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A New Species of Liquidambar (Altingiaceae) from the Late Eocene of South China

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A New Species of Liquidambar (Altingiaceae) from the Late Eocene of South China Journal of Plant Research https://doi.org/10.1007/s10265-019-01091-0 REGULAR PAPER A new species of Liquidambar (Altingiaceae) from the late Eocene of South China Natalia P. Maslova1,2 · Tatiana M. Kodrul1,3 · Alexei B. Herman3 · Ming Tu1 · Xiaoyan Liu1 · Jianhua Jin1 Received: 27 June 2018 / Accepted: 27 January 2019 © The Botanical Society of Japan and Springer Japan KK, part of Springer Nature 2019 Abstract A new fossil leaf species, Liquidambar bella (Altingiaceae), is described from the lower part of the Eocene Huangniuling Formation, Maoming Basin, South China. Suprabasal venation in the fossil lobed Liquidambar leaves is reported for the first time. The new species provides additional palaeobotanical evidence on the morphological variability of this genus sup- porting the idea of combining the genera Liquidambar, Semiliquidambar and Altingia into the single genus Liquidambar as proposed based on molecular markers. Keywords Altingiaceae · Eocene · Huangniuling Formation · Liquidambar · Maoming Basin · South China Introduction type (Melchior 1998; Wheeler et al. 2010) and Liquidam- bar pollen grains (Graham 1965; Kuprianova 1960; Muller Extant species of the genus Liquidambar L. (Altingiaceae) 1981) are dated as Paleocene. Liquidambar-type leaves are possess a disjunctive pan-continental natural distribution in well represented in the Cenozoic of Asia (e.g., Dong et al. eastern and western Asia, Europe and North and Central 2018; Endo and Morita 1932; Huzioka 1972; Huzioka and America (e.g., Wen 1998, 1999). Study of related fossil taxa, Uemura 1979; Maslova 1995, 2003; Maslova et al. 2015; which can provide evidence about the diversification and Onoe 1974; Ozaki 1991; Suzuki 1961; Uemura 1983), evolutionary history of the genus, are therefore of particular Europe (e.g., Ferguson 1971, 1989; Koch et al. 1973; Mar- interest. Fossils of Altingiaceae have been reported from the tinetto 1998; Worobiec et al. 2012), and North America Cretaceous, Paleogene and Neogene of Asia, Europe and (e.g., Brown 1933; Knowlton 1902; MacGinitie 1941; Smi- North America. The earliest fossil record of reproductive ley et al. 1975; Stults and Axsmith 2011; Wolfe and Tanai structures similar to those typical of this family is known 1980). from the upper Turonian (Zhou et al. 2001). Permineralized In China, several fossil occurrences of Liquidambar have female reproductive structures unequivocally assigned to been recognized based on leaves and/or reproductive struc- Liquidambar were described from the Miocene (Pigg et al. tures: Eocene L. maomingensis N. Maslova, Kodrul, Song et 2004). The first appearance of the Liquidambar foliar mor- Jin from the Huangniuling Formation, Guangdong Province phological type (Maslova 1995), wood of the Altingiaceae (Maslova et al. 2015) and two Miocene species L. miosinica Hu et Chaney from the Shanwang Formation in Shandong Province (Hu and Chaney 1940) and the Shengxian Forma- * Xiaoyan Liu [email protected] tion in Zhejiang Province (Xiao et al. 2011, 2013, 2015) as well as L. fujianensis J.L. Dong et B.N. Sun from the Fotan * Jianhua Jin [email protected] Group in Fujian Province (Dong et al. 2018). Apart from these, several Eocene fossil leaves have been ascribed to 1 State Key Laboratory of Biocontrol, Guangdong Key genus level only (Dalianhe Formation, Heilongjiang Prov- Laboratory of Plant Resources and School of Life Sciences, ince, He and Tao 1997; Huadian flora, Jilin Province; Man- Sun Yat-sen University, Guangzhou 510275, China chester et al. 2005). 2 Borissiak Paleontological Institute, Russian Academy Taxonomic revision of the modern Liquidambar (Ickert- of Sciences, Moscow 117647, Russia Bond and Wen 2013), adopting a broad circumscription of 3 Geological Institute, Russian Academy of Sciences, the genus and subsuming the previously recognized genera Moscow 119017, Russia Vol.:(0123456789)1 3 Journal of Plant Research Altingia Noronha and Semiliquidambar H.-T. Chang, shows New finds of fossil Liquidambar species in China are there- that the genus includes 15 species, 11 of which grow in fore of an exceptional importance for a reconstructing the China. They are: L. acalycina H. T. Chang (Anhui, Guang- phylogeny of the genus, particularly in view of recent taxo- dong, Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, nomic changes (Ickert-Bond and Wen 2013). Sichuan and Zhejiang provinces), L. caudata (H. T. Chang) Ickert-Bond et J. Wen (Fujian and Zhejiang provinces), L. chinensis Champ. (Guangdong, Guangxi, Guizhou, Hainan Materials and methods provinces and Hong Kong), L. chingii (Metcalf) Ickert- Bond et J. Wen (Fujian, Guangdong, Guangxi, Guizhou The plant fossils examined were collected in the Jintang and Jiangxi provinces), L. excelsa (Noronha) Oken (Yun- opencast mine (21°42′50″N, 110°52′35″E) of the Maoming nan Province), L. formosana Hance (Anhui, Chongqing, Basin located northwest of Maoming City in southwestern Fujian, Gansu, Guangdong, Guangxi, Guizhou, Hainan, Guangdong Province, China (Fig. 1). The stratigraphic suc- Hunan, Hubei, Jiangsu, Jiangxi, Shanxi, Sichuan, Zhejiang cession of this locality is composed of the Youganwo and provinces, Taiwan and Hong Kong), L. gracilipes (Hemsl.) Huangniuling formations. The Youganwo Formation con- Ickert-Bond et J. Wen (Fujian, Guangdong, Zhejiang prov- sists of terrigenous coal-bearing deposits in its lower part inces and Hong Kong), L. multinervis (Cheng) Ickert-Bond and oil shales in the upper part. The overlying Huangniul- et J. Wen (Guizhou Province), L. obovata (Merrill et Chun) ing Formation is composed mainly of fluvial sandstones, Ickert-Bond et J. Wen (Hainan Province), L. siamensis siltstones, and conglomerates with extended lenses of mud- (Craib) Ickert-Bond et J. Wen (Guangdong and Yunnan stones. Details of the Jintang opencast mine section are provinces), L. yunnanensis (Rehder et Wilson) Ickert-Bond given in Aleksandrova et al. (2015). The upper part of the et J. Wen (Yunnan Province) (Ickert-Bond and Wen 2013). Youganwo Formation has been dated as late Eocene on the Hence, China should be the place where the maximal num- basis of the vertebrate fossils (Averianov et al. 2017; Jin ber of naturally growing representatives of the Liquidambar 2008). In a recent palynological study the Youganwo and occur today. Huangniuling formations are considered to be Lutetian-Bar- Here we introduce new species of Liquidambar from the tonian and Priabonian in age, respectively (Aleksandrova Eocene Huangniuling Formation (Maoming Basin, Guang- et al. 2015). Therefore, the new species of Liquidambar is dong Province) with brief discussion on its palaeoecology supposed to be of a late Eocene age. and habitat conditions. These fossils are from stratigraphi- About 145 fossil leaf impressions of the new species cally lower beds than the previously described species, L. were recovered from a lens of mudstone in the lower part maomingensis which also occurs in the Huangniuling For- of the Huangniuling Formation (locality MMJ2-2) which mation. We demonstrate similarities in leaf form of the new is approximately 20 m below the occurrence of previ- species to the species of the former genus Semiliquidambar. ously reported Liquidambar maomingensis from the Upper Fig. 1 Outline map showing the fossil locality 1 3 Journal of Plant Research Huangniuling flora (Maslova et al. 2015). Plant megafossil suprabasal or basal actinodromous, with three primary assemblage of the Lower Huangniuling flora contains a few veins; lateral primary veins straight or curved, extending taxa of conifers (Pinaceae, Podocarpaceae) but is dominated outward or upward, often diverging from midvein subop- by angiosperms of the Lauraceae, Fagaceae, Hamameli- positely at various angles. Secondaries up to 6–10 pairs, daceae, Altingiaceae, Myrtaceae, Juglandaceae, Aceraceae, alternate to subopposite, often irregular, semicraspedodro- Dipterocarpaceae, Rhamnaceae, Fabaceae, and Celastraceae mous or festooned semicraspedodromous, arching upward families. Fagaceae, along with Altingiaceae, are the most near the margin; branches from loops terminating in teeth; common components of this flora (Spicer et al. 2016, 2017). intersecondaries weak. Tertiary veins alternate percurrent The specimens are housed at the Museum of Biology to mixed opposite–alternate percurrent. Fourth order veins of Sun Yat-sen University, Guangzhou, China. Images of alternate percurrent. the leaves were taken using digital cameras: an Olympus E-500 and a Panasonic GX7 with a Leica DG MacroElmarit Description 1:2.8/45 mm macro lens. For the leaf descriptions, we used terms from the Manual of Leaf Architecture (Ellis et al. Leaves are simple, palmately three-lobed, with petioles. The 2009). The length to width (L/W) ratio of the lamina was longest preserved fragment of petiole is 20 mm. Leaves are calculated using the maximum lamina length and width. The symmetric or asymmetric with different depths of sinuses L/W ratio for incompletely preserved leaves was calculated between the lobes (Figs. 2c, 4e, f) and medial asymmetry using the reconstructed lamina. Biotic damage types on the (Figs. 3a, 4e, f). Laminar bases are also symmetrical or Liquidambar leaves were considered using the “Guide to asymmetrical in insertion and/or width (Figs. 3a, d, 4g). Insect (and Other) Damage Types on Compressed Fossil Insertion points of lamina base on either side of the petiole Plants” (Labandeira et al. 2007). are occasionally separated by 1–2 mm (Figs. 3a, b, 4g). The Comparative material of extant Liquidambar caudata lamina length varies from 37 to 80 mm, lamina width—from was obtained from the South China Botanical Garden, 19 to 140 mm. The length of the central lobe varies from Guangzhou. 15 to 50 mm, width—from 10 to 28 mm. The shape of the central lobe is triangular (Figs. 2h, 4a, b, d, h) or frequently narrowly triangular (Figs. 2a–e, 3a, 4c), with nearly paral- Systematics lel margins at the lobe base (Figs. 2a, 3a, 4c) or narrowing towards the base (Figs. 2c, d, 3f). Lobe apices are acute Family: Altingiaceae Horan. 1841 (Figs. 3g, 4d, h) and frequently strongly acuminate (Figs. 2a, Genus: Liquidambar L. 1753 c, d, 4a, b). In some leaves lateral lobes extend obliquely Species: Liquidambar bella N. Maslova et Kodrul, sp.
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