Mud-packing frog: a novel breeding behaviour and parental care in a stream dwelling new species of Nyctibatrachus (Amphibia, Anura, Nyctibatrachidae).

Reproductive modes are diverse and unique in anurans. Selective pressures of evolution, ecology and environment are attributed to such diverse reproductive modes. Globally forty different reproductive modes in anurans have been described to date. The genus Nyctibatrachus has been recently revised and belongs to an ancient lineage of frog families in the Western Ghats of India. Species of this genus are known to exhibit mountain associated clade endemism and novel breeding behaviours. The purpose of this study is to present unique reproductive behaviour, oviposition and parental care in a new species Nyctibatrachus kumbara sp. nov. which is described in the paper. Nyctibatrachus kumbara sp. nov. is a medium sized stream dwelling frog. It is distinct from the congeners based on a suite of morphological characters and substantially divergent in DNA sequences of the mitochondrial 16S rRNA gene. Males exhibit parental care by mud packing the egg clutch. Such parental care has so far not been described from any other frog species worldwide. Besides this, we emphasize that three co-occurring congeneric species of Nyctibatrachus, namely N. jog, N. kempholeyensis and Nyctibatrachus kumbara sp. nov. from the study site differ in breeding behaviour, which could represent a case of reproductive character displacement. These three species are distinct in their size, call pattern, reproductive behaviour, maximum number of eggs in a clutch, oviposition and parental care, which was evident from the statistical analysis. The study throws light on the reproductive behaviour of Nyctibatrachus kumbara sp. nov. and associated species to understand the evolution and adaptation of reproductive modes of anurans in general, and Nyctibatrachus in particular from the Western Ghats.

Diversifying in the mountains: spatiotemporal diversification of frogs in the Western Ghats biodiversity hotspot.

Mountain ranges are hotspots of biodiversity. However, the mechanisms that generate biodiversity patterns in different mountainous regions and taxa are not apparent. The Western Ghats (WG) escarpment in India is a globally recognized biodiversity hotspot with high species richness and endemism. Most studies have either invoked paleoclimatic conditions or climatic stability in the southern WG refugium to explain this high diversity and endemism. However, the factors driving macroevolutionary change remain unexplored for most taxa. Here, we generated the most comprehensive dated phylogeny to date for ranoid frogs in the WG and tested the role of paleoclimatic events or climatic stability in influencing frog diversification. We found that the diversity of different ranoid frog clades in the WG either accumulated at a constant rate through time or underwent a decrease in speciation rates around 3-2.5 Ma during the Pleistocene glaciation cycles. We also find no significant difference in diversification rate estimates across elevational gradients and the three broad biogeographic zones in the WG (northern, central, and southern WG). However, time-for-speciation explained regional species richness within clades, wherein older lineages have more extant species diversity. Overall, we find that global paleoclimatic events have had little impact on WG frog diversification throughout most of its early history until the Quaternary and that the WG may have been climatically stable allowing lineages to accumulate and persist over evolutionary time.

Discovery and description of a new species of burrowing frog Sphaerotheca Gnther, 1859 (Anura: Dicroglossidae) from the suburban landscapes of Bengaluru, India.

Recent resolution of prevailing taxonomic ambiguities in the genus Sphaerotheca and new species discoveries from urban/suburban landscapes highlight the need for attention to non-forested habitats for amphibian conservation. In this paper, we review the status of the members of the genus Sphaerotheca and justify the synonymy of Sphaerotheca magadha as a junior synonym of Sphaerotheca swani. The prospects of resurrection of Sphaerotheca swani (herein preliminarily referred to as Sphaerotheca cf. breviceps [swani]) are discussed. In addition, we describe a new species Sphaerotheca varshaabhu sp. nov. from the suburban region of Bengaluru, India. We employ an integrative taxonomic approach to characterize the new species using molecular phylogeny, genetic distance, morphological characters, and geographical isolation as lines of evidence. We also provide a description of vocal repertoire of Sphaerotheca varshaabhu sp. nov. and provide comparative bioacoustics data for four species. This previously undescribed species from the suburban areas of Bengaluru described herein as Sphaerotheca varshaabhu sp. nov. forms a genetically divergent lineage and its genetic distance varied from 3.6% to 12.2% for 16S rRNA with respect to other species of Sphaerotheca. Our phylogenetic analysis for the genus including the new species confirms the synonymy of one recently described species, resulting in 10 valid species in the genus Sphaerotheca. These results emphasize the need for utilizing an integrative taxonomic approach for uncovering hidden diversity of suburban areas. Given these recent discoveries, we advocate for more robust surveys in human dominated areas, so that these amphibians may receive more attention.

Discovery of a new species of Euphlyctis (Anura: Dicroglossidae) from the western coastal plains of peninsular India.

The aquatic frog genus Euphlyctis is now considered under two separate genera- Euphlyctis Fitzinger and Phrynoderma Fitzinger, with four valid species in each genus, respectively. The current distribution range of the genus Phrynoderma is restricted to peninsular India and Bangladesh, with a probability of availability in Sri Lanka. The genus Euphlyctis has a wide range of distribution from the Arabian Peninsula and the South Asian countries, with a few historical descriptions treated as junior synonyms of the species Euphlyctis cyanophlyctis in India. A new species of skittering frog, Euphlyctis jaladhara sp. nov. is described here from the surroundings of the Thattekad Bird Sanctuary of Kerala, India, which has a predominant distribution in the western coastal plains of peninsular India and some parts of the Western Ghats. The new species has distinct morphological characters and a high genetic distance that distinguishes it from the sympatric species. Discussions are made concerning the validity of the species E. ghoshi and E. kalasgramensis.

Coexisting good neighbours: acoustic and calling microhabitat niche partitioning in two elusive syntopic species of balloon frogs, Uperodon systoma and U. globulosus (Anura: Microhylidae) and potential of individual vocal signatures.

BACKGROUND: Most amphibians use a repertoire of acoustic signals to propagate signals in social contexts. The description of these repertoires provides a key towards the understanding of the behaviour of individuals and the evolutionary functions of calls. Here, we assessed the variations in advertisement calls within and between two fossorial sympatric species, Uperodon systoma and Uperodon globulosus, that share their breeding season and breeding sites. For each species, we applied Beecher's index of total information capacity (HS) for the individual vocal signature, determined the difference in call properties and demonstrated the segregation in the calling microhabitat niche between the two species. RESULTS: Our results demonstrated that the advertisement calls of U. systoma are pulsatile with a call rate of 3.00 ± 0.97 calls per second while those of U. globulosus are not pulsatile with a lower call rate of 0.53 ± 0.22 calls per second. For both species, the variations in call properties among individuals was higher than that within individual, a pattern consistent with that of other fossorial anurans. The body condition and air temperature did not significantly impact the call properties of either species. The outcome of the Beecher's index (HS) showed that the calls of U. systoma can be used to identify 14 different individuals and the calls of U. globulosus can be used to identify 26 different individuals. The statistical analyses on the advertisement call of the two species showed a significant difference in the temporal properties as the call duration, and fall time and rise time were significantly different between the two species. Lastly, we successfully demonstrated that there is a clear segregation in calling site microhabitat between the two species, where U. globulosus calls floating close to the bank of the waterbody while U. systoma calls floating further away from the bank. CONCLUSION: This study highlights the potential for pre-mating isolation, character displacement and assortative mating in two syntopic fossorial anurans, leading to association between acoustic, calling microhabitat niche and body index divergence as important behavioural and ecological traits.

Morphological groupings within Euphlyctis (Anura: Dicroglossidae) and description of a new species from the surroundings of Thattekad Bird Sanctuary, Kerala, India.

The skittering frog genus Euphlyctis is known to have a distribution range from Arabian Peninsula to Bangladesh through India and Sri Lanka. Although species descriptions were initiated about two centuries ago in the genus with a small number of species described, taxonomic identities and genetic information for many species are still incomplete / inaccurate. Here, we report the systematic status of this group, discuss a series of taxonomic issues, and describe a new species. Based on morphological characters and colour patterns, and their reciprocal monophyly in the molecular tree, two morphological groups, 'cyanophlyctis group' and 'hexadactylus group' are established within the genus Euphlyctis. We discuss the genetic identity for E. cyanophlyctis, described 220 years ago from the Indian subcontinent, from the surroundings of its type locality. E. mudigere is proposed as a junior synonym of E. cyanophlyctis as it was described from populations here considered to originate from the general distribution area of the latter taxon, and without substantial genetic divergence from other populations here attributed to E. cyanophlyctis. The potential of resurrection of Euphlyctis seistanica due to distinct genetic population in Iran is discussed; this lineage appears to be genetically rather similar to E. kalasgramensis, and the species distinctness of these two lineages requires further study. The validity of E. kalasgramensis is also discussed in the context of the historically earlier nomina, Rana bengalensis and Rana leschenaultii due to distribution range overlaps. The prospects of establishing either Rana cyanophlictis variety fulvus or Rana cyanophlictis variety flavens from Sri Lanka are also mentioned. Based on a comparative account, we report the discovery of a new species of skittering frog Euphlyctis kerala sp. nov. which has a restricted distribution in the western coastal plains of India, south of the Palghat gap. [http://zoobank.org/urn:lsid:zoobank.org:act:02B1DE04-C40C-4AAD-AC87-EF97673A4BF5].

Phenotypic divergence in large sized cricket frog species that crossed the geographical barriers within peninsular India.

A good number of new species are being described and included in the genus Fejervarya / Minervarya from South Asia and Southeast Asia in the recent past. But, the classification or consideration of the genus Fejervarya / Minervarya is only phylogenetic, where it is difficult to morphologically identify the genus in the overlapping range of geographical distribution. Recently a large sized new species of cricket frog Fejervarya kalinga has been described, which is thought to be endemic to the Eastern Ghats (a unique geographical landscape running parallel to the East Coast of India). Here we report a morphologically variable and distinct population of F. kalinga from the Western Ghats (an amphibian diversity hotspot running parallel to the West Coast of India). We discuss the importance of genetic studies in handling such morphologically highly variable populations of the species and their adaptation to different habitats and their implications in taxonomic identity.

Distribution status of the Western Burrowing Frog, Sphaerotheca pashchima in India.

In a recent study, Hortal et al. (2015) recognized seven shortfalls of knowledge that hamper our understanding of biodiversity. Three of these are of particular importance for drawing conservation strategies for ecological indicator species, among which we can count the amphibians (Simon et al. 2011): the Linnean shortfall describes 'the knowledge incongruity between the described species and the remaining species yet to be described' (Brown Lomolino 1998); the Wallacean shortfall defines 'paucity of knowledge about the actual distribution range of the species' (Lomolino 2004); and the Darwinian shortfall refers to 'lack of knowledge about the tree of life and evolution of lineages, species and traits' (Diniz-Filho et al. 2013).

Parapsilorhynchus alluriensis, a new species of cyprinid fish (Teleostei: Cyprinidae) from the Eastern Ghats, of India.

A new species of cyprinid fish, Parapsilorhynchus alluriensis, is described from the Alluri Hills of Andhra Pradesh State, India. It is distinguished from its congeners by the combination of the following characters: A poorly-developed callous pad present behind lower lip; head deep (depth at occiput 47.3-72.3% head length, HL); body stout and deep (depth at dorsal fin origin 17.3-21.7 % standard length, SL); gape width 27.3-32.8% HL; inter-orbital space 33.9-43.2% HL; lower lip rounded; mouth opening situated very close to the anterior tip of snout; upper lip concealed by a poorly-developed rostral fold which is slightly fringed; minute papillae on rostral fold; 3 simple pectoral-fin rays; 33-34 lateral-line scales. A partial phylogeny based on the DNA Barcode COI gene suggests a sister-group relationship between the new species and P. prateri. Till date, the genus consists of four species from the Western Ghats, three from the Eastern Ghats, including the species described herein totaling it to seven from the peninsular India.

A new species of Sphaerotheca Günther, 1859 (Anura: Dicroglossidae) from the degraded urban ecosystems of Bengaluru, Deccan Plateau, India.

The dicroglossid genus Sphaerotheca was erected by Günther in 1859, with its range of distribution in South Asia. Since then, 13 species have been described and 10 species are considered valid. Many of these descriptions were from low to mid-elevation ranges of homestead areas, agroecosystems and degraded landscapes with a few from forested areas. In the present account, a new species of Sphaerotheca, a genetically distinct lineage with a morphological character set distinguishable from its congeners, is described from the surroundings of Bengaluru city, India. Based on evidence for the origin of the original specimen, we also designated a neotype for Rana variegata Gravenhorst, 1829, a junior primary homonym of Rana variegata Linnaeus, 1758, and permanently invalid name, so that it is now a subjective synonym of Rana (Tomopterna) breviceps rolandae Dubois, 1983.

Importance of genetic data in resolving cryptic species: A century old problem of understanding the distribution of Minervarya syhadrensis Annandale 1919, (Anura: Dicroglossidae).

Frogs of the genus Minervarya are cryptic and widely distributed in South Asia. However, many of them lack information about the precise type locality, genetic data, and distribution range. The present study aimed to examine the genetic affinities of a widely distributed species Minervarya syhadrensis around its type locality in the northern Western Ghats (Pune, Maharashtra). We studied the type specimen of M. syhadrensis and collected similar sized Minervarya frogs from Pune district. In the field, we observed two different calls from morphologically similar (M. syhadrensis like) males suggesting the sympatric occurrence of two cryptic species (that we initially named Minervarya species A and Minervarya species B). We analyzed morphology, call pattern, and mitochondrial 16S rRNA gene sequence of both species. Minervarya species A has a long call with a low pulse repetition rate and higher dominant frequency compared to that of the Minervarya species B. These species cannot be differentiated based on morphometric data. However, they can be sorted out using morphological characters such as the presence of longitudinal skin folds on the dorsal side (Minervarya species A) and differences in foot webbing. DNA sequences of Minervarya species A and Minervarya species B are matching with those of M. caperata and M. agricola respectively. After studying the type specimens of M. syhadrensis and M. caperata, we found morphological similarities (longitudinal skin folds) with the samples of Minervarya species A collected during the present study. Based on the results of our study (morphology and genetic) and available literature, we propose to redefine M. syhadrensis as applying to the lineage initially named Minervarya species A, and to treat the species M. caperata as a junior synonym of M. syhadrensis. Our study will be helpful in further taxonomic revision of the genus, and provides natural history information for M. syhadrensis and M. agricola.

A deeply divergent lineage of Walkerana (Anura: Ranixalidae) from the Western Ghats of Peninsular India.

The frog family Ranixalidae is endemic to the Western Ghats of Peninsular India and contains two genera, Indirana and Walkerana. The three known species of Walkerana are restricted to different hill ranges south of the Palghat Gap, an ancient valley in the Western Ghats. In this study, we report the discovery of a deeply divergent lineage of Walkerana from the high elevations of the Elivalmalai hill range. This finding extends the geographic range of the Walkerana clade to the north of the Palghat Gap. The new species Walkerana muduga sp. nov. is genetically and morphologically divergent, and geographically isolated from its sister lineages. We also recovered a potential new lineage in the adjoining hill ranges suggesting the presence of additional new species in this genus north of the Palghat Gap.

A new species of Fejervarya Bolkay, 1915 (Anura: Dicroglossidae) from the northern Western Ghats parts of Maharashtra, India.

In the recent past the systematic position and taxonomy of genus Fejervarya, Bolkay is undergoing changes in its systematic position due to fairly good amount of phylogenetic resolution, cryptic morphological characters and lack of systematic sampling for phylogenetic studies across the range of distribution. In our sampling in the northern Western Ghats, we encountered a new lineage sister to the phylogenetic cluster which comprises the 'Rufescens complex'. This new lineage is described here as new species Fejervarya marathi sp. nov. based on distinctness in a combination of morphological characters, genetic distance and geography. The problems in morphological groupings for the Fejervarya frogs of the Western Ghats in the recent studies are discussed with the sub-clade composition based on geography in the phylogenetic tree.

A new ancient lineage of frog (Anura: Nyctibatrachidae: Astrobatrachinae subfam. nov.) endemic to the Western Ghats of Peninsular India.

The Western Ghats (WG) is an escarpment on the west coast of Peninsular India, housing one of the richest assemblages of frogs in the world, with three endemic families. Here, we report the discovery of a new ancient lineage from a high-elevation massif in the Wayanad Plateau of the southern WG. Phylogenetic analysis reveals that the lineage belongs to Natatanura and clusters with Nyctibatrachidae, a family endemic to the WG/Sri Lanka biodiversity hotspot. Based on geographic distribution, unique morphological traits, deep genetic divergence, and phylogenetic position that distinguishes the lineage from the two nyctibatrachid subfamilies Nyctibatrachinae Blommers-Schlösser, 1993 and Lankanectinae Dubois & Ohler, 2001, we erect a new subfamily Astrobatrachinae subfam. nov. (endemic to the WG, Peninsular India), and describe a new genus Astrobatrachus gen. nov. and species, Astrobatrachus kurichiyana sp. nov. The discovery of this species adds to the list of deeply divergent and monotypic or depauperate lineages with narrow geographic ranges in the southern massifs of the WG. The southern regions of the WG have long been considered geographic and climatic refugia, and this new relict lineage underscores their evolutionary significance. The small range of this species exclusively outside protected areas highlights the significance of reserve forest tracts in the WG in housing evolutionary novelty. This reinforces the need for intensive sampling to uncover new lineages and advance our understanding of the historical biogeography of this ancient landmass.

Systematic status of Fejervarya ((Amphibia, Anura, Dicroglossidae) from South and SE Asia with the description of a new species from the Western Ghats of Peninsular India.

We carried out a large-scale phylogenetic analysis of fejervaryan (dicroglossid frogs with 'Fejervaryan lines' on the ventral side of the body) frogs, distributed in South and SE Asia, using published and newly generated sequences of unidentified individuals from the northern Western Ghats. The results corroborate the presence of a larger fejervaryan clade with a sister relationship to a clade composed of Sphaerotheca. Two sister clades could be discerned within the lager fejervaryan clade. The unidentified individuals formed a monophyletic group and showed a strong support for a sister relationship with Minervarya sahyadris. The species was found to be highly divergent (16S rRNA-4% and tyr-1%) from its sister lineage Minervarya sahyadris, and the clade composed of these two lineages were found to be deeply nested within the larger clade of Fejervarya. Based on this, the genus Minervarya Dubois, Ohler and Biju, 2001 is synonymized under the genus Fejervarya Bolkay, 1915. The unidentified lineage is recognized, based on phylogenetic position, genetic divergence and morphological divergence, as a distinct species and named here as Fejervarya gomantaki sp. nov. The presence of rictal glands was observed to be a synapomorphic character shared by the nested clade members, Fejervarya sahyadris and Fejervarya gomantaki sp. nov. Based on the presence of rictal gland and small size, Minervarya chilapata, a species from a lowland region in the Eastern Himalayas, is synonymized under Fejervarya and evidence for morphological separation from the new species, Fejervarya gomantaki sp. nov. is provided. For fejervaryan frogs, three generic names (Frost, 2015) are currently in use for two phylogenetic subclades; the genus Fejervarya Bolkay, 1915 for species distributed in South East Asia; the genus Zakerana Howlader, 2011 for species distributed in South Asia and the genus Minervarya Dubois, Ohler and Biju, 2001 nested within the 'Zakerana clade'. In the phylogenetic analysis Minervarya sahyadris and the new species described herein as Fejervarya gomantaki sp. nov. are nested within the 'Zakerana clade'. If the 'Zakerana clade' for the fejervaryan frogs distributed in South Asia is assigned a generic status, the nomen 'Minervarya' should be used as per the principle of priority of the ICZN Code. Taking into consideration the overlapping distribution ranges of members of the sister clades within the larger fejervaryan clade and the absence of distinct morphological characteristics, we also synonymize the genus Zakerana Howlader, 2011, a name assigned to one of the sister clades with members predominantly distributed in South Asia, under the genus Fejervarya Bolkay, 1915. We discuss the need for additional sampling to identify additional taxa and determine the geographical ranges of the members of the sister clades within Fejervarya to resolve taxonomy within this group. [Corrected]

Lineage delimitation and description of nine new species of bush frogs (Anura: Raorchestes, Rhacophoridae) from the Western Ghats Escarpment.

Bush frogs of the genus Raorchestes are distributed mainly in the Western Ghats Escarpment of Peninsular India. The inventory of species in this genus is incomplete and there is ambiguity in the systematic status of species recognized by morphological criteria. To address the dual problem of taxon sampling and systematic uncertainty in bush frogs, we used a large-scale spatial sampling design, explicitly incorporating the geographic and ecological heterogeneity of the Western Ghats. We then used a hierarchical multi-criteria approach by combining mitochondrial phylogeny, genetic distance, geographic range, morphology and advertisement call to delimit bush frog lineages. Our analyses revealed the existence of a large number of new lineages with varying levels of genetic divergence. Here, we provide diagnoses and descriptions for nine lineages that exhibit divergence across multiple axes. The discovery of new lineages that exhibit high divergence across wide ranges of elevation and across the major massifs highlights the large gaps in historical sampling. These discoveries underscore the significance of addressing inadequate knowledge of species distribution, namely the "Wallacean shortfall", in addressing the problem of taxon sampling and unknown diversity in tropical hotspots. A biogeographically informed sampling and analytical approach was critical in detecting and delineating lineages in a consistent manner across the genus. Through increased taxon sampling, we were also able to discern a number of well-supported sub-clades that were either unresolved or absent in earlier phylogenetic reconstructions and identify a number of shallow divergent lineages which require further examination for assessment of their taxonomic status.