Early Oligocene Itea (Iteaceae) Leaves from East Asia and Their Biogeographic Implications
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Plant Diversity 43 (2021) 142e151 Contents lists available at ScienceDirect Plant Diversity journal homepage: http://www.keaipublishing.com/en/journals/plant-diversity/ http://journal.kib.ac.cn Review article Early Oligocene Itea (Iteaceae) leaves from East Asia and their biogeographic implications ** * Yi-Min Tian a, Jian Huang b, , Tao Su b, c, Shi-Tao Zhang a, a Kunming University of Science and Technology, Kunming, Yunnan, 650500, China b Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, Yunnan, 666303, China c University of the Chinese Academy of Sciences, Beijing, 100049, China article info abstract Article history: Compressed materials of fossil foliage described here as Itea polyneura sp. nov. (Iteaceae) were collected Received 10 May 2020 from the Oligocene of Wenshan, Yunnan Province, southwestern China. The identification is based on the Received in revised form following characters: eucamptodromous secondary veins, strict scalariform tertiary veins, irregular tooth 8 September 2020 with setaceous apex. The leaf morphology of all modern and fossil species was compared with the new Accepted 10 September 2020 species from Wenshan and show that I. polyneura is most similar to the extant East Asian species Itea Available online 3 October 2020 omeiensis, which inhabits subtropical forests of southern China. This discovery represents the first un- ambiguous leaf fossil record of Itea in East Asia. Together with other species in the Wenshan flora and Keywords: fl fi East Asia evidence from several other ora in southern China, these ndings demonstrate that Itea from East Asia Itea arose with the Paleogene modernization. Iteaceae Copyright © 2020 Kunming Institute of Botany, Chinese Academy of Sciences. Publishing services by Leaf fossil Elsevier B.V. on behalf of KeAi Communications Co., Ltd. This is an open access article under the CC BY- Oligocene NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). Southwestern China 1. Introduction southern China (Quan et al., 2016). They show a high degree of similarity to the extant East Asian evergreen broad-leaved forests, In the early Cenozoic, the Northern Hemisphere may have been which offers an excellent opportunity to understand the origin and covered by vast subtropical broad-leaved evergreen forests (Wolfe, evolution of modern flora in East Asia. 1985; Milne and Abbott, 2002). A large number of fossil species Itea L. are small trees, shrubs or sometimes scramble plants that (e.g., Fagaceae Dumort., Lauraceae Juss., Theaceae Mirb., Magno- grow in forests and thickets (Jin and Ohba, 2001; Kubitzki, 2007). liaceae Juss.) found in North America and Europe, ranging from the Itea belongs to Iteaceae J. Agardh, Saxifragales Bercht. & J. Presl, Paleocene to Miocene, are almost morphologically identical to although it was formerly included in Escalloniaceae R. Br. ex extant taxa in southern China (Mai, 1995; Manchester, 1999). Dumort., Grossulariaceae DC. or Saxifragaceae. Molecular analyses Because few Paleogene fossil localities have been reported in East place it as an independent family into the Core Saxifragales sister to Asia, the biogeographical histories of these taxa have mostly been Pterostemon S. Schauer (Savolainen et al., 2000; Fishbein et al., interpreted on the basis of materials from North America and 2001). There are 21 species in the genus, and most of them are Europe. Researchers have speculated that East Asia is a ‘refugium’ evergreen and distributed in the subtropical to tropical regions of rather than the source of the Northern Hemisphere's biodiversity East Asia (Guo and Ricklefs, 2000; Kubitzki, 2007). Thus, the sub- (Axelrod et al., 1998; Tiffney and Manchester, 2001). In recent years, tropical evergreen broad-leaved forest region in southern China is many fossil floras from the Paleogene have been discovered in Itea's modern biodiversity center (Guo and Ricklefs, 2000). The distribution of Itea displays an East Asian-eastern North American- southeastern African disjunction (Fig. 1), which is unusual among * Corresponding author. Faculty of Land Resource Engineering, Kunming Uni- genera with a North American-East Asian disjunction pattern (Wen, versity of Science and Technology, Kunming, Yunnan 650093, China. Fax: þ86 0871 1999). Although the genus is primarily subtropical, two species of 65177163. deciduous trees inhabit the temperate forests of Japan and eastern ** Corresponding author. Fax: þ86 0691 8713226. E-mail addresses: [email protected] (J. Huang), [email protected] North America (Kubitzki, 2007). Thus, the distribution pattern of (S.-T. Zhang). Peer review under responsibility of Editorial Office of Plant Diversity. https://doi.org/10.1016/j.pld.2020.09.006 2468-2659/Copyright © 2020 Kunming Institute of Botany, Chinese Academy of Sciences. Publishing services by Elsevier B.V. on behalf of KeAi Communications Co., Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). Y.-M. Tian, J. Huang, T. Su et al. Plant Diversity 43 (2021) 142e151 Fig. 1. Map showing the modern distribution of Itea and the locations from which Itea fossils were recovered. Itea provides an opportunity to study the origin and differentiation of Northern Hemisphere temperate forests (Folk et al., 2019). Itea pollen grain can easily be distinguished by its unique bilaterally symmetrical and diporate morphology, and fossil re- cords of iteoide pollen grains are common in the Cenozoic of Europe and North America (Hermsen, 2013). In addition, its distinctive infructescence structure is highly recognizable for its identification, and fossil fruits of Itea europaea are also widely re- ported in Europe from the Eocene to Pliocene (Mai, 2001; Hermsen, 2013). However, confirmed leaf fossils of Itea are rare and only known from western North America (Wolfe and Wehr, 1987; Hermsen, Personal communication). Nevertheless, leaves of Itea species can be easily identified because of their unique character- istics (e.g., eucamptodromous secondary veins, strict scalariform tertiary veins, irregular teeth). In this paper, we identified fossil leaves of Itea from the early Oligocene of Wenshan Basin, Yunnan, southwestern China, which constitutes the first reliable leaf fossil record of the genus from East Asia and Eurasia. This study establishes a new species of Itea to accommodate these new fossils with distinct and diagnostic leaf architecture, and compares it with all extant and fossil species of the genus. Furthermore, this study discusses the implications this fi ‘ nding has for our understanding of the histories of the Tertiary Fig. 2. Location of the fossil site (leaf symbol). relic flora’ (Milne and Abbott, 2002) and East Asian-North American disjunct distributions. situ volcanic ash reveals a much earlier geological time, the early 2. Materials and methods Oligocene (~30/32 Ma), not the Miocene as previously suggested (Tian et al., unpublished). 2.1. Fossil materials 2.2. Extant materials and morphological observation Compressed leaf fossils studied here were collected from the Wenshan Basin, Yunnan Province, southwestern China (23200N; Herbarium specimens of extant Itea species used for compar- 104170E, 1271 m a.s.l.) (Fig. 2). Wenshan is a fault basin in a group ison were accessed from online data sources such as JSTOR Global of the Wenshan-Funing fold that belongs to the southeast Yunnan Plants (http://plants.jstor.org/) and the Chinese Virtual Herbarium fold belt of the China Caledonian Fold System; the Wenshan Basin is (http://www.cvh.org.cn/). Cleared leaf materials of extant species divided into the Yangtze platform by geotectonic characteristics. were collected from the Xishuangbanna Tropical Botanical Garden Leaf fossils were found in a grey, light grey, and greyish yellow (XTBG). These leaves were cleared following the protocol devel- mudstone and marlstone. The environment of deposition is oped by Li (1987). Several photographs of cleared leaves of extant interpreted to be lacustrine facies. The fossil-bearing stratum Itea species were downloaded from the Smithsonian Cleared Leaf belongs to the Huazhige Formation. This outcrop was previously Collection, and their contrasts were adjusted using Adobe Pho- assigned to different periods of the Miocene by biostratigraphic toshop. All fossil and modern specimens were photographed us- correlation and magnetostratigraphic study (Lebreton-Anberree ing a digital camera (Nikon D700). Further, the details of the et al., 2016). However, a recent UePb dating of zircon from in fossils were photographed using a stereomicroscope (Leica 143 Y.-M. Tian, J. Huang, T. Su et al. Plant Diversity 43 (2021) 142e151 S8APO). In addition, line-drawings and plates were created using Repository: Paleoecology Collections, Xishuangbanna Tropical CorelDRAW X7. Digital photographs of extant and fossil specimens Botanical Garden, Chinese Academy of Sciences. were measured using ImageJ 1.45s software. The leaf architecture Type locality: Dashidong Village, Wenshan, Yunnan, China. terminology followed Ellis et al. (2009). Stratigraphy: Oligocene, Huazhige Formation. Diagnosis: Leaf oblong, size ~4.5e~8.0 cm long  2.5e3.4 cm wide; secondary veins eucamptodromous, ~7e8 pairs; inter- secondaries absent; tertiary veins densely scalariform; margin 3. Results with crenate to serrate teeth, spacing irregular, tooth apex simple or setaceous (Fig. 3). 3.1. Systematic palaeobotany Description: Isolated leaves, simple, petiolate.