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The Out-Of-India Hypothesis: What Do Molecules Suggest? 687

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The Out-Of-India Hypothesis: What Do Molecules Suggest? 687 The Out-of-India hypothesis: What do molecules suggest? 687 The Out-of-India hypothesis: What do molecules suggest? ANIRUDDHA DATTA-ROY and K PRAVEEN KARANTH* Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560 012, India *Corresponding author (Email, [email protected]) 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 confi ned 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 oversimplifi ed yet explicit phylogenetic predictions. These predictions can be tested through the use of molecular phylogenetic tools in conjunction with palaeontological and geological data. [Datta-Roy A and Karanth K P 2009 The Out-of-India hypothesis: What do molecules suggest?; J. Biosci. 34 687–697] DOI 10.1007/s12038-009-0057-8 1. Introduction ferry model” was proposed, according to which the rafting Indian plate carried ancient Gondwanan forms to Asia (Mani Peninsular India was part of the Gondwana supercontinent 1974; McKenna 1995; Hedges 2003; Briggs 2003a; Bossuyt about 200 million years ago (mya) (Mani 1974; Briggs et al. 2006). Upon collision with Asia these Gondwanan 1989, 2003a; Hedges 2003). During the Jurassic period, forms dispersed out of India and into Asia (also called approximately 158-160 mya, the Indo-Madgascar plate as “Out-of-India” hypothesis). However, Briggs (2003a) drifted away from East Africa, followed by the separation suggested that peninsular India was not completely isolated of peninsular India from Madagascar around 84-96 mya during its northward journey and that faunal links were (Briggs 2003a). The geomorphological, geological, and maintained with Africa and Madagascar until it collided geophysical data supporting these events have been with the Eurasian plate, at least with respect to salt water discussed in detail by previous workers (Krishnan 1974; resistant faunal elements. Chatterjee and Scotese (1999) also Chatterjee and Scotese 1999; Ali and Aitchison 2008). discussed fossil evidence that supports a Late Cretaceous After the separation from Madagascar, the Indian plate dispersal of dinosaurs and other vertebrates from Africa supposedly underwent a period of isolation for about 30- and Europe. Rage (1996) even depicted an end Cretaceous 40 million years (my), before colliding with the Eurasian arrangement of tectonic plates, in which the northern part plate around 40-50 mya. Given this long period of isolation of peninsular India was attached to the Eurasian plate and experienced by the Indian plate it is expected that the fauna the southern part to the Madagascar plate. Thus the limited and fl ora of peninsular India would be unique and highly fossil data suggests that during its northward journey, the endemic (Kaest 1971). With the collision of the Indian and Indian plate has remained close to Africa and Madagascar Eurasian plates there was exchange of biota between Asia even when it began to contact Eurasia (Briggs 2003a). and peninsular India (Mani 1974). Consequently, the “Biotic Nevertheless, numerous recent molecular phylogenetic Keywords. Gondwanan biogeography; India; molecular dating; molecular phylogeny http://www.ias.ac.in/jbiosci J. Biosci. 34(5), November 2009, 687–697, © IndianJ. Biosci. Academy 34(5), of November Sciences 2009 687 688 Aniruddha Datta-Roy and K Praveen Karanth studies have supported the Out-of-India hypothesis Out-of-India hypothesis, the Gondwanan taxa distributed (see Karanth 2006 and references therein). outside of India should be nested within a clade consisting As mentioned above, since the Indian plate collided of Gondwanan taxa from India. In recent times, molecular with Eurasia, not only did potentially Gondwanan elements phylogenetic approaches have been increasingly used to disperse out of India and into Asia, but also many Asian and address biogeographical problems, such as Gondwanan African elements dispersed into India. Mani (1974) points biogeography. In this paper we review studies undertaken on out that these relatively young intrusive elements are largely a range of taxa of supposedly Gondwanan origin, to better derived from the Indo-Chinese and Malayan subregion understand this remarkable pattern. Additionally, we discuss of tropical Asia. Numerous authors have commented the utility of molecular phylogenetic approaches in the study on the overall faunal similarity between peninsular India of biogeography of the Indian subcontinent given the above and Southeast Asia (Blanford 1901; Hora 1949; Jayaram mentioned requirements. 1949; Mani 1974). This out of Asia and into India scenario has been discussed in mammals based on palaeontological 2. Flora evidence (Clyde et al. 2003). Hora (1949) also discusses the Southeast Asian origin of freshwater, torrential fi shes 2.1 Asian dipterocarps of Western Ghats and their possible dispersal routes. In case of plants, the possibility of an out of Asia and into One of the fi rst plant systems studied explicitly in a India origin has been discussed by Lakhanpal (1970) and phylogenetic framework in order to test for the Out-of- Bande (1992). In this regard, a recent molecular study of India hypothesis were the Asian dipterocarps. Work done freshwater gastropods of supposedly Gondwanan origin is by Dayanandan et al. (1999) on this family revealed that of much relevance (Köhler and Glaubrecht 2007). Results they formed a monophyletic group with the Sarcolaenaceae, from this work indicate that the gastropods of Southern India a tree family endemic to Madagascar. The most likely way are nested within a clade consisting of Southeast Asian taxa. by which the Dipterocarpaceae would have reached the It is clear that this group in fact dispersed out of Asia and Asian mainland is via peninsular India acting as a biotic into India. ferry, followed by dispersal out of India. However, the split Much of the evidence for the Gondwanan origin of some between Malagasy and Indian taxa has not been dated. Thus, Asian biota and the Out-of-India hypothesis is based on more recent transoceanic dispersal cannot be ruled out as fossil data and indirect evidence from species distribution alternative explanation. Transmarine dispersal appears ranges. As has been shown by the gastropod work (Köhler unlikely as these families also share similar ectomycorrhizal and Glaubrecht 2007), extant faunal distributions cannot symbiosis, which is thought to have evolved in their be taken as evidence supporting Gondwanan origin and common ancestor (Ducousso et al. 2004). Ectomycorrhizal Out-of-India hypothesis. Another confounding factor in symbiosis occurs in only 3% of angiosperms. Therefore, it our attempt at understanding Gondwana biogeography is is unlikely that similar mycorrhiza in the two families could the situation wherein classifi cation of a taxonomic group have originated independently. does not correspond to its evolutionary relationship, i.e, when unrelated taxa have be erroneously placed in the same taxonomic group (Karanth 2006). Thus to rigorously 2.2 Crypteroniaceae test these hypotheses the knowledge of phylogenetic relationships of the taxa of interest is vital. The family Crypteroniaceae is distributed in tropical Asia In order to confi rm that a taxonomic group is of and their morphology-based taxonomy suggests affi nities Gondwanan origin and that vicariance due to continental drift with African (western Gondwanan) taxa. This implies that is responsible for its present distribution, three requirements this group may have reached Asia rafting on the drifting must be met. Firstly, the taxonomic group should be Indian plate. Molecular work done by Conti et al. (2002) monophyletic and distributed predominantly in some (if not confi rmed their close relationship with African species all) of the former Gondwanan fragments of South America, and molecular date estimates by Rutschmann et al. Africa, Madagascar, peninsular India and Australia. (2004) suggest that Crypteroniaceae diverged from their Secondly, in the phylogenetic trees of Gondwanan taxa, west Gondwanan sister clade in the early to middle the split between taxa from various Gondwanan fragments Cretaceous. The molecular dates are concordant with an should mirror the sequence of geophysical events that led ancient Gondwanan origin of the Crypteroniaceae, and the to the breakup of the Gondwana. Thirdly, the estimated hypothesis that they were transported by the drifting Indian divergence
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