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Fossil Palm Fruits from India Indicate a Cretaceous Origin of Arecaceae Tribe Borasseae

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applyparastyle “fig//caption/p[1]” parastyle “FigCapt” Botanical Journal of the Linnean Society, 2019, 190, 260–280. With 6 figures. Fossil palm fruits from India indicate a Cretaceous origin of Arecaceae tribe Borasseae Downloaded from https://academic.oup.com/botlinnean/article-abstract/190/3/260/5518348 by University of Michigan user on 13 July 2019 KELLY K. S. MATSUNAGA1, , STEVEN R. MANCHESTER2, RASHMI SRIVASTAVA3, Keywords=Keywords=Keywords_First=Keywords 4 1* HeadA=HeadB=HeadA=HeadB/HeadA DASHRATH K. KAPGATE and SELENA Y. SMITH HeadB=HeadC=HeadB=HeadC/HeadB 1Earth and Environmental Sciences, University of Michigan, Ann Arbor, MI, USA HeadC=HeadD=HeadC=HeadD/HeadC 2Florida Museum of Natural History, Gainesville, FL, USA Extract3=HeadA=Extract1=HeadA 3Birbal Sahni Institute of Palaeosciences, Lucknow, India REV_HeadA=REV_HeadB=REV_HeadA=REV_HeadB/HeadA 4 REV_HeadB=REV_HeadC=REV_HeadB=REV_HeadC/HeadB Department of Botany, J.M Patel College, Bhandara, Maharashtra, India REV_HeadC=REV_HeadD=REV_HeadC=REV_HeadD/HeadC REV_Extract3=REV_HeadA=REV_Extract1=REV_HeadA Received 26 November 2018; revised 3 March 2019; accepted for publication 20 April 2019 BOR_HeadA=BOR_HeadB=BOR_HeadA=BOR_HeadB/HeadA BOR_HeadB=BOR_HeadC=BOR_HeadB=BOR_HeadC/HeadB BOR_HeadC=BOR_HeadD=BOR_HeadC=BOR_HeadD/HeadC The fossil record of palms (Arecaceae) is essential for understanding the deep evolutionary and geographical history BOR_Extract3=BOR_HeadA=BOR_Extract1=BOR_HeadA of the family. We studied palm fruit fossils from the c. 67–64-Myr-old Deccan Intertrappean Beds of India to infer EDI_HeadA=EDI_HeadB=EDI_HeadA=EDI_HeadB/HeadA the systematic relationships of the fossils and their relevance to palm evolution. Using X-ray micro-computed EDI_HeadB=EDI_HeadC=EDI_HeadB=EDI_HeadC/HeadB tomography, physical sectioning techniques and a total-evidence phylogenetic analysis, we show that these fossils EDI_HeadC=EDI_HeadD=EDI_HeadC=EDI_HeadD/HeadC represent a crown group member of subtribe Hyphaeninae (tribe Borasseae, subfamily Coryphoideae) allied EDI_Extract3=EDI_HeadA=EDI_Extract1=EDI_HeadA with the extant genera Satranala and Bismarckia, now endemic to Madagascar. These fossils, synonymized here CORI_HeadA=CORI_HeadB=CORI_HeadA=CORI_HeadB/HeadA as Hyphaeneocarpon indicum, provide evidence for the existence of crown group Hyphaeninae during the late CORI_HeadB=CORI_HeadC=CORI_HeadB=CORI_HeadC/HeadB Maastrichtian–early Danian. This pre-dates prior age estimates for the Hyphaeninae crown node by nearly 40 Myr CORI_HeadC=CORI_HeadD=CORI_HeadC=CORI_HeadD/HeadC and implies an earlier divergence of Borasseae. The presence of Hyphaeneocarpon in India shows that Borasseae CORI_Extract3=CORI_HeadA=CORI_Extract1=CORI_HeadA have persisted in the Indian Ocean region for > 64 Myr. This study illustrates the utility of palm fruit characters ERR_HeadA=ERR_HeadB=ERR_HeadA=ERR_HeadB/HeadA for placing fossils in a phylogenetic context and has important implications for understanding the evolution and ERR_HeadB=ERR_HeadC=ERR_HeadB=ERR_HeadC/HeadB diversification of Borasseae and the palaeobiogeography of palms. ERR_HeadC=ERR_HeadD=ERR_HeadC=ERR_HeadD/HeadC ADDITIONAL KEYWORDS: biogeography – Deccan – Maastrichtian – MrBayes – palaeobotany – Palaeocene – ERR_Extract3=ERR_HeadA=ERR_Extract1=ERR_HeadA palm phylogeny – X-ray micro-computed tomography (µCT). INRE_HeadA=INRE_HeadB=INRE_HeadA=INRE_HeadB/HeadA INRE_HeadB=INRE_HeadC=INRE_HeadB=INRE_HeadC/HeadB INRE_HeadC=INRE_HeadD=INRE_HeadC=INRE_HeadD/HeadC INRE_Extract3=INRE_HeadA=INRE_Extract1=INRE_HeadA INTRODUCTION 2009; Pross et al., 2012; Suan et al., 2017). The fossil record of palms thus represents an important source of App_Head=App_HeadA=App_Head=App_HeadA/App_Head Palms (Arecaceae) are found today throughout tropical data for understanding the deep evolutionary history BList1=SubBList1=BList1=SubBList regions worldwide occupying a variety of environments of the family and terrestrial environments of the BList1=SubBList3=BList1=SubBList2 ranging from arid deserts to tropical rainforests geological past. SubBList1=SubSubBList3=SubBList1=SubSubBList2 (Dransfield et al., 2008). Currently, Arecaceae comprise The Maastrichtian–Danian (c. 67–64 Ma) Deccan SubSubBList3=SubBList=SubSubBList=SubBList c. 2600 species classified into five subfamilies and Intertrappean Beds of India host plant fossil SubSubBList2=SubBList=SubSubBList=SubBList 181 genera (Baker & Dransfield, 2016). Additionally, assemblages with numerous palm macrofossils (Bonde, SubBList2=BList=SubBList=BList they have a rich fossil record extending back to the 2008; Kapgate, 2009; Srivastava, 2011). Located Late Cretaceous; unequivocal palm fossils first primarily in central India, these localities preserve appear during the Turonian (c. 94–90 Ma) and are the remains of palm stems, leaves, roots, pollen, geographically widespread by the Maastrichtian inflorescences and fruits, indicating that palms were (c. 72–66 Ma; Gee, 2001; Harley, 2006; Dransfield an important component of the vegetation of central et al., 2008). Subsequently, they achieved a global India during the Late Cretaceous and Palaeocene, distribution, extending into high-latitude regions during which time India was geographically isolated such as Alaska and Antarctica during the warm and from other major landmasses (Ali & Aitchison, 2008; equable climatic conditions of the Eocene (Sluijs et al., Chatterjee, Goswami & Scotese, 2013). Today palms do not comprise a significant component of the vegetation of central India. Although the flora of India includes *Corresponding author. E-mail: [email protected] © 2019 The Linnean Society of London, Botanical Journal of the Linnean Society, 2019, 190, 260–280 260 FOSSIL FRUITS OF BORASSEAE FROM INDIA 261 c. 96 species in 20 genera (Kulkarni & Mulani, 2004), such as placement of fossils in the stem or crown of Modern palms in India are thought to have descended a group and alliances with particular extant taxa. from relatively recent colonizations rather than an This information can facilitate inferences of historical ancient flora (Baker & Couvreur, 2013). However, little biogeography and character evolution and inform is known about the taxonomic composition of historical node calibrations in future dating analyses, providing palm assemblages and the role of India in the valuable information on the diversification of palms in evolutionary and biogeographical history of Arecaceae. deep time. Downloaded from https://academic.oup.com/botlinnean/article-abstract/190/3/260/5518348 by University of Michigan user on 13 July 2019 Over the course of nearly a century of study on the Deccan flora, c. 168 fossil species have been assigned to Arecaceae, including 85 species based on MATERIAL AND METHODS stem specimens, 37 on fruits, 28 on leaves, roots and inflorescence axes, and 18 on palynomorphs (Bonde, LOCALITY AND AGE 2008; Kapgate, 2009). These diverse assemblages The Deccan Volcanic Province (DVP) comprises a could be essential for understanding the evolutionary sequence of continental flood basalts (traps) formed tempo in palm diversification, historical biogeography during the late Maastrichtian to early Danian (c. of palm lineages and transitions in the terrestrial 67–64 Ma, chrons 30N–29N; Hooper, Widdowson vegetation of India through time. For example, & Kelley, 2010; Renne et al., 2015; Schoene et al., recent re-examination of Palmocarpon drypeteoides 2015), exposed across central and western peninsular (Mehrotra, Prakash & Bande) Manchester, Bonde, India. Intertrappean sedimentary layers, which occur Nipunage, Srivastava, Mehrotra & Smith revealed between some basalt flows and represent quiescent morphological characters diagnostic of subtribe intervals between volcanic episodes, frequently Attaleinae, which is currently most diverse in South contain permineralized plants preserved three- America and with no representatives in India or dimensionally in chert deposits. More than 50 plant elsewhere in Asia (Manchester et al., 2016). However, fossil-bearing localities have been discovered during the systematic affinities of most of the palm fossils in the last century, most of which are concentrated the Deccan Intertrappean beds are poorly understood, in central India in the states of Maharashtra and as is the extent to which the number of described Madhya Pradesh and occur primarily in the north- palms accurately represents the true species richness eastern portion of the Deccan Main Plateau or in the in these fossil assemblages. Morphological studies Mandla subprovince (Fig. 1; Kapgate, 2009; Smith and taxonomic revisions are therefore essential for et al., 2015). Where magnetostratigraphic data are understanding the Deccan floras and applying them to available, many of these localities fall within Chron broader questions of palm evolution. 29R, which straddles the Cretaceous–Palaeogene In this study we re-examined five previously described (K-Pg) boundary (M. Widdowson, pers. comm., 2018). fossil palm species, in light of new specimens recovered Although more precise ages of most macrofossil from the Deccan Intertrappean Beds at Dhangaon, localities are poorly constrained, many are considered Keria and Mohgaonkalan: Hyphaeneocarpon either late Maastrichtian or early Danian depending indicum Bande, Prakash & Ambwani; Palmocarpon on their location in the DVP, stratigraphic continuity arecoides Mehrotra; Arecoidocarpon kulkarnii Bonde; with dated outcrops and palynomorph content A. palasundarensis Bonde; and Pandanusocarpon (Samant & Mohabey, 2009). Specifically,
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