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Nelumbo Jiayinensis Sp. Nov. from the Upper Cretaceous Yong'ancun Formation in Jiayin, Heilongjiang, Northeast China

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Nelumbo Jiayinensis Sp. Nov. from the Upper Cretaceous Yong'ancun Formation in Jiayin, Heilongjiang, Northeast China Cretaceous Research 84 (2018) 134e140 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes Short communication Nelumbo jiayinensis sp. nov. from the Upper Cretaceous Yong'ancun Formation in Jiayin, Heilongjiang, Northeast China * Fei Liang a, b, Ge Sun a, b, c, , Tao Yang a, b, Shuchong Bai a, b a College of Paleontology, Shenyang Normal University, Shenyang 110034, China b Key Laboratory of Evolution of Past Life in NE Asia, Ministry of Land ＆ Resources, Shenyang 110034, China c Key Laboratory for Evolution of Past Life and Environment in NE Asia, MOEC, Jilin University, Changchun 130026, China article info abstract Article history: Fossil leaves of Nelumbo jiayinensis sp. nov. are described from the Yong'ancun Formation (Santonian) in Received 6 July 2017 Jiayin, Heilongjiang, Northeast China. This is the first report on fossil Nelumbo from the Upper Cretaceous Received in revised form of China. The leaves are simple with entire margins, orbicular or sub-orbicular in shape, with a peltate 18 October 2017 and symmetric lamina. The venation is actinodromous with 20e25 primary veins, the secondary and Accepted in revised form 7 November 2017 tertiary veins are poorly developed and interweave into meshes. The areoles are irregularly polygonal in Available online 8 November 2017 shape and well developed both in upper and lower surfaces. Combined with data derived from associ- ated fossil mega-plants, spores and pollen, bivalves and dinosaur footprints, the aquatic angiosperm Keywords: fl Nelumbo Nelumbo existed in a uviolacustrine environment under a warm but seasonal temperate climate. © New species 2017 Elsevier Ltd. All rights reserved. Upper Cretaceous Santonian Jiayin 1. Introduction Many fossil leaves, fruits and pollen of Nelumbo have been described from Early Cretaceous and younger deposits (Kuprianova The earliest known flowering plants from the fossil records seem and Tarasevich, 1983; Erickson, 1991; Tao, 2000; Johnson, 2002; to be aquatic and herbaceous in nature (Sun et al., 1998, 2002; Gandolfo and Cuneo, 2005; He et al., 2010; Gabrielyan et al., Dilcher et al., 2007), which prompted us to pay more attention to 2012; Li et al., 2014). Additionally, fossil Nelumbo leaves from the the significance of aquatic plants Nelumbonaceae in regard to the Asuwa flora (Upper Cretaceous) have been described from Japan, early evolutionary trends of angiosperms. Nelumbo is the sole extant which is so far the earliest record in eastern Asia (Matsuo, 1954). genus of the family Nelumbonaceae, and consists of two extant However, in China, most Nelumbo fossils are reported from species Nelumbo nucifera Gaertner and N. lutea Willdenow Eocene deposits including the Changchang Formation of Chang- (Nicolson, 1991; Ni et al., 1995). According to the previous records chang Basin, Hainan Island (Guo, 1979; He et al., 2010; Li et al., the earliest fossil Nelumbo emerged in the Albian (Early Cretaceous), 2014) and the Dalianhe Formation in Yilan of Heilongjiang Prov- yet the systematic position, the evolution and phytogeography of ince (Tao, 2000). this genus have long been intensively discussed by both botanists In this paper, we describe a new species, Nelumbo jiayinensis sp. and paleobotanists (Crabtree, 1987; Upchurch et al., 1994; Parkinson nov., preserved as leaf remains collected from the Upper Creta- et al., 1999; Xue et al., 2012). However, as an aquatic angiosperm, ceous Yong'ancun Formation in the Jiayin area of NE China. The age fossil Nelumbo is a clearly indicator of non-marine aquatic envi- of the Yong'ancun Formation is assigned to the Santonian based on ronments, such as lakes, wetlands and slow flowing rivers, and likely the combined data mainly of megafossils and palynological as- indicates a warm climate, which is significant for understanding semblages (Sun et al., 2007, 2011; Markevich et al., 2011; Liang past environments and climates. et al., 2015a). Ostracods, bivalves, conchostracans, dinosaur foot- prints (Jiayinosauropsis johnsoni) in addition to many aquatic an- giosperms have been found in this formation (Dong et al., 2003; * Corresponding author. College of Paleontology, Shenyang Normal University, Sun et al., 2007, 2011; Markevich et al., 2011; Liang and Sun, Shenyang 110034, China. 2015b), which indicates abundance of fresh water environments. E-mail address: [email protected] (G. Sun). https://doi.org/10.1016/j.cretres.2017.11.007 0195-6671/© 2017 Elsevier Ltd. All rights reserved. F. Liang et al. / Cretaceous Research 84 (2018) 134e140 135 2. Materials and methods Type locality: East hill of Yong'ancun village, Jiayin, Heilongjiang, China. The specimens of Nelumbo jiayinensis sp. nov. described here Geological horizon: Yong'ancun Formation (Santonian) were collected from the upper part of the Yong'ancun Formation Derivation of specific name: From the Jiayin Basin where the fossils (Fig. 1), Jiayin County of Heilongjiang Province. Outcrops of the collected. Yong'ancun Formation (Fig. 2) are mainly distributed along the Diagnosis. Simple leaves, orbicular or sub-orbicular with entire Heilongjiang (Amur) River near the Yong'ancun village. The Yon- margin, primary veins strong, almost straight, 20e25 in number, g'ancun Formation is mainly composed of alternating yellow- radiating from the center at acute angles, secondary veins forming brown cross-bedded sandstones, greyish-brown siltstones and meshes in the intercostal area, finer veins usually forming small mudstones, intercalated with pebbly sandstone, representing a quadrangular or pentagonal areoles. Epidermal cells of leaves basin dominated by alluvial-lacustrine (Bureau of Geology and usually polygonal and regular, while the petiole cells show an Mineral Resources of Heilongjiang Province, 1993; Sun et al., 2011, elongate and compact arrangement. 2016). The age of the Yong'ancun Formation is Santonian mainly Description. Leaves are simple, nearly orbicular or sub-orbicular in based on palynological studies (Sun et al., 2007, 2011; Markevich shape, with peltate and symmetric lamina (Fig. 3, 4, 5). Even though et al., 2011; Liang et al., 2015a). The terminology we use for the preserved fossil lamina fragments are 9e12 cm long and almost describing leaf morphology follows Ellis et al. (2009). 11 cm wide in size, the material suggests the whole orbicular leaves Six specimens of Nelumbo in total were collected from the fossil could approach 15e24 cm in diameter. The lamina surface is site (Fig. 2), which is located in the upper part of the Yong'ancun membranous in texture and the margin is entire or undulates Formation. The fossil leaves of Nelumbo are preserved as impres- slightly, but is incompletely preserved (Fig. 3A, B, C, D, E; Fig. 4; sions and compressions, and were photographed using a Nikon Fig. 5). The petiole is about 6e10 mm in diameter, situated in the digital camera (Nikon D600). Details of the leaf structures were center of the lamina (Figs. 3A, 4A, B), and the epidermal cells of the observed and photographed using 3D super depth microscopy petiole are highly elongate (Fig. 4C). In the preserved specimens, systems (Keyence VHX-600). All specimens described here are the leaf venation shows actinodromous branching, which is com- housed in the Paleontological Museum of Liaoning (PMOL) in mon in living and fossil species of Nelumbo. Primary veins or ribs Shenyang, China. are 20e25 in number, almost straight or bend slightly and radiate from the leaf center at an angle of 5e15 on the upper surface (Fig. 3A, B) while on the lower surface the primary veins are clearer 3. Systematic paleontology and more prominent (Fig. 4A; Fig. 5A). In the intercostal zones of primary veins the secondary vein branches from the primary veins Family: Nelumbonaceae with an angle of 40e60 and interweave into meshes (Fig. 3B, C, D; Genus: Nelumbo Adanson, 1763 Fig. 5B), in these meshes, tertiary veins become feeble and form Species: Nelumbo jiayinensis Liang, Sun et Yang, sp. nov. into irregular and polygonal areoles, usually 0.5e1.0 mm in diam- eter and 3- to 5-sided (Figs. 3E and 4D, E; Fig. 5C, D, E, F, G). Holotype: specimens YX-B-300, Yong'ancun village of Jiayin County, NE China, the Yong'ancun Formation, Upper Cretaceous d Epidermal cells of the upper surface are relatively regular in size Fig. 3 and shape, which are generally polygonal and usually 20e50 mmin Paratype: specimens YX-B-301d Fig. 4 diameter; the anticlinal cell wall is generally regular and almost Material: samples YX-B-300, YX-B-301, YX-B-302, YX-B-304, YX-B- straight (Fig. 3D, E; Fig. 5G). On the lower surface, the epidermal 307a, YX-B-307b cells are more or less same as those of the upper surface in size and shape, usually regular and polygonal, the average size of cells is 40 mm long by 30 mm wide (Figs. 3E and 4D, E; Fig. 5C, D, E, G). Stomata were not observed. 4. Discussion 4.1. Comparisons The features of the monotypic family Nelumbonaceae include a simple, orbicular lamina with an entire margin, actinodromous veins, a central petiole and the hexagonal areoles (Hickey and Wolfe, 1975; Upchurch et al., 1994). It is clear that the new spe- cies shares many such characters and so should be assigned to the family Nelumbonaceae. Obviously, the new species differs from the extant species Nelumbo nucifera and N. lutea in the number of the primary veins and the lamina size, and Hollick (1894) noticed that it was possible that Nelumbo leaf size could progressively increase over geological time. Up to now, many of fossil species of Nelumbo have been re- ported, and most of fossil specimens represent only leaf portions. Nevertheless many of the descriptions and illustrations could provide valuable information allowing comparisons with the present material (Matsuo, 1954; Gandolfo and Cuneo, 2005; He et al., 2010; Li et al., 2014). The radial vein number and the leaf margin of the present specimens are similar to those of Nelumbo Fig. 1. Location of Jiayin, China (the arrow shows the fossil site).
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