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Singpuria, a New Genus of Eudicot Flower from the Latest Cretaceous Deccan Intertrappean Beds of India

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Singpuria, a New Genus of Eudicot Flower from the Latest Cretaceous Deccan Intertrappean Beds of India Acta Palaeobotanica 60(2), 323–332, 2020 e-ISSN 2082-0259 https://doi.org/10.35535/acpa-2020-0017 ISSN 0001-6594 Singpuria, a new genus of Eudicot flower from the latest Cretaceous Deccan Intertrappean Beds of India DEEPAK RAMTEKE 1, STEVEN R. MANCHESTER 2*, VAISHALI D. NAGRALE 3 and SELENA Y. SMITH4 1 Gondia Education Society High School and Junior College, Mohadi. Dist., Gondia, 4416014-M.S., India; e-mail: [email protected] 2 Florida Museum of Natural History, University of Florida, Gainesville, Florida, 32611-7800, USA; e-mail: [email protected] 3 Block Resource Center, P.S. Sakoli, Dist., Bhandara, 441802, M.S. India; e-mail: [email protected] 4 Department of Earth & Environmental Sciences and Museum of Paleontology, University of Michigan, Ann Arbor, MI, 48019, USA; e-mail: [email protected] Received 25 February 2020; accepted for publication 6 November 2020 ABSTRACT. A new bisexual flower, Singpuria kapgatei, gen. et sp. nov., is described from chert of latest Creta- ceous age from the Deccan Intertrappean Beds at Singpur, Madhya Pradesh, in central India. The hypogynous, actinomorphic flower is ~1.6 mm wide, with distinct sepals and petals. The androecium consists of 18 tetraspo- rangiate, dithecal, basifixed anthers borne in radial pairs on nine bifurcate filaments. The gynoecium is superior and syncarpous, with pentagonal symmetry. Pollen from the stamens is tricolporate and microreticulate. The combined morphological features of this flower indicate that Singpuria is a eudicot with affinities in the Penta- petalae, but we have been unable to make a more precise assignment. Nevertheless, we consider it useful to place this rare fossil flower on record as an exemplary extinct member of the Deccan biota. Singpuria may represent a clade that was isolated on the Indian subcontinent and became extinct in response to environmental changes at the K/Pg boundary, or later in the Cenozoic as the land mass moved northward through new climate zones and collided with Eurasia. KEYWORDS: Deccan chert, Singpur, paleobotany, bifurcate stamens, tricolporate microreticulate pollen INTRODUCTION Fossil plants preserved in the Deccan Inter- flowerSahnianthus Shukla (1944; syn. Chital- trappean Beds of central India provide insights eypushpam Paradkar, Deccananthus Chitaley into the biota that inhabited the Indian subcon- et Kate, Flosfemina Kar, Ambwani, A. Sahni tinent when it was isolated from other major et Sharma, Lythraceopushpam Narkhede, land masses in the late Cretaceous and early Bhowal et Meshram, Raoanthus Chitaley et Cenozoic. The late Maastrichtian to Paleocene Patel) is an exception, being common at sev- flora of the Deccan Intertrappean Beds is rich eral (at least seven) localities and in various in silicified woods, fruits and seeds (reviewed, states of maturation. Very small flowers (1.0 Kapgate, 2005; Smith et al., 2015; Wheeler to 1.5 mm diameter) include those described et al., 2017) but the flowers that have been as Chenopodianthus VD Kapgate, DK Kap- recovered are low in diversity. The myrtalean gate et Sheikh (2006), Tetraplasandranthus VD Kapgate, DK Kapgate et Sheikh (2009) * Corresponding author and Flosvirulus Kar, Ambwani, A. Sahni et © 2020 W. Szafer Institute of Botany, Polish Academy of Sciences This is an Open Access article, distributed under the terms of the Creative Commons Attribution license (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited. 324 D. Ramteke et al. / Acta Palaeobotanica 60(2): 323–332, 2020 Sharma (2003) from the classic locality of and carefully applying an acetate sheet to bond with Mohgaonkalan. Here we recognize a new kind the etched surface. When dry, the peels were detached, of flower with intact stamens and pollen from cut with scissors, mounted on glass slides and covered by a glass coverslip affixed with Canada balsam. The a site near Singpur, northeast of Nagpur in peels were photographed on a Nikon Labophot micro- central India. scope and imported to Avizo 9.0 Lite (FEI Visualiza- The late Maastrichtian Singpur locality is tion Science Group, Bordeaux, France) for alignment well known for its flora preserved in chert, and 3-D reconstruction. Succesive peel images and a slice movie of the holotype are accessible at Morpho- including pollen and spores (Samant et al., source.org (key word = Singpuria). 2008) and fruits and seeds, such as Baccato- Pollen was studied and photographed by trans- carpon Bhowal et Sheikh (2004; Manchester mitted light microscopy of the peels with a Nikon et al., 2019), Drupaceocarpon Nandeshwar et Eclipse compound microscope. In addition, one of the Narkhede (2016), Pantocarpon Kapgate, Patil, peels was cleaned in hydrofluoric acid, mounted on an aluminum stub (etched surface upward) and ana- Ilamkar et Ramteke (Kapgate et al., 2007; lyzed with a Hitachi SU5000 field-emission scanning Manchester et al., 2020; syn. Verbenaceocarpon electron microscope (SEM) at the Interdisciplinary Dhabarde, Sheikh, et Kolhe), Phyllanthocar- Center for Biotechnology Research at the University pon Mistri, Kapgate et Sheikh ex Kapgate of Florida. et Manchester (Kapgate et al., 2017), Rama- The floral diagram was generated with automatic rendering from http://kvetnidiagram.8u.cz/index_ konospermus Matin et Juneja (Sheikh and en.php (Kebert, 2014–2020). In attempting to place Bhowal, 2003), Singhpurospermum Nandesh- the flower systematically, we entered morphological war et Narkhede (2017) and Viracarpon Sahni characters of the fossil into the web-based key, The (Matsunaga et al., 2018). In addition, we have Families of Angiosperms (Watson and Dallwitz, 1992 recognized specimens of Enigmocarpon Sahni onwards), to obtain lists of extant families conform- ing to those characters, and considered each of those and Graminocarpon Chitaley et Sheikh in results in light of additional characters not available collections from this site, supplementary to in the key program. Additional floral features were the original occurrence reports of these taxa investigated with reference to those described in the from Mohgaonkalan (Sahni, 1943; Chitaley The Families and Genera of Flowering Plants (e.g. and Sheikh, 1971), a paleobotanically rich site Bayer, 2003; Bayer and Kubitzki, 2003; Anderberg, 2004; Fritsch, 2004; Stevens, 2004; Stevens et al., 64 km northeast of Singpur. 2004a,b; Horn, 2007; Daly et al., 2011; Kubitzki et al., The new permineralized flower presented 2011; Pell et al., 2011) and other literature. Pollen was here adds to the diversity of angiosperms compared with that of extant taxa treated in system- known from the Deccan Intertrappean Beds. atic literature as cited, and with reference to the SEM Analysis of successive serial peels facilitated survey of Li et al. (2010). Terminology for pollen fol- lows Halbritter et al. (2018). three-dimensional (3-D) conceptualization of the specimen, and an examination of pollen within the anthers by light and scanning elec- SYSTEMATICS tron microscopy yielded additional characters. Here we document the morphology and con- sider its systematic placement among eudicot Clade. Pentapetalae angiosperms. Family. Incertae Sedis MATERIAL AND METHODS Genus. Singpuria The flower was discovered in a piece of black chert Ramteke, Manchester, et Nagrale gen. nov. collected from the edge of a farming field near Singpur Pls. 1, 2 (alternate spelling Singhpur), ~65 km NNW of Nag- pur in Chhindwara District, Madhya Pradesh, India E t y m o l o g y. The generic name refers to the (21°36.958′N, 78°43.827′E). The Singpur locality is interpreted to be latest Maastrichtian, based on strati- town of Singpur (also known as Singhpur), close graphic correlations of pollen (Samant et al., 2008) and to the location where this fossil was collected. megafossils (Smith et al., 2015). The flower was exposed in a natural fracture of the Type species. Singpuria kapgatei Ramteke, chert and was serially sectioned by the acetate peel Manchester et Nagrale sp. nov. method. Successive cellulose acetate peels were pre- pared by etching first with hydrofluoric acid, washing Plant Fossil Names Registry. PFN001778 in water, squirting acetone across the etched surface, (for new genus). D. Ramteke et al. / Acta Palaeobotanica 60(2): 323–332, 2020 325 Species: Singpuria kapgatei Ramteke, in the outer whorl and the other in the inner Manchester et Nagrale sp. nov. whorl. Thus, while there are 18 anthers in total, each dorsiventral pair shares a common E t y m o l o g y. The specific epithet honors Prof. filament. The anthers are 0.3 mm wide and Dashrath K. Kapgate, whose research and 1.9 mm deep, with four pollen sacs dehiscing mentorship have greatly improved our knowl- by longitudinal slits, attached to a connective edge of the Deccan Intertrappean flora. that is 0.12 to 0.15 mm in diameter (Pl. 1, figs H o l o t y p e. UF19278-62141. 13, 14; Pl. 2 figs 1, 2). Successive serial peels reveal that the anthers are basifixed (Pl. 1, R e p o s i t o r y. Paleobotanical Collection, Flor- figs 4, 5). The pollen grains are prolate, elon- ida Museum of Natural History, University of gate, 32–35 µm long, 19–23 µm in equatorial Florida, Gainesville, Florida, USA diameter, tapered at the poles, and tricolpo- rate (Pl. 2, figs 2–5). They are psilate by LM T y p e l o c a l i t y. Singpur, Madhya Pradesh, (Pl. 2, fig. 2), with microreticulate ornamen- India; Late Maastrichtian. tation visible at higher magnification by SEM Plant Fossil Names Registry. PFN001779 (Pl. 2, fig. 5). The style is ~0.2 mm in diam- (for new species). eter and pentagonal in cross section (Pl. 1, figs 13, 14), suggesting a pentamerous gynoecium. Generic and specific diagnosis. Flower The apical-most part of the flower is missing, bisexual, actinomorphic, perianth in two so we were unable to observe whether there whorls, petals imbricate, merosity estimated is a single stigma or more.
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