Research Article |
Corresponding author: Ramesh Aggarwal ( rameshka9@gmail.com ) Academic editor: Franco Andreone
© 2016 S. R. Chandramouli, Karthikeyan Vasudevan, S Harikrishnan, Sushil Kumar Dutta, S Jegath Janani, Richa Sharma, Indraneil Das, Ramesh Aggarwal.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Chandramouli SR, Vasudevan K, Harikrishnan S, Dutta SK, Janani SJ, Sharma R, Das I, Aggarwal RK (2016) A new genus and species of arboreal toad with phytotelmonous larvae, from the Andaman Islands, India (Lissamphibia, Anura, Bufonidae). ZooKeys 555: 57-90. https://doi.org/10.3897/zookeys.555.6522
|
A new bufonid amphibian, belonging to a new monotypic genus, is described from the Andaman Islands, in the Bay of Bengal, Republic of India, based on unique external morphological and skeletal characters which are compared with those of known Oriental and other relevant bufonid genera. Blythophryne gen. n. is distinguished from other bufonid genera by its small adult size (mean SVL 24.02 mm), the presence of six presacral vertebrae, an absence of coccygeal expansions, presence of an elongated pair of parotoid glands, expanded discs at digit tips and phytotelmonous tadpoles that lack oral denticles. The taxonomic and phylogenetic position of the new taxon (that we named as Blythophryne beryet gen. et sp. n.) was ascertained by comparing its 12S and 16S partial genes with those of Oriental and other relevant bufonid lineages. Resulting molecular phylogeny supports the erection of a novel monotypic genus for this lineage from the Andaman Islands of India.
Amphibian, bufonid, tadpole, rRNA, molecular phylogeny, skeletal characters
Neobatrachian anurans of the family Bufonidae Gray, 1845 are represented in the Oriental portion of Asia by 14 genera (Table
Members of the Neobatrachian anurans of the family Bufonidae Gray, 1845 represented in the Oriental portion of Asia.
Genus | Number of species | Distribution | |
---|---|---|---|
1 | Adenomus Cope, 1860 | 2 | Sri Lanka |
2 | Ansonia Stoliczka, 1870 | 28 | Sundaland and Philippine archipelago |
3 | Bufoides Pillai & Yazdani, 1973 | 1 | Khasi Hills, Meghalaya, India |
4 | Duttaphrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green & Wheeler, 2006 | 29 | Eastern Africa to Papua New Guinea; 25 species are known from India and south east Asia |
5 | Ingerophrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green & Wheeler, 2006 | 12 | Indochina and the Sundaland |
6 | Leptophryne Fitzinger, 1843 | 2 | Sundaland |
7 | Parapelophryne Fei, Ye & Jiang, 2006 | 1 | Indochina |
8 | Pedostibes Günther, 1876 “1875” | 5 | Western Ghats and Eastern Himalayas, India; Malay Peninsula, Borneo & Sumatra |
9 | Pelophryne Barbour, 1938 | 11 | Sundaland and the Philippines Archipelago |
10 | Phrynoidis Fitzinger, 1843 | 2 | Indochina-Sundaland |
11 | Pseudobufo Tschudi, 1838 | 1 | Sundaland |
12 | Sabahphrynus Matsui, Yambun & Sudin, 2007 | 1 | Borneo |
13 | Xanthophryne Biju, Van Bocxlaer, Giri, Loader & Bossuyt, 2009 | 2 | Northern Western Ghats, India |
14 | Ghatophryne Biju, Bocxlaer, Giri, Loader & Bossuyt, 2009 | 2 | Western Ghats, India |
Of all the above, Duttaphrynus melanostictus (Schneider, 1799) is the only bufonid reported from the Andaman Islands (Sarkar 1990;
Specimens were hand-collected, euthanised and fixed in absolute ethanol for a minimum of 24 hours, and eventually transferred to 60% ethanol for preservation. Tissue samples were extracted and stored in absolute ethanol (prior to specimen fixation) for phylogenetic analyses. Tadpoles were collected and reared for preservation of samples across developmental stages in 4% formalin solution. Conspecificity between tadpoles and the adults was confirmed by rearing them to metamorphosis, as well as matching 16S ribosomal DNA sequences to those of the adults. Staging of tadpoles follow
Skeletal characters of a paratype were examined under a microscope by clearing using trypsin and 0.5% potassium hydroxide solution and staining with alcian blue and alizarin red dye, following
Tadpoles were described based on collections made in May 2011, from a phytotelm, located ca. 1.3 m above the ground. The clutch was monitored continuously till complete transformation. The observed eggs got transformed into pale white embryos on 2 May 2011; subsequently, tadpoles at different developmental stages were collected and preserved in 5% formalin. Tail tips of these individuals were collected and preserved in absolute ethanol for DNA barcoding studies before the tadpoles were preserved in formalin.
Abbreviations and definitions of morphometric measurements made on adult, metamorph of frogs and tadpole. Measurements made only on tadpoles are indicated by an asterisk after the abbreviation.
AG | Distance from posterior point of the forelimb at its insertion into the body to the anterior point of the hindlimb insertion |
BL* | Distance from snout tip to the point of the initiation of tail from the body |
BW | Distance at the broadest point at the trunk |
DFH* | Height of the fin measured at the place of the maximum height of the dorsal fin |
DL fold | Length of dorso-lateral fold |
ED | Horizontal diameter of the orbit |
EN | Distance between anterior border of the eye to posterior edge of the nostril |
ES | Distance between anterior border of the eye to the snout tip |
ETY | Distance between posterior border of the eye to anterior margin of the tympanum |
f1 to 4 | Distance measured from the fork of the fingers to the tip of the finger disc for fingers 1 to 4 |
FEL | Distance measured from the cloaca to the tip of the knee |
FOL | Distance measured from the anterior end of the tarsus to the tip of the fourth toe |
HD | Height of the head measured at the post-orbital region before the parotoid gland |
HL | Distance from the tip of the snout to the posterior edge of the mandible |
HW | Width of the head measured at the jaw angle |
IN | Closest distance between the nares |
IND* | Distance between the external nares |
IO | Distance between the anterior margins of the upper eyelids |
IOL* | Distance between the two orbits |
LAL | Distance measured from the elbow to the base of the outer metacarpal tubercle; palm length |
MBW* | Distance measured at point of the maximum width of the body |
MTH* | Distance measured at the point of the maximum height of the tail by laterally positing the tadpole |
MTMW* | Distance measured on the tail at the point of initiation of the tail from the body where the tail width is maximum |
NA | Not measured |
NED* | Distance between nostril and eye |
NSD* | Distance from the snout to the eye |
ODD* | Oral disc diameter |
PAL | Distance measured from the posterior border of the outer metacarpal tubercle to the tip of the third finger |
PL | Length of the parotoid gland |
PW | Maximum width of the parotoid gland |
SS* | Distance from snout to the spiracle |
SV* | Distance from snout to the vent |
SVL | Distance from tip of the snout till the cloaca |
t1 to 5 | Distance measured from the fork of the toe to the tip of the toe disc for toes 1 to 5 |
TBL | Distance from the knee to the obtuse margin of the tibia |
TL* | Distance from the point of initiation of tail till the tip of the tail |
TMH* | Distance measured on the tail at the point where the tail muscle reaches maximum height |
TYH | Horizontal diameter of the tympanum |
TYV | Vertical diameter of the tympanum |
UAL | Distance measured from the point of insertion of the forelimb to the trunk to elbow |
UEW | Maximum width of the upper eyelid |
VFH* | Ventral fin height measured at the place of the maximum height of the ventral fin |
VTL* | Vent tube length |
Molecular phylogeny. Total genomic DNA was extracted from the alcohol-preserved soft tissue (muscle), taken from the holotype, following the standard procedure of SDS & proteinase-K lysis, followed by chloroform-isoamyl extraction method. The taxonomic position of the toad was ascertained by rDNA typing of both 16S and 12S rDNA genes of the mitochondrial genome broadly following the method as described earlier by
Taxon sampling for phylogenetic analysis of selected Oriental members of the Bufonidae.
Taxon | Range/ Collection location | NCBI Acc. No. | Tree_7a | Subtree_7b | Subtree_7c | Reference |
---|---|---|---|---|---|---|
Blythophryne beryet gen. et sp. n. | India (A&N Islands) | KT991336, KT991347 | + | + | + | This study |
Adenomus kelaartii | Sri Lanka | FJ882780 | + | + | + |
|
Amietophrynus brauni | Tanzana | DQ158437 | + |
|
||
Amietophrynus gracilipes | Equatorial Guinea | DQ158456 | + |
|
||
Amietophrynus gutturalis | Kenya | DQ158460 | + |
|
||
Amietophrynus poweri | Namibia | DQ158482 | + |
|
||
Amietophrynus steindachneri | Kenya | DQ158488 | + |
|
||
Ansonia hanitschi | Malaysia | FJ882794 | + |
|
||
Ansonia longidigita | Malaysia | KT991329, KT991340 | + | This study | ||
Bufo bufo | Turkey | DQ158438 | + |
|
||
Bufoides meghalayanus | India | KT991331, KT991342 | + | + | + | This study |
Duttaphrynus atukoralei | India | FJ882835 | + | + |
|
|
Duttaphrynus brevirostris | India | FJ882786 | + | + |
|
|
Duttaphrynus crocus | India | FJ882789 | + | + |
|
|
Duttaphrynus dhufarensis | India | FJ882837 | + | + |
|
|
Duttaphrynus himalayanus | India | KT991334, KT991345 | + | + | + | This study |
Duttaphrynus hololius | India | FJ882781 | + | + |
|
|
Duttaphrynus melanostictus | India | KT991335, KT991346 | + | + | + | This study |
Duttaphrynus parietalis | India | FJ882784 | + | + |
|
|
Duttaphrynus scaber | India | KT991332, KT991343 | + | + | + | This study |
Duttaphrynus stomaticus | India | KT991333, KT991344 | + | + | + | This study |
Duttaphrynus stuarti | India | FJ882788 | + | + |
|
|
Ghatophryne ornata | India | FJ882797 | + | + |
|
|
Ingerophrynus divergens | Malaysia | KT991328, KT991339 | + | This study | ||
Ingerophrynus galeatus | Laos | DQ158452 | + |
|
||
Ingerophrynus macrotis | Laos | DQ158468 | + |
|
||
Leptophryne borbonica | Malaysia | FJ882799 | + |
|
||
Mertensophryne micranotis | Tanzania | FJ882821 | + |
|
||
Mertensophryne uzunguensis | Tanzania | FJ882819 | + |
|
||
Nectophryne afra | Cameroon | DQ283360 | + |
|
||
Nectophryne batesi | Gabon | DQ283169 | + |
|
||
Nectophrynoides minutus | Tanzania | FJ882814 | + |
|
||
Nectophrynoides tornieri | Tanzania | DQ283413 | + |
|
||
Pedostibes hosii | Malaysia | KT991330, KT991341 | + | This study | ||
Pedostibes tuberculosus | India | FJ882793 | + | + |
|
|
Pelophryne api | Malaysia | KT991326, KT991337 | + | This study | ||
Phrynoidis asper | Brunei | DQ158431 | + |
|
||
Phrynoidis juxtasper | Malaysia | KT991327, KT9913387 | + | This study | ||
Sabahphrynus maculatus | Malaysia | AB331718 | + |
|
||
Schismaderma carens | Zimbabwe | DQ158424 | + |
|
||
Vandijkophrynus robinsoni | Namibia | GU183857 | + |
|
||
Xanthophryne koynayensis | India | FJ882782 | + | + | + |
|
Rhaebo guttatus | Brazil | DQ158459 | + |
|
For each of the phylogenetic analysis, the concatenated 12S+16S sequence alignment was first used to find the best fitting DNA substitution model using Akaike Information criterion (AIC), as implemented in jModelTest2 (
Blythophryne beryet gen. et sp. n. by monotypy (Fig.
Morphometric measurements of the holotype and paratype series of adult and two gravid (g) individuals of Blythophryne beryet gen. et sp. n.
ZSI A-12521 | ZSI A-12524 | ZSI A-12522 | ZSI A-12523 | ZSI A-12526 | ZSI A-12529 | ZSI A-12527 | ZSI A-12530 | ZSI A-12528 | ZSI A-12525 | |
---|---|---|---|---|---|---|---|---|---|---|
Sex | ♀ | ♀(g) | ♂ | ♀(g) | ♂ | ♂ | ♀ | ♂ | ♂ | ♂ |
SVL | 27.4 | 25.5 | 25.5 | 25.2 | 24.5 | 23.0 | 22.7 | 22.3 | 22.2 | 21.8 |
AG | 10.6 | 9.2 | 9.8 | 12.5 | 8.0 | 7.3 | 8.0 | 6.7 | 6.5 | 8.5 |
HL | 7.7 | 7.5 | 8.2 | 6.9 | 7.5 | 7.9 | 7.5 | 7.6 | 7.6 | 7.1 |
HW | 7.9 | 7.6 | 8.1 | 6.8 | 8.0 | 7.8 | 7.6 | 7.7 | 7.4 | 7.2 |
HD | 4.3 | 3.5 | 3.9 | 3.4 | 3.9 | 3.4 | 3.2 | 3.2 | 3.0 | 3.2 |
BW | 9.9 | 10.3 | 9.1 | 11.8 | 8.3 | 7.1 | 6.1 | 7.5 | 6.3 | 9.8 |
EN | 2.2 | 2.3 | 1.7 | 1.9 | 1.9 | 2.3 | 2.1 | 2.0 | 2.1 | 2.2 |
ES | 3.4 | 3.4 | 3.3 | 3.1 | 3.5 | 3.3 | 3.2 | 3.2 | 3.3 | 3.1 |
ETY | 0.5 | 0.7 | 0.7 | 0.7 | 0.7 | 0.7 | 0.7 | 0.6 | 0.6 | 0.5 |
UEW | 1.9 | 1.5 | 2.0 | 1.8 | 1.8 | 1.7 | 1.7 | 1.5 | 1.9 | 1.8 |
IO | 3.8 | 3.8 | 3.5 | 3.4 | 3.5 | 3.8 | 3.4 | 3.5 | 3.6 | 3.4 |
IN | 2.2 | 2.1 | 2.2 | 2.2 | 1.8 | 2.1 | 2.1 | 1.9 | 2.0 | 1.9 |
TYH | 1.6 | 1.6 | 1.9 | 1.6 | 1.6 | 1.4 | 1.5 | 1.4 | 1.8 | 1.5 |
TYV | 1.8 | 1.7 | 1.9 | 1.6 | 1.6 | 1.6 | 1.6 | 1.5 | 1.8 | 1.5 |
UAL | 5.1 | 4.7 | 4.4 | 5.2 | 4.3 | 4.3 | 4.3 | 4.6 | 4.3 | 4.1 |
LAL | 5.8 | 5.4 | 5.6 | 5.5 | 5.6 | 5.5 | 5.4 | 5.3 | 5.4 | 5.3 |
PAL | 6.2 | 5.7 | 6.2 | 6.8 | 5.8 | 5.9 | 5.9 | 5.8 | 5.9 | 6.1 |
FEL | 9.2 | 7.5 | 7.7 | 7.2 | 9.3 | 8.2 | 9.5 | 8.2 | 8.6 | 8.5 |
TBL | 10.6 | 8.0 | 9.4 | 8.4 | 9.1 | 7.9 | 9.0 | 8.1 | 8.3 | 8.5 |
FOL | 9.6 | 9.7 | 9.4 | 8.3 | 9.4 | 8.0 | 9.2 | 8.3 | 8.7 | 8.3 |
ED | 2.8 | 2.5 | 2.4 | 2.6 | 2.5 | 2.1 | 1.9 | 2.4 | 2.1 | 2.3 |
DL fold | 13.3 | 12.2 | 11.3 | 11.9 | 12.0 | 12.3 | 11.7 | 12.0 | 11.4 | 11.9 |
PL | 6.1 | 5.9 | 6.5 | 6.0 | 4.5 | 3.7 | 4.0 | 5.9 | 3.2 | 3.9 |
PW | 1.4 | 1.4 | 1.3 | 1.6 | 1.0 | 1.0 | 0.9 | 1.3 | 0.9 | 0.9 |
f1 | 1.8 | 1.2 | 1.1 | 1.5 | 1.6 | 1.3 | 2.0 | 1.2 | 1.6 | 0.9 |
f2 | 1.9 | 1.4 | 1.6 | 1.7 | 2.2 | 1.8 | 2.2 | 1.4 | 1.9 | 1.6 |
f3 | 3.1 | 3.0 | 2.9 | 2.8 | 2.8 | 2.6 | 2.8 | 2.9 | 2.4 | 2.9 |
f4 | 2.2 | 1.9 | 1.8 | 2.1 | 2.2 | 1.9 | 2.1 | 1.8 | 1.6 | 1.8 |
t1 | 1.1 | 1.1 | 1.2 | 1.1 | 1.3 | 1.0 | 1.3 | 1.2 | 1.0 | 1.1 |
t2 | 1.4 | 1.7 | 1.4 | 1.4 | 1.7 | 1.4 | 1.5 | 1.1 | 1.5 | 1.5 |
t3 | 2.6 | 2.0 | 2.0 | 2.7 | 2.3 | 2.1 | 2.6 | 2.1 | 2.2 | 1.9 |
t4 | 4.7 | 4.1 | 3.9 | 4.9 | 4.4 | 2.9 | 4.6 | 3.7 | 3.0 | 4.0 |
t5 | 3.0 | 2.1 | 2.3 | 2.6 | 2.5 | 2.1 | 2.5 | 2.1 | 2.0 | 1.9 |
A single species is currently known.
Holotype ♀ : ZSI_A-12521(Fig.
Morphological characters of the Blythophryne beryet gen. et sp. n.: a dorso-lateral view b dorsal view c ventral view d ventral view of left palm e ventral view of left foot of the adult female holotype (ZSI_A-12521) in life f adult female holotype in preservation g dorsal view of the male paratype (ZSI_A-12529) in life showing inverted-V shaped markings and the inter-ocular band on the dorsum.
The generic name is a patronym, coined in appreciation of Edward Blyth (1810–1873), the first curator of the Asiatic Society of Bengal, who initiated herpetological studies in the Andaman and Nicobar Islands, through his phenomenal, pioneering paper “Notes on the fauna of the Nicobar islands” (Blyth 1846).
This currently monotypic genus and species is diagnosed by the following suite of external morphological and osteological characters: small adult size (mean SVL 24.0 mm; range 21.8–27.4 mm); distinct tympanum, slightly smaller than eye; absence of cephalic ridges; absence of vomerine teeth; presence of a single, median, external vocal sac in males; presence of elongated pair of parotoid glands; absence of enlarged, keratinised tubercles on dorsum; presence of well developed, sheath-like webbing on fingers and on toes; digit tips dilated to discs, lacking circum-marginal grooves; presence of six presacral vertebrae; urostyle lacking lateral dilations; absence of omosternum and presence of arciferal pectoral girdle. Mature ova small (0.62 mm mean diameter), yolky and unpigmented; tadpoles lacking keratodont.
A small bufonid (mean SVL 24.2 ± 0.6 mm), with depressed, moderately robust (AG:BW 1.0) habitus (Fig.
Dorsum reddish-brown, with two feeble dark brown inverted ‘V’ shaped markings which fail to reach flanks, interorbital band indistinct, canthus dark chocolate brown, colour extending a little beyond tympanum, subequal to half-length of parotoid gland; forearm and hind limbs barred, one each on thigh, shank and tarsus. Venter heavily speckled with dark brown spots, throat dark brown, lower lip spotted with white and brown, pupil large, horizontally elliptical.
Dorsum drab brown with indistinct ‘inverted-V’ shaped pattern, darker bands on limbs, venter cream, with black mottled pattern, throat black throughout (Fig.
(based on paratype ZSI_A12527). Axial and appendicular skeleton composed primarily of bony elements; cartilaginous elements not observed. Atlas (the first vertebra) with rudimentary hypapophysis and not fused to axis, presacral vertebrae six in number, Vertebrae II–V bearing horizontally elongate hypapophyses, those on Vertebrae II and V oriented anteriorly; Vertebrae III–IV oriented horizontally; sacral diapophysis laterally dilated; coccyx not fused to sacrum; articulating with former by a double condyle and lacking lateral expansions, omosternum absent, pectoral girdle arciferal, with epicoracoids united to each other anteriorly and overlapping posteriorly (Fig.
Adult females and males range between 25.2–27.4 mm and 21.8–25.5 mm, respectively. Measurements of paratypes are provided in Table
(Macaulay Library, Cornell Lab of Ornithology; voucher no: ML 174095). A calling male was observed on 24 November 2010 on the surface of leaves within bushes. Calls were composed of continuous syllables of “pip-pip-pip-pip-pip-” at a constant frequency of 8 kHz, without pause, lasting for 23 seconds, with mean amplitude of -3 db / 20 kU (Fig.
This species has been documented from five islands of the Andaman archipelago, namely, the South Andaman (Mt. Harriet), Rutland, Little Andaman, Havelock Island in the Ritchie’s Archipelago and North Andaman (Saddle Peak) (Fig.
‘Andaman bush toad’ is proposed as the common English name for this new species, indicating its arboreal habit and restricted distribution as understood currently.
The new species is often seen on surface of leaves of herbaceous bushes. It is nocturnal and regularly seen year round. It was the third most common anuran in the islands (
The Andaman bush toad emits a white, viscous, pungent smelling secretion from the parotoid glands when handled (Fig.
a Eggs and hatchling tadpoles of Blythophryne beryet gen. et sp. n. b, c endotrophic larvae of Blythophryne beryet gen. et sp. n. showing pale white abdominal yolk d Lateral view of a Stage 43 tadpole of Blythophryne beryet gen. et sp. n. e Oral disc of a Stage 35 larva of Blythophryne beryet gen. et sp. n., showing absence of keratodont and the presence of keratinised jaw sheaths f a metamorph of Blythophryne beryet gen. et sp. n. showing initiation of tail absorption.
The Andaman bush toad is widely distributed in islands where it occurs, and occupies forested habitats from 29–250 m asl, more common above 100 m asl and rarer at lower altitudes. The forest types in this elevation range include littoral, moist-deciduous, giant evergreen and montane stunted evergreen forests (
The Andaman bush toad is known from five islands: North Andaman (Saddle Peak National Park only), South Andaman, Rutland, Havelock (only in a small patch of wet forest towards the south of the island) and Little Andaman. Based on searches carried out using 21 bounded quadrats of 100 m2 each in these islands, the new species occurs at densities of 1.1 ± 0.37 toads per 100 m2 of forest floor (unpublished data). It is considered ‘Endangered’ based on IUCN Ver. 3.1. Second Edition (
(Fig.
Morphometric measurements of tadpoles of Blythophryne beryet gen. et sp. n.
Stage | IOL | IND | NED | NSD | SS | SV | BL | TL | MBW | MTH | MTMW | TMH | ODD | VTL | DFH | VFH |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
30 | 1 | 0.8 | 0.4 | 0.7 | 3.6 | 1.6 | 5 | 11.1 | 2.8 | 2.5 | 1.1 | 1.3 | 1.2 | 0.7 | 0.6 | 0.4 |
35 | 1.1 | 1.1 | 0.4 | 0.8 | 5 | 3.3 | 6.8 | 12.4 | 3.7 | 3.5 | 1.3 | 2 | 1 | 1.7 | 0.9 | 0.7 |
41 | 1 | 1 | 0.4 | 0.6 | 3.9 | 2.4 | 6.1 | 11.8 | 3.8 | 2.8 | 1.3 | 1.8 | 1.5 | 0.9 | 0.7 | 0.7 |
42 | 1.3 (±.20) | 1.1 | 0.4 (±.05) | 0.7 (±.10) | NA | NA | 7.3 (±.05) | 13.6 (±.05) | 3.5 (±.05) | 3.2 (±.10) | 1.8 (±.20) | 1.9 (±.15) | 1.3 (±.25) | NA | 0.9 (±.05) | 0.7 (±.05) |
43 | 1.5 (±.30) | 1.1 | 0.8 (±.20) | NA | NA | NA | 7.0 (±.10) | 9.4 (±1.05) | 3.8 (±.20) | 1.9 (±.35) | 1.2 (±.10) | 1.6 | 1.7 (±.10) | NA | 0.4 (±.05) | 0.3 |
(Stage 35). Body tubular in dorsal and ovoid in lateral views, respectively (Fig.
Oral disc positioned at terminal portion of body opening antero-ventrally (Fig.
Measurements (in mm; mean shown without parentheses and standard errors are shown in parentheses): Measurements of the seven tadpoles of various stage of development (Stages 30, 35, 41, 42 and 43) are presented in Table
In life, dorsally, outer integument brown, with no melanopores. Ventrally, integument translucent but the gut was not visible; throat speckled. Both tail fins transparent with few melanophores. Laterally, tail muscle white with a few brown spots spread mainly at anterior region of tail. A completely transformed metamorph (SVL 10.6 mm; HL 4.23 mm) resembles adult in morphology, with an evident inverted ‘V’ mark on dorsum and transverse crossbars on limbs.
Morphological and osteological characteristics of this new taxon are compared with members of other known Oriental bufonid genera below. The new taxon described here differs from the following known genera thus (only opposing character states in the genera being compared are mentioned):
Parapelophryne Fei, Ye & Jiang, 2003: type species– Nectophryne scalptus [current name combination: Parapelophryne scalpta (
Pedostibes Günther, 1875: type species – Pedostibes tuberculosus Günther, 1875: Larger adult size (SVL 36.6–38.5 mm), presence of eight presacral vertebrae; short, rounded parotoid glands; tips of fingers dilated into truncated discs; small, numerous pigmented ova laid in strings, as in members of the genus Duttaphrynus and exotrophic larvae (Günther 1875,
Bufoides Pillai & Yazdani, 1973: type species– Ansonia meghalayana [current name combination: Bufoides meghalayanus (Yazdani & Chanda, 1971); currently monotypic, but additional, unnamed species recognised;
Sabahphrynus Matsui, Yambun & Sudin, 2007: type species– Nectophryne maculata [current name combination: Sabahphrynus maculatus (Mocquard, 1890)]: Larger adult size (41.21 ± 2.5, 30.4–52.6), presence of eight presacral vertebrae, absence of tympanum and parotoid glands, absence of webbing between the fingers, over 50 eggs/ovary and absence of an external vocal sac in males (
Duttaphrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green & Wheeler, 2006: type species– Bufo melanostictus [current name combination: Duttaphrynus melanostictus (Schneider, 1799)]: Large adult size (mean SVL 43.7 mm), presence of eight presacral vertebrae, presence of keratinised cephalic ridges in some species, presence of large, keratinised warts on the dorsum, absence of expanded discs in finger and toe tips, absence of webbing between the fingers, numerous black pigmented ova laid in long, continuous strings, exotrophic larvae and terrestrial habit (
Ansonia Stoliczka, 1870: type species – Ansonia penangensis Stoliczka, 1870: small to medium adult size (35–40 mm), absence of (or rudimentary) webbing between the fingers, presence of eight presacral vertebrae, absence of dilations in finger and toe tips, absence of parotoid glands, exotropic larvae with prominent oral discs and torrential stream dwelling habit (
Adenomus Cope, 1861: type species– Adenomus badioflavus Cope, 1861, a junior synonym of Bufo kelaartii [current name combination: Adenomus kelaarti (Günther, 1858)]: The genus Adenomus was resurrected from the synonymy of ‘Bufo’ by
Ghatophryne Biju, Bocxlaer, Giri, Loader & Bossuyt, 2009: type species– Ansonia ornata [current name combination: Ghatophryne ornata (Günther, 1876)]: larger adult size (up to 35 mm SVL), characteristic reddish dorsal and ventral colouration, absence of parotoid glands, absence of webbing between the fingers, finger tips not dilated to discs and torrential stream dwelling habit (
Xanthophryne Biju Bocxlaer, Giri, Loader & Bossuyt, 2009: type species– Bufo koynaensis [current name combination: Xanthophryne koynaensis (Soman, 1963)]: Larger adult size (up to 35.3 mm SVL), presence of characteristic chrome yellow patches along the flanks and sides of the abdomen, indistinct tympanum, weak, rounded parotoid glands, absence of webbing in fingers and discs in toes and fingers; large, pigmented ova laid in stagnant puddles on the ground (
Leptophryne Fitzinger, 1843: type species – Bufo cruentatus [current name combination: Leptophryne cruentata (Tschudi, 1838)]:
Pseudobufo Tschudi, 1838: type species – Pseudobufo subasper Tschudi, 1838: Large body size, stout habitus; presence of seven presacral vertebrae (vs. six in Blythophryne gen. n.) completely (to the tip of Toe IV) webbed feet (vs. incomplete toe webbing in Blythophryne beryet gen. et sp. n.), fingers basally webbed; parotoid glands absent; dorsal, lateral and ventral skin surfaces with fine spinules, dorsoventrally depressed body with large, round warts and dorsally positioned nostrils (vs. lateral) distinguish it from the new genus described here (
Ingerophrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green & Wheeler, 2006: type species– Bufo biporcatus [current name combination: Ingerophrynus biporcatus (Gravenhorst, 1829)]: Presence of seven presacral vertebrae (vs. six); absence of lateral dilations in the digit tips (vs. present); absence of webbing between the fingers (vs. present) and endotrophic (vs. exotrophic) larvae distinguish Blythophryne beryet gen. et sp. n. from Ingerophrynus. Distribution: Southern Yunnan, Indochina, the Malay Peninsula, the islands of Indo-Malaya, and Philippines.
Phrynoidis Fitzinger, 1843: type species – Bufo asper [current name combination: Phrynoidis asper (Gravenhorst, 1829)]: Large adult size (up to 100 mm SVL) presence of an omosternum, (vs. absent); presence of seven presacral vertebrae (vs. six); absence of lateral dilations of digit tips (vs. present) and exotrophic (vs. endotrophic) larvae distinguish this genus from the new genus Blythophryne gen. n. Distribution: Myanmar through western and peninsular Thailand, the Malay Peninsula, Sumatra, Borneo, and Java.
Apart from the above bufonid genera known from Oriental Asia, the new taxon described herein differs from the following central-west African genera:
Nectophryne Buchholz & Peters, 1875: type species – Nectophryne afra Buchholz & Peters, 1875 by the presence of eight presacral vertebrae (vs. six in Blythophryne beryet gen. et sp. n.); presence of lamelliform subdigital pads – a character unique to Nectophryne which is absent in the new taxon described here. Oriental forms including members of the genera Pedostibes and Pelophryne were attributed to Nectophryne earlier (
Nectophrynoides Noble, 1926: type species – Nectophryne tornieri [current name combination: Nectophrynoides tornieri (Roux, 1906)]: The comparisons made here are restricted to the type species of Nectophrynoides because the genus is poorly defined and is composed of representatives with a broad spectrum of morphological and developmental characteristics. Though unique among bufonids in possessing an omosternum and a direct developmental mode (in N. viviparus), members of this genus are poorly diagnosed with respect to other genera (
Multiple sequence alignment of the 16S homologous regions resulted in 498 conserved sites and 246 parsimoniously informative sites. In the phylogenetic analysis using both Maximum likelihood and Bayesian inference, the focal taxon showed a unique taxonomic position. The phylograms of both inference methods were similar (Fig.
Phylogenetic position of Blythophryne beryet gen. et sp. n., inferred from concatenated partial 12S and 16S rDNA sequences. The posterior probabilities for Bayesian Inference (BI) and the bootstrap support values for the ML are given as (BI /ML) above and below the branch nodes. a The tree was generated using 36 species related to 21 genera, and was rooted using Rhabeo gutattus as outgroup b the subclade containing the Indian and Sri Lankan toads (7 genera, 17 species) rooted using Ghatophryne ornata as outgroup; and c the subclade containing the Indian and Sri Lankan toads (5 genera, 15 species) rooted using Adenomus kelaartii as outgroup.
The average uncorrected K2p distance estimates across genera of Bufonidae based on partial 16S (lower triangular matrix) and 12S (upper triangular matrix) rDNA sequences, calculated using MEGA6.06; within genus K2p distances are given along the diagonal (left value for 16S and right value for 12S).
16S/12S k2p uncorrected pair-wise distance estimates | ||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Taxa (Genus*/Species) | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | |
1 | Amietophrynus | 0.068/0.044 | 0.060 | 0.049 | 0.072 | 0.067 | 0.094 | 0.066 | 0.059 | 0.074 | 0.071 | 0.066 | 0.049 | 0.077 | 0.043 | 0.100 | 0.103 | 0.067 | 0.063 | 0.043 | 0.037 | 0.071 |
2 | Ansonia | 0.106 | 0.066/0.060 | 0.061 | 0.086 | 0.083 | 0.115 | 0.079 | 0.066 | 0.081 | 0.073 | 0.065 | 0.071 | 0.104 | 0.061 | 0.114 | 0.097 | 0.067 | 0.067 | 0.055 | 0.063 | 0.085 |
3 | Duttaphrynus | 0.092 | 0.089 | 0.049/ 0.038 | 0.070 | 0.063 | 0.099 | 0.067 | 0.060 | 0.065 | 0.076 | 0.065 | 0.054 | 0.084 | 0.045 | 0.098 | 0.099 | 0.074 | 0.054 | 0.045 | 0.049 | 0.077 |
4 | Ingerophrynus | 0.090 | 0.084 | 0.074 | 0.056/0.077 | 0.077 | 0.102 | 0.075 | 0.082 | 0.073 | 0.085 | 0.071 | 0.081 | 0.093 | 0.065 | 0.113 | 0.111 | 0.078 | 0.071 | 0.062 | 0.075 | 0.085 |
5 | Mertensophryne | 0.095 | 0.100 | 0.099 | 0.090 | 0.059/ 0.061 | 0.098 | 0.079 | 0.066 | 0.075 | 0.083 | 0.075 | 0.079 | 0.094 | 0.067 | 0.108 | 0.114 | 0.083 | 0.079 | 0.059 | 0.065 | 0.073 |
6 | Nectophryne | 0.143 | 0.141 | 0.127 | 0.139 | 0.140 | 0.089/0.049 | 0.092 | 0.095 | 0.098 | 0.107 | 0.093 | 0.098 | 0.108 | 0.073 | 0.146 | 0.136 | 0.102 | 0.083 | 0.090 | 0.102 | 0.081 |
7 | Nectophrynoides | 0.097 | 0.091 | 0.077 | 0.082 | 0.099 | 0.132 | 0.021/0.015 | 0.068 | 0.084 | 0.069 | 0.069 | 0.069 | 0.085 | 0.051 | 0.119 | 0.089 | 0.061 | 0.077 | 0.034 | 0.077 | 0.069 |
8 | Pedostibes | 0.091 | 0.086 | 0.066 | 0.083 | 0.104 | 0.115 | 0.074 | 0.076/0.065 | 0.075 | 0.071 | 0.067 | 0.063 | 0.085 | 0.051 | 0.130 | 0.107 | 0.068 | 0.069 | 0.048 | 0.063 | 0.071 |
9 | Phrynoidis | 0.086 | 0.084 | 0.073 | 0.082 | 0.102 | 0.135 | 0.080 | 0.080 | 0.039/0.085 | 0.089 | 0.079 | 0.091 | 0.108 | 0.063 | 0.123 | 0.119 | 0.085 | 0.069 | 0.071 | 0.085 | 0.081 |
10 | Ghatophryne ornata | 0.092 | 0.104 | 0.075 | 0.086 | 0.103 | 0.148 | 0.088 | 0.083 | 0.087 | n/a | 0.061 | 0.081 | 0.102 | 0.053 | 0.124 | 0.097 | 0.081 | 0.077 | 0.057 | 0.065 | 0.086 |
11 | Leptophryne borbonica | 0.103 | 0.107 | 0.095 | 0.096 | 0.117 | 0.121 | 0.099 | 0.082 | 0.099 | 0.092 | n/a | 0.073 | 0.085 | 0.045 | 0.106 | 0.085 | 0.069 | 0.053 | 0.061 | 0.061 | 0.069 |
12 | Vandijkophrynus robinsoni | 0.083 | 0.102 | 0.063 | 0.071 | 0.082 | 0.132 | 0.077 | 0.077 | 0.073 | 0.064 | 0.102 | n/a | 0.072 | 0.053 | 0.110 | 0.097 | 0.073 | 0.072 | 0.037 | 0.049 | 0.085 |
13 | Schismaderma carens | 0.096 | 0.099 | 0.076 | 0.088 | 0.086 | 0.132 | 0.082 | 0.079 | 0.095 | 0.097 | 0.107 | 0.092 | n/a | 0.089 | 0.110 | 0.110 | 0.089 | 0.069 | 0.077 | 0.081 | 0.094 |
14 | Bufo bufo | 0.102 | 0.113 | 0.078 | 0.107 | 0.116 | 0.134 | 0.085 | 0.088 | 0.082 | 0.087 | 0.105 | 0.076 | 0.105 | n/a | 0.110 | 0.085 | 0.057 | 0.057 | 0.034 | 0.045 | 0.057 |
15 | Sabahphrynus maculatus | 0.092 | 0.085 | 0.077 | 0.071 | 0.097 | 0.124 | 0.093 | 0.079 | 0.081 | 0.082 | 0.093 | 0.087 | 0.082 | 0.104 | n/a | 0.145 | 0.114 | 0.110 | 0.102 | 0.106 | 0.123 |
16 | Pelophryne api | 0.108 | 0.105 | 0.102 | 0.092 | 0.100 | 0.155 | 0.106 | 0.097 | 0.106 | 0.100 | 0.113 | 0.100 | 0.122 | 0.126 | 0.112 | n/a | 0.089 | 0.093 | 0.068 | 0.101 | 0.093 |
17 | Bufoides meghalayanus | 0.088 | 0.091 | 0.055 | 0.080 | 0.109 | 0.133 | 0.073 | 0.064 | 0.069 | 0.078 | 0.095 | 0.062 | 0.087 | 0.071 | 0.087 | 0.105 | n/a | 0.073 | 0.041 | 0.069 | 0.069 |
18 | Adenomus kelaartii | 0.091 | 0.091 | 0.065 | 0.084 | 0.107 | 0.136 | 0.070 | 0.071 | 0.070 | 0.066 | 0.098 | 0.059 | 0.090 | 0.083 | 0.085 | 0.103 | 0.062 | n/a | 0.061 | 0.069 | 0.065 |
19 | Xanthophryne koynayensis | 0.084 | 0.078 | 0.054 | 0.070 | 0.098 | 0.129 | 0.072 | 0.056 | 0.077 | 0.073 | 0.087 | 0.059 | 0.082 | 0.078 | 0.078 | 0.083 | 0.039 | 0.057 | n/a | 0.049 | 0.049 |
20 | Rhaebo guttatus | 0.111 | 0.085 | 0.092 | 0.088 | 0.104 | 0.136 | 0.089 | 0.083 | 0.086 | 0.087 | 0.105 | 0.092 | 0.088 | 0.114 | 0.094 | 0.112 | 0.092 | 0.085 | 0.075 | n/a | 0.086 |
21 | Blythophryne beryet gen. et sp. n. | 0.103 | 0.119 | 0.089 | 0.098 | 0.118 | 0.165 | 0.098 | 0.102 | 0.088 | 0.099 | 0.112 | 0.092 | 0.101 | 0.105 | 0.106 | 0.112 | 0.080 | 0.082 | 0.075 | 0.109 | n/a |
The small-sized bush toad described here is an interesting new find from the Andaman Islands, in the Bay of Bengal, Republic of India. It has a number of unique external morphological and skeletal characters, in comparison to known Oriental and other relevant bufonid genera. Its distinctiveness and unique taxonomic position (warranting the erection of a monotypic genus), is also robustly supported by phylogenetic reconstruction carried out using partial 16S and 12S gene sequences and showing its position relative to other Asian and African bufonids (
Biogeographic remarks.Bufonidae is a species-rich family, with nearly cosmopolitan distribution around the globe (
The herpetofauna of Andaman and Nicobar Islands are considered to be of either Indo-Chinese or Indo-Malayan affinities (
The submerged chain of mountains referred to as the “Burma arc” was formed at the same time as the main Himalayan chain, during the late Cretaceous (
The new taxon described here is characterised with a small adult body size, semi-arboreality high specificity for larval microhabitat niche, absence of inguinal fat bodies, relatively low number of mid-sized ova and a narrow distributional range. Further, it seems to be an exception in possessing parotoid glands, which was a character associated with widely distributed bufonid species (
Likewise, the larvae of this new taxon with a moderate, intermediate clutch size and a high specificity towards the site of oviposition (i.e., phytotelms) explain its limited range of distribution as currently understood. Further studies in the Andaman archipelago are needed to understand the identity and origins of its fauna.
We thank Wildlife Institute of India (
List of comparative material of Oriental and other relevant members of the Bufonidae examined.
Adenomus ‘dasi’ WHT 2267–69; Adenomus kandianus BMNH 1947.2.62–63; Adenomus kelaartii