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Wood of Oleaceae from the Latest Cretaceous of India – the Earliest Olive Branch?

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Wood of Oleaceae from the Latest Cretaceous of India – the Earliest Olive Branch? SrivastavaIAWA Journal et al. 36 – Oleoxylon(4), 2015: from443–451 India 443 WOOD OF OLEACEAE FROM THE LATEST CRETACEOUS OF INDiA – THE EARLiEST OLiVE BRANCH? Rashmi Srivastava1,*, Elisabeth A. Wheeler2, Steven R. Manchester3, and Pieter Baas4 1Birbal Sahni Institute of Palaeobotany, 53 University Road, Lucknow 226 007, India 2Department of Forest Biomaterials, N.C. State University, Raleigh, NC 27695-8005, U.S.A. 3Florida Museum of Natural History, Gainesville, FL 32611-7800, U.S.A. 4Naturalis Biodioversity Center, PO Box 9517, 2300 RA Leiden, The Netherlands *Corresponding author; e-mail: [email protected] Abstract The wood of Oleoxylon deccanense, reported informally in 1981 from the Dec- can Intertrappean Beds of central India, is re-examined. We provide a formal diagnosis for the species and a more detailed description. The similarity to wood from species groups of the modern genera Chionanthus and Olea leads us to infer that this fossil taxon probably belongs to the monophyletic drupaceous subtribe Oleinae of the olive family, Oleaceae (Lamiales), although affinities with Rhamnaceae and Rutaceae cannot be wholly excluded. Since the fossil is from a late Maastrichtian-Danian horizon (65–67 MY BP) this would imply that a member of the Oleaceae was part of the flora that inhabited India several million years prior to the tectonic impact of India with Asia. The seemingly mod- ern appearance of this and other Deccan fossil woods is briefly discussed. Keywords: Deccan Intertrappean Beds, Maastrichtian, Paleocene, India, wood anatomy, Oleaceae, Rhamnaceae, Rutaceae. Introduction In 1981, Trivedi and Srivastava reported a fossil wood Oleoxylon deccanensis (erron. for deccanense), showing resemblance with the genus Olea from the Deccan Intertrap- pean Beds of Jheria village about 3 km south-west of Mohgaon Kalan, Chhindwara District, Madhya Pradesh, India. However, they did not provide a diagnosis and holo- type number for the fossil and thus, according to the rules of nomenclature, this name is not valid (McNeill et al. 2012). We recently had the opportunity to re-examine this fossil wood, and herein describe it in more detail and compare it with more recent comprehensive information on modern and fossil woods in general (InsideWood 2004-onwards; Wheeler 2011) and the Oleaceae family in particular (Baas & Zhang Xinying 1986; Baas et al. 1988). The original specific name assigned to this fossil is retained and validated in the present communication by providing a diagnosis and holotype number for the specimen. The collecting locality is close to the town of Keria, adjacent to which several petrified woods have been collected and described previously (reviewed in Kapgate © International Association of Wood Anatomists, 2015 DOI 10.1163/22941932-20150113 Published by Koninklijke Brill NV, Leiden Downloaded from Brill.com10/07/2021 11:35:16AM via free access 444 IAWA Journal 36 (4), 2015 2005). These sites are apparently continuous with the classic Mohgaon Kalan fossil forest horizon, which is treated as Late Maastrichtian based on stratigraphic correla- tion and palynology (Samant & Mohabey 2013). These are interbeds of the great sequence of basalt flows known as the Deccan Traps. The area occupied by the Dec- can Traps today is about 500,000 km2 in peninsular India while the original stretch may have been three times greater, including sediments found in the Arabian Sea to the west of Mumbai (Jay & Widdowson 2008). The outpourings of lava were as- sociated with the northward drifting of the Indian Plate after it was separated from Gondwana during the Early Cretaceous and moved over the volcanic hot spot over which Reunion island is now positioned in the Indian Ocean (Chatterjee et al. 2013). Recent studies based on radiometric dating and planktonic foraminifera, indicate that the age of the intertrappean sediments ranges from upper Maastrichtian to Danian (Keller et al. 2009). The precision of whole-rock basalt radiometric dating is insuf- ficient to resolve whether many of the sites are latest Maastrichtian or earliest Paleo- cene. However, palynological and faunal assemblages suggest a Maastrichtian age for most of the intertrappean exposures with few Paleocene indicators (Kar & Srinivasan 1998; Samant & Mohabey 2009, 2013). Smith et al. (2015) give an integrated review of the geology and paleobotany of the Late Cretaceous-Paleocene transitions of India that also highlights the remaining uncertainties of the precise dating of the macrofossils. Paleobotanical and biogeographical studies suggest that some angiosperm taxa might have an Indian origin because their earliest records are known from these Maastrichtian- Danian sediments. Numerous petrified woods have been reported from the Deccan intertrappean sediments since the 1920s (compiled in Lakhanpal et al. 1976; Srivastava 1991; Kapgate 2005; Srivastava & Guleria 2006; Srivastava 2011). We have recently re-examined many of the original prepared slides of silicified wood specimens stored at the Birbal Sahni Institute of Paleobotany as part of a collaborative project to facilitate access to data and images on the internet (via InsideWood). In this article we elaborate on the fossil record of Oleoxylon, analyze its anatomical character combination in comparison with a global dataset of modern and fossil woods, and discuss its signifi- cance in the context of the fossil record of Oleaceae and the floristic composition and biogeographic importance of the Deccan flora. Systematic description Core – Eudicots Order – Lamiales Family – Oleaceae Subfamily – Oleoideae Genus – Oleoxylon Dupéron 1973 emend. Heteroxylous wood with characters of the family Oleaceae. Diffuse porous wood. Vessels with tangential diameters < 200 µm, solitary and in radial multiples. Perfora- tions simple. Intervessel pits alternate, minute to medium-sized. Axial parenchyma not abundant (scanty paratracheal-vasicentric). Rays narrow and low, usually 2–3 cells wide, heterocellular. Fibers with simple pits, non-septate. Longitudinal canals, radial canals, storied structure, oil/mucilage cells, and cambial variants absent. Downloaded from Brill.com10/07/2021 11:35:16AM via free access Srivastava et al. – Oleoxylon from India 445 The diagnosis is emended, broadening the size range of intervessel pitting so as to be similar to that of present-day Oleoideae, and specifying diffuse porous wood, scanty paratracheal to vasicentric axial parenchyma, nonseptate fibers, and heterocellular rays (features included in the original description of Oleoxylon), and adding vessel- parenchyma pits similar to intervessel pits, which is consistent with Dupéron’s “Bois hétéroxylé possédant les caractères de la famille de Oleaceae” and necessary for this generic diagnosis to be unique. Species – Oleoxylon deccanense Srivastava, Wheeler, Manchester & Baas ex Trivedi & Srivastava. Holotype – BSIP Museum No. 40570. Stratigraphic horizon – Deccan Intertrappean Beds, Dhuma Formation of Amarkan- tak Group. Locality – Jheria, about 3 km southwest of Mohgaon Kalan, Chhindwara District, Madhya Pradesh, India. Age – Maastrichtian-Danian. Material – The description is based on a well-preserved piece of petrified wood meas- uring about 4.9 cm in width and 15 cm long. Diagnosis – Diffuse porous. Vessels often in long radial multiples of >4. Perforation plates simple. Intervessel pits alternate, 8–10 µm; vessel-ray pits similar to interves- sel pits. Fibers with inconspicuous pits, nonseptate. Parenchyma scanty paratracheal to vasicentric and marginal. Rays 1–3-seriate, heterocellular. Storied structure absent. Description (Fig. 1–6): Wood diffuse porous. Growth rings indistinct, but one (possibly traumatic?) ring boundary present and demarcated by marginal parenchyma preceded by weakly flattened latewood fibers and differential distribution of smaller vessels. Vessels diffuse, 25–55 (average 32)/mm2, 14–20% solitary, remainder in ra- dial multiples of 2–8 (rarely up to 17), round to oval in cross section, those in multiples slightly angular and flattened at the place of contact, tangential diameter 20–118 (average 59, SD 21) µm, radial diameter 30–150 (average 76, SD 30) µm. Vessel member length 225–530 (mostly 350–450) µm; perforations simple and oblique, often tailed end walls; intervessel pits alternate, medium-sized, 8–10 µm in diameter. Vessel-ray pits poorly preserved, at places seemingly similar to intervessel pits in shape and size. Vascular tracheids intergrading with narrow vessel elements in radial multiples very infrequent- ly present. Fibers angular in cross section, thin-walled to medium thick-walled, walls 3–4 µm thick, pits not conspicuous, nonseptate, 12–18 µm in diameter. Axial paren- chyma scanty paratracheal to vasicentric (only one, probably traumatic, seemingly marginal band of 2–4 or more cells wide present in the cross section examined), cells 20–32 µm in diameter, 3–4 cells per parenchyma strand. Rays heterocellular, 8–12 rays per mm, 1–3- (mostly 1–2-)seriate; 5–32 cells or 234–814 (mostly up to 580) µm high, made up of procumbent cells in the central portion with 1–4 marginal rows of upright or square cells at one or both ends; uniseriate rays made up of either upright or both procumbent and upright cells, about 3–10 cells or 134–580 µm tall. Codified description according to the IAWA Hardwood List (IAWA Committee 1989): 2 5 10 13 22 23v 26 30 41 42 48 49 60v 61 66 69 78 79 89? 92 97 106 107 115. Downloaded from Brill.com10/07/2021 11:35:16AM via free access 446 IAWA Journal 36 (4), 2015 1 2 3 4 5 6 Figure 1–6. Oleoxylon deccanense. – 1–3: Diffuse porous wood, TS. – 1 & 2: Radial multiples of 4 or more common, axial parenchyma scanty paratracheal. – 3: Region where vessel multiples of 4 or more are not common (post-injury?). – 4: Rays 1–3-seriate, mostly 2-seriate, TLS. – 5: Simple perforation plate and ray composition, RLS. – 6: Crowded alternate intervessel pits, TLS. — Scale bars: 200 µm in 1 & 3; 100 µm in 2, 4 & 5; 20 µm in 6. Downloaded from Brill.com10/07/2021 11:35:16AM via free access Srivastava et al. – Oleoxylon from India 447 Discussion Affinities of the fossil We used the InsideWood database, the literature, and the Leiden slide collection (Lw) for searching the best matches of the fossil.
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