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ALYTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

May 2009 Volume 27, N° 1

Alytes, 2009, 27 (1): 1-12.

Ingerana borealis (Annandale, 1912):

a new record from Mizoram (India),

with notes on its systematic position

and natural history

Saipari SAILO”, H. T. LALRE MSANGA .R. N. K. HO0RO0",

LALROTLUANGA"”" & A. OHLER

<spsailo@yahoo.co.in>

y. Aizawl 796012, Mizoram, India

*** Reptiles et Amphibiens, DSE, MNHN, CNRS, 25 rue Cuvier, 75005 Paris, France

<ohler@mnhn.fr>

** Department of Zoology, Mizoram Univ

A study on the distribution of Ingerana borealis was carried out in

Mizoram. Based on morphological comparison with type species of the

genera Ingerana, Occidozvga and Phrynoglossus and data on behaviour,

Micrixalus borealis is allocated to the genus Ingerana Dubois, 1987. The

study also reveals its occurrence up to 1000 m asl, previously unrecorded

for this species. The species lives in lotic habitat and is well adapted not only

to the slow-moving waters, but also to the fast flowing permanent streams

in and near forests. The holophoront of Micrixalus borealis is redescribed

and recently collected specimens from Mizoram allocated to this species are

described.

INTRODUCTION

ANNANDALE (1912) described Micrixalus borealis from the collections of the Abor

Expedition based on 11 specimens. It was only Dumois (1987) who reconsidered generic

allocation of this species using the character states given in original description. He allocated

the species to Phrynoglossus Peters, 1867 as it shares with this genus the absence of vomerine

tecth, the enlarged, rounded finger and toe tips without grooves and the indistinct tympanum.

Morphological data and recent phylogenetic studies showed that Phrynoglossus and Occido-

zyga Kuhl& Van Hasselt, 1822 are members of a monophyletic elade (MARMAYOU et al., 2000:

Bibliothèque Centrale Muséum

ILES

3 3001 00327664 8 Source : MNHN, Paris

ALYTES 27 (1)

KosucH et al., 2001) transcribed by Fer et al. (1990) and Duois (2005) as the subfamily

Occinozvensr Fei, Ye & Huang, 1990, and by others (INGER, 1993; Frosr et al., 2006) by

synonymisation of Phrynoglossus with Occidozyga. Thus, Micrixalus borealis was listed as

Occidozyga borealis by FRosr et al. (2006). The OccipozrGNar might include further ranoid

groups as molecular data (Bossuyr et al., 2006) indicate close relationship of Occidozyga lima

(Gravenhorst, 1829), ©. laevis (Günther, 1859) and /ngerana tenasserimensis (Sclater, 1892).

In the last decade, more specimens of Micrixalus borealis from a large range including

Bangladesh, Bhutan, NE India and Myanmar have been collected and observed in the wild

and the onomatophore deposited in the collection of Zoological Survey of India, Kolkata,

has been studied. At present, M. borealis is known from several north-eastern states of India,

including Arunachal Pradesh, Meghalaya (MATTHEW & SEN, 2003) and Nagaland (Ao et al.,

2003). Outside India, it has also been recorded from Bhutan (DuBois, pers. comm.) and from

the Rangamati Hill district in Bangladesh and the Arkan state in Myanmar (FROST et al.,

2006). We report here its occurrence in the Mizoram state of north-eastern India.

Recent observation of new specimens and behavioral data from the field made generic

allocation to either Phrynoglossus or Occidozyga somehow doubtful. Therefore Micrixalus

borealis is compared to specimens of the genera Occidozyga, Phrynoglossus and Ingerana as

defined by their onomatophores and a new generic allocation is proposed. We provide a

description of these newly collected specimens, along with diagnostic characters for the

speci s distribution and ecological notes. For the sake of completene: redescription of

the onomatophore of Micrixalus borealis is given.

MATERIAL AND METHODS

Museum abbreviations. - BMNH, Natural History Museum, London, United Kingdom:

MNHN, Muséum National d'Histoire Naturelle, Paris, ZMB, Zoologisches Museum

Berlin, Berlin, Germany; ZSI, Zoological Survey of India, Kolkata, India.

Specimens studied. — M1ZORAM. Aizawl districr: MZU-ZOO A.108, AIS, A.II8, 3 ©,

Mizoram University Campus (23°43.08 1°N, 92°43.668°E; 915 m MZU-ZO0 A.36, A.38,

14,1 ®,Tamdil National Wetland (23°44.399°N, 92° 57.313°E; 745 m asl); MZU-ZOO A.43,

A.46, 1 4,1 %, Durlui (23°53.508°N, 92°29.056'E; 110 m asl). Kolusib district: MZU-ZOO

A.31,A.33-34, 1 4,2 ©, Tuitun stream (23°58.213-58.402°N, 92°41.055°-41.104°E; 300-325

m asl); MZU-ZOO A.122, 1 4, Zanlawntlang Reserved Forest (23°56.116°N, 92°43.465'E;

748 m asl).

Our specimens were collected and preserved in 8 % formalin. Ecological and morphome-

tic data were collected for allspecimens. Voucher specimens were catalogued in the Museum of

Zoology department, Mizoram University (MZU-ZOO). Selected morphometrie measure-

ments were made to the nearest 0.01 mm with dial calipers (Mitutoyo series No. 505-671).

Morphometric and meristie characters largely followed those of SENGUPTA (2008). Onomato-

phore description follows thestandards of OnLer & Dusois (2006) and BoRboLotetal. (2007).

Other specimens used for comparisons. — Ingerana borealis (Annandale, 191 1 16932,

holophoront, sex not investigated, Rotung (1300 ft = 400 m asl; 28°07°N, 95°12'E), South of

Source : MNHN, Paris

SAILO, LALREMSANGA, HOOROO, LALROTLUANGA & OHLER 3

Yembung, Arunachal Pradesh, India; MNHN 1994.6489-6492, 4 G, Deothang (1000 m asl:

226°53°N, 91°30°E), Samdrup Jongkhar District, Bhutan. /ngerana tenasserimensis (Sclater,

1892): ZSI 10429, symphoront, sex not investigated, Tenasserim expedition (near 12°04'N,

99°0l'E), Myanmar, MNHN 1989.0718-0719, Banthat Mts. (7°30°N, 99°40'E), Thailand.

Occidozyga lima (Gravenhorst, 1829): MNHN 1999.6112-6132, Luang Prabang (19°53°N,

102°08'E), Laos. Phrynoglossus martensii Peters, 1867: ZMB 5645, holophoront, young ©

Bangkok (13°34°N, 100°13°E), Thailand; MNHN 1997.5359-5369,9 d,2 ©, Ben En National

Park (19°35°N, 105°28'E), Thanh Hoa Province, Vietnam.

Measurements. — SVL, snout-vent length; SL, snout length; EN, distance from eye to nostril;

NS, distance from nostril to snout; TE, distance from tympanum to eye; IN, internarial dis-

tance; IOD, interorbital distance; TUE, distance between upper eyelids: UEW, upper eyelid

width; ED, eye diameter; HTYD, horizontal tympanic diameter; HL, head length; HW, head

width; HDN, head depth at nostril; MN, distance from back of mandible to nostril; MFE, dis-

tance from back of mandible to front of eye; MBE, distance from back of mandible to back of

eye; IFE, distance between front of eyes; IBE, distance between back of eyes; FLL, forelimb

length; HAL, hand length: F,, length of first finger; F, length of second finger: F,, length of

third finger; F,, length of fourth finger; HLL, hindlimb length; TL, shank length; TW, shank

width; FL, thigh length; T,, length of first toe; T, length of second toe; T,, length of third toe;

T,, length of fourth toe; T,;, length of fifth toe; IMT, length of inner metatarsal tubercle; F,D,

width of first finger disc; F,D, width of third finger disc; T,D, width of first toe disc; T,D, width

of fourth toe disc; MTTF, distance from distal edge of metatarsal tubercle to maximum incur-

vation of web between third and fourth toe; TFTF, distance from maximum incurvation of web

between third and fourth toe to tip of fourth toe; MTFF, distance from distal edge of metatar-

sal tubercle to maximum incurvation of web between fourth and fifth toe; FFTF, distance from

maximum incurvation of web between fourth and fifth toe to tip of fourth toe.

GENERIC ALLOCATION

Comparison of morphological characters (tab. 1) of specimens allocated to Micrixalus

borealis and specimens of /ngerana, Occidozyga and Phrynoglossus present quite a complex

pattern of variation. In fact some character states, as the absence of vomerine teeth, apply to

all species observed and might be variable within genera, as /ngerana tasanae (Smith, 1921), a

larger species allocated to the genus /ngerana, has vomerine teeth (TAYLOR, 1962). Occido

lima can be distinguished by a series of character states that should be autapomorphies in

cladistic analysis, such as the presence of an outer metatarsal tubercle and of a tarsal tubercle,

the pointed tongue shape, the pointed tit-like subarticular tubercles, the complete webbing,

the pearly tipped tubercles on dorsal and ventral skin and the colour pattern on posterior

shank which shows continuous dark and light transve: tripes (Dusois & OHLER, 2001).

These character states cannot be observed in either /ngerana tenasserimensis, Phrynoglossus

martensii or Micrixalus borealis. Phrynoglossus martensii can be distinguished from /ngerana

by the length of first finger (longer than second), the absence of grooves on discs, the presence

of a metatarsal web, and the skin bearing glandular warts. It shares with Occidozyga lima the

presence of metatarsal web.

Source : MNHN, Paris

4 ALYTES 27 (1)

Table 1. - Comparison of 22 characters, including data on behaviour, observed on specimens

of the type species of the gencra /ngerana, Occidozyga and Phrynoglossus with the

holophoront (holotype) of Micrixalus borcalis.

Genus Occidozyga Phrynoglossus Ingerana Micrixalus borealis

Type species or Rana lima Phrynoglossus Rana tenasserimensis | ZSI 16932

holophoront Gravenhorst, 1829 martensii Peter Sclater, 1892

| | 1867

| Vomerine tecth | Absent Absent Absent Absent

Tongue Pointed Rounded Notched Notched

Tympanum Indistinet Indistinet Distinct Indistinct

l'ympanic fold Poorly di Distinct Distinct Distinct

h of finger 1 Equal to 11 Longer than II Much shorter than 11 Much shorter than 11

Tips of fingers Pointed Rounded Dilated Dilated

| Grooves on digital Absent Absent Present Absent

dises

| Subarticular tuberles | Prominent, pointed Rounded Rather large, Mat, Large, distinct

| indistinet

Outer metatarsal Absent Absent Absent

tubercle

Tarsal tubercle Absent Absent Absent

Webbing Complete Large Small Moderate

Metatarsal web Present Present Absent Absent

Tarsal fold Long, distinct Short, distinct Short, indistinet Short distinet

Dorsal skin Pcarly tipped Scattered tubereles Fine net-like skin Fine net-like skin

tubercles or spines ridges ridges

| Ventral skin Pearly tipped Smooth Smooth Smooth

tubercles or spines

Lateral line system | Present Absent Absent Absent

Coloration Greyish brown with Greyish brown with Usually light Greyish brown,

darker pattern mid-dorsal darker dorsolateral stripes: darker pattern

forming two more or | band outlined by brown with darker including band

less distinct lighier lines spots and scattered between eyes and

dorsal S white blotches chevron: hter

dorsolateral lines

Posterior shank Continuous dark Whitish with a Brown, slightly

light transverse peppering of darkish | lighter mottled

Stripes pigment

Vocal sacs Internal vocal sac Internal vocal sacs | Not known

Nuptial spines Present Present Not known

ï |

Colour of egg | Brown and white Whitish Whitish

Amplexus | Axillary Lumbar Not observed Axillary

Habitat Ponds, paddy field | Ditches, marshes Streams, cascades | Streams, cascades

If we try to allocate Micrixalus borealis using these characters, it does not share tongue

shape, extension of web, presence of pearl-tipped tubercles nor colour pattern of posterior

shank with Occidozyga lima. does not share longer finger I, presence of metatarsal web and

glandular warts on skin with Phr) Micrixalus borealis has a notched

tongue, a finger 1 much shorter than finger IE, its shank shows not longitudinal pattern, its

roglossus martensii.

Source : MNHN, Paris

SAILO, LALREMSANGA, HOOROO, LALROTLUANGA & OHLER ÿ

metatarsal web is absent and its skin shows fine net-like skin ridges. It shares the absence of

metatarsal web and the presence of these particular skin ridges with /ngerana tenasserimensis.

Considering behavioral characters, Micrixalus borealis specimens were collected in lotic

habitat as were specimens of /ngerana tenasserimensis. Occidoz ga lima is often found calling

in ponds and paddy fields in open habitat. Phrynoglossus martensii is present in ditches and

marshes in open habitat or secondary forests. Phrynoglossus martensii was described from

Bangkok, its tadpole was first described from this town and it still can be found there

everywhere. The ecological niche of Micrixalus borealis clearly is distinct from that of

Phrynoglossus and Occidozyga, but corresponds to the niche of /ngerana. This supports

change of generic allocation.

On behalf of these morphological and behavioral characters, Micrixalus borealis should

be allocated to the genus /ngerana as Ingerana borealis. H is distinct from the two species

presently allocated to this genus by its smaller size, absence of grooves on tip of fingers and

toes and larger webbing.

SPECIFIC ALLOCATION

Comparison of the newly collected specimens with the onomatophore of Micrixalus

borealis (ZSI 16932; see below) confirms specific allocation. The specimens from Mizoram

correspond in general shape and size. They share the presence of a tooth like projection on

lower jaw, rounded finger tips, relatively small webbing. Some character states differ. In the

onomatophore, the tympanum is indistinct and no measurements have been taken, whereas in

the specimens from Mizoram it is not very distinct but measurements could be taken.

Differences in toe tip shape and presence of short groove may be consequences of poor

conservation conditions of the onomatophore. Nevertheless some of the differences might

indicate intraspecific variation over the range of /ngerana borealis. More comparative studies

are needed to test this hypothesis.

DiAGNosis

Ingerana borealis is characterized by a combination of the following characters. Body

and head: small-sized frog; head small and triangular; snout bluntly rounded, projecting in

profile; canthus rostralis obtuse; loreal region concave: width of upper eyelid less than

interorbital distance; tympanum rounded; supratympanic fold present: vomerine teeth

absent; apical projection at the lower jaw prominent, pointed: a white spot between eye and

upper jaw; à dark bar between eyes. Legs: fingers free: toes half webbed; metatarsal web

absent; larger subarticular tubercles on toes than on fingers: outer metatarsal tubercle absent;

inner metatarsal tubercle elongated, followed by tarsal fold; tips of digits dilated, rounded,

grooves absent. Skin and colour: dorsum and flanks with net-like ridges and with tiny

glandular warts: venter smooth, whitish yellow: dorsal colour grey or brown (in 8 % formalin)

with light dorsolateral band. Sex characters: calling male with a pair of indistinct gular

pouches on throat; females with 58-72 unpigmented eggs.

Source : MNHN, Paris

Table 2. - Measurements and some natural history parameters of /ngerana borealis from Mizoram, India.

number AS A3 A A36 AS A4 A46 A.108 AUS AUS A2

Sex Female Male Female Female Male Female Female Female Female Male

Locality | Tuitun Tüitun Tuitun FTamdil National!" Durlui Durlui | MZU Campus | MZU Campus MEU %, Faslavotiang

| Wetland Campus

SE | 219 ET] 47 19.96 EE 2808

SL 700 330 ‘4 300 EN 361 385

EN 1 162 1.60 ï 1.90 168 147

NS 1 1.60 1 ES 132 200 Ls0 155

TE 9 094 n 074 08 10 LS 135

IN 2 267 2 268 20 275 298 316

10D Z 330 z 29 54 8 207

UEW 2 30 ï Er 16 338 23 506

ED z 35 3 306 296 Fe 336 ER

HIYD ï 216 1 210 190 24 230 218

HL 600 6 650 68 774 #40 775

HW ë 7.96 7 Reel 96 sé 906 93

HDN | "> 28 z 293 248 288 200

MN 3 507 5 527 4 53 540 53

MFE Ie: 37 as 430 ; 415 405

MBE 2 200 26s 258 220 248

IFE ; 303 39 El 360 300

IBE (BR 596 60 98 664 69

FLL De 129 1242 13.70 15.06 1514

Fi |. 3% 408 cer] 426 360 ca

F 30 480 394 48 5

5 520 59 546 650 7

F4 458 510 330 se 546 614

FD 05 073 065 ü65 07 083

FLD 054 086 080 001 066 0.90

HLL 3340 3618 3406 3510 4120 4428

TL 1158 DT 1147 13:00 lis 1407

TK 38 367 390 4.60 468 455

Tl 430 507 3 6 540

Le 586 &ss 6 3 720

Le 840 928 286 1048

1 1072 184 il 12. 1521

T$ 74 9.10 85 557 102

TD 078 108 086 102

TD 084 100 1:00 107

IMT 12 136 180 138

Tibio-tarsal | Anterior | Posterior | Middle Middle Anterior Between eyes] Between eyes Anterior Anterior | Anterior

isopteran | Isoptera, nymphs (2) | Hymenopiera wings |worker. winged| (2). isopteran nymph.

nymph Diptera Isoptera (| “small Myrmicidae

Norofeges | 68 = G = = = 5 G G =

Size ofeggs [143-220mm] = [1H224mm = = [425 mm 12026 mm) 138278mm [I25288mm) -

(D) LT SALATV

Source : MNHN, Paris

SAILO, LALREMSANGA, HOOROO, LALROTLUANGA & OHLER di

DESCRIPTION OF HOLOPHORONT OF MICRIXALUS BOREALIS ANNANDALE, 1912

ZSI 16932, sex not studied. Rotung (1300 ft = 400 m), South of Yembung, Arunachal

Pradesh, India.

(A) Size and general aspect. (1) Specimen of small size (SVL 20.4 mm), body stout.

(8) Head. — (2) Head moderate, wider (HW 8.4 mm) than long (HL 7.4 mm; MN 6.6 mm;

MFE 5.6 mm; MBE 3.0 mm), convex. (3) Snout rounded, not protruding; its length

(SL 3.18 mm) longer than horizontal diameter of eye (EL 2.87 mm). (4) Canthus rostralis

indistinct, loreal region concave, obtuse in cross section. (5) Interorbital space flat, less broad

QUE 1.83 mm) than upper eyelid (UEW 1.67 mm) but narrower than internarial distance

(IN 2.49 mm); distance between front of eyes (IFE 3.8 mm) about three fourth of distance

between back of eyes (IBE 6.2 mm). (6) Nostrils rounded, without flap of skin laterally,

closer to eye (EN 1.17 mm) than to tip of snout (NS 1.86 mm). (7) Pupil oval, horizontal.

(8) Tympanum indistinct. (9) Pineal ocellus absent. (10) Vomerine ridge and teeth absent.

(11) Tongue moderate, rounded, emarginate; median lingual process absent. À prominent

unique tooth-like projection medially on maxilla. (12) Supratympanic fold poorly distinct,

from eye to level of lips.

5.6 mm), not enlarged. (14) Fingers short, thin (F; 2.47 mm). (15) Relative length of fingers,

shortest to longest: F, < F, < F, < F;. (16) Tips of fingers I to IV rounded, not enlarged.

(17) Finger II with dermal fringe; webbing absent. (18) Subarticular tubercles indistinct,

rounded, single, tubercles of finger IV absent. (19) Prepollex indistinct; palmar tubercles

indistinct; supernumerary tubercles absent.

(D) Hindlimbs. — (20) Shanks three times longer (TL 11.8 mm) than wide (TW 3.6 mm),

longer than thigh (FL 10.8 mm) and distance from base of internal metatarsal tubercle to tip

of toc IV (FOL 10.9 mm). (21) Toes long, thin, toe IV (T, 7.40 mm) longer than third of

distance from base of tarsus to tip of toe IV (TFOL 15.7 mm). (22) Relative length of toes,

shortest to longest: T, < T, < T; < T; < T,. (23) Tips of toes rounded, slightly enlarged.

(24) Webbing moderate: [0-1 110-21112-21V 3-1 V(MTTF 6.84 mm; MTFF 6.36 mm:

FTET 3.66 mm; FFTF 4.77 mm). (25) Dermal ridge along toe V absent. (26) Subarticular

tubercles indistinct, oval, simple, all present. (27) Inner metatarsal tubercle indistinct, short,

its length (IMT 1.36 mm) 1.83 times in length of toe I (T, 2.49 mm). (28) Tarsal fold absent.

(29) Outer metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent.

(E) Skin. —(30) Dorsal and lateral parts of head and body: snout smooth: between eyes, side

of head, back and flank roughly shagreened. (31) Cephalic ridges absent. (32) Dorsolateral

folds absent; “Fejervaryan” line absent; lateral line system absent. (33) Dorsal parts of limbs:

forelimb shagreened; thigh and leg roughly shagreened; tarsus smooth. (34) Ventral parts of

head, body and limbs: throat with indistinct glandular warts; chest and upper belly smooth:

posterior part of belly smooth. (35) Macroglands absent.

(F) Coloration in alcohol. - (36) Dorsal and lateral parts of head and body: dorsal parts of

head and body brown with a dark band between eyes and a dark chevron between shoulders:

Source : MNHN, Paris

8 ALYTES 27 (1)

flank brown with darker spots: loreal region and upper lip brown with ivory white spots:

tympanic region brown. (37) Dorsal parts of limbs: brown with indistinct darker brown

bands: posterior part of thigh brown. (38) Ventral parts of head, body and limbs: throat and

its margin dirty white covered with brown shade:; chest ivory-white with brown spots on sides,

belly dirty white; thigh dirty white with brown spots on border; webbing dirty white with few

brown spots.

(G) Sexual characters. — Not observed.

DESCRIPTION OF SPECIMENS FROM MIZORAM

Size. À small-sized frog (fig. 1), mature males SVL 19.96-23.95 mm (x = 4); gravid females

SVL 23.14-28.12 mm (n = 7).

Head. — Head shorter than broad, small triangular. Snout blunt in sl

projecting beyond mouth; canthus rostralis obtuse; loreal region concave: width at angle of

s around 2!/, times distance from tip of snout to anterior corner of eye. Nostril equidistant

from tip of snout and eye; length of snout slightly longer than diameter of eye; internarial

distance larger than interorbital distance; tympanum round, not distinct and nearly half of

eye diameter: supratympanic fold starting from posterior corner of eye and ending at level of

forelimb insertion. Upper jaw bearing teeth; vomerine ridge absent; choanae situated far

forward, partially covered by musculature. A tooth-like structure at apex of lower jaw,

prominent and pointed: tongue small, free and bifid.

ape, convex at tip

Forelimb. - Moderately strong, more than half of snout-vent length; fingers thin, small,

without webbing between them: first finger slightly shorter than second: no inner metacarpal

tubercle on hand. Fingers dilated into rounded discs; discs longer than broad; subarticular

tubercles of hand distinct and prominent, smaller than subarticular tubercles of feet. Relative

length of fingers, shortest to longest: F, <F,<F,<F;.

Hindlimb. — Hindlimbs stout: exceeding total body length; tibiotarsal articulation reaching

eyes. Heels slightly overlapping when hindlimbs folded at right angle to body. Tibia short, half

of snout-vent length; its width about !/, of its length. Toes slender; an elongated inner

metatarsal tubercle present at base of first toe, followed by tarsal fold; outer metatarsal

tubercle absent. Relative length of toes, shortest to longest: T, <T,<T;<T;,<T,. Dises of

toes small, bearing a narrow median groove. Subarticular tubercles moderately prominent.

Webbing small, not quite reaching discs of first and fifth toes. Webbing moderate: 1 1-1!/,

I 1-1, D 1-21, IV 3-1 V.

Skin. — Dorsum as well as flank region rough with numerous tiny glandular warts forming

net-like ridges; skin of snout, between eyes, upper eyelids and lateral side of head above the

supratympanic fold uniformly granular, below fold smooth; throat and chest smooth to

shagreened; belly and inner part of limbs smooth but outer limbs with minute warts;

glandular warts on the outer shank larger than those of outer thigh: skin around anus

shagreened to granular; anus unmodified, directed posteriorly, at upper level of thighs.

Source : MNHN, Paris

SAILO, LALREMSANGA, HOOROO, LALROTLUANGA & OHLER 9)

Fig. 1. /ngerana borealis (MZU-ZOO A.46), SVL 24.35 mm, female, at Durlui (Mizoram: 23°53.508°N,

92°29.056E; 110 m asl).

Colour. Dorsal colour varying from light grey to light brown spotted with dark grey to dark

brown; pupil rounded and bluish-black:; flanks darker, throat and chin tinted with light grey:

belly white or slightly light yellow without any markings: inner limbs including hands and feet

light yellow; limbs and digits speckled with faint dark bands. A faint black line running from

throat region to level of groin ventro-medially. À dark brown canthal stripe from snout to eye;

one prominent white spot present between eye and upper jaw. A dark bar present at posterior

region of interorbital space, snout darker. À prominent light yellow dorsolateral stripe

extending from posterior corner of eye to level above insertion of hindlimb, demarcating

lighter grey or light brown dorsum from darker flanks. Webbing whitish yellow.

Sexual dimorphism. — (a) Males: comparatively smaller in size; a pair of indistinct gular

pouches present on throat of calling male. (b) Female: comparatively larger in size; wider

abdomen during breeding season.

Distribution. — ANNANDALE (1912) collected the type specimens of Micrixalus borealis from

close to Mouling National Park (Arunachal Pradesh) and after nine decades PaWAR &

BiRAND (2001) rediscovered the species in Pakhui Wildlife Sanctuary and Nameri National

Park. MATTHEW & SEN (2003) reported it from Baghmara Reserve Forest, South Garo Hills

and Narpuh Reserve Forest, Jainta Hills (Meghalaya). Ao et al. (2003) recorded it from

Source : MNHN, Paris

10 ALYTES 27 (1)

Medziphema (Nagaland) and AsMaAT et al. (2003) from Rangamati Hill district in Bangla-

desh. The present study recorded this species from the two districts of Mizoram, i.e., Aizawl

district and Kolasib district. In Aizawl district they were collected in Lawibual stream

(23°43.193/N, 92°44.328"E; 743 m asl), Mizoram University Campus (23°43.08l°N,

92°43.668'E; 915 m asl), Tamdil National Wetland (23°44.399°N, 92° 57.313'E; 745 m asl),

Durlui (23°53.508°N, 92°29.056'E; 110 m asl). In Kolasib district they were found in Tuitun

stream (23°58.213°-58.402°N, 92°41.055°-41.104°E; 300-325 m asl) and Zanlawntlang Re-

served Forest (23°56.116°N, 92°43.465'E; 748 m asl). The present records add not only new

data to the distributional information of the species in north-eastern India, but also to the

altitudinal distribution. It is notable that many of these specimens, except those collected in

Durlui and Tuitun stream, came from elevations above 740 m asl and up to 915 m asl; our

Bhutan specimens were collected at 1000 m asl, thus extending the elevational range much

above the 400 m asl given in the literature (STUART et al., 2008). This species is not rare in the

two districts of Mizoram where it could be easily observed during our study. Further

exploration will probably reveal that Z. borealis is present in other parts on this region, and its

range may be throughout the state.

Natural history. — Ingerana borealis is a widely distributed species in our current study sites,

found in and around lotic water sources mostly in canopy area, marshes and wetlands. During

our survey, most of the specimens were found at night among litter and boulder in trickling

and cascading streams in and around forests. Most of them were seen perching on the wet

rocks within the stream or on the streamside rock ledges, about 1-5 m away from the water

surface. In Mizoram, this frog can be observed from the onset of monsoon onwards. During

day time, the frogs remained inside the crevices of rock as well as under moist leaf litter and

bushes. Other amphibian species found in sympatry include Amolops marmoratus (Blyth,

1855), Fejervarya limnocharis group, Polypedates leucomystax group, Nasirana alticola (Bou-

lenger, 1882), Sylvirana cf. nicobarensis (Stoliczka, 1870), Kaloula pulchra Gray, 1831, Micro-

hyla berdmorei (Blyth, 1856), Leptobrachium smithi Matsui, Nabhitabhata & Panha, 1999 and

Xenophrys parva (Boulenger, 1893).

During the present study, /ngerana borealis was found from April to August, which

indicates that the species is a seasonal breeder. We observed that breeding in this species is

related to rainfall with the breeding peak taking place from May to July. The male calls from

outside water and amplexus is axillary. Frogs in amplexus are seen from around 22:30 h on the

margin of water, near the bank. No eggs and larvae of this species have been recorded.

Remarkably, in laboratory, maintained amplecting pairs did not lay eggs.

CHANDA (2002) reported that no data were available on the wild status of the species and

may warrant inclusion in the Indian Wildlife (Protection) Act, 1972. The new data published

here may fill this gap. As indicated from range extension and abundance in the field, this

species might be changed to LC (Least Concern) in IUCN categories.

ACKNOWLEDGEMENTS

This study is based on work supported by the UGC's Rajiv Gandhi National Fellowship granted to

SS. We thank the department of Zoology, Mizoram University, in particular Prof. G.C. Jagetia, head of

Source : MNHN, Paris

SAILO, LALREMSANGA, HOOROO, LALROTLUANGA & OHLER 11

department, for providing us with laboratory facilities. We also thank Freddie Lalrinawma and Jerry

Ramherliana for their outstanding field assistance. Shyamal Kumar Chanda, curator at the Zoological

Survey of India, Barry T. Clarke, curator at the Natural History Museum (United Kingdom), Rainer

Günther, former curator at the Zoologisches Museum Berlin (Germany), Alain Dubois and the staff of

the Reptiles and Amphibians Collection of the Muséum national d'Histoire naturelle (Paris, France) are

thanked for giving access to specimens. Stéphane Grosjean helped for the work in the Kolkota Museum.

Work in India for AO and SG was financed by the PPF “Etat et structure phylogénétique de la biodiversité

actuelle et fossile”.

LITERATURE CITED

ANNANDALE, N., 1912. -Zoological results of the Abor Expedition (1911-1912). I. Batrachia. Rec. Indian

Mus., 8 (1): 7-36.

A0, J. M., BORDOLOI, S. & OHLER, A., 2003. - Amphibian fauna of Nagaland with nineteen new records

from the state including five new records for India. Zoos’ Print Journal, 8 (6): 1117-1125.

ASMAT, G.S. M., BANU, M. Q., ISLAM, M. A., AHSAN, F. & CHAKMA, S., 2003. - Amphibian fauna from

Chittagong and Chittagong Hill-tracts, Bangladesh. Uni. J. Zool., Univ. Rajshahi, Bangladesh,

22: 141-143.

BoRDOLOI, S., BORTAMULI, T. & OuLer, A., 2007. - Systematics of the genus Rhacophorus (Amphibia,

identity of red-webbed forms and description of a new species from Assam. Zootuxa,

F., Hicuis, D. M., CANNATELLA, D. €. & MiLINKOVITCH, M. C., 2006. —

Phylogeny and biogeography of a cosmopolitan frog radiation: Late Cretaceous diversification

resulted in continent-scale endemism in the family Ranida

CHaNDA, S. K., 2002. — Indian Amphibians. Kolkata, Zoological Survey of India i-vii] + 1

Duois, A., 1987. - Miscellanea taxinomica batrachologica (1). Alytes, 1986”, 5 (1-2): 7-9

2005.— Amphibia Mundi. 1.1. An ergotaxonomy of recent amphibians. Alytes, 23 (1-2):

Dunois, A. & OHLER, A., 2001. — A new genus for an aquatic ranid (Amphibia, Anura) from Sri Lanka.

Alytes, 19 (2-4): 81-106.

Fe, L., VE, C. & HUANG, Y., 1990. - Key 10 Chinese Amphibia, Chongging, Editions of Sciences and

Techniques: [i-v] + 1-2 + 1-364. [In Chinese].

FRosT, D. R., GRANT, T., FAIVOVICH, J., BAZIN, R. H., Haas, A. HaDDAD, C. F. B., DE S4, R. O..

CHANNING, A., WILKINSON, M., DONNELLAN, S. C., RAXWORTHY, C. J., CAMPBELL, J. A., BLOTTO,

B. L., MOLER, P., DREWES, R. C., NUSSBAUM, R. A., LYNCH, J. D., GREEN, D. M. & WHEELER, W.

C., 2006. - The amphibian tree of life. Bull. amer. Mus. nat. Hist., 297: 1-370.

INGER, R. F., 1993. - Commentary on a proposed classification of the family Ranidac. Herpetologica, 52:

241-246.

KosucH, J., VENCES, M., DURoIS, A., OHLER, A. & BÔHME, W., 2001. - Out of Asia: mitochondrial DNA

evidence for an Oriental origin of tiger frogs, genus Hoplobatrachus. Mol. Phyl. Evol, 21 (3):

398-407.

MARMAYOU, J., DUBOIS, A., OHLER, A., PASQUET, E. & TILLIER, A., 2000. - Phylogenetic rc

the Ranidae (Amphibia, Anura): independent origin of direct development in the gen.

rana and Philautus. C. r. Acad. Sci, Paris, 323: 1-11.

MATTHEW, R. & SEN, N., 2003. - Rediscovery of Micrixalus borealis Annandale, 1912 (Ranidae: Anura:

Amphibia) from Meghalaya, India. Cobra, 53: 1

Our, A. & Durois, A., 2006. - Hyla reimvardtit Schlegel, 1840 as a nomen protectum. 4lyres, 23 (3-4):

123-132.

PAWAR, & BIRAND, A., 200

Centre for Ecological Res

SENGUPT

335.

— A survey of amphibians. reptiles, and birds in Northeast India. Mysore,

arch and Conservation, CERC Technical Report No. 6: 1-126

S., 2008. — Study of herpetological specimens for research. Baripada, Orissa, Abstracts of

C 1% School in Herpetology, 26.12.2007 to 8.1.2008, North Orissa University.

Source : MNHN, Paris

12 ALYTES 27 (1)

STUART, S., HOFFMANN, M., CHANSON, J. S$., COX, N. A., BERRIDGE, R. J., RAMANI, P. & YOUNG, B. E.,

(ed.), 2008. — Threatened amphibians of the world. Lynx Edicions (Spain), IUCN (Switzerland) &

Conservation International (USA): i-xv + 1-758.

TAYLOR, E. H., 1962. - The amphibian fauna of Thailand. Univ. Kansas Sci. Bull., 63: 265-599.

Corresponding editor: Alain Dunois.

Source : MNHN, Paris

Alytes, 2009, 27 (1): 13-24. 13

Fejervarva triora (Amphibia, Ranidae):

first description of the adult male and

recent distribution records

Yodchaiy CHUAYNKERN* **, Nakorn SALANGSINGHA***, Sunchai MAKCHAI**,

Chantip INTHARA* **** & Prateep DUENGKAE***

* Muséum national d'Histoire naturelle, Département de Systématique et

Evolution, UMR 7205 CNRS OSEB, Reptiles et Amphibiens,

25 rue Cuvier, CP 30, 75005 Paris, France

<ychuaynkern@yahoo.com>

** National Science Museum, Thailand Natural History Museum, Technopolis,

Khlong 5, Khlong Luang, Pathum Thani 12120, Thailand

**# Kasetsart University, Faculty of Forestry, Department of F

Khen, Bangkok 10900, Thailand

rest Biology, Bang

#### Khon Kaen University, Faculty of Science, Department of Biology, Muang.

Khon Kaëen 40002, Thailand

We provide the first description of the adult male of Fejervarya triora

Stuart, Chuaynkern, Chan-ard & Inger, 2006, based on a single specimen

collected from Ubon Ratchathani Province, northeastern Thailand. Addition-

al records of its distribution, based on voucher specimens, are given and a

recent distribution map is provided. Our specimens represent new provin-

cial records.

INTRODUCTION

The frog species Fejervarya triora was recently described from Ubon Ratchathani Prov-

ince (type locality: “Phu Jong-Na Yoi National Park”, Na Chaloey District, northeastern

Thailand), based on a series of adult females and unsexed juvenile specimens (STUART et al.,

2006). The holotype description was made from an adult female (FMNH 266172/THNHM

05325). Previously, even though CHAN-ARD (2003) provided a short description of the species

in Thai as well as a life photo and a distribution map in his field guide, no information on male

specimens was published. In general, local people from places surrounding the type locality

are familiar with both sexes of this frog as they use it for consumption.

During a herpetological survey in 2005, we conducted field trips in several areas of

Thailand. We thus obtained twenty-two specimens which we identified as Fejervarva triora.

The frogs show external morphological characters similar to the original description given by

STUART et al. (2006). This series of specimens included an adult male, which allows us to

provide here the first description of an adult male of this species, accompanied by a recent

distribution map.

Source : MNHN, Paris

14 ALYTES 27 (1)

MATERIAL AND METHODS

The specimens were caught in the field by hand, preserved in 10 % buffer formalin, and

later transferred to 70 % ethanol. Before the specimens were fixed in formalin, tissue samples

were taken by preserving pieces of liver in 95 % ethanol. The specimens in this study are

catalogued and deposited in the collection of the Thailand Natural History Museum

(THNHM), Pathum Thani, Thailand. The description format was based on the works of

Ohler (e.g., OHLER, 1996; OuLer & Dumois, 1999; OuLer et al., 2000, 2002). Data concerning

the type series of the species were obtained by YC. The criterion used for determination of the

sex was the presence of vocal sac openings (HEYER, 2005). Webbing formula is given according

to MYERs & DUELLMAN (1982). The illustration of nuptial pad morphology was made by CI

using a Leica MSS stercomicroscope with a camera lucida attachment, at the Laboratoire des

Reptiles et Amphibiens, Muséum national d'Histoire naturelle (MNHN), Paris, France.

Measurements were made with digital calipers to the nearest 0.1 mm. Abbreviations used

for measurements are:

SVL, snout vent length.

Head: HW, head width; HL, head length (from back of mandible to tip of snout); MN, distance from

back of mandible to nostril, M ance from back of mandible to front of eye; MBE, distance from

back of mandible to back of ey 3, distance between fronts of eyes: IBE, distance between backs of

eyes: IN, internarial sp: EN, distance from front of eye to nostril; EL, eye length: SN, distance from

nostril to tip of snout; tance from front of eye to tip of sno: YD, greatest tympanum diameter;

TYE, distance from tympanum to back of eye; TUE, minimum distance between upper eyelids; UEW,

maximum width of inter upper eyelid.

Forearm: HAL, hand length (from base of outer palmar tubercle to tip of third finger); FLL, forelimb

length (from elbow to base of outer palmar tubercle): TFL, third finger length (from base of first

subarticular tubercle); fd1-fd4, width of pads of fingers 1 to 4; fwl-fw4, width of fingers 1 to 4.

Hindlimb: FL, femur length (from vent to knee): TL, tibia length; FOL, foot length (from base of inner

metatarsal tubercle to tip of fourth toe); FTL, fourth toe length (from base of first subarticular tubercle);

td1-1d5, width of pads of toes ! to 5; twl to tw5, width of toes 1 to 5; IMT, length of inner metatarsal

tubercle: ITL, inner toe length.

Webbing: MTTE, distance from distal edge of metatarsal tubercle to maximum incurvation of web

between third and fourth toe; TFTF, distance from maximum incurvation of web between third and

fourth toe to tip of fourth toe; MTFF, distance from distal edge of metatarsal tubercle to maximum

incurvation of web between fourth and fifth toe; FFTF, distance from maximum incurvation of web

between fourth and fifth toe to tip of fourth toe; WTF, webbing between third and fourth toe (from base

of first subarticular tubercl webbing between fourth and fifth toe (from base of first subarticular

tubercle); WI, webbing between third and fourth toe when folded along fourth toe (from base of first

subarticular tubercle); WIT, webbing between fourth and fifth toe when folded along fourth toc (from base

of first subarticular tubercle).

Source : MNHN, Paris

CHUAYNKERN, SALANGSINGHA, MAKCHAI, INTHARA & DUENGKAE 15

RESULTS

Fejervarya triora Stuart, Chuaynkern, Chan-ard & Inger, 2006

(fig. 1-3)

arya Sp.: CHAN-ARD, 2003: 110.

Fejervarya triora Stuart, Chuaynkern, Chan-ard & Inger, 2006: 11.

Material examined. - THNHM 09052-65, four adult females and eight juveniles, collected by

N. Salangsingha and S. Makchai between 14 and 21 September 200$ at Mukdahan National

Park, Muang District, Mukdahan Province, Thailand; THNHM 09069-76, an adult male and

an adult female, six juveniles, collected by N. Salangsingha and S. Makchai between 23 and 25

September 2005 at Pha Tam National Park, Khong Chiam District, Ubon Ratchathani

Province, Thailand.

Comparative material. — See STUART et al. (2006).

Description of male specimen. - THNHM 09074 (field number YC 0117), an adult male, Pha

Tam National Park, Khong Chiam District, Ubon Ratchathani Province, northeastern

Thailand. The specimen was collected by N. Salangsingha and S. Makchai on 25 September

2005. AI measurements below are in millimetres.

(A) Size and general aspect. — (1) Frog of moderate size (SVL 45.3), body slender.

(B) Head. — (2) Head of moderate size, about as broad as long (HW 17.7; HL 17.6; MN 15.5;

MFE 12.5; MBE 7.3), flat above. (3) Snout obtusely pointed in dorsal view, rounded in lateral

view, slightly projecting beyond lower jaw, its length (SL 7.4) longer than horizontal diameter

of eye (EL 6.0). (4) Canthus rostralis rounded, loreal region obtuse in cross section. (5)

Interorbital space flat, narrower (TUE 2.2) than upper eyelid (UEW 4.6) and than internarial

distance (IN 3.4); distance between fronts of eyes (IFE 7.9) 1.6 times in distance between

backs of eyes (IBE 12.4). (6) Nostrils rounded with flap of skin laterally, closer to tip of snout

{NS 2.8) than to eye (EN 4.2). (7) Pupil not observed in this specimen. (8) Tympanum (TYD

4.1) distinct, rounded, 69 % of eye diameter; tympanum-eye distance (TYE 1.6) 39 % of

tympanum diameter. (9) Pineal ocellus absent. (10) Vomerine ridge present, bearing few small

teeth (7 = 4), with an angle of 45° to body axis, less cl ch other, longer

than distance between them. (11) Tongue large, oval, emarginated, bearing no median lingual

process; tooth-like projections on lower jaw absent.

9.5), not enlarged. (13) Fingers L, IT and IV short and thin: finger [I long and thin (TFL 5.6)

(14) Relative length of fingers: I < IV << HI. (15) Tips of fingers rounded, not enlarged,

without grooves, narrow compared to finger width (fd1 0.9, fw1 0.9; fd2 0.9, fw2 1.0: AA

fw3 0.9; fd4 0.7, fw4 0.9). (16) Dermal fringe on inner si IT and IT indistin

webbing on fingers absent. (17) Subarticular tubercles strongly prominent, oval, single, all

present. (18) Prepollex distinct, oval; palmar tubercles not separated into inner and outer

metacarpal tubercles: no supernumerary tubercle.

e 10 choanae than to e

Source : MNHN, Paris

16 ALYTES 27 (1)

Fig. L.— Adult male specimen of Fejervarya triora (THNHM 09074, SVL 45.3 mm.). Left, dorsal view:

right, ventral view: scale bar, 10 mm.

(D) Hind limbs. — (19) Hind limbs moderately long, heels overlapping when limbs are folded

at right angles to body: tibia 3.3 times longer (TL 21.3) than wide (TW 6.6), longer than thigh

(FL 20.7) and longer than distance from base of internal metatarsal tubercle to tip of toe IV

(FOL 21.0). (20) Toes long and thin, toe IV (FTL 12.3) longer than one third of distance from

base of tarsus to tip of toe IV (TFOL 30.2). (21) Relative length of toes: I < IT < V < III < IV.

(22) Tips of all toes rounded, not enlarged:; di bsent on toes I-V, without grooves, narrow

compared to toe width (td1 0.7, tw1 0.7; td2 0.6, tw2 0.7; td3 0.6, tw3 0.7; td4 0.7, tw4 0.7; td5

0.5, tw5 0.6). (23) Webbing present, rudimentary: 1 1-2" 111-2"%1II11-31V3-1 V(WTF

6, WFF 3.1; WI 3.7, WI 3.1; MTTF 9.5, MTFF 9.3, TFTF 8.4, FFTF 9.6). (24) Dermal

ridge along toe V present from tip of toe to distal outer metatarsal tubercle, well developed.

(25) Subarticular tubercles strongly prominent, oval, simple, all present. (26) Inner metatarsal

tubercle distinct, elongated: its length (IMT 3.1) 1.4 times in length of toe I (ITL 4.2). (27)

Ta fold absent. (28) Outer metatarsal tubercle present, rounded; supernumerary tubercles

and tarsal tubercle absent.

(E) Skin. — (29) Snout and skin between eyes smooth, side of head smooth with few horny

spinules on the area of upper lip. Anterior part of back smooth, posterior part shagreened

with horny spinules. Upper part of flank, from line from insertion of arm to groin, granular

with horny spinules, lower part of flank granular. (30) Dorso-lateral folds absent; supratym-

panic folds present and strong, from posterior edge of eyelids to shoulders; parotoid glands

Source : MNHN, Paris

CHUAYNKERN, SALANGSINGHA, MAKCHAI, INTHARA & DUENGKAE 17

Fig. 2. - The external vocal sac (wrinkled skin pointed to by arrow) of male Fejervarya triora (THNHM

09074).

absent; cephalic ridges absent; co-ossified skin absent; Fejervaryan line indistinct. (31) Dorsal

parts of forelimb, leg and tarsus smooth with few horny spinules; dorsal part of thigh with

feeble glandular folds bearing few horny spinules. (32) Throat, chest, belly and anterior

ventral part of thighs smooth; posterior ventral part of thigh with tree-frog belly skin. (33)

Macroglands absent.

(F) Coloration (in alcohol). — (34) Dorsal parts of head and dorsum dark grey with dark

brown marbling, “V” shape band extending from edge of eyelid to each other; flank dark

brown with light marbling; loreal region and tympanic region dark grey; upper lip dark grey

with four feeble dark brown bands. (35) Dorsal parts of forelimb dark grey with dark brown

crossbars on forearm; dorsal parts of thigh, leg and foot dark grey with black crossba

Source : MNHN, Paris

18 ALYTES 27 (1)

Fig. 3.- Hand of male Fejervarya triora (THNHM 09074) showing nuptial spines on first finger in dorsal

(left, drawn from right hand) and dorsolateral (right, drawn from left hand) views. Scale bar,

30 mm.

posterior part of thigh dark brown with light vermiculations. (36) Throat and its margin dark

mottled on approximately 2/3 of throat; vocal sacs with dark mottling (fig. 1-2); chest and belly

greyish white with more mottlingin the area of breast and lateral sides of belly; webbing brown.

(G) Male secondary sexual characters - (37) Two pads in contact, oval shaped nuptial pad on

finger L, with small, translucent spines (fig. 3). (38) Vocal sacs present, distinct on throat as wrin-

kled skin beside corners of jaws (fig. 1-2); paired openings of vocal sacs distinct, slit-like, at

anteriorcorner of jaw. (39) Other secondary sexual characters absent.

Natural history notes. — Frogs were collected at night starting at 19h00 in deciduous forest. At

Mukdahan National Park, the substrate where the frogs were found is very large igneous rock

with several small hollows (containing water only in the rainy season), whereas at Pha Tam

National Park it is sandstone bedrock where erosion formed several small temporary hollows.

The frogs were found sitting on the ground, on dead leaves, grass, or sittingin the hollows. From

the observation of NS and SM, only the juvenile frogs (tentatively so defined by their small size)

were found in daytime but never adult frogs (tentatively so defined by their larger size). Adult

frogs could be observed when the sun went down. Individuals of very large size were found

more frequently at night when it did not rain, whereas adult male frogs (large size but smaller

than females and calling) were mostly found duringrainy nights.

Source : MNHN, Paris

CHUAYNKERN, SALANGSINGHA, MAKCHAI, INTHARA & DUENGKAE 19

Tab. 1.— Selective measurements in millimetres (mean + standard deviation, and min-max) of

adult Fejervarya triora from Phu Jong-Na Yoi National Park (10 adult females,

holotype and paratypes), Mukdahan National Park (4 adult females) and Pha Tam

National Park (an adult female and an adult male). Measurements are defined in the

Material and methods. Some measurements of the type series are not available,

indicated as n.a., because of use of different measurement methods by YC in 2005.

PRE | Mukdahan | PhaTam PRE | Mukdahan | PhaTam

females females [female] male fmales females [female] male

SVL | 582417 | 575424 | 556 | 45.3 | FLL | 141406 | 134200 | 133 | 95

(55.5-60.8) (13-15.1) (12.4-14.1)

Hw | 242413 219 [177 | TrL | 68203 | 72403 | 69 | 56

(28-272) (6.3-7.3) 7-7.)

HL 21.140.6 22.5 17.6 FL 28.5 + 0.9 25.5+1.1 25.6 20.7

(20.22.1) (78-303) | (246-269)

MN na. 196 | 15.5 | TL | 30221 | 283408 | 264 | 213

(183-203) 932.1) | (27.7-29.4)

ME | na. 144408 | 145 | 12.5 | roL | 29441 | 279408 | 267 | 21

(13.5-15.2) @7.5-31.1) | (26.9-28.8)

MBE | na. 88407 | 94 | 73 | rrL | 168206 | 16203 | 153 | 123

(8.1-9.7) (5.7-17.8) | (158-164)

IFE na 93404 | 93 | 79 |imr | 38402 | 38403 | 35 | 313

(9.1-9.8) (3.7-4.1) (3:5-4.1

IBE na. 148 40.6 138 124 ITL 63403 6.240.1 62 42

(14.1-15.5) (5.5-6.6)

IN | 44403 | 47402 | 43 | 34 | rw | 99408 19 | 66

(4-48) (44-49) (8.3-11.1)

EN | 5403 56402 | 53 | 42 |TroL| 41341 | 40:05 | 385 | 302

(4-55) | (53-58) (99-43) | (39.4-40.4)

EL | 67402 | 654202 | 65 | 6 [mrrr| na. 134202 | 126 | 95

(6.5-7) (6.3-6.8) (32-135)

NS na. 4.14#+04 33 28 |MTFF na. 13.24 0.3 12.9 9.3

G.7-4.6) (28-135)

SL 9+0.4 9.2+04 8.6 74 TFTF na. 10.4 40.7 LA 84

(8.3-9.4) (8.8-9.8) (94-111)

TYD 4.8 40.2 44403 44 4.1 FFTF na. 11.7+ 0.2 112 9.6

(4.3-5) G.1-4.7) QL5-11.9)

TYE 15+03 17402 17 16 WTF na. 6+09 5 3.6

(1:5-1.8) (5.1-7.1)

IUE na. 32403 | 33 | 22 | wrr na. 52404 | 48 | 31

C.8-3.5)

EUW na. 5.6+0.6 5 47 WI na. 52 3.7

(5-62)

HAL | 12404 | 121404 | 19 | os | w2 na. 46 | 31

GL5-12.7) | G1.7-127)

Comparisons. — Our single male specimen has morphological characters that agree with the

females described by SruaART et al. (2006) in having a very broad head and a broad supratym-

panic fold obscuring the dorso-posterior margin of tympanum. Yet, this specimen diflers

from these females in its smaller size (SVL 45.3 mm, vs. 55.5-60.8 mm in 10 adult females,

average 58.2 + 1.7). This appears to be sex dimorphism in size, known in other

Source : MNHN, Paris

20 ALYTES 27 (1)

species of Fejervarya (DUBois & OHLER, 2000; Veith et al., 2001). The pineal ocellus, visible in

the female specimens studied by SruART et al. (2006), is absent in our male specimen, so this

character shows intraspecific variation. The male specimen observed exhibits two palmar

tubercles in contact (not separated into inner and outer metacarpal tubercles), as mentioned

in the description of STUART et al. (2006). The difference might be due to preserving

preparation. The skin on top of the head in our male specimen seems to be smooth, versus

shagreened in the specimens described by SruaART et al. (2006). This also might be caused by

different conservation conditions. According to SrUART et al. (2006), the type series and other

specimens they examined are in good condition whereas our specimens are rather slightly stiff.

Due to the poor condition of our cimen, we could not observe the flap of skin on the

preaxial side of fingers IT and III which were described by SruART et al. (2006) in their females.

Adult female specimens from Mukdahan and Pha Tam national parks show dorsum pattern

{in preservative) variable in amount of blotches or spots, as noted by STUART et al. (2006).

Measurements data show that the adult females from those three localities are in the same

range (tab. 1). More specimens, especially adult males, are further required to test sexual

dimorphism in this species. Nevertheless, we consider that the differences observed result

either from sex dimorphism or from fixation and conservation conditions, and we assign this

Frog to Fejervarya triora.

Distribution. — THAILAND: UBON RATCHATHANI , Na Chaloey District, Phu Jong-Na Yoi

National Park (STUART et al., 2006); Khong Chiam District, Pha Tam National Park (this

study); MUKDAHAN PROVINCE, Muang District, Mukdahan National Park (this study).

Prior to this study, Fejervarya triora had only been known from the type locality. Our

specimens represent the first provincial record for Mukdahan Province and also an additional

provincial record for Ubon Ratchathani Province. The distribution map of this species is

shown in fig. 4. The species Æcjervarya triora is currently known only from Thailand, but

might be expected in Laos.

DISCUSSION

Recently, FROST et al. (2006) proposed a new taxonomy for living amphibians based on

combined anatomical and molecular data. The family Ranidae Rafinesque-Schmaltz, 1814

Was partitioned into eleven families to avoid paraphyly with regard to the families Rhacopho-

ridae and Mantellidae. Among them, the Dicroglossinae (sensu DuBois, 1992, 2005) were

elevated to family status. The genus Fejervarya Bolkay, 1915 was referred to this family. In this

paper we adopt a conservative attitude in using the family Ranidae in the traditional sense

(Dusois, 1992, 2005; Bossuyr et al., 2006; OnLEr & DuBois, 2006). Within this frame, the

genus Fejervarya is a member of the subfamily Dicroglossinae Anderson, 1871.

As a result of this study, the male of Fejervarya triora is shown to have secondary sex

characters similar to those indicated by DuBo)is et al. (2001), who gave diagnostic morpholog-

ical characters of nine genera of the subfamilies Dicroglossinae and Raninae of the family

Ranidae. For the genus Fejervarya Bolkay, 1915, they described the vocal sac in males as

“Marked by darker coloration, and sometimes also by longitudinal folds, on sides of throat”,

Source : MNHN, Paris

CHUAYNKERN, SALANGSINGHA, MAKCHAI, INTHARA & DUENGKAE 21

MES

Thailand

z FA

{

{ N

Cambodia ) 4

5 |

z $ Z

Pur

° 100 200 300

Kilometers

1OB"E

Fig. 4. Mapof the Indochinese peninsula showing localities of distribution of Fejervarva triora reported

in this study and in Sruarr et al. (2006). 1, Phu Jong-Na Yoi National Park (type locality), Na

Chaloey District, Ubon Ratchathani Province; 2, Pha Tam National Park, Khong Chiam District,

Ubon Ratchathani Province; 3, Mukdahan National Park, Muang District, Mukdahan Province

Source : MNHN, Paris

22 ALYTES 27 (1)

and the nuptial pads in male as “Present on prepollex and finger [”. Our adult male specimen

agrees well with this information (fig. 1-3).

The male specimen was caught in September while it was calling. In northeastern

Thailand, this month is within the period of rainy season. The type series and tadpoles studied

by STruaRT et al. (2006) had also been caught in September. These data suggest that Fejervarya

triora is mating during the rainy season, in contrast with Fejervarya limnocharis, whose

tadpoles can be found throughout the year (INTHARA et al., 2005).

In the areas of study, Fejervarya triora is common, as also noted by STUART et al. (2006).

This species was much more frequently encountered than the sympatric species Fejervarya

limnocharis. Both are consumed by local people. Specimens collected for food mainly came

from populations in rice fields surrounding areas protected by Thai law, such as a National

Park. The distribution area of Fejervarya triora might be much larger as currently known and

we suspect that this species might be found in areas of neighbouring countries, i.e., southern

Laos and northern Cambodia (see fig. 4). We need a better understanding of the natural

history and ecology of the species to ascertain its potential threats and managing its habitats

and populations. The problems of habitat loss or land use are probably among the main

causes for amphibian declines (COLLINS & STORFER, 2003). It is widely assumed that the

majority of amphibian populations’ declines throughout the world result from habitat

destruction, but this assumption is very difficult to test, because of the nature of many

amphibian habitats (HALLIDAY, 2005). Attention should also be paid to the effect of harvest-

ing on amphibian populations (VerrH et al., 2000) such as Fejervarya triora.

During the last decade, the herpetofauna of the Indochinese region has been given

significant attention. Several new species and additional records were published by various

authors (e.g. TEYNIÉ et al., 2004; STUART, 2005; MATSUI & PANHA, 2006; STUART & EMMETT,

2006: GRisMeR et al., 2007a-b; MCLEoD, 2008). The knowledge on these animals grows

rapidly. Species accounts summarizing our knowledge in the form of checklists are published

regularly (KHONSUE & THIRAKHUPT, 2001; ORLOV et al., 2002; NABHITABHATA et al., 2004;

TEYNIÉ & DAvip, 2007). However, further work on this fauna is still necessary requiring long

term study as well as study on the identities of previously named taxa.

ACKNOWLEDGEMENTS

We are gre

naturelle (Pari:

eful to our institutes for facilitating this study: Muséum national d'Histoire

France), National Science Museum (Pathum Thani, Thailand), Faculty of

estry, Kasetsart University (Bangkok, Thailand) and Faculty of Science, Khon Kaen

University (Khon Kaen, Thailand). The field work was conducted during the project “Bio-

diversity Hot Spot of Thailand” supported by funding from the Office of Environmental

Policy and Planning (OEPP) (Bangkok, Thailand) to PD, and permitted by National Park,

Wildlife and Plant Department, Ministry of Natural Resource and Environment (Bangkok,

Thailand). Field work of YC and CI was partly financed by the PPF “Etat et structure

phylogénétique de la biodiversité actuelle et fossile”. We thank Annemarie Ohler (MNHN)

for comments. We especially thank Jean-Pierre Brutus and Sutee Duangjai for the corrections

of the text and providing some references, respectively.

Source : MNHN, Paris

CHUAYNKERN, SALANGSINGHA, MAKCHAI, INTHARA & DUENGKAE 23

LITERATURE CITED

BossuyT, F., BROWN, R. M., Hicuis, D. M., CANNATELLA, D. C. & MILINKOVITCH, MC. eau.

Phylogeny and biogeography of a cosmopolitan frog ation: late Cretaceous diveï i

resulted in continent-scale endemism in the family Ranidae. Syst. Biol, 55: 579-594.

CHan-aRD, T., 2003. — À photographic guide to amphibians in Thailand. Bangkok, Darnsutha Press Co.,

Ltd: 1-175. [In Th:

CoLLis, J. P. & STORFER, A., 2003. — Global amphibian declines:

Distributions, 9: 89-98.

Dusois, A., 1992. - Notes sur la class

Lyon, 61: 305-352.

2005. — Amphibia mundi. 1.1. An ergotaxonomy of recent amphibians. Alytes, 23: 1-24.

Dusois, A. & OHLER, A., 2000. — Systematics of Fejervarya limnocharis (Gravenhorst, 1829) (Amphibia,

Anura, Ranidae) and related species. 1. Nomenclatural status and type-specimens of the nominal

species Rana limnocharis Gravenhorst, 1829. Alytes, 18: 15-50.

Durois, A., OHLER, A. & Buu, S. D., 2001. - À new genus and species of Ranidae (Amphibia, Anura)

from south-western India. Alytes, 19: 53-79.

Frost, D. R., GRANT, T., FAIVOVICH, J, BAIN, R. H., HaSs, A., HabDAD, C. F. B.. DE S4, R. O..

CHANNING, A., WILKINSON, M., DONNELLAN, S. C. RAxWORTHY, C. J., CAMPBELL, J. A., BLOTTO,

B. L., MOLER, P., DREWE u

2006. — The amphibian tree of

GRISMER, L. L., CHAV, T., NE, Woop, Jr., P. L., GRis n

Day, J. C. & KaIsER, H., 20074. - The herpetofauna of the Phnom Aural Wild il

and checklist of the herpetofauna of the Cardamom mountains, Cambodia. Hamadryad, 31:

216-241.

GRISMER, L. L., THy, N., CHAv, T. & HOLDEN, J., 2007b. - A new species of Chiromantis Peters 1854

: Rhacophoridae) from Phnom Samkos in the northwestern Cardamom mountains, Cam-

Herpetologica, 63: 392-400.

Hauuinay, T., 2005. - Diverse phenomena influencing amphibian population declines. In: M. LANNOO

{ed.), Amphibian Declines: The Conservation Status of United States Species, Berkeley, University

of California Press: 3-6.

Hever, W. R., 2005. - Variation and taxonomic clarification of the large species of the Leprodactylus

pentadactylus species group (Amphibia: Leptodactylidae) from Middle America, northern South

America, and Amazonia. Arquivos de Zoologia, 37: 269-348.

INTHARA, C., LAUHACHINDA, V., NABHITABHATA, JL, CHUAYNKERN, Ÿ. & KUMTONG, P., 2005. - Mouth

part Structures and distribution of some tadpoles from Thailand. Thailand nat. Hist. Mus. JL, :

55-78.

KHONSUE, W. & THIRAKHUPT, K.,

Chulalongkorn Univ., 1: 69-8

McLro, D.S., 2008. - A new species of big-headed fanged dicroglossine frog (genus Linmmonectes) from

Thailand. Zootaxa, 1807: 26-46.

Matsut, M. & PANHA, S., 2006. — À new species of Rhacophorus from eastern Thailand (Anura:

Rhacophoridae). Zool. Sci., 23: 477-481.

Myers, C. W. & DuëLLMAN, W. E., 1982. - À new species of //yla from Cerro Colorado, and other tree

frog records and geographical notes from western Panama. Am. Mus. Novit.. 2152: 1-32.

NABHITABHATA, J, CHAN-ARD, T. & CHUAYNKERN, Y., 2004. — Checklist of amphibians and reptiles in

Thailand. Bangkok, Ofice of Environmental Policy and Planning, “2000: 1-152.

Our, A., 1996. — Systematies, morphometrics and biogeography of the genus Aubria (Ranidae,

Pyxicephalinae). Alytes, 13: 141-166.

Our, A. & Dunois, A., 1999. - The identity of Elachyglossa gyldenstolpei Anderson, 1916 (Amphibia,

Ranidae), with comment on some aspects of statistical support to taxonomy. Zoologica scripta, 28:

269-279,

es 2006. — Phylogenetic relationships and generic taxonomic of the tribe Paini (Amphibia. Anura,

Ranidae, Dicroglossinae), with diagnoses of two new genera. Zoospstema, 28: 769-784.

rting the hypotheses. Diversity &

fication des Ranidae (Amphibiens Anoures). Bull. mens. Soc. linn.

. Bull. amer. Mus. Nat. Hi

2001. — A checklist of the amphibians in Thailand. Nat. Hist. J

Source : MNHN, Paris

24 ALYTES 27 (1)

OHLER, A., MARQUIS, O., SWAN, S. R. & GROSIEAN, S., 2000. - Amphibian diversity of the Hoang Lien

Nature Reserve (Lao Cai Province, northern Vietnam), with a description of two new species.

Herpetozoa, V3: 71-87.

ù R. & Darrry, J C., 2002. - A recent survey of the amphibian fauna of the

mom Mountains, southwest Cambodia with descriptions of three new species. Raflles Bull.

Zool., 50: 465-481

ORLOV, N. L., MURPHY, R. W., ANANIEVA, N. B., RYABOV, S. A. & CUC, H. T., 2002. - Herpetofauna of

Vietnam, a Checklist. Part L. Amphibia. Russ. J Herp., 9: 81-104.

Sruarr, B. L., 2005. - New frog records from Laos. Herp. Rev., 36: 473-479.

SruarT, B. L., CHUAYNKERN, Y., CHAN-ARD, T. & INGER, R. F,, 2006. - Three new species of frogs and a

new tadpole from castern Thailand. Fieldiana: Zool., (n.s.), HE: 1-19.

Sruart, B. L. & Emmeërr, D. A., 2006. — A collection of amphibians and reptiles

Mountains, southwestern Cambodia. Fieldiana: Zool., (n.s.), 109: 1-27.

TEyNié, A. & Davib, P., 2007. - Additions to the snake fauna of southern Laos, with the second Laotian

specimen of Naja siamensis (Laurenti, 1768) and the first country record of Oligodon taeniatus

(Günther, 1861) (Squamata, Serpentes). Russ. J_ Herp., 14: 39-44.

TEYNIÉ, A. DAVID, P., OuLER, A. & LUANGLATH, K., 2004. - Notes on a collection of amphibians and

reptiles form southern Laos, with a discussion of the occurrence of Indo-Malayan species.

Hamadryad, 29: 33-62

Varru, M. KOSUCH, JL, FELDMANN, R., MARTENS, H. & Srrrz, A., 2000. — A test for correct species

declaration of frog legs imports from Indonesia into the European Union. Biodiversity & Conser-

vation, 9: 333-341.

Verrn, M. KoSUCH, J., OHLER, A. & Dumois, A., 2001. - Systemati

horst, 1829) (Amphibia, Anura, Ranidae) and related spi

variation from the Great Sunda Islands

Alytes, 19: 5-28.

from the Cardamom

of Fejervarva limnocharis (Graven-

2. Morphological and molecular

Sumatra, Java, Borneo) with definition of two species.

Corresponding editor: Alain Dunots.

‘© ISSCA 2009

Source : MNHN, Paris

Alytes, 2009, 27 (1): 25-37. Book review

Threatened amphibians

Annemarie OHLER & Alain DUBoIs

Reptiles et Amphibiens, UMR 7205 OSEB, Département Systématique & Évolution.

Muséum national d'Histoire naturelle, CP 30, 35 rue Cuvier.

75005 Paris, France

<ohler@mnhn.fr>, <adubois@mnhn.fr>

Simon N. STUART, Michael HOFFMANN, Janice S. CHANSON, Neil A. Cox, Richard J. BERRIDGE, Pavithra

RaMaNI & Bruce YOUNG, B. E.. (ed.), 2008. - Threatened amphibians of the world. Barcelona (Lynx

Edicions), Gland (IUCN) & Arlington (Conservation International): i-xv + 1-758.

1. Threatened amphibians of the world, and much more

Annemarie OHLER

When we started working on the project to establish the conservation status of all known amphibian

species of the world, it seemed à rather impossible goal. AII of us who are working on these species are

aware of how little we actually know. But it was an exciting goal and 1 went to the workshops on Asian

amphibians. It was fun, hard work, many people to meet, knowledge to share. And then we realized that

there is much more information in the peoples’ mind than in all the books. And this is the strength of this

opus: the knowledge of those 500 listed in the dedication, put together under the perceptive and generous

guidance of Simon Stuart and his team, resulted in this monumental book of about 800 pages.

Why à book in the age of internet? Just go to the IUCN website [http//wwwiucnredlist.org/

amphibians] and you will find what you search! Most of scientific knowledge today is accessible on the

web only. Nevertheless the book is much more. Open it and frogs come into your life, Turn the pages and

many more. And not only to the life of herpetologists, conservation specialists, but to any politician,

administrative person, journalist or just reader who takes the book on a shelf will dive immediately in the

exciting diversity of amphibians, as Julia Marton-Lefèvre says in her foreword: “Afïer reading this book

and contemplating all these pictures, no one will look at amphibians with the same eves”. And here is the

strength of a book, in its physical existence, the smell of the printed pages, the brilliance of the paper, the

layout that guides you, the information that is concentrated in one page, giving with a glimpse à nam

photo, a distribution, a conservation status on a species and by turning the pages you see the colout

shapes, the richness and diversity.

, à

; the

The book has three major parts. The introductory chapter presents the amphibians, the Global

Amphibian Assess. ate of knowledge on these animals by biogeographic regions, closing

by a chapter on amphibian conservation, The ma entation of all species according to their

status of threat. The book then gives a list of ries of append oncerning the IUCN

categories and lists of species presented under various criteria, a glossary of terms and websites and

amphibian-related resource:

Source : MNHN, Paris

26 ALYTES 27 (1)

À book review clearly cannot summarize all the subjects raised in this book. Just take the chapter

that introduces the amphibians. You can find distribution maps of all amphibian families. Such maps have

not been published since ViaL's book in 1973 and here you can find them based on the latest ations

proposed and with very fine di stribution gaps, if existing. Similarly, maps on

endemism, data on median range size of amphibian species or maps showing the world's countries

according to their relative proportions of world's amphibians or relative proportions of threatened

amphibians showing a large bulging south and central America and reducing Eurasia and north America

Lo à fine stripe. These are only examples of how the data available have been exploited and presented in

tables, figures and maps. A nice idea is the sections called “Essays” that initiate to specific subjects

concerning amphibian biodiversity, aspects particular for the different biogcographic realms and conserv-

ation study.

have to es Cons dered extinct. Species are listed in alphabetical order. For every species its conservation

ribution and current population trend are given with a map and photo of

ble Then the geographic range, the information available on population status, the

habitat and ecology, major threats and conservation measures are described. Sources indicate

bibliography and persons that provided the data presented. What is lacking is size indication for the

as some àre à 20 mm others might have more than 100 mm.

In the appendices, many complementary data can be found. The IUCN critera are presented and

explained. The reader finds the Amphibian Conservation Action Plan and conservation required for

every species concerned. The status of species according to their IUCN red list is analysed by genus and

by country. Other information given in tables, such as the list of rapidly declining species or the list of

critically endangered species, helps to point to conservation issues. The list of species that need imme-

diately ex situ conservation action may help immediate action plans. To be complete the list of L

concern and data deficient species as listed in 2007 is given. The species evaluated differently by the Brazil

GAA workshop and the team that checked for consistency of the GAA work are listed in order not to wipe

outsuchdifferences, Finally,aftera glossaryof somekeytermsandalistof websitesonamphibians,thebook

closes with an index Lo extinct, threatened and near threatened species by genus and species name.

This book should be present in any office dealing with conservation issues, in public and private

libraries but also on the desk of researchers and field workers. It may guide political decisions, but also

help preparing research and conservation plans. It is both useful and nicely done. To use without

moderation!

Many people from all over the world worked together to complete the book but all this effort to make

people participate could not hide an important gap. When checking for the origin of the authors, it is

quite nice to see that many authors come from the countries concerned. An exception is still Africa. In

herpetology it is reduced to South Africa and since the First World Congress of Herpetology in 1989,

when L'entered herpetologists” world, this did not change. Here clearly is a need for capacity building, so

that conservation could be more focused on high biodiversity regions and threatened species that are in

southern countries (BRrTo, 2008).

And to finish with good news and to go on adding data, Conraua derooi Hulselmans, 1972, listed as

critically endangered, possibly extinct, isstillalive. Gabriel Segniagbeto and myself saw in May 2007 sever-

aladultspecimensof this frogin two locationsof thestreamat Missahohéin Togo. ltsoccurrenceislimited, it

must be considered critically endangered, but forest habitat is still available. It is still time to act.

2. The special message from threatened amphibians

Alain Dusois

Many books, reports and papers are available concerning threatened birds and mammals, many

actions taken for ex situ and in situ conservation of some of the species of these two groups, a number of

Source : MNHN, Paris

Dugois 27

Table 1. - Average ycearly increase in species number recognized by taxonomists over 10 periods

from 1768 to 2002 (DuBois, 2004) and from 2003 10 2008 (DUBOIS, unpublished). The

values marked * only give the number of new taxa described over this period but do not take

synonymisations into account, so they are certainly slightly overestimated. The value for

2008 is that given on the website AmphibiaWeb on 21 January 2009.

Total number | Average yearly increase Total number | Average yearly increase

Date ofspecies | in species number since | Date ofspecies | in species number since

recognized preceding date recognized preceding date

1768 57 — 1992 4522 634

1854 234 2: 1997* 4975 90.6

1882 1003 27.5 2002* 5472 99.4

1969 3343 26.9 2007* 6116 128.8

1984 4015 44.8 2008 6452 336.0

which are widely known even among the public at large. In contrast, “lower vertebrates”, not to mention

“invertebrates”, have until now attracted much less attention from laymen, conservationists and even

biologists. The impressive book Threatened Amphibians of the World (TAW below) therefore fills a large

gap: hopefully, amphibians will now be fully considered when conservation plans are prepared and when

research priorities are decided upon. Beside all the positive aspects of this book stressed above by

Annemarie, this book also sends us several important messages, some of which concern only amphibians

whereas others are more general.

The models of conservation biology developed for birds and mammals do not have generality over

the whole animal kingdom. For amphibians, it is clear that ex situ conservation can only be very marginal.

It can apply only to a few species that accept captivity and for which we know the basic needs and are able

to cover them. Furthermore, it is of limited interest whenever the habitat where these perse lived ha:

disappeared. Are we going to keep captive populations of amphibians in z0os, aqua-te: “tanks

for decades, without any possibility to put them back in their natural ecosystems? This may be meaningful

and worthy the effort for a few species used as experimental material in research, or for very beautiful or

unusual species, but it cannot apply to the thousands of species of amphibians that are going to get extinct

in the current and next decades. This is even worse for many invertebrates, for which we only have a very

limited experience of captive breeding, and the number of species threatened with extinction is greater by

several orders of magnitude.

For most amphibians, just like for many other “small” continental organisms, the main factor of

extinction is not direct destruction by man (through hunting, fishing or voluntary destruction), as the

latter concern only a few “well-known” species of large size, widely distributed and often living in

anthropic habitats. Even the peculiar ecological or genetical factors of threat identified and usually

highlighted by conservation biologists, like fr gmentation of metapopulations, consanguinity, road

trafic, predation by and competition with in species or domestic anima

ultraviolet or other radiations, pal ï except for chytridiomyc n some areas), play

only a marginal role in amphibian extinctions at world-scale level, The most important factor is by f

“simply” the complete destruction of habitats, especially humid zones and forests. The case of the

tropical forests of Sri Lanka, more than 95% of which have been wiped out in the last century

CMANAMENDRA-ARACHCHI & PETHIYAGODA, 2005), is exemplary. As most amphibian species have a very

limited distribution range, destruction of any tropical forest results in the immediate extinction of

numerous species, many of which are still unknown to science. Unfortunately, worldwide destruction of

habitats is a phenomenon on which “conservation biology” can only very marginally act. National and

other natural parks can only protect some particularly exceptional or “spectacular"” habitats, but only a

fraction of the total number of amphibian species.

The book puts focus on several regions and taxa, but this is only one part of the story. The most

important problem is perhaps that the species and habitats for which we have enough data to make precise

Source : MNHN, Paris

28 ALYTES 27 (1)

estimates of the status and thr nd to propose conservation actions, are only a small minority of the

total. In this respect, the situation in amphibians is drastically different from what it is in birds, the animal

group which has attracted the most important conservation efforts until now, because in birds a lot more

is known about the species, their needs and threats. A first strong conclusion of this book is how little we

know about the current situation of many amphibian species of the planet, despite the enormous amount

of work devoted to the task of building the database which made TAW possible. Of a total of 5915 s à

surveyed in the book, 1382, 23.4, are considered “data deficient” at the end of this survey. If one

considers a rate of error of 5 % as a standard yardstick for acceptable errors and missing data in ie

research, then it is clear that our current knowledge of amphibian taxa and populations is insufficient to

qualify as a ctory scientific knowledge. Furthermore, this rate is still an underestimation, both

quantitatively and qualitatively.

The book reviews data bearing 6n almost 6000 speci

sive, this number is certainly much below the number of

of 2008, taxonomists recognized about 6500 amphibian species (this is only a rough estimate, because

disagreements exist between specialists about the status of many taxa). This number was about 5000 in

1997 and about 4000 in 1984. Many new species are described each year, and even if a few of the new

nomina are later shown to be synonyms, the mean increase has been around 100 new species per year in

the last decade. Is this number progressively decreasing? No, much less to the contrary, as shown in table

1. In 2008, for the first time in the history of zoology, more than 300 new amphibian species were

described by taxonomists. No indication exists that the curve of increase in the number of known species

of amphibians is approaching an inflexion point that would precede a stabilisation of this curve

{STEyskAL, 1965; Dumois, 1977, 1998). It is much more likely that, after having surpassed the number of

species of mammals (GLAW & KÔHLER, 1998), the number of known species of amphibians will soon

xceed that of reptiles, and several authors (DuBois, 2004: Parka et al., 2005: 45) have recently

expressed the idea that the number of amphibian species described is probably only half or less than half

the number of spe actually still present on the planet. However, no precise estimate exists for the total

number of amphibian species still living on earth today, and of those that were living, say, one or two

centuries ago, before large deforestations took place in most tropical regions: the latter number was

probably significantly higher.

of Amphibia worldwide. Although impres

mphibian species stif{ living on earth. At the end

Where are the new species currently being described? Table 2 provides data on this question. Thi

data are approximate, especially as different taxonomic approaches result in rather different estimates of

the numbers of species by countries, as shown in this table by the figures borrowed from the online

database Amphibia Web (AW) [htip:/amphibiaweb.org/] and from TAW. However, they clearly show that

the “taxonomic effort” is very unevenly distributed on the planet, The highest proportional increase in the

number of known species over the period 1998-2007 was in Sri Lanka (about 40 %), then in Chile,

Madagascar, Papua New Guinea, Peru and Vietnam (about 25%). These are mostly tropical countries

with still important forest remnants. The amphibian specific richness can be correlated with various

ccological and geographical parameters (GALLANT et al., 2007: TAW), but, 10 put the things very bricfy,

itis above all correlated with latitude. Table 3 shows that, over the whole planet, the mean species density

of known species is higher than 330 species by 1,000,000 km? in the ribbon of 20 degrees of latitude on

each side of the equator, and may be much higher in this intertropical zone in some realms (Australasian,

Neotropical and Oriental). Then this density goes decreasing rather regularly towards either the North or

the South.

However, the number of newly described species does not follow exactly this pattern (table 3), the

highest number being in the latitudinal band 10-19S. This is in part because more than half of the species

described from 1998 to 2007 were from the Neotropical realm. This region is currently the matter of much

more Laxonomic effort for amphibians than the other tropical areas. In contrast not a single new species

was described from 1998 to 2007 in 12 species-rich countries (where about 50 species or more have already

been reported): (numbers between parentheses: total species number

of country according to AIT) Congo Democratic Republic (214/211), Nigeria (108/103), Angola

(90/99). Ghana (76/76), Malawi (64/76), Central African Republic (57/62), rra Leone (50/55),

Zimbabwe (47/63), Liberia (44/56) and E d Guinea (29/70). Two of these countries are in the

Neotropical realm: Paraguay (76/74) and Haiti (57/51). There is no such country in the Australasian,

Source : MNHN, Paris

Table 2. - Total amphibian species known (SN) in 2008 and increase in species (and subspecies) number (SI) from 1998 to 2007 in various

countries and the six biogeographic realms of the world. The value of SN is given according to AW and to TAW (see text), respectively as

SN(AW) and SN(TAW). No value is given in AW for the realms. The value of SI is given according to DUBOIS et al. (2005 and

unpublished). New species and subspecies are here referred only to the country including their onymotope (type-locality) but their

distribution may be wide. The rate of increase, RI (100 SI / SN) is given according to both values of SN. The mention nd means “no

data”.

Surface ” SN(AW) SN(TAW) SI RI(AW) RI(TAW)

Ken Country (km°) Lsntade 2008 2008 | 1998-2007 Ce) Ce)

Affotropical Cameroon 46940 | OGOUN 195 196 12 62 61

Ethiopia 1.119.683 | 08°00°N 65 6 2 31 32

Gabon 257,667 | 01°00°S 4 87 4 91 46

Guinea 245.857 TISO0N 63 74 2 3.2 2.7

Ivory Coast 318.000 O08°00°N 57 91 6 10.5 6.6

Kenya 569.250 | 01°00°N 105 96 3 29 31

Madagascar 581,540 | 20°00°S 244 227 54 2.1 238 D

Mozambique 784,090 | 18158 6 67 1 16 1.5 ë

Namibia 825.418 | 22°00'S 38 48 1 26 2.1 5

Sao Tome & Principe 1,001 | 01°00°N 7 6 1 143 16.7 o

Seychelles 455 049358 12 12 à 16.7 16.7

| South Africa 1219912 | 29°00°S 117 ns s 43 43

Tanzania 886.037 | 06°00°S 183 162 31 169 191

Uganda 199.710 | 01°00°N 52 6 1 19 16

Zambia 740.724 | 15008 | 85 | 84 1 12 12

a 17200°N ’ =

All other countries (34) 14,552,463 _29°30'S 0-214 O-211 0 0 0

TOTAL ArROrROmMCAL (49) | 22.771.247 | ONU nd 969 126 nd 13.0

| Australasia ” Papua New Guinea 452,860 | 06°00'S 294 247 69 23.5 27.9

| New Zealand 268.021 | 41°00°S 7 7 1 143 143

Australia 7,617,930 | 27°00'S 226 218 il 49 50

1 1S12N

All other countries (7) 78,061 PE LS 0-20 0 0 0

TOTAL AUSTRALASIA (10) | 8.416.872 | SIN nd 544 si nd 149

= 41900" »

Source : MNHN, Paris

Table 2. — (continued 1).

. Surface SN(AW) | SN(TAW) SI | RI(AW) | RITAW)

BA Cointy (km°) pie 2008 2008 | 1998-2007 | (%) (%)

Nearctie Canada 9.093.507 | GOOON 4 48 Q Q 0

United States of America | 9.161.923 | 38*00°N | 292 279 26 | 89 93

ToraL NEarcnic (3) | 18255483 | OUN | nd | 327 2 | 8.0

Neotropieal | Argentina 2736690 | 34°00S TSS | 157 6 38 38

Bolivia 1.084390 | 17°00°S 25 209 41 182 196

Brazil 8456,510 | 10°00° 806 753 125 16.6

Chile 748,800 | 30,00S 55 56 14. | 25.0

Colombia 1,038,700 | 04*00°N 736 699 78 112

Costa Rica 50.660 | 10°00°N | 187 18 ul 6.1 >

Cuba 110.860 | 2130N | 64 60 8 133 Ê

Ecuador 276.840 | 02°00°S 474 45 3 69 5

El Salvador 20.720 | 13°50N | 28 | 32 1 3. 5

French Guiana 83.534 | 04°00° 97 | 105 6 57 a

Guatemala 108.430 | 1530°N u6 | 138 14 10.1 S

Guyana 196.850 | 05°00°N 122 119 15 12.6 =

Honduras 111,890 | 15°00°N 103 118 4 119

Mexico 1,923,040 | 23*00N | 357 365 4 118

Nicaragua 120,254 | 13°00°N 67 69 3 43

Panama 75.990 | 09°00°N 197 197 nl 56

Peru 1,280,000 | 10°00°S 460 413 3 | 274

Puerto Rico 8870 | 1815N 25 24 1 42

Suriname 161.470 | 04°00°N 86 103 1 1.0

Trinidad & Tobago 5,128 | 11°00N 35 34 1 29

Uruguay 173,620 | _33°00S 4 | MS 1 23

Venezuela 882,050 | O8*00"N | 321 302 45 149

> = 24°15N

Al other countries (23) 525554 | PEN | 1-76 0-74 0 0 0

ToraL Nrorronical (45) | 20,180,850 | AN nd 2916 585 | nd 20.0

Source : MNHN, Paris

Table 2. (continued 2).

Surface F SN(AW) | SN(TAW) SI RI(AW) | RITAW)

| Counuy (km°) Latitude 2008 2008 1998-2007 (%) C6)

Oriental Burma (Myanmar) 657,740 | 2200 N #4 79 3 3.6 38

Cambodia 176,250 | 13°00°N 44 44 6 13.6 13.6

China 9.326410 | 35°00°N 345 330 49 142 148

India 2.973.190 | 20°00°N 263 237 54 20.5 228

Indonesia 1,826,440 | 05°00S 306 354 59 193 16.7

Laos 230,800 | 18°00°N 82 73 12 14.6 164

Malaysia 328,550 | 02°30°N 210 205 14 67 68

Nepal 143,181 | 28°00°N 44 46 3 68 6.5

Pakistan 778,720 | 30°00°N 20 18 2 10.0 on

Philippines 298,170 | 13°00°N 102 103 12 118 11.7

Sri Lanka 64740 | 07:00 N 107 102 4 393 412

Thailand 511,770 | 15°00°N 124 130 14 113 10.8

Vietnam 325,360 | 16°00°N 165 145 39 236 269 D

" 27930°N [e}

All other countries (9) 235,500 | ogos | 0-4 1-54 0 0 0 =

ToraL ORtEntaL (22) | 17877091 | NX nd 999 309 nd 30.9 …

Palacarctic | Israel” 20,330 | 31°30°N 8 7 Î 12.5 143

Italy 294,020 | 42°50N 48 40 2 42 50

| Japan 374,744 | 36°00°N 65 64 2 31 31

Morocco 446.300 | 32°00°N 14 12 2 143 16.7

| Portugal 91951 | 39°30°N 20 22 [ 5.0 45

Serbia 77474 | 44°00°N 21 21 1 48 48

South Korea 98.190 | 37°00°N 17 16 2 1L8 125

Spain 499,542 40°00'N | 32 44 4 12.5 9.1

Turkey 770.760 | 39°00°N 29 26 1 34 38

; 64°00°N

Allother coumries (71) | 40058282 | SÉgx | 0-40 0-39 0 0 0

| TOTAL PALAFARCTIC (80) | 42,731,593 Ent nd 468 16 nd 34

TOTAL | 209 COUNTRIES 130,233,136 | SPOON | y 6223 1141 nd 183

L 4100 | ww

Source : MNHN, Paris

Lei

Table 3. - Mean amphibian species density (by 1.000.000 km°). computed from the total number of species by area according to AW, and number of

species and subspecies described as new from 1998 to 2007. in countries of surface above 50,000 km, according to their “mean” latitude and in

the six biogeographical realms (from the data of tab. 2). New species and subspecies are here referred only to the country including their

onymotope (type-locality) but their distribution may be wider. The mean species density (MSD) is given first, followed after oblique bar by the

number in bold of species described as new (NS)

Latitude Afrotropical | Australasia Nearctic Neotropical Oriental Palearctic AI realms

60-69° N - - 528/0 - - 3.80/0 >

50-59 N : = L | . 5: 107.12/0 &

40-49 N - = 2 | L = 58.06 / 7 =

30-39 N ë - 30.45 /26 | 2 3444/51 3531785 w

20-29 N - _ _- 211.31/81 109.57 / 60 61.77/111 S

10-19 N 45.92/2 | 982.32/0 e | 1478.74/42 | 32088/83 | 152.06/ 127 S

00-09 N 176.31/24 | 218341/0 - 625.36/156 | 976.12/56 - 333.62 / 236

00-09 S 24931/38 | 555.79 / 69 - | 1629.10/31 | 195.50/59 | 353.33 / 194

10-19 S 114.08 /2 65.64/0 = | 127.07/279 - - 124.22 / 281

20-29 153.93/60 | 28.68/11 . 186.26 / 0 - = 73.17/7

30-395 - = | e 69.96 / 21 = | - 69.96 / 21

40-495 = 2 | = = = = 26.12/1

Mean MSD / Total NS | 120.81/123 59. | 1791/26 | 5/580 | 113.55/309 23.38/15 | 8846/1134

Source : MNHN, Paris

Dugois 33

Fig. 1. — Chaparana ( Paa) minica (Dubois, 1975). Adult 4, MNHN 1989.2078, Dobhi (1520-1550 m),

between Katrain (32°07°N, 77°08'E) and Raison, Himachal Pradesh, India, 4 August 1977 (photo A.

Dubois).

Fig. 2. - “Ombrana sikimensis (Jerdon, 1870)”. Young d, MNHN 1974.1504, Namdu Khola (27°06°N.

87°48'E, 2000-2060 m), between Kalikharka and Ito, Nepal, 15 May 1973 (photo A. Dubois).

Nearctic, Oriental and Palaearctic realms, but several other species-rich countries in these realms have

witnessed a very low rate of taxonom e in this period (see table 2 for details). These data suggest

that some parts of the world, especially in tropical Africa but also elsewhere, are currently almost

completely abandoned by amphibian taxonomists and field naturalists, mostly for political reasons (wars,

dictatorships, domestic troubles, economic problems).

Finally, even for species that have been described, for which there seems to exist no important

taxonomic problem, and sometimes mentioned a good number of times in the literature, the information

available on their distribution, habitat and mode of life is too scanty for allowing a proper estimate of

their conservation status. Let us just consider the stream frogs of the tribe Parvr of the Ranr4r

DicroëLossiN4E (OnLER & Duois, 2006). AI these frogs are strictly dependent on running water for

breeding, larval development and most of their life (Liu, 1950; Dumois, 1976) and they are therefore

threatened by any change in water regime in streams, e.g.. caused by deforestation, dam construction or

water exploitation and management by man (DUBots, 1980, 2000). As such, all the 42 known species of

this tribe should be considered at least as “vulnerable”. However, the choice of species in TAW

(p. 490-492) is difficult to understand. Four species are listed as “endangered” and eight as “vulnerable”

although some of these have a rather wide distribution. Apparently some were considered “endangered

for being consumed by man, but others that are also harvested (e.g.. Chaparana licbigii) were not lis

The species Chaparana minica (fig. 1) is considered as “’vulnerable” although it is present in most wat

courses between 1000 and 2500 m in a rather large area, from western Nepal to Himachal Prad

through Himalayan regions of Uttar Pradesh (Dumois, 1976) and most likely present in the Nanda Devi

Biosphere Reserve of the latter state (Dunois & DinesH, 2007). In contrast, the species listed by FRosr et

(2006) as Ombrana sikimensis (fig. 2), but not mentioned in TAW, has a range of similar size (from

central Nepal to Sikkim), but in this region it has a very scattered distribution, being restricted to a few

ict forested areas at rather low altitudes (Dunois, 1976) and is therefore very vulnerable. Some Par

s which are known only from a few specimens (Chaparana arnoldi, Chaparana ercepeue) are listed in

TAW as “near threatened”, but others are not at all mentioned in the book although they are known only

from a few specimens and one or a few localities, such as Chaparana aenea (see Durois & OHLER, 2005),

Chaparana rarica or Gynandropaa bourreti, or even from a single holotype (Gynandropaa feae).

Source : MNHN, Paris

34 ALYTES 27 (1)

Of course, it would be easy to say that, if 1 had attended the Global Amphibian Assessment

meetings, where 1 was invite, 1 could have modified the situation by providing unpublished information,

and I certainly regret not to have been able to do so, but this example among many other possible ones

shows how fragile is the information on which many decisions taken in this book are based. In numerous

cases, only one or à few persons are aware of some unpublished information that may be crucial for

ascertaining the conservation status of a spe In amphibians we are not in a situation like in birds,

where many field naturalists, amateur ornithologists, “bird-watchers”, observe many species cach year,

thus providing much useful information and being able to call almost immediately the attention to sudden

problems concerning a species. Hopefully, “frogs and salamanders watching” will develop, in part thanks

to the many colourful field guides that are now regularly published in various countries, but we are still far

from having a complete network of competent and vigilant batrachologists all around the world. This will

probably be long and hard to establish. The field identification of amphibians is much more difficult than

that of birds, because visual clues (colours, conspicuous behaviours) are much less abundant and reliable

in frogs, as they played a minor role in speciation in this group (see Duois, 1988).

As well stated by Parka et al. (2005), and briefly reminded in TAW (p. 18, 45), taxonomy plays a

major role in species conservation. A strategy of conservation biology that would rely on a grossly

incomplete or faulty taxonomy would not only be useless but might be harmful. The taxonomy of

amphibians is currently in a very incomplete stage, covering perhaps less than half the species of the

planet. Any biologist who fecls concerned with the need to protect and conserve these extraordinary

animals should therefor feel concerned with supporting taxonomic research. This may require some

change in the priorities of the international community of batrachologists. For the time being, this

community has put an enormous amount of collective effort into providing information about phyloge-

netic relationships between major amphibian groups (e.g., Frosr et al., 2006) and in compiling an

enormous database regarding the conservation status of many known species (TAW). These works are

extremely useful, but they won't be enough for an efficient action for the conservation of as many of these

animals as possible for the future generations.

Furthermore and unfortunately, in many cases such an action will prove impossible, whenever

truction of the habitat is unavoidable. In such situations, it would be of the greatest importance at least

ickled” specimens, tissues, observations, photographs, call recordings, etc. (DUBoIs, 2003).

cases, this may be more realistic than claiming that we will “save” il the amphibians. The

community of batrachologists should be careful in well defining its priorities, not only regarding the

geographical focuses of our works (BkrTo, 2008), but more generally in giving taxonomic rescarch,

collection, study, description and naming of species preeminence in the works to be carried out in this

century. An important problem that will have to be solved soon is the conflict between taxonomy and

conservation biology regarding the possibility to collect specimens in the field for taxonomic research: as

long as conservation biology acts as a break against the collection of specimens, it will act as a break

against taxonomic research, thus against knowledge and, in the long run, against the conservation of

biodiversity itself (Dusois, 2003; Dunots & NEMÉSIO, 2007).

If taxonomists continue their descriptive work at a “usual” pace, without a strong voluntary action

10 à rate it, centuries may be necessary to complete the taxonomic inventory of animal species on

earth (GONZALEZ-OREJA, 2008). Compared to other groups however, the situation may be rather

favourable in amphibians. These are relatively large animals, that live in habitats usually accessible to

man, and it should be possible, if our community decided to make it a major priority, to develop an

international, collaborative strategy, for carrying out as quickly as possible an “exhaustive taxonomy” of

this group (Dunois, 2008). IF indecd there are 10-15,000 spe of amphibians on earth, and if 300 of

them are described cach year from now on, including in regions that have so lar been neglected, a

subcomplete inventory of this group should be feasible within a span of 20-30 years. But time is running,

short and extinctions are going quicker than our decisions. If an exhaustive taxonomy of amphibians is

not considered a scientific priority and implemented by the coming generation of batrachologists, it will

be 100 late for ever for many species.

Source : MNHN, Paris

DuBois 35

Brro, D., 2008. - Amphibian conservation: are we on the right track? Biological conservation, 41:

917

1976. - Les Grenouilles du sous-genre Paa du Népal (famille Ranidae, genre Rand). Cahiers

népaluis. Documents, Paris, |

Ben: 1977. - Les problèmes de l'espèce chez les Amphibiens Anoures. /n: C. BOCQUET, J. Gé

LaMOTTE (ed.), Les problèmes de l'espèce dans le règne animal, 2, Mémoires de la Société =oologique

de France, 39: 161-284.

1980. — L'influence de l'homme s

r la répartition des Amphibiens dans l'Himalaya central et

occidental. Comptes rendus des Séances de la Soc de Biogéographie, 85: 155-178.

---- 1988. - The genus in zoology: a contribution to the theory of evolutionary systematics. Mémoires du

Muséum national d'Histoire naturelle, (A), 140: 1-123.

La 1998. — List of European species of amphibians and reptiles:

Amphibia-Reptilia, 19 (1): 1-28.

000. — The influence of man on the distribution of amphibians in the Himalayas of Nepal: an

example of critical evaluation of biogeographical data. Jr: G. Miëur & Y. ZHANG (ed.). Envin

mental changes in high Asia, Marburger geographischer Schrifien, 135: 326-345

2003. - The relationships between taxonomy and conservation biology in the century of extinctions.

Comptes rendus Biologies, 326 (suppl. 1): S9-S21.

os 2004. - Developmental pathway, speciation and supraspecific taxonomy in amphibians. 1. Why are

there so many frog species in Sri Lanka? A/rtes, 22 (1-2): 19-37.

2008. — I had a dream... Alytes, 25 (3-4): 89-02.

Dugois, A., CROMBIE, R. I. & GLAW, EF, 200$. - Amphibia Mundi. 1.2. Re

infrageneric taxonomic additions (1981-2002). A/ytes, 23 (1-2): 25-69.

Duois, A. & DINESH, K. P., 2007. - The status of the nomen Rana ( Paa) dhakuriensis Ray, 1997 (Anura,

Ranidae), and comments on the Amphibia reported from the Nanda Devi Bi

(Uttar Pradesh, India). Alvtes, 25 (1-2): 75-82.

Dunois, À. & NEMÉsIO, AÀ., 2007. — Docs nomenclatural availability of nomina of new species or

subspecies require the deposition of vouchers in collections? Zootaxa, 1409: |

Duois, A. & OuLER, A., 2005. - Taxonomic notes on the Asian frogs of the tribe Paini (Ranidae.

Dicroglossinae). 1. Morphology and synonymy of Chaparana aenea (Smith, 1922), with proposal

will we soon be reaching “stability”?

nt amphibians: generic and

here Reserve

of anews al method for testing homogencity of small samples. Journal of natural Historr, 39

(20): 1759-1778

FRoST, D. R., GRANT, T., FaivoviCH, EL, BAIN, R. H., Haas, A. HaDbbab, C. F. R. O. br,

CHANNING, A, WILKINSON, M., DONNELLAN, S. C., RAXWORTHY, C. I, CAMPBELL, a) Ra BLOTTO,

B. L., MOLER, P., DREWES, R. C., N LR. A. LYNCH, JL D., GREEN, D. M. & WHEELER, WC.

2006. - The amphibian tree of life. Bulletin of the American Museum of natural History, 297: 1-370.

GALLANT, A. L., KLAVER, R. W., CASPER, G.S. & LANNOO, M. J., 2007. — Global rates of habitat loss and

implications for amphibian conservation. Copeia, 2007: 967-979.

Graw, E & KOHLER, J., 1998. — Amphibian species diversity excecds that of mammals. Herpetological

Review, 29 (1): 11-12

GONZALEZ-OREJA, J. A., 2008. — The Encyclopedia of L the Brochure of Life: exploring the

onships between the extinction of species and the inventory of life on Earth. Zootaxa, 1965:

61-68.

Liv, C 1950. - Amphibians of western China. Fieldiana: Zoology Memoirs, 2 1-400, pl. 1-10.

MANAMENDRA-ARACHCH, K. & PETIHIYAGODA, R., 2005. - The Sri Lankan shrub-frogs of the genus

Philautus Gistel, 1848 (Ranidae: Rhacophorinae), with description of 27 new spi The Raffles

Bulletin of Zoology, suppl. 12: 163-303

OuLEr, À. & Dunois, À., 2006. — Phylogenctic relationships and

(Amphibia, Anura, Ranidae, Dicroglossinae), with diagnose:

(3): 769-784.

enerie taxonomy of the tribe Paini

pl two new gencra. Zousrstema, 28

Source : MNHN, Paris

10 FEV. 2010

36 ALYTES 27 (1)

Parka, G.. BROWN, R.. HANKEN, J., HEDGES, B., HEYER, R., KUZMIN, S., LAVILLA, E., LÔTTERS, S..

PIMENTA, B., RICHARDS. RôDEL, M. O., DE SA, R. O: & Wake, D., 2005. - Systematics and

conservation, Ju: C. GascoN, J. P. COLLINS, R. D. Moor, D. R. CHuRen, J. E. MCKAY & 1. R

MENDELSON LL (ed.), Amphibian Conservation Action Plan, Gland (Switzerland) & Cambridge

(UK), IUCN/SSC Amphibian Specialist Group: 45-48.

ca, G. €. 1965. - Trend curves of the rate of species description in zoology. Science, 149: 880-882.

. L. (ed.), 1973. — Evolutionar biology of the Anurans. Columbia, University of Missouri Press:

i-xiit + 1-470.

Source : MNHN, Paris

AINTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD

Chief Editor: Stephane GRosIAN (Reptiles et Amphibiens, UMR 7205 OSEB, Département de Systématique &

Evolution, Muséum national d'Histoire naturelle, CP 30, 25 rue Cuvier, 75005 Paris, France;

<sgrosja@mnhn.fr>)

Deputy Editor: Franco ANDREONE (Museo Regionale di Scienze Naturali, Via G. Giolitti 36, 10123 Torino, Italy:

<fandreone@libero.it>).

Alytes Editorial Board: Ariadne ANGULO (Toronto, Canada); David C. BLacxBuRN (Kansas, USA); Lauren E.

Brown (Normal, USA); Angus I. CARPENTIER (Norwich, UK); Ignacio DE LA Riva (Madrid, Spain);

Rafael O. DE SA (Richmond, USA); Alain Dugors (Paris, France): W. Ronald HEvr (Washington, USA):

Rafael MatquEz (Madrid Spain) Masafumi Marsut (Kyoto, pan), Annemarie One (Paris, France);

Mark-Oliver RôDEL (Berlin, Germany); Miguel VEnCES (Braunschweïg, Germany).

Amphibia Mundi Editorial Board. Alain Dunois, Chief Editor (Paris, France), Ronald I. CROMBIE (San

Francisco, USA); Stéphane GROSEAN (Paris, France); W. Ronald HevEr (Washington, USA); JIANG

Jianping (Chengdu, China); Esteban O. LAvILLA (Tucumän, Argentina); Jean-Claude RAGE (Paris,

France): David B. Wake (Berkeley, USA).

Technical Editorial Team (Paris, France): Alain Dunots (texts); Roger BouR (tables); Annemarie OHLeR (figures).

Book Review Editor: Annemarie OuLER (Paris, France).

SHORT GUIDE FOR AUTHORS

(for more detailed Instructions 10 Authors, see Alvtes, 1997, 14: 175-200)

Alytes publishes original papers in English, French or Spanish, in any discipline dealing with amphibians.

Beside articles and notes reporting results of original research, consideration is given for publication to synthetic

review articles, book reviews, comments and replies, and to papers based upon original high quality illustrations

such as colour or black and white photographs), showing beautiful or rare species, interesting behaviours, etc.

The title should be followed by the name(s) and address(es) of the author(s). The text should be typewritten

or printed double-spaced on one side of the paper. The manuscript should be organized as follows: English

abstract, introduction, material and methods, results, discussion, conclusion, French or Spanish abstract,

acknowiedgements, literature cited, appendix.

Figures and tables should be mentioned in the text as follows: fig. 4 or tab. 4. Figures should not exceed 16

X 24 cm. The size of the lettering should ensure its legibility after reduction. The legends of figures and tables

should be assembled on a separate sheet. Each figure should be numbered using a pencil.

References in the text are to be written in capital letters (BOURRET, 1942; GRAF & POLLS PELAZ, 1989; INGER

et al., 1974). References in the Literature Cited section should be presented as follows:

BouRRET, R., 1942. - Les batraciens de l'Indochine. Hanoï, Institut Océanographique de l'Indochine: i-x + 1-547,

GRAF, L-D. & PouLs PeLaz, M., 1989. Evolutionary genetics of the Rana esculenta complex. In: R. M. DAWLEY

&1 P Bogaer (ed) Évoltion and ecology of unisexual vertebrates, Albany, The New York Siate Museum:

INGER, R. F., Voris, H. K. & Voris, H. H., 1974. - Genetic variation and population ecology of some Southeast

Asian frogs of the genera Bufo and Rana. Biochem. Gener., 12: 121-145.

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