K. Praveen Karanth

Molecular phylogeny and biogeography of langurs and leaf monkeys of South Asia (Primates: Colobinae)

The two recently proposed taxonomies of the langurs and leaf monkeys (Subfamily Colobinae) provide different implications to our understanding of the evolution of Nilgiri and purple-faced langurs. Groves (2001) [Groves, C.P., 2001. Primate Taxonomy. Smithsonian Institute Press, Washington], placed Nilgiri and purple-faced langurs in the genus Trachypithecus, thereby suggesting disjunct distribution of the genus Trachypithecus. [Brandon-Jones, D., Eudey, A.A., Geissmann, T., Groves, C.P., Melnick, D.J., Morales, J.C., Shekelle, M., Stewart, C.-B., 2003. Asian primate classification. Int. J. Primatol. 25, 97-162] placed these langurs in the genus Semnopithecus, which suggests convergence of morphological characters in Nilgiri and purple-faced langurs with Trachypithecus. To test these scenarios, we sequenced and analyzed the mitochondrial cytochrome b gene and two nuclear DNA-encoded genes, lysozyme and protamine P1, from a variety of colobine species. All three markers support the clustering of Nilgiri and purple-faced langurs with Hanuman langur (Semnopithecus), while leaf monkeys of Southeast Asian (Trachypithecus) form a distinct clade. The phylogenetic position of capped and golden leaf monkeys is still unresolved. It is likely that this species group might have evolved due to past hybridization between Semnopithecus and Trachypithecus clades.

Molecular phylogeny of Hemidactylus geckos (Squamata: Gekkonidae) of the Indian subcontinent reveals a unique Indian radiation and an Indian origin of Asian house geckos

Represented by approximately 85 species, Hemidactylus is one of the most diverse and widely distributed genera of reptiles in the world. In the Indian subcontinent, this genus is represented by 28 species out of which at least 13 are endemic to this region. Here, we report the phylogeny of the Indian Hemidactylus geckos based on mitochondrial and nuclear DNA markers sequenced from multiple individuals of widely distributed as well as endemic congeners of India. Results indicate that a majority of the species distributed in India form a distinct clade whose members are largely confined to the Indian subcontinent thus representing a unique Indian radiation. The remaining Hemidactylus geckos of India belong to two other geographical clades representing the Southeast Asian and West-Asian arid zone species. Additionally, the three widely distributed, commensal species (H. brookii, H. frenatus and H. flaviviridis) are nested within the Indian radiation suggesting their Indian origin. Dispersal-vicariance analysis also supports their Indian origin and subsequent dispersal out-of-India into West-Asian arid zone and Southeast Asia. Thus, Indian subcontinent has served as an important arena for diversification amongst the Hemidactylus geckos and in the evolution and spread of its commensal geckos.

Phylogeny and evolution of Malagasy plated lizards

The endemic plated lizards (Gerrhosauridae) of Madagascar are one of the most diverse groups of lizards on the island (19 species) and are found in all ecoregions. On an island that presents so many interesting biological questions, plated lizards are an ideal group for examining patterns of diversification due to their high (but tractable) diversity and wide distribution. To resolve the taxonomy and evolutionary relationships of Malagasy plated lizards, and to explore their biogeographic history, we have generated a molecular phylogeny based on >2kb of both mitochondrial (cob, 1142 bp) and nuclear (NT-3, 419 bp; c-mos, 542 bp) DNA sequences. Phylogenetic relationships and divergence-time estimates indicate Zonosaurinae are monophyletic, arising from a single colonization event, likely from Africa to Madagascar in the Paleogene (Paleocene/early Eocene), with subsequent radiation. Furthermore, our results reveal widespread mischaracterization of species delimitation and assignment based on phenotypic characteristics. Paraphyly of a number of zonosaurine species was strongly supported by our dataset, revealing not only instances of likely cryptic species (phenotypic conservatism) but also phenotypic/genotypic discordance in the delimitation of species (phenotypic variability). These results clearly demonstrate the complex history of biotic diversification on Madagascar and provide novel insight into biogeographic patterns on the island.

The Out-of-India hypothesis: What do molecules suggest?

The remarkable geological and evolutionary history of peninsular India has generated much interest in the patterns and processes that might have shaped the current distributions of its endemic biota. In this regard the “Out-of-India” hypothesis, which proposes that rafting peninsular India carried Gondwanan forms to Asia after the break-up of Gondwana super continent, has gained prominence. Here we have reviewed molecular studies undertaken on a range of taxa of supposedly Gondwanan origin to better understand the Out-of-India scenario. This re-evaluation of published molecular studies indicates that there is mounting evidence supporting Out-of-India scenario for various Asian taxa. Nevertheless, in many studies the evidence is inconclusive due to lack of information on the age of relevant nodes. Studies also indicate that not all Gondwanan forms of peninsular India dispersed out of India. Many of these ancient lineages are confined to peninsular India and therefore are relict Gondwanan lineages. Additionally, for some taxa an “Into India” rather than “Out-of-India” scenario better explains their current distribution. To identify the “Out-of-India” component of Asian biota it is imperative that we understand the complex biogeographical history of India. To this end, we propose three oversimplified yet explicit phylogenetic predictions. These predictions can be tested through the use of molecular phylogenetic tools in conjunction with palaeontological and geological data.

Primate numts and reticulate evolution of capped and golden leaf monkeys (Primates: Colobinae)

A recent phylogenetic study of langurs and leaf monkeys of South Asia suggested a reticulate evolution of capped and golden leaf monkeys through ancient hybridization between Semnopithecus and Trachypithecus. To test this hybridization scenario, I analysed nuclear copies of the mitochondrial cytochrome b gene (numts) from capped, golden and Phayre’s leaf monkeys. These numts were aligned with mitochondrial cytochrome b sequences of various species belonging to the genera Semnopithecus and Trachypithecus. In the phylogenetic tree derived from this alignment, the numts fell into three distinct clades (A, B and C) suggesting three independent integration events. Clade A was basal to Semnopithecus, and clades B and C were basal to Trachypithecus. Among the numts in clades A and C were sequences derived from species not represented in their respective sister mitochondrial groups. This unusual placement of certain numts is taken as additional support for the hybridization scenario. Based on the molecular dating of these integration events, hybridization is estimated to have occurred around 7.1 to 3.4 million years ago. Capped and golden leaf monkeys might have to be assigned to a new genus to reconcile their unique evolutionary history. Additionally, northeast India appears to be a ‘hot spot’ for lineages that might have evolved through reticulate evolution.

Insights into Himalayan biogeography from geckos: A molecular phylogeny of Cyrtodactylus (Squamata: Gekkonidae)

The India-Asia collision profoundly influenced the climate, topography and biodiversity of Asia, causing the formation of the biodiverse Himalayas. The species-rich gekkonid genus Cyrtodactylus is an ideal clade for exploring the biological impacts of the India-Asia collision, as previous phylogenetic hypotheses suggest basal divergences occurred within the Himalayas and Indo-Burma during the Eocene. To this end, we sampled for Cyrtodactylus across Indian areas of the Himalayas and Indo-Burma Hotspots and used three genes to reconstruct relationships and estimate divergence times. Basal divergences in Cyrtodactylus, Hemidactylus and the Palaearctic naked-toed geckos were simultaneous with or just preceded the start of the India-Asia collision. Diversification within Cyrtodactylus tracks the India-Asia collision and subsequent geological events. A number of geographically concordant clades are resolved within Indo-Burmese Cyrtodactylus. Our study reveals 17 divergent lineages that may represent undescribed species, underscoring the previously undocumented diversity of the region. The importance of rocky habitats for Cyrtodactylus indicates the Indo-Gangetic flood plains and the Garo-Rajmahal Gap are likely to have been important historical barriers for this group.

Cretaceous–Tertiary diversification among select Scolopendrid centipedes of South India

Given that peninsular India was part of the Gondwanan super continent, part of its current biota has Gondwanan origin. To determine the Gondwanan component of the peninsular Indian biota, a large number of species spanning diverse taxonomic groups need to be sampled from multiple, if not all, of the former Gondwanan fragments. Such a large scale phylogenetic approach will be time consuming and resource intensive. Here, we explore the utility of a limited sampling approach, wherein sampling is confined to one of the Gondwanan fragments (peninsular India), in identifying putative Gondwanan elements. To this end, samples of Scolopendrid centipedes from Western Ghats region of peninsular India were subjected to molecular phylogenetic and dating analyses. The resulting phylogenetic tree supported monophyly of the family Scolopendridae which was in turn split into two clades constituting tribes Otostigmini and Scolopendrini-Asanadini. Bayesian divergence date estimates suggested that the earliest diversifications within various genera were between 86 and 73mya, indicating that these genera might have Gondwanan origin. In particular, at least four genera of Scolopendrid centipedes, Scolopendra, Cormocephalus, Rhysida and Digitipes, might have undergone diversification on the drifting peninsular India during the Late Cretaceous. These putative Gondwanan taxa can be subjected to more extensive sampling to confirm their Gondwanan origin.

A phylogeny of the only ground-dwelling radiation of Cyrtodactylus (Squamata, Gekkonidae): diversification of Geckoella across peninsular India and Sri Lanka.

The subgenus Geckoella, the only ground-dwelling radiation within Cyrtodactylus, closely overlaps in distribution with brookii group Hemidactylus in peninsular India and Sri Lanka. Both groups have Oligocene origins, the latter with over thrice as many described species. The striking difference in species richness led us to believe that Geckoella diversity is underestimated, and we sampled for Geckoella across peninsular India. A multi-locus phylogeny reveals Geckoella diversity is hugely underestimated, with at least seven undescribed species, doubling previously known richness. Strikingly, the new species correspond to cryptic lineages within described Indian species (complexes); a number of these endemic lineages from the hills of peninsular India outside the Western Ghats, highlighting the undocumented diversity of the Indian dry zone. The Geckoella phylogeny demonstrates deep splits between the Indian species and Sri Lankan G. triedrus, and between Indian dry and wet zone clades, dating back to the late Oligocene. Geckoella and brookii group Hemidactylus show contrasting diversification patterns. Geckoella shows signals of niche conservatism and appears to have retained its ancestral forest habitat. The late Miocene burst in speciation in Geckoella may be linked to the expansion of rain forests during the mid-Miocene climatic optimum and subsequent fragmentation with increasing late Miocene aridification.

Taxonomic Implications of a Field Study of Morphotypes of Hanuman Langurs (Semnopithecus entellus) in Peninsular India

The Hanuman langur is one of the most widely distributed and morphologically variable non-human primates in South Asia. Even though it has been extensively studied, the taxonomic status of this species remains unresolved due to incongruence between various classification schemes. This incongruence, we believe, is largely due to the use of plastic morphological characters such as coat color in classification. Additionally these classification schemes were largely based on reanalysis of the same set of museum specimens. To bring greater resolution in Hanuman langur taxonomy we undertook a field survey to study variation in external morphological characters among Hanuman langurs. The primary objective of this study is to ascertain the number of morphologically recognizable units (morphotypes) of Hanuman langur in peninsular India and to compare our field observations with published classification schemes. We typed five color-independent characters for multiple adults from various populations in South India. We used the presence-absence matrix of these characters to derive the pair-wise distance between individuals and used this to construct a neighbor-joining (NJ) tree. The resulting NJ tree retrieved six distinct clusters, which we assigned to different morphotypes. These morphotypes can be identified in the field by using a combination of five diagnostic characters. We determined the approximate distributions of these morphotypes by plotting the sampling locations of each morphotype on a map using GIS software. Our field observations are largely concordant with some of the earliest classification schemes, but are incongruent with recent classification schemes. Based on these results we recommend Hill (Ceylon Journal of Science, Colombo 21:277-305, 1939) and Pocock (Primates and carnivora (in part) (pp. 97–163). London: Taylor and Francis, 1939) classification schemes for future studies on Hanuman langurs.

Molecular systematics and conservation of the langurs and leaf monkeys of South Asia.

Numerous morphology-based classification schemes have been proposed for langurs and leaf monkeys of South Asia but there is very little agreement between them. An incorrect classification scheme when used as a basis for biogeographic studies can support erroneous hypotheses. Further, lack of taxonomic resolution will also confound conservation efforts, given that conservation biologists use traditional morphology-based-classification schemes to prioritize species for conservation. Here, I have revisited recent molecular phylogenetic studies done on langurs and leaf monkeys of South Asia. Results from these studies are in turn used to derive a rational and scientific basis for prioritizing species for conservation. Molecular data support the classification of langurs of the Indian subcontinent—Hanuman, Nilgiri and purple-faced langurs—in the genus Semnopithecus, whereas Phayre’s leaf monkey along with other Southeast Asian leaf monkeys form another distinct clade (Trachypithecus). The phylogenetic position of capped and golden langurs remains unresolved. Molecular data suggest that they are closely related to each other but this group might have evolved through past hybridization between Semnopithecus and Trachypithecus. Additionally, genetic data also support the splitting of the so-called Hanuman langurs into at least three species. The scores for taxonomic uniqueness of langurs and leaf monkeys of South Asia were revised using this molecular phylogeny-based classification. According to the revised scores, Phayres leaf monkey and golden langur are priority species for conservation followed by capped and Nilgiri langurs.Numerous morphology-based classification schemes have been proposed for langurs and leaf monkeys of South Asia but there is very little agreement between them. An incorrect classification scheme when used as a basis for biogeographic studies can support erroneous hypotheses. Further, lack of taxonomic resolution will also confound conservation efforts, given that conservation biologists use traditional morphology-based-classification schemes to prioritize species for conservation. Here, I have revisited recent molecular phylogenetic studies done on langurs and leaf monkeys of South Asia. Results from these studies are in turn used to derive a rational and scientific basis for prioritizing species for conservation. Molecular data support the classification of langurs of the Indian subcontinent—Hanuman, Nilgiri and purple-faced langurs—in the genus Semnopithecus, whereas Phayre’s leaf monkey along with other Southeast Asian leaf monkeys form another distinct clade (Trachypithecus). The phylogenetic position of capped and golden langurs remains unresolved. Molecular data suggest that they are closely related to each other but this group might have evolved through past hybridization between Semnopithecus and Trachypithecus. Additionally, genetic data also support the splitting of the so-called Hanuman langurs into at least three species. The scores for taxonomic uniqueness of langurs and leaf monkeys of South Asia were revised using this molecular phylogeny-based classification. According to the revised scores, Phayres leaf monkey and golden langur are priority species for conservation followed by capped and Nilgiri langurs.