The 18-Electron Rule for Main-Group Alkaline Earth Octacarbonyl
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10 Natl Sci Rev, 2019, Vol. 6, No. 1 PERSPECTIVES GEOSCIENCES Natatanuran frogs used the Indian Plate to step-stone disperse and radiate across the Indian Ocean Downloaded from https://academic.oup.com/nsr/article-abstract/6/1/10/5090988 by University of Kentucky Libraries user on 29 April 2019 Zhi-Yong Yuan1,2,†, Bao-Lin Zhang1,3,†, Christopher J. Raxworthy4, David W. Weisrock5, Paul M. Hime5,6, Jie-Qiong Jin1,7, Emily M. Lemmon8, Alan R. Lemmon9, Sean D. Holland8, Michelle L. Kortyna8, Wei-Wei Zhou1,7, Min-Sheng Peng1,10,JingChe1,7,11,∗ and Elizabeth Prendini4 Investigating the evolutionary history [6–9]. Previous studies addressed some data set assembled for this group. Sam- of widespread higher taxa, subjected to of these issues from palaeobiogeograph- ples include all major lineages, 85 Natata- multiple tectonic events, can provide ical and evolutionary perspectives, but nura species and 20 outgroup taxa (Sup- evidence for or against various palaeo- were inconclusive due to the selection of plementary Data). The novel evidence geographical models of early Earth taxa that did not include all landmasses or reveals how Natatanuran frogs inter- history [1,2]. Contemporary biotic dis- the limited recovery of evolutionary rela- changed between Laurasia and Gond- tributions have been strongly influenced tionships due to the use of sequence data wana around the Indian Ocean during by events associated with the breakup from only a few genes [10–13]. the Cretaceous–Palaeogene, challenging of Gondwana into present-day Africa, The neobatrachian clade Natatanura recent biogeographical assumptions and Antarctica, Australia, South America, are an ideal group to infer Gondwanan providing new insights into Indian Ocean Madagascar and India, during the Late geological and environmental history due biotic exchanges. Mesozoic and Early Palaeogene [2]. The to their ancient origins (divergence from fragmentation of Gondwana and subse- Afrobatrachia at around 100 Ma), high quent tectonic drift ultimately allowed species diversity (>1500 extant species), biotic exchanges between Laurasia and almost cosmopolitan distribution (ab- Gondwana [2,3], influencing the global sent only from Antarctica), general low distributions of many taxa. terrestrial vagility and poor overwater The relative positions of the post- dispersal capabilities [14]. Previous breakup Gondwanan landmasses during studies suggested the divergence of the Late Cretaceous, especially of the In- Natatanura was characterized by a dian and Australian plates around the historical association with the breakup of Indian Ocean, are highly debated [1]. Gondwanan plates [12]. These frogs are The plate reshuffling was probably ac- thus an appropriate group of organisms companied by the formation of multi- to test hypotheses of Cretaceous– KP ple temporary land bridges and involved Palaeogene biotic exchanges between GR biotic exchange among the plates. Al- Laurasia and Gondwana around the though most models agree that the In- Indian Ocean. However, prior studies dian Plate carried a biotic ‘ferry’ of taxa that have included Natatanura failed to (both plants and animals) to Asia after resolve the major nodes in its phylogeny Figure 1. Schematic representation of differ- it broke away from other Gondwanan or suffered from incomplete lineage ent hypotheses regarding land connections and landmasses from about 88 to 55 Ma sampling, which, until now, hampered corridors for dispersal among the landmasses [2,4], both geological and paleontologi- conclusive testing of these hypotheses around the Indian Ocean from 88 to 55 Ma. cal data also support land bridges or mi- [12,15]. (1) Africa and India were reconnected to each nor marine barriers that permitted biotic Here, we integrate phylogenetic, other directly [3,5]; (2) Asia and Madagascar exchanges with other Gondwanan land- biogeographic and molecular dating were linked by India, with possible disper- masses (e.g. Africa, Madagascar; Fig. 1) methods to reconstruct the spatiotem- sal between Asia and Madagascar over India [3,5]. There is also debate about whether poral diversification of Natatanura (see and the Seychelles plateau [5]; (3) Antarctic– Australia–New Guinea and Madagascar were Antarctica–Australia–New Guinea was Supplementary Data). Results resolve connected to: (i) the Indian Plate via the connected by the Gunnerus Ridge (GR) [6,7]; the evolutionary history of Natatanuran (4) Antarctic–Australia–New Guinea and India Kerguelen Plateau (KP) land bridge and frogs (Supplementary Figs 1–3), based were connected by the Kerguelen Plateau (KP) (ii) Madagascar via the Gunnerus Ridge on molecular data from 376 nuclear loci, [13]. Paleo-reconstructions are modified from (GR) land bridge in the Late Cretaceous representing by far the largest molecular Briggs [5] and Bossuyt et al. [12]. PERSPECTIVES Yuan et al. 11 STEPPING-STONE ROLE OF THE from the Indian landmass [18] and that scenario will also be recovered for other INDIAN PLATE FOR BIOTIC the Indian Plate would have been well nonvolant organisms with Indian Ocean EXCHANGE BETWEEN AFRICA, placed to minimize oceanic dispersal dis- distributions. ASIA AND MADAGASCAR tances between Asia and Madagascar, the Indian Plate could have served as a step- Using the traditional Gondwana and ping stone for long-distance dispersal, as Laurasia model, it has been commonly Downloaded from https://academic.oup.com/nsr/article-abstract/6/1/10/5090988 by University of Kentucky Libraries user on 29 April 2019 suggested previously [13,15]. DISPERSALS WITH assumed that the Indian Plate was an Geological and paleontological ev- ∼ AUSTRALIA–NEW GUINEA isolated island between 88–55 Ma, and idence has previously challenged the the Indian landmass served as an ‘ark’ ‘Indian-biotic Ark’ model. For example, AND ASIA to transport lineages from various biotic Briggs [5] proposed that India was in The Antarctica–Australia–New Guinea groups ‘Out-of-India’ into Asia, after close proximity to other landmasses Plate has been proposed to have been India broke away and drifted northward during its journey northwards because a connected to the Indian Plate by the KP from Gondwana in the Late Cretaceous significant endemic biota did not evolve land bridge, or connected to Madagascar [4]. According to this model, there was on the Indian Plate during this period. by the GR land bridge (Fig. 1)inthe no biotic exchange (which would have Ali and Aitchison [8] proposed the exis- Late Cretaceous [6,7], although these required crossing marine barriers) be- tence of a palaeogeographic connection land bridges have been disputed due to a tween India and its nearby landmass after between Madagascar and India in the lack of evidence that they were sub-aerial ∼ it separated from Madagascar 88 Ma, Late Cretaceous, which may have been during this period [8,9]. Concerning and dispersal events only resumed after formed by the Seychelles–Mascarene Natatanura diversification, we find no India’s collision with Asia in the early Plateau. More recently, Chatterjee et al. sister relationships between frogs from ∼ Eocene ( 65–55 Ma, [16]). ‘Rafting’ [3] argued that biotic links were possibly Antarctica–Australia–New Guinea and of the flora and fauna on the Indian re-established between India and Africa either India or Madagascar (Fig. 2), and Plate enabled unidirectional migration of during the Late Cretaceous, during In- thus find no support for biota exchanges Gondwanan taxa into Asia. However, our dia’s collision with the Kohistan–Ladakh among these landmasses via a KP and phylogenomic results reject this model. Arc along the Indus Suture in the Late GR land bridge. In addition, we date We do not find any periods between 88 Cretaceous. Sister relationships and Natatanura dispersal into Australia–New and 55 Ma, when there was no biotic divergence dates for some vertebrate Guinea to be much later than these exchange occurring between Africa fossil groups (Supplementary Data) also hypothetical Late Cretaceous land and India, India and Asia, or India and support stepping-stone biotic exchange bridges. Madagascar. In contrast, our ancestral via the Indian Plate, consistently with Bossuyt et al.[12] suggested that reconstruction suggests Natatanura our results. However, we did not find Australia–New Guinea acted as a raft, en- originated in Africa and then dispersed any studies of nonvolant extant groups abling Gondwanan Natatanura frogs to ∼ to Asia through India 75.6–72.8 Ma that provide substantial evidence for colonize Southeast Asia, although most (Fig. 2 and Supplementary Fig. 3). It using the Indian Plate as a stepping-stone of their basal relationships were not is unlikely that frogs could cross a large route among these three plates from 88 well resolved. Our genomic-based esti- saltwater barrier, although a few extant to 55 Ma. Although several plant and mates of phylogeny, divergence times species may have made more modest animal groups exhibit sister relationships and biogeographic reconstruction cast Ptychadena oceanic dispersals (e.g. between India (or Asia) and Madagascar doubt on this dispersal route. We found mascareniensis ,[17]). Briggs [5] and (or Africa) (e.g. [19,20]), their deep strong support that the two Australia– et al Chaterjee .[3] suggested a geograph- divergences (>88 Ma) attribute this New Guinea clades, Cornufer and Papu- ical model in which there were ‘corridors’ relationship to ancient vicariance coin- rana, were embedded within Asian clades or ‘landspans’ that reconnected Africa ciding with the breakup of Gondwana.