A blue-eyed Leptobrachium (Anura: Megophryidae)

Transcription

Terms of use: for private research use; for a high quality printable version,
see http://www.mapress.com/zootaxa/list/2011/2912.html
Zootaxa 2912: 28–36 (2011)
www.mapress.com / zootaxa/
Copyright © 2011 · Magnolia Press
ISSN 1175-5326 (print edition)
Article
ZOOTAXA
ISSN 1175-5334 (online edition)
A blue-eyed Leptobrachium (Anura: Megophryidae)
from Arunachal Pradesh, India
SANJAY SONDHI1,3 & ANNEMARIE OHLER2
1
TITLI TRUST. Villa # 49, Rajpur Road Enclave, Dhoran Khas, Dehradun, Uttarakhand, 248001. India. www.titlitrust.com.
E-mail: sanjay.sondhi1@gmail.com
2
UMR 7205 OSEB, Département de Systématique et Evolution, Muséum National d’Histoire Naturelle, 25 rue Cuvier, CP 30, 75005
Paris, France
3
Corresponding author
Abstract
A new species of Megophryidae, Leptobrachium bompu is described from Eaglenest Wildlife Sanctuary, Arunachal
Pradesh, India. It is a medium sized member of the genus Leptobrachium. In life, the frog is distinguished by the combination of a series of morphological characters from its congeners. Its entirely greyish-blue coloured eyes differentiate the
species from all other species in the genus but Leptobrachium waysepuntiense. The morphological comparison is based
on a table of 16 characters discriminant in Leptobrachium species.
Key words: Leptobrachium, Leptobrachium bompu, Megophryidae, Eaglenest, description
Introduction
The genus Leptobrachium described by Tschudi (1838) and belonging to the family Megophryidae Bonaparte,
1850 consists of 27 known species worldwide (Frost 2010). Traditionally this genus has been divided into two
groups, either recognized as genera (Fei et al. 2009) or as subgenera of a single genus (Dubois & Ohler 1998). In
these classifications, species with males bearing horny spines on upper lip have been allocated to Vibrissaphora
Liu, 1945 whereas the species without such a secondary sexual character were grouped into Leptobrachium.
However, findings by Zheng & Fu (2008) on the phylogeny of species allocated to the subgenera Vibrissaphora and Leptobrachium have revealed that the species allocated to these two subgenera, do not form monophyletic groups; hence Vibrissaphora should be considered a synonym of Leptobrachium. The recent work of Matsui et
al. (2010) confirms this result inasmuch as species with males bearing horny spines are part of a clade that also
includes species without this character. They recognize, for the time, two groups, clade A and clade B that can be
attributed the two names. But for these clades no morphological apomorphies are available, nor is a diagnosis possible. Therefore here we do not use subgeneric allocation, and if we use “Vibrissaphora”, we explicitly refer to the
species of Leptobrachium bearing spines on upper lip of adult males, not to a formal taxon.
Two species from the genus Leptobrachium are reported to be found in India: Leptobrachium smithi Matsui,
Nabhitabhata & Panha, 1999 (Sengupta et al. 2001) and Leptobrachium hasseltii Tschudi, 1838 (BCCP CAMP
Report 1998; Chanda 2002). While Leptobrachium smithi has been reported from Meghalaya, India (Ahmed et al.
2009), neighbouring Bangladesh (Reza 2009) and Myanmar (Das & Chanda 2004), records of Leptobrachium hasseltii are doubtful. Chanda (2002) reports specimens under this name from Meghalaya. This name has been largely
used, but in recent works it applies to a species from Java and Bali (Brown et al. 2009). Hence the status of the species published as Leptobrachium hasseltii from India clearly needs further investigation.
While most of the species belonging to the spine-bearing species (“Vibrissaphora”) are from China and Indochina, those devoid of such spines (“Leptobrachium”) have a more southern distribution including Sunda region.
Four species of Leptobrachium are reported from neighbouring Myanmar: Leptobrachium chapaense, Leptobrachium hasseltii, Leptobrachium pullum and Leptobrachium smithi (Zug et al. 2001; Das & Chanda 2004).
28
Accepted by M. Vences: 3 Mar. 2011; published: 9 Jun. 2011
We here report the presence of a third species of Leptobrachium from the Indian region which has a very particular eye coloration. For species belonging to Leptobrachium eye colour can be a very useful discriminant character in the field (Ohler et al. 2004). The specimen described here as a new species has an entirely greyish-blue eye
and differs from all other species in the genus, other than Leptobrachium waysepuntiense, a species recently
described from Sumatra (Hamidy & Matsui 2010).
Material and methods
All measurements were carried out with slide calipers to the nearest 0.1 mm. A field microscope was used to examine the specimen for spines on its upper lip and determine its sex. An effort has been made to follow the same format and methodology adopted to describe other Leptobrachium species by Ohler et al. (2004) for the purpose of
consistency. The list of measurements (as given for the holotype in Table 1) and the abbreviations used are mentioned below:
Body: SVL, snout-vent length.
Head: EL, eye length; EN, distance from anterior corner of eye to nostril; HL, head length (from posterior corner of mandible to tip of snout); HW, head width, at the angle of jaws; IBE, distance between posterior corner of
eyes; IFE, distance between anterior corner of eyes; IN, internarial distance; IUE, minimum distance between
upper eyelids; MBE, distance from posterior corner of mandible to posterior corner of eye; MFE, distance from
posterior corner of mandible to anterior corner of eye; MN, distance from posterior corner of mandible to nostril;
NS, distance from nostril to tip of snout; SL, distance from anterior corner of eye to tip of snout; UEW, maximum
width of upper eyelid.
Forelimb: FLL, forelimb length (from elbow to base of outer palmar tubercle); HAL, hand length (from base
of outer palmar tubercle to tip of third finger); TFL, third finger length (from the distal part to the articulation
between proximal phalange and metacarpal bone of the third finger); Fw3, width of third finger measured at the
halfway length of the finger.
Hindlimb: FFTF, distance from maximum incurvation of web between fourth and fifth toe to tip of fourth toe,
toes being spread; FL, femur (thigh) length (from vent to knee); FOL, foot length (from base of inner metatarsal
tubercle to tip of fourth toe); FTL, fourth toe length (from the distal part to the articulation between proximal phalange and metatarsal bone of the fourth toe); IMT, length of inner metatarsal tubercle; ITL, inner toe length; MTFF,
distance from distal edge of metatarsal tubercle to maximum incurvation of web between fourth and fifth toe, toes
being spread; MTTF, distance from distal edge of metatarsal tubercle to maximum incurvation of web between
third and fourth toe, toes being spread; TFOL, length of tarsus and foot (from base of tarsus to tip of fourth toe);
TFTF, distance from maximum incurvation of web between third and fourth toe to tip of fourth toe, toes being
spread; TL, tibia (shank) length; TW, maximum tibia (shank) width; Tw4, width of fourth toe, measured at the halfway length of the toe.
Eyes of Leptobrachium show variable colour patterns that concern iris or sclera. Sclera is the opaque outer
layer of eyeball (Duellman & Trueb 1985) that in the human eye is usually white. In Leptobrachium, sclera can be
white, black, light blue or orange-red (Table 2). In addition, the iris can be either uniformly coloured or bi-coloured
with the lower part black or dark brown and the upper third red, blue, white, lime or greenish. We distinguish these
two anatomical structures in the description of the eye colour.
The table of descriptive characters has been prepared using Xper2 programme (LIS 2010). Xper2 is a plate-form
dedicated to taxonomic descriptions and computer-aided identification. Descriptions of species were entered into
the database creating 52 characters with variable numbers of character states. Numerous characters are missing in
some species description as authors do not always follow the same standard. Using the programme we chose 16
characters with the highest Xper2 discriminant power (0.70 to 0.88) to prepare the table. This discriminant index is
calculated as the ratio of the number of couples of species with no common values by the total number of couples
(LIS 2010). An index close to the value 1.0 means a good discriminant power whereas low values indicate feeble
discriminant power of a character.
Nomenclatural terms follow Dubois (2000, 2005). They are employed to avoid using the term “type” and terms
derived from it. This term carries a “typological” meaning, a confusion which should be avoided. The function of
the name-bearers is not to define any systematic taxon (group of animals), but to objectively allocate names
(nomina) to taxa recognized by taxonomists.
A BLUE-EYED LEPTOBRACHIUM FROM INDIA
Zootaxa 2912 © 2011 Magnolia Press ·
29
TABLE 1. Measurements of the holophoront of Leptobrachium bompu.
Variable
mm
Variable
mm
EL
6.5
IUE
5.4
EN
4.8
MBE
5.0
FFTF
9.6
MFE
11.0
FL
24.4
MN
13.5
FLL
12.2
MTFF
1.8
FOL
19.8
MTTF
4.3
FTL
8.7
NS
5.5
Fw3
1.2
SL
7.6
HAL
12.8
SVL
47.0
HL
16.9
TFL
6.4
HW
18.8
TFOL
27.3
IBE
14.0
TFTF
10.4
IFE
6.3
TL
20.9
IMT
3.0
TW
5.7
IN
4.7
Tw4
1.3
ITL
3.4
UEW
3.9
Leptobrachium bompu n. sp.
Material examined. One adult male, holophoront (holotype), no. KA0001/200905, locality: Bompu (27º06’61” N,
92º 40’64” E), altitude of 1940 m, coll. by Sanjay Sondhi on 22 May 2009 at 2000 hours, deposited in the State
Forest Research Institute, Itanagar, Arunachal Pradesh. A tissue sample of this specimen has been taken for molecular phylogenetic analysis.
Two other frogs caught and examined but not collected, showed little variation from the holophoront in terms
of size and coloration.
Etymology. This species is named after the type locality that it was discovered in, i.e. Bompu at Eaglenest. By
naming it after the location it was found in, it is hoped that the local community in the area will continue with their
significant efforts to conserve the forests in the area. The specific epithet is a noun in apposition, thus invariable.
Diagnosis. Leptobrachium bompu was placed in the genus Leptobrachium because of the following diagnosis:
femoral gland present; oval flat axillary glands on flanks posterior to arm insertion present; rictal glands and ventrolateral glandular ridges absent; inner palmar tubercle circular of similar size to outer tubercle; vomerine teeth
absent; snout and dermal palpebral projections absent; skin with fine ridges forming reticulum. It can be distinguished from the other species of Leptobrachium by the combination of the following characters; spines on upper
lip absent in adult males; head wider than long; tympanum indistinct; second finger shortest; finger tips swollen;
tibia longer than 40 % of snout-vent length; toe tips swollen; webbing leaving 3 ½ phalanges of toe IV and 2 phalanges of toe V free; length of inner metatarsal tubercle almost equals first toe length; dorsal pattern with irregular
distinct darker markings; ventrum brown (in alcohol), dark purple (in life) with small white spots; iris of uniform
grey-blue coloration; sclera black. The new species differs from all congeners by a combination of various morphological characters, besides its body and eye coloration; for comparisons, see Table 2.
Onymotope. Bompu (27°06’61” N, 92°40’64” E, altitude of 1940 m) in the Eaglenest Wildlife Sanctuary,
West Kameng district, Arunachal Pradesh, India.
Description of the holophoront (Figure 1–2, Table 1): Size and general aspect: (1) Specimen of moderate
size (SVL 47.0 mm), body rounded.
30 · Zootaxa 2912 © 2011 Magnolia Press
SONDHI & OHLER
FIGURE 1. Images of Leptobrachium bompu in life. a. Dorso-lateral view, b. Dorsal view, c. Ventro-lateral view.
Head: (2) Head large, wider (HW 18.8 mm) than long (HL 16.9 mm; MN 13.5 mm; MFE 11.0 mm; MBE 5.0
mm), flat. (3) Snout rounded, not protruding; its length (SL 7.6 mm) longer than horizontal diameter of eye (EL 6.5
mm). (4) Canthus rostralis distinct, not prominent, loreal region flat; obtuse in cross section. (5) Interorbital space
flat, larger (IUE 5.4 mm) than upper eyelid (UEW 3.9 mm) and than internarial distance (IN 4.7 mm); distance
between front of eyes (IFE 6.3 mm) about two fifth of distance between back of eyes (IBE 14.0 mm). (6) Nostrils
rounded, without flap of skin; slightly closer to eye (EN 4.8 mm) than to tip of snout (NS 5.5 mm). (7) Pupil oval,
vertical. (8) Tympanum indistinct, not measured. (9) Pineal ocellus absent. (10) Vomerine ridge absent. (11)
Tongue large, cordate, emarginate; median lingual process absent. (12) Supratympanic fold prominent from posterior corner of eye to forelimb.
31
FIGURE 2. Images of Leptobrachium bompu holophoront, KA0001/200905. a. Dorso-lateral view; b. Dorsal view; c. Ventral
view.
Forelimbs: (13) Arm long, thin; fore-arm (FLL 12.2 mm) about as long as hand (HAL 12.8 mm), not enlarged.
(14) Fingers I, II and IV short, thin, subequal in length; finger III long, thin (TFL 6.4 mm). (15) Relative length,
shortest to longest: II < I < IV < III. (16) Tips of fingers I to IV rounded, slightly enlarged, without grooves. (17)
Fingers with thick dermal fringes; webbing absent. (18) Subarticular tubercles absent, but callous ridges beneath all
fingers and metacarpus present. (19) Prepollex rounded, prominent; a single, round, prominent palmar tubercle;
supernumerary tubercles absent.
Hindlimbs: (20) Shanks rather long (tibiotarsal articulation reaching posterior corner of eye), four times longer (TL 20.9 mm) than wide (TW 5.7 mm), shorter than thigh (FL 24.4 mm) but longer than distance from base of
internal metatarsal tubercle to tip of toe IV (FOL 19.8 mm). (21) Toes short, thin, toe IV (FTL 8.7 mm) longer than
third of distance from base of tarsus to tip of toe IV (TFOL 27.3 mm). (22) Relative length of toes, shortest to longest: I < II < V < III < IV. (23) Tips of toes rounded, slightly enlarged, without grooves. (24) Webbing small: I 1 –
2 II 1 – 2½ III 1 – 3½ IV 3½ – 2 V (MTTF 4.3 mm; MTFF 1.8 mm; TFTF 10.4 mm; FFTF 9.6 mm). (25) Dermal
fringe present along the entire length of toe V (26) Subarticular tubercles absent, but callous ridges beneath all toes
present. (27) Inner metatarsal tubercle long, distinct, its length (IMT 3.0 mm) is 0.9 times the length of toe I (ITL
3.4 mm). (28) Tarsal fold absent. (29) Outer metatarsal tubercle absent; supernumerary tubercles and tarsal tubercle
absent.
Skin: (30) Dorsal and lateral parts of head and body: snout, between eyes, side of head, back and flank covered
with fine ridges forming reticulum. (31) Dorsal parts of limbs: covered with longitudinal folds. (32) Ventral parts of
head, body and limbs: covered with dense, small glandular warts, coarser on throat. (33) Macroglands: pectoral
glands small, rounded, on side of chest, posterior to arm insertion; small white femoral gland, on posterior shank,
closer to knee than to vent.
SONDHI & OHLER
33
Coloration in alcohol. (34) Dorsal and lateral parts of head and body: brown with irregular distinct darker
markings; loreal region lighter brown with dark bands; a dark spot on canthus near nostril; tympanic region brown
with a dark line under tympanic fold; upper lip continuous to canthal and tympanic region. (35) Dorsal parts of
limbs: brown with dark brown narrow bands; posterior part of thigh light brown with reduced white spotting. (36)
Ventral parts of head, body and limbs: throat, margin of throat and chest light brown; belly brown with white spots;
webbing dark brown.
Coloration in life. (37) Dorsum and head greyish-brown getting lighter on flanks; temporal region dark greyish-brown; sides of head and body with brownish shine, indistinct in natural light, but quite visible in artificial
light; upper and lower lip whitish; upper side of forearms and legs grey with darker transversal bands; ventral surface of belly and legs dark purple with small white spots on glandular warts. Iris entirely greyish-blue with vertical
black pupil.
Male secondary sexual characters. (38) Vocal sacs: not distinct externally; openings as small slits, positioned
near the proximal part of the tongue, surrounded by skin folds. (39) Nuptial spines absent.
Ecology and behaviour. The species was observed on 21, 22 and 26 May 2009 after sundown (1830 to 2000
hours). The weather was cloudy and it was raining heavily during the study period. On the first and second day, the
frogs were heard calling upstream, approximately 150 m from the road, but on the third visit, after very heavy rainfall, the frog's calls were heard further downstream only. It is possible that the frogs had moved downstream with
the force of the water or because suitable microhabitats were no longer available.
The frogs were passive and allowed themselves to be picked up without a struggle. They sat in a crouch. When
disturbed or agitated, they moved or rather crawled laboriously. Three separate specimens were found and photographed. One of these specimens was collected on 22 May and is described here as the holophoront.
Vocalisation. The loud call of the male is a croaking “kek-kek-kek-kek”. The frogs continued calling at night
when in the bag.
Discussion
Arunachal Pradesh ranks second in India for the fraction of state’s land area with 81.2% of its total geographic area
under forest cover (FSI 2003). It has a steep topology with a wide altitudinal range, a gradient in precipitation
extending to very high rainfall, largely intact forests, and a special location at the junction of the Palearctic and the
Indo-Malayan biogeographic realms (in fact Arunachal is also at the junction of the Indian and Indo-Burmese subregions) (Athreya 2006). Bordered by Bhutan on the west, China on the north and east, Myanmar on the south-east
and the state of Assam on the south—all places known for their rich herpetofauna—we can expect to find new species with distributions and ranges beyond our current knowledge. The Eaglenest Wildlife Sanctuary is in the West
Kameng district of western Arunachal Pradesh. During the Eaglenest Biodiversity project undertaken by Ramana
Athreya between 2003 and 2006 (http://www.iiserpune.ac.in/~rathreya/documents.html), many new taxa were
described. The highlight was the description of the Bugun Liocichla Liocichla bugunorum, the first new bird species to be described from India in more than fifty years (Athreya 2006). Amongst herpetofauna, the Dafla Mountain Agamid Mictopholis austeniana, known from a type specimen collected by Col. Godwin-Austen during the
Dafla expedition in 1874 was re-discovered at Eaglenest and Sessa Orchid wildlife sanctuaries (Agarwal et al.
2010). The Darjeeling False-wolfsnake Dinodon gammiei known to science from only 5 specimens from Darjeeling, West Bengal was discovered here (Agarwal et al. 2010). While conducting a herpetofaunal survey at Eaglenest
during 2006–2008, four Indian researchers reported the first Indian record of the Jerdon’s Red Spotted Pitviper
Protobothrops jerdonii xanthomelas (Zambre et al. 2009). These findings demonstrate that the area's herpetofauna
is not well studied and new discoveries are only to be expected.
The spot where the frogs were located was near a campsite called Bompu in the Eaglenest Wildlife Sanctuary.
The frogs were at a stream approx. 1.14 km north-west of Bompu, at an altitude of 2000 m, close to the only road in
the forest. The forest type at Bompu transitions between East Himalayan subtropical wet hill forest 8B/C1 at lower
altitudes and East Himalayan wet temperate forest 11B/C1 at higher altitudes (forest types as per Champion et al.
1968).
The holophoront and two other frogs were found under leaf litter at a small, slow-flowing perennial stream,
close to the road. There were more frogs calling both up and downstream. The frogs were only heard calling from
the area close to the stream, at an estimated distance of no more than 150 meters upstream and downstream.
SONDHI & OHLER
Numerous species in Leptobrachium have coloured eyes, and this feature is useful to their identification but it
is not sufficient (Ohler et al. 2004; Matsui et al. 2010). The only species in Leptobrachium without spines that has
entirely blue eyes is the recently described Leptobrachium waysepuntiense from Sumatra, Indonesia.
The new species Leptobrachium bompu differs from L. waysepuntiense in the coloration in life: L. bompu lacks
the black reticulations in the blue eye, it has black bands on the limbs, hands, feet, digits and upper lip, and has
irregular dark markings on the dorsal surface, which are all absent in L. waysepuntiense. L. bompu lacks the Vshaped brownish orange inter-orbital and parietal marking and the white and orange spotting on the lateral and ventral surfaces, as well as on the toes. Leptobrachium bompu also has distinctly wrinkled skin, while L. waysepuntiense has nearly smooth skin.
Many Leptobrachium species have the upper third of their eye brightly coloured, while the rest of the eye is
dark coloured (Table 2). Leptobrachium smithi, which is reported from the Indian region has its upper third of the
eye scarlet or yellow (Matsui et al. 1999). The other species with eyes that are scarlet or yellow are L. pullum and
L. hendricksoni (Ohler et al. 2004). Leptobrachium abbotti, L. chapaense, L. gunungense, L. hasseltii, L. lumadorum, L. mangyanorum, L. montanum, L. mouhoti, L. nigrops and L. tabanorum have black, dark brown or dark grey
eyes (Hamidy & Matsui 2010). Leptobrachium lumadorum, L. mangyanorum and L. tabanorum have a light blue
sclera. L. mouhoti has black eyes with an orange-red sclera (Stuart et al. 2006). Leptobrachium hainanense and L.
huashen have the upper third of the eye blue, while the rest of the eye is black, quite unlike the species described
here. L. buchardi has upper third of the eye pale green. Leptobrachium banae, L. guangxiense and L. xanthospilum
have white coloured eyes (Table 2).
The seven species of Leptobrachium with males bearing horny spines are Leptobrachium ailaonica, L. boringii, L. echinata, L. liui, L. liushanense, L. ngoclinhense and L. promustache. The new species does not show such
spines. In all these species, males are much larger than in the species described here. When compared with L. promustache (Table 2), the holophoront of L. bompu looks similar in coloration, has similar markings, and similar
morphological characters, such as the metatarsal tubercles on the hind feet. However, L. promustache has a black
eye with a white line on the upper eyelid, and unlike L. bompu, it does not have a greyish-blue eye. The males of L.
promustache are much larger than the holophoront (62 mm vs 47 mm) and all other measurement exceed those of
the holophoront. Finally, L. promustache has white spines or sockets on the upper lip (Rao et al. 2006; Bain et al.
2009), which L. bompu lacks. Furthermore, no Leptobrachium species, other than L. waysepuntiense has an
entirely greyish-blue eye. In addition to coloration, there are numerous other characters which differentiate L.
bompu from each nominal species of Leptobrachium (Table 2). Hence this frog is a new species, Leptobrachium
bompu.
Acknowledgements
SS thanks the Eaglenest Biodiversity Project initiated by Ramana Athreya (Kaati Trust, and Indian Institute of Science Education and Research (IISER, Pune) for logistical support which made this work possible in the remote wilderness of Eaglenest. We would like to acknowledge the Arunachal Forest Department for their support and
permission to conduct a long term study and collect specimens (vide letter reference No. CWL/G/13 (17)/06-07/1214). SS would also like to acknowledge Dr. Karthikeyan Vasudevan's (Wildlife Institute of India, Dehradun) assistance in reading through the manuscript draft and assisting with examining the specimen in the lab. He acknowledges Shashank Dalvi who helped preserve the specimen in the field. Thanks are also due to Dr. Deepak Barua,
Sachit, Goswami and Reddy of IISER, Pune, who helped with measurements of the specimens in the field. Big
thanks to his family, wife Anchal and son, Yash, who wholeheartedly allow his nocturnal sojourns. Last, but not the
least, Indi Glow and his support staff at Eaglenest Wildlife Sanctuary, who helped to provide access remote areas
like Bompu, even in the monsoons.
References
Ahmed, M.F., Das, A. & Dutta, S.K. (2009) Amphibians and Reptiles of North East India-A Photographic Guide. Aaranyak,
Assam, pp. 13.
Agarwal, I., Mistry, V.K, & Athreya, R. (2010) A preliminary checklist of the reptiles of Eaglenest Wildlife Sanctuary, West
Kameng District, Arunachal Pradesh. Russian Journal of Herpetology, 17, 81–93.
35
Athreya, R. (2006) A new species of Liocichla (Aves: Timaliidae) from Eaglenest Wildlife Sanctuary, Arunachal Pradesh,
India. Indian Birds, 2, 82—94.
Athreya, R. (2006) Eaglenest Biodiversity Project (2003–2006). Conservation Resources for Eaglenest Wildlife Sanctuary.
Kaati Trust, Pune.
Bain, H.R., Truong, N. & Doan, K.V. (2009) First record of Leptobrachium promustache from Vietnam. Herpetology Notes, 2,
27—29.
BCPP CAMP Report. (1998) Amphibians of India. Zoo Outreach Organisation, India, p. 8, 14.
Bonaparte, C.L. (1850) Conspectus Systematum Herpetologiae et Amphibiologiae. Brill, Lugduni Batavorum, 1 plate.
Brown, R.N., Siler, C.D., Diesmos, A.C. & Alcala, A.C. (2009) Philippine frogs of the genus Leptobrachium (Anura; Megophryidae): Phylogeny-based species delimitation, taxonomic review, and descriptions of three new species. Herpetological
Monographs, 23, 1–44.
Champion, F.W. & Seth, S.K. (1968) A Revised Survey of the Forest Types of India. Nasik, India: Manager. Government of
India Press, Delhi, India, 404 pp.
Chanda, S.K. (2002) Handbook-Indian Amphibians. Zoological Survey of India, Kolkata, 7 pp.
Das, I. & Chanda, S.K. (2004) Leptobrachium smithi Matsui, Nabitabhata, and Panha, 1999 (Anura: Megophryidae), an addition to the fauna of Myanmar (Burma). Asiatic Herpetological Research, 10, 245—246.
Dubois, A. (2000) Synonymies and related lists in zoology: general proposals, with examples in herpetology. Dumerilia, 4, 33–
98.
Dubois, A. (2005) Proposed Rules for the incorporation of nomina of higher-ranked zoological taxa in the International Code
of Zoological Nomenclature. 1. Some general questions, concepts and terms of biological nomenclature. Zoosystema, 27,
365–426.
Dubois, A. & Ohler, A. (1998) A new species of Leptobrachium (Vibrissaphora) from northern Vietnam, with a review of the
taxonomy of the genus Leptobrachium (Pelobatidae, Megophryinae). Dumerilia, 4, 1–32.
Duellman, W.E. & Trueb, L. (1985) Biology of amphibians. New York, McGraw-Hill, “1986”, xix + 670 pp.
Fei, L., Hu, C., Ye, C.-Y. & Huang, Y. (2009) Fauna Sinica. Amphibia. Volume 2. Anura. Beijing: Chinese Academy of Science. Science Press.
Frost, D.R. (2010) Amphibian Species of the World: an Online Reference. Version 5.4 (8 April, 2010). Electronic Database
accessible at http://research.amnh.org/vz/herpetology/amphibia/American Museum of Natural History, New York, USA.
(accessed 3 December 2010)
Forest Survey of India (2003) Website at http://www.fsi.nic.in/sfr_2003.htm. (Accessed 3 December 2010)
Hamidy, A. & Matsui, M. (2010) A new species of blue-eyed Leptobrachium (Anura: Megophryidae) from Sumatra, Indonesia.
Zootaxa, 2395, 34—44.
LIS (2010) Xper²: an open tool to manage descriptive data! http://lis-upmc.snv.jussieu.fr/lis/?q=en/resources (accessed 10
December 2010)
Liu, C.-c. (1945) New frogs from West China. Journal of the West China Border Research Society, Series B, 15, 28–44.
Matsui, M., Nabhitabhata, J. & Panha, S. (1999) On Leptobrachium from Thailand with a description of a new species (Anura:
Pelobatidae). Japanese Journal of Herpetology, 18, 19–29.
Matsui, M., Hamidy, A., Murphy, R.W., Khonsue, W., Yambun, P., Shimada, T., Ahmad, N., Belabut, D.M. & Jiang, J.-P.
(2010) Phylogenetic relationships of megophryid frogs of the genus Leptobrachium (Amphibia, Anura) as revealed by
mtDNA gene sequences. Molecular Phylogenetics and Evolution, 56, 259–272.
Ohler, A., Teynie, A. & David, P. (2004) A green-eyed Leptobrachium (Anura: Megophryidae) from Southern Laos. The Raffles Bulletin of Zoology, 52, 696—700.
Rao, D.,Wilkinson, J.A. & Zhang, M. (2006) A new species of the genus Vibrissaphora (Anura: Megophryidae) from Yunnan
Province, China. Herpetologica, 62, 90—95.
Reza, A.H.M.A. (2009) Leptobrachium smithi (Smith's Litter Frog). Defense and Pigmentation. Herpetological Review, 40, 1.
Sengupta, S., Choudhury N.K. & Das, I. (2001) Leptobrachium smithi Matsui, Nabhitabhata and Panha, 1999 (Anura: Megophryidae), a new record for India. Journal of the Bombay Natural History Society, 98, 289—291.
Stuart, B.L., Sok, K. & Neang, T. (2006) A collection of amphibians and reptiles from hilly eastern Cambodia. The Raffles Bulletin of Zoology, 54, 129—155.
Tschudi, J.J. (1838) Classification der Batrachier, mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien.
Petitpierre, Neuchâtel, ii + 98 pp., 6 pl.
Zambre, A., Sheth, C., Dalvi, S. & Kulkarni, N. (2009) First record of Protobothrops jerdonii xanthomelas (Günther, 1889)
from Eaglenest Wildlife Sanctuary, India. Journal of the Bombay Natural History Society, 106, 325—327.
Zheng,Y., Li, S. & Fu, J. (2008) A phylogenetic analysis of the frog genera Vibrissaphora and Leptobrachium, and the correlated evolution of nuptial spine and reversed sexual size dimorphism. Molecular Phylogenetics and Evolution, 46, 695—
707.
Zug, G., Slowinski, J. & Wogan, G. (2001) The herpetofauna of Myanmar. Checklist of the Amphibians and Reptiles of Myanmar. California Academy of Sciences. http://research.calacademy.org/redirect?url=http://researcharchive.calacademy.org/
research/herpetology/myanmar/project.html (Accessed 10 December 2010).
SONDHI & OHLER

A blue-eyed Leptobrachium (Anura: Megophryidae)

Transcription

Similar documents

Lichens new to the Azores collected on Terceira