case, honouring Sir Everard F. im Thum (1852-1932),

British colonial official, author, explorer, botanist, and photographer. Everard im Thum was the first to

climb a major tepui (Mount R or a im a in December 1884), along with British surveyor Harry Perkins, a

Pomeroon Amerindian named Gabriel, and five other u nn amed Am erindians (Dalziell 2007). Im Thum’s

expedition on Roraima and his numerous discoveries were partly eclipsed by the popular novel that they

inspired: “The Eost World” by Sir Arthur Conan Doyle (see Introduction; Dalziell 2007).

Material examined

Holotype

VENEZUEEA: an adult male collected by Philippe J. R. Kok, 20 Jun. 2012 at 20h37, summit of Ptari-

tepui, Bolivar state, 5°46’09” N, 61°49’02” W, 2471 m elevation, IRSNB 4165 (field number PK 3671).

Fig. 8. Pristimantis imthurni sp. nov. in life. A. Dorsolateral view of the S holotype (IRSNB 4165).

B. Ventral view of the same specimen. Photographs by the author.

KOK P.J.R., Two new Pristimantis from Pantepui

Morphological comparisons with congeneric species

For the same reasons mentioned above, eomparisons with eongeners only foeus on the 21 Pristimantis

speeies known to oeeur in the Pantepui region.

Pristimantis imthurni sp. nov. is most easily distinguished from P abakapa, P aureoventris, P avius,

P cantitans, P. dendrobatoides, P guaiquinimensis, P marmoratus, P memorans, P pruinatus, P

pulvinatus, P saltissimus, P sarisarinama, P. vilarsi, P yuruaniensis, and P zeuctotylus by laeking a

differentiated tympanie membrane and an external tympanie annulus (at least one of these struetures is

deteetable in the latter 15 speeies). It further differs from P vilarsi and P. zeuctotylus in having Finger I

< II (Finger I > II in P vilarsi and P. zeuctotylus).

Pristimantis imthurni sp. nov. is immediately distinguished from P. auricarens, P jester, P. marahuaka,

P muchimuk, and P yaviensis in having lateral fringes on most fingers and toes (absent or limited to a

weak keel in the latter 5 speeies).

By laeking a differentiated tympanie membrane and an external tympanie annulus, and in having

lateral fringes on fingers and toes, Pristimantis imthurni sp. nov. is morphologieally most similar to

P. jamescameroni sp. nov., deseribed earlier in this paper. These two speeies are mainly distinguished

based on distinet dorsal and ventral eolour patterns (eompare Fig. 2 with Fig. 8), skin texture (smooth

in P. imthurni sp. nov. v^. shagreen in P. jamescameroni sp. nov.), eondition of the supratympanie fold

(ineonspieuous inP imthurni sp. nov. v^. eonspieuous inP. jamescameroni sp. nov.), and head proportion

(shorter, with a more rounded snout in P imthurni sp. nov.). Besides being morphologieally reliably

diagnosable, the two speeies are eurrently geographieally isolated (isolated lineage segments).

Fig. 9. Pristimantis imthurni sp. nov. Ventral view of left hand and left foot of the S holotype (IRSNB

4165). Photographs by the author.

European Journal of Taxonomy 60: 1-24 (2013)

Description of the holotype

An adult male 22.9 mm SVL (Figs 8-10, see Table 1 for measurements), in very good eondition, exeept

a large sear under the right thigh where a pieee of musele was removed prior to preservation. Head

slightly longer than wide (HW 97.7% of HL), wider than body; HW 37.1% of SVL; HL 38.0% of SVL;

eranial erests absent. Snout longer than eye length (SL 112.1% of EL), broadly rounded in dorsal view

and in profile; canthus rostralis nearly straight in dorsal view, rounded in eross seetion, loreal region

eoneave, fiaring slightly at upper lip; eye-naris distanee shorter than eye length (EN 72.7% of EE).

Nares slightly protuberant, direeted posterolaterally, visible in frontal and dorsal views. Widest upper

eyelid width narrower than interorbital distanee (upper eyelid width 81.5% of 10), upper eyelid smooth

with one barely visible enlarged tuberele on eaeh eyelid. Tympanum absent (tympanie membrane not

differentiated and tympanie annulus not visible externally); tiny pharyngeal ostia present. Supratympanie

fold ineonspieuous in life, barely visible and fragmented in preservative, slightly arehed, apparently

originating at posterior eomer of eye, failing to reaeh shoulder; a few low and small post-rietal tubereles.

Choanae very small, round; dentigerous proeesses of vomers ineonspieuous, ovoid, posterior and medial

to ehoanae, eaeh bearing 3-4 teeth. Tongue eordiform, slightly wider than long, rounded posteriorly,

posterior one-third free. Voeal slits and voeal sae absent.

Dorsal skin smooth, ineluding on head; middorsal raphe barely deteetable in preservative, well visible in

life; no dermal folds or ridges visible on dorsal surfaee; fianks slightly granular. Throat and upper ehest

surfaees smooth; no thoraeie fold deteetable; belly skin eoarsely areolate; no diseoidal fold deteetable;

posteroventral thigh and eloaeal region eoarsely areolate; eloaeal sheath absent, two distinetly enlarged

tubereles below eloaea.

Hand length 27.1% of SVE. Finger 1 88.6% of 11. Relative length of fingers 111 > IV > 11 > 1; adpressed

Finger 1 fails to reaeh proximal edge of digital pad of Finger 11; adpressed Finger IV reaehes proximal

edge of digital pad of Finger 111. Two large, broadly eonneeted, non-spinous, mostly unpigmented

(translueent when wet, white when dry) nuptial pads on eaeh thumb, extending from the preaxial surfaee

of the thenar tuberele and invading most of it, along the dorsal and the preaxial surfaee of the thumb.

Eateral fringes on all fingers, best developed preaxially on Fingers 11-111 (Fig. 9). Finger dises expanded,

elliptieal, broader than long, eireumferential groove eonspieuous, distal edge of dise rounded; dise of

Fig. 10. Pristimantis imthurni sp. nov. in preservative. A. Dorsol view of the S holotype (IRSNB 4165).

B. Ventral view of the same speeimen. Photographs by the author.

KOK P.J.R., Two new Pristimantis from Pantepui

Finger III 1.8 times wider that the distal end of the adjaeent phalanx. Palmar tuberele large, well defined,

fully pigmented, deeply bifid; thenar tuberele large, protuberant, ovoid; supernumerary tubereles few,

of variable sizes, slightly protuberant; subartieular tubereles large, round and protuberant, one eaeh on

FI and FII, two eaeh on Fill and FIV. Ulnar tubereles absent; presenee of a small antebraehial tuberele.

Hind limbs moderate in length, heels slightly overlap when held at right angles to sagittal plane; TIL

50.2% of SVL; FL 40.6% of SVL. Relative length of Toes IV > V > III > II > I; tip of Toe V extends

to the distal edge of the distal subartieular tuberele on Toe IV; tip of Toe III extends to the distal

edge of the penultimate subartieular tuberele on Toe IV. Lateral fringes on all toes, best developed

preaxially on Toes III-IV (Fig. 9); webbing basal between Toe IV-V. Toe dises slightly larger in size

than finger dises, WTDAVFD = 1.08; toe dises expanded, elliptieal, broader than long, eireumferential

groove eonspieuous, distal edge of dise rounded. Inner metatarsal tuberele elongate, oval, about four

times the size of the projeeting, round outer metatarsal tuberele; subartieular tubereles round, large and

protuberant; supernumerary plantar tubereles small, low and round, inereasing in size distally. Single,

ineonspieuous, very small, round ealear tuberele; no outer tarsal tubereles deteetable; inner tarsal fold

not deteetable.

Colour of holotype in life (see Fig. 8)

Dorsal baekground eolouration vivid yellow, top of head greyish brown; ehoeolate brown middorsal

stripe and ehoeolate brown band on fianks; ill-defined, irregular reddish orange dorsolateral line; a

few reddish orange blotehes on the posterior part of the dorsum; a few reddish orange minute speekles

on fianks; brown faeemask; enlarged tuberele on eyelid whitish. Lower fianks yellow; groins, anterior

thighs, and armpits brown. Arms and legs greyish brown with a few ill-defined white and dark yellow

blotehes. Ventral surfaees translueent greyish brown with a few irregular white spots and fieeks; posterior

surfaee of thighs and eloaeal area brown. Upper faee of hands and feet greyish brown with a few white

and yellow markings; upper faee of tip of fingers white; upper faee of tip of Toes I-III white, brown on

Toes IV-V. Palms, soles, undersurfaee of fingers and toes brown; dises greyish brown. Iris silver with

dark brown venation and an ill-defined broad horizontal brownish stripe.

Colour of Holotype in preservative (see Fig. 10)

After 13 months in 70% ethanol (July 2013). Vivid yellow faded to pale yellow. Choeolate brown and

brown faded to light brown; top of head is dark brown; ill-defined white and dark yellow blotehes on

arms and legs are slightly more eonspieuous and form irregular, ill-defined transverse bands; dorsal

pattern turned generally more eonspieuous. Ventral eolouration is light grey with a few white spots and

fieeks. Granules in the eloaeal area, as well as subartieular and supernumerary tubereles beeame dark

brown. Dises are grey.

Distribution and ecology

Pristimantis imthurni sp. nov. is only known from the summit of Ptari-tepui (Figs IB, 6B, 11), Bolivar

state, Venezuela, where it oeeurs at 2471 m elevation. Ptari-tepui lies in the Ptari massif, in the Los

Testigos subdistriet (MeDiannid & Donnelly 2005). It is loeated ca. 41 km NE of the Chimanta massif,

and ca. 22 km SE of Kamarkawarai-tepui (airline). Aeeording to the GPS, Ptari-tepui reaehes a maximal

elevation of about 2471 m above sea level. The summit area of Ptari-tepui is ca. 1 km^ and is dominated

by fiat open roek surfaees and small islands of low meadow-like tepui vegetation (Huber 1995; Fig.

IIB).

The male holotype - and single speeimen eolleeted - was found at night while ealling partly eoneealed in

the muddy soil in a small vegetation pateh. A few other males were heard ealling very sporadieally from

other small vegetation islands by day or night, but eould not be loeated. Pristimantis imthurni sp. nov.

European Journal of Taxonomy 60: 1-24 (2013)

Fig. 11. A. Ptari-tepui, photographed from the surrounding savannah looking to the NE (21 Jun. 2012).

B. Summit of Ptari-tepui (21 Jun. 2012). Photographs by the author.

KOK P.J.R., Two new Pristimantis from Pantepui

is not abundant. The only other anuran reported from the summit of Ptari-tepui is Tepuihyla rimarum

Ayarzagtiena, Senaris & Gorzula, 1993, whieh apparently also oeeurs in low density (pers. obs.).

Discussion

AXthou^ Pristimantis imthurni sp. nov. andP jamescameroni sp. nov. are not the geographieally elosest

tepui summit speeies (both new speeies oeeur on very isolated tepui summits separated from eaeh other

by a distanee of ca. 50 km airline), they are reeovered as sister speeies in preliminary phylogenetie

analyses based on mtDNA (pers. obs., August 2012). Unfortunately, the laek of genetie data from

several tepui summit Pristimantis speeies (e.g., P auricarens, P. muchimuk) prevents any eoneluding

phylogenetie inferenees at this point. It should be noted that DNA sequenees of P. jamescameroni sp.

nov. have been used in previous moleeular phytogenies under the name ''Pristimantis sp” (Hedges et

al. 2008) and "Pristimantis sp Aprada” (Kok et al. 2012), both eonfirming the generie alloeation of the

speeies.

MeDiarmid & Donnelly (2005) listed no less than 20 undeseribed Pristimantis speeies from the Guiana

Shield highlands (under the genus name Eleutherodactylus, from sp. A to T). Sinee that publieation,

only five of these 20 speeies have been named: Pristimantis sp. C from Auyan-tepui was deseribed as P.

auricarens by Myers & Donnelly in 2008, R sp. E from Yuruani-tepui was deseribed as P. yuruaniensis by

Rodder & Jungfer in 2008, P. sp. G from Sarisarinama-tepui was deseribed as P. sarisarinama by Barrio-

Amoros & Brewer-Carias in 2008, P. sp. S from Guaiquinima-tepui was deseribed as P. guaiquinimensis

by Sehluter & Rodder in 2007, and P sp. T from Cerro Marahuaka was deseribed as P marahuaka by

Fuentes-Ramos & Barrio-Amoros in 2004 (thus slightly before the opus of MeDiarmid & Donnelly was

published). Three new speeies, not listed in MeDiarmid & Donnelly (2005), were reeently deseribed

from additional loealities (P. abakapa, P aureoventris, and P. muchimuk). The present paper deseribes

two of the still unnamed speeies listed in MeDiarmid & Donnelly (2005), P. sp. B and P. sp. H, still

leaving 13 speeies to be named from the Pantepui region. Ten of these speeies oeeur in the Amazonas

state of Venezuela (seven are reported from Cerro La Neblina, a huge massif loeated at the border with

Brazil), three are from tepuis in the Bolivar state of Venezuela (the deseriptions of two of them being in

progress, Barrio-Amoros, pers. eomm.).

The number of deseribed Pristimantis speeies oeeurring exelusively on tepui (and faunistieally related

granitie mountains) summits and upper slopes now reaehes eleven.

Although there is eurrently no evidenee of eontinuing deeline, given their apparent very restrieted

ranges {ca. 1 km^ fox Pristimantis imthurni sp. nov., less than 5 km^forE jamescameroni sp. nov.) and

the reported sensitivity of tepui eeosystems to global wanning (see Nogue et al. 2009), it is suggested

that Pristimantis imthurni sp. nov. and P. jamescameroni sp. nov. be elassified as “Endangered” (EN) in

aeeordanee with eriteria B1 a-b (iii) and B2 a-b (iii) of the lUCN Red List of Threatened Speeies (2001).

Acknowledgements

The partial finaneial support of the non-profit organisation “les Amis de ITnstitut Royal des Seienees

Naturelles” is aeknowledged. I thank B. Wilson (Atlanta, USA) for field eompanionship and C. Brewer-

Carias (Caraeas, Venezuela) for invaluable help with logisties in Venezuela and useful elarifieations

about some tepui names. I also thank B. Means (Coastal Plains Institute and Land Conservaney, Florida,

USA) for the gift of eomparative material, and the following persons who loaned speeimens from their

respeetive institutions or provided aeeess to eomparative museum speeimens and working spaee during

my visit: B. Clarke and C. MeCarthy (BMNH), R. Ernst (SNSD), and A. Sehluter (SMNS). The author

was granted a postdoetoral fellowship from the Ponds voor Wetensehappelijk Onderzoek Vlaanderen

(FW012A7614N).

European Journal of Taxonomy 60: 1-24 (2013)

References

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Manuscript received: 5 August 2013

Manuscript accepted: 27 September 2012

Published on: 22 October 2012

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT eonsortium: Museum National d’Histoire Naturelle, Paris, Franee; National Botanie Garden

of Belgium, Meise, Belgium; Royal Museum for Central Afriea, Tervuren, Belgium; Natural History

Museum, London, United Kingdom; Royal Belgian Institute of Natural Seienees, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.

Appendix: Additional material examined

Pristimantis abakapa'. VENEZUELA, Estado Bolivar, Abakapa-tepui, IRSNB 15868 (topotypie

speeimen).

Pristimantis aureoventris: GUYANA, Cuyuni-Mazaruni Distriet, Wei-Assipu-tepui, IRSNB 4152

(holotype), IRSNB 4153-54 (paratopotypes).

Pristimantis dendrobatoides: GUYANA, Potaro-Siparuni Distriet, Wokomung Massif, IRSNB 15918

(topotypie speeimen).

Pristimantis jester. GUYANA, Cuyuni-Mazaruni Distriet, slopes of Maringma-tepui, IRSNB 15867,

IRSNB 15916-17.

Pristimantis marmoratus: GUYANA, Cuyuni-Mazaruni Distriet, foot of Mount Roraima, BMNH

1947.2.16.92 (formerly 99.3.25.19) (holotype).

Pristimantispulvinatus: VENEZUEEA, Estado Bolivar, Ea Esealera, IRSNB 15654.

Pristimantis saltissimus: GUYANA, Potaro-Siparuni Distriet, Wokomung Massif, IRSNB 15644-53

(topotypie speeimens).

Pristimantis yuruaniensis: VENEZUEEA, Estado Bolivar, Yuruani Tepui, IRSNB 15638-41 (topotypie

speeimens), SMNS 09855 (paratype).