Table of Contents

01 March 2019, Volume 57 Issue 2

Cover illustration: Hypothesized fruit morphological evolution and biogeographic history of Cedrelospermum. Image from Lin-Bo Jia et al. See Jia et al., pp. 94–104 in this issue.

  

Editoral
• Ecological and Biogeographic Implications of Asian Cenozoic Fossil Floras

  Xiaoyan Liu and Steven R. Manchester

  J Syst Evol. 2019, 57(2): 91-93.
  https://doi.org/10.1111/jse.12488

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Research Articles
• First fossil record of Cedrelospermum (Ulmaceae) from the Qinghai–Tibetan Plateau: Implications for morphological evolution and biogeography

  Lin-Bo Jia, Tao Su, Yong-Jiang Huang, Fei-Xiang Wu, Tao Deng, and Zhe-Kun Zhou

  J Syst Evol. 2019, 57(2): 94-104.
  https://doi.org/10.1111/jse.12435

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  Cedrelospermum Saporta is an extinct genus in the Ulmaceae with abundant fossil records in North America and Europe. However, so far, fossil records of this genus from Asia are sparse, which limits the interpretations of the morphological evolution and biogeographical history of the genus. Here we report well‐preserved fruits (Cedrelospermum tibeticum sp. nov.) and a leaf (Cedrelospermum sp.) of Cedrelospermum from the upper Oligocene Lunpola and Nyima basins in the Qinghai–Tibetan Plateau (QTP). This is the first fossil record of Cedrelospermum in the QTP, showing that this genus grew in this region during the late Oligocene. Cedrelospermum tibeticum fruits are double‐winged, morphologically similar to the Eocene and Oligocene double‐winged Cedrelospermum species from North America. This supports the hypothesis that Cedrelospermum migrated to Asia from North America by way of the Bering Land Bridge. Given that Cedrelospermum was a typical element of Northern Hemispheric flora in the Paleogene and Neogene, the presence of this genus indicates that the central region of the QTP was phytogeographically linked with other parts of the Northern Hemisphere during the late Oligocene. The morphological observations of C. tibeticum fruits and other double‐winged Cedrelospermum fruits suggest an evolutionary trend from obtuse to acute apex for the primary wing. Cedrelospermum tibeticum likely had warm and wet climatic requirements. This type of an environment possibly existed in the central QTP in the late Oligocene, thereby supporting the survival of C. tibeticum.
• Leaf and infructescence fossils of Alnus (Betulaceae) from the late Eocene of the southeastern Qinghai–Tibetan Plateau

  He Xu, Tao Su, and Zhe-Kun Zhou

  J Syst Evol. 2019, 57(2): 105-113.
  https://doi.org/10.1111/jse.12463

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  Plant fossils from the Qinghai–Tibetan Plateau (QTP), China are critical to understand not only the diversification history of plants there, but also the paleoenvironmental conditions. Alnus are deciduous trees, mainly distributed in temperate and subtropical regions of Eurasia and North America, and they are well known in the fossil records throughout the Cenozoic in the Northern Hemisphere. We collected numerous well‐preserved Alnus leaf and infructescence fossils from the Lawula Formation (～34.6 Ma with ⁴⁰Ar/³⁹Ar dating) at the present elevation of 3910 m a.s.l. in the southeastern QTP. Based on detailed morphological comparisons with existing and fossil species, these fossils show closest affinity to Alnus ferdinandi‐coburgii C. K. Schneid., and we refer to these fossils as A. cf. ferdinandi‐coburgii. These specimens comprise the oldest megafossil record of Alnus in the QTP, and provide solid evidence for the distribution of Alnus there as early as the late Eocene. Extant A. ferdinandi‐coburgii is distributed in areas with mean annual temperature values between 9.7 °C and 16.9 °C, and mean annual precipitation values ranging from 896.2 mm to 1161.2 mm; therefore, fossils of A. cf. ferdinandi‐coburgii suggest a much warmer and wetter climate during the late Eocene than today in the southeastern QTP. This finding is consistent with other evidence for continued uplift of the southeastern QTP after the late Eocene that might be due to the eastward extension of the QTP.
• Oligocene–Neogene fossil history of Asian endemic conifer genera in Japan and Korea

  Atsushi Yabe, Eunkyoung Jeong, Kyungsik Kim, and Kazuhiko Uemura

  J Syst Evol. 2019, 57(2): 114-128.
  https://doi.org/10.1111/jse.12445

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  Temporal and spatial changes of ten conifer genera that are endemic to East Asia were analyzed based on fossil data from humid temperate forests in the Japanese Islands and Korean Peninsula to elucidate the phytogeographic history, and to understand differences between those genera eliminated from the Japanese Islands and those that remained extant. All these genera, except for Thujopsis, have existed in the area since the Paleogene and remained in the Japanese islands after initial separation from the continent at the early–middle Miocene. Fossil representatives of locally extinct six genera have tendencies to adapt to wider ranges of climatic conditions than their modern relatives. Metasequoia, Glyptostrobus, and Taiwania began to change their distributions since the late Miocene possibly through habitat partitioning. Keteleeria, Pseudolarix, and Cunninghamia appeared to have expanded their habitat toward warmer conditions during the mid‐Miocene Climatic Optimum and then became restricted to warmer forest vegetation by the end of Pliocene. Overall changes in their distribution can be explained by climatic effects. On the contrary, three genera endemic to Japan (Sciadopitys, Cryptomeria, and Thujopsis) followed clearly different trends from the others. Cryptomeria and Thujopsis were especially adapted to cooler‐temperate climate and they retained their habitat areas in the northern part of Japan. During the late Miocene–Pliocene, the islands connected with the Eurasian continent again, which probably acted as a corridor for warm‐adapted genera to disperse southwest. Current data suggest that ecological requirements of each genus might be essential to determine whether they could survive on the Japanese Islands.
• Dynamics and evolution of Turgay‐type vegetation in Western Siberia throughout the early Oligocene to earliest Miocene—a study based on diversity of plant functional types in the carpological record

  Svetlana Popova, Torsten Utescher, Dmitry Gromyko, Volker Mosbrugger, Louis François

  J Syst Evol. 2019, 57(2): 129-141.
  https://doi.org/10.1111/jse.12420

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  Based on ecospectra of 66 published carpofloras we study dynamics and evolution of Turgay vegetation in Western Siberia during the early Oligocene to earliest Miocene. The ecospectra are obtained using a Plant Functional Type (PFT) classification system comprising 26 herbaceous to arboreal PFTs. The carpofloras originate from seven floristic levels covering the time‐span from the Rupelian to early Aquitanian. Key elements of these levels are documented based on original collection materials. Although impacted by local edaphic conditions, the ecospectra can be interpreted in terms of changing vegetation. Our data show that warm temperate mesophytic, mixed conifer‐broad‐leaved deciduous forest assemblages persisted throughout the Oligocene and earliest Miocene in this core area of Turgai type vegetation. This is in line with comparatively stable climate conditions persisting in the studied time‐span, showing a minor temperature decline and coeval moderate increase in seasonality and precipitation. Concurrently, the reconstructed ecospectra contradict significant continental drying throughout the Oligocene and earliest Miocene. Spatial variability of the proportions of PFTs within the single floristic horizons primarily reflects local edaphic conditions. High diversities of PFTs characteristic for swamp vegetation are mainly confined to the early Oligocene and have a regional focus. Our results indicate that taxonomical diversity, particularly concerning mesic herbs and deciduous shrubs and trees, increased towards the end of the Oligocene. This increase in biodiversity probably can be attributed to an increase in rainfall and extension of terrestrial habitats after the final retreat of the Paratethys.
• Early Oligocene Calocedrus (Cupressaceae) from the Maoming Basin, South China, and its paleogeographic and paleoclimatic implications

  Yan Wu, Jian-Hua Jin, Nan Li, Hui-Min He, Ting Chen, and Xiao-Yan Liu

  J Syst Evol. 2019, 57(2): 142-152.
  https://doi.org/10.1111/jse.12424

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  The genus Calocedrus Kurz is characterized by flattened branches covered with decussate and dimorphic leaves. In this study, we describe fossil foliage of Calocedrus discovered in the Shangcun Formation (early Oligocene) of the Maoming Basin, South China. The fossils have wedge‐shaped branchlet segments and scale‐like strongly dimorphic leaves, similar to the fossil species Calocedrus lantenoisi (Laurent) Tao. There have been no detailed studies of the morphological features and cuticle structures of C. lantenoisi. Therefore, a newly emended diagnosis of the species C. lantenoisi is given based on a detailed study of leaf morphology and cuticular characters exhibited by the Maoming fossils. This is one of the earliest fossil records of Calocedrus in the world, providing additional evidence for the early biogeographic history of this genus and supporting the inference that eastern Asian Calocedrus is primitive among all the living species. The extant species of Calocedrus are mainly distributed in mountainous regions. On the basis of the “nearest living relative” analysis, we propose that the Maoming Basin was adjacent to a mountainous region during the early Oligocene.
• An early Oligocene fossil acorn, associated leaves and pollen of the ring‐cupped oaks (Quercus subg. Cyclobalanopsis) from Maoming Basin, South China

  Xiao-Yan Liu, Sheng-Lan Xu, Meng Han, and Jian-Hua Jin

  J Syst Evol. 2019, 57(2): 153-168.
  https://doi.org/10.1111/jse.12450

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  The ring‐cupped oaks (Quercus subg. Cyclobalanopsis) characterized by united and concentric ring‐cupped acorns, are mainly distributed in broad‐leaved evergreen forests in tropical and subtropical regions of East and Southeast Asia. Their geological history has been traced from the middle Eocene to Pleistocene. Most reports of the subgenus have been on the basis of leaves, whereas the acorns or cupules, more valuable evidence for the inter‐ and intrageneric classifications, have been relatively poorly documented. Here, we describe a new species, Quercus shangcunensis sp. nov., based on a ring‐cupped fossil acorn and pollen on the acorn, as well as Quercus sp. associated leaves, recovered from the early Oligocene Shangcun Formation of Maoming Basin, Guangdong Province, South China. The morphological and anatomical structures of these fossils confirm their attribution to the subg. Cyclobalanopsis. Quercus shangcunensis sp. nov. represents the world's first Oligocene fossil acorn and the earliest fossil acorn in China for the subgenus, indicating that the subg. Cyclobalanopsis arrived in the low latitude area of South China at least by the early Oligocene. Our fossil provides evidence that the subgenus was present in one of its modern distribution centers in the Oligocene, suggesting that the modern distribution patterns of the subg. Cyclobalanopsis most likely originated during or prior to the Oligocene.
• Occurrence of Christella (Thelypteridaceae) in Southwest China and its indications of the paleoenvironment of the Qinghai–Tibetan Plateau and adjacent areas

  Cong-Li Xu, Tao Su, Jian Huang, Yong-Jiang Huang, Shu-Feng Li, Yi-Shan Zhao, and Zhe-Kun Zhou

  J Syst Evol. 2019, 57(2): 169-179.
  https://doi.org/10.1111/jse.12452

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  The uplift of the Qinghai–Tibetan Plateau dramatically changed the regional topography and climate, profoundly impacting the distribution of many plant lineages. Plant responses to environmental changes are particularly prominent in lineages that require ecological factors differentiated from those present before the uplift of the QTP. Two fossil occurrences of Christella H. Lév., Fl. Kouy–Tchéou (Thelypteridaceae), a fern genus now distributed mainly at low elevations of the pantropics with warm and moist habitats, are described based on fossilized Cenozoic leaf fronds recovered from SW China: late Paleocene Christella nervosa (J. R. Tao) C. L. Xu, T. Su & Z. K. Zhou comb. nov. found in Liuqu, southern Tibet and middle Miocene Christella sp. recovered from the Jinggu Basin in western Yunnan. The frond fossils from both sites share key morphological characteristics that diagnose these fossils as Christella. After detailed comparisons, we further clarified Christella papilio (C. Hope) Holttum, a species distributed in warm, humid habitats at altitudes no more than 1300?m, as the nearest living relative of C. nervosa. This finding suggested that southern Tibet had not reached its present elevation during the late Paleocene (ca. 56 Ma). We propose that the uplift, accompanied by severe cooling and aridification after the late Paleocene, caused the disappearance of Christella in southern Tibet, whereas paleoenvironmental conditions enabled the genus to survive in Yunnan. Our study provides the first example of distributional constraints of ferns in SW China in response to paleoenvironmental changes in the Qinghai–Tibetan Plateau and nearby areas.
• Fruit fossils of Rosoideae (Rosaceae) from the late Pliocene of northwestern Yunnan, Southwest China

  Yong-Jiang Huang, Hai Zhu, Arata Momohara, Lin-Bo Jia, and Zhe-Kun Zhou

  J Syst Evol. 2019, 57(2): 180-189.
  https://doi.org/10.1111/jse.12443

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  The subfamily Rosoideae Focke (Rosaceae) has a good fossil record in the Northern Hemisphere, but these fossil records are confined mainly to a few genera, whereas the majority, in particular those with herbaceous members, are still under‐represented. In this study, we describe new fruit fossils of Rosoideae, including Fragaria achenes and Rubus endocarps, from the late Pliocene of northwestern Yunnan, Southwest China. These fossils add new accounts to the fossil archive of Rosoideae and provide the first fossil record of Fragaria in East Asia. The new fossil findings provide a historical backdrop for the modern diversity and distribution of the subfamily in northwestern Yunnan, a topographically complex area accommodating a high diversity for many plant groups. Our Rubus fossils, in combination with other nearby coeval occurrences of the genus, suggest that Rubus was already establishing its modern diversity in northwestern Yunnan during the late Pliocene. This finding enriches our knowledge of the post‐Neogene diversification of flowering plants in northwestern Yunnan, which is thought to be largely driven by dramatic mountain uplifts and environmental complications associated with the southeastern extension of the Tibetan Plateau.
• New pollen classification of Chenopodiaceae for exploring and tracing desert vegetation evolution in eastern arid central Asia

  Kai-Qing Lu, Min Li, Guo-Hong Wang, Lian-Sheng Xu, David K. Ferguson, Anjali Trivedi, Jing Xuan, Ying Feng, Jin-Feng Li, Gan Xie, Yi-Feng Yao, and Yu-Fei Wang

  J Syst Evol. 2019, 57(2): 190-199.
  https://doi.org/10.1111/jse.12462

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  Members of the Chenopodiaceae are the most dominant elements in the central Asian desert. The different genera and species within this family are common in desert vegetation types. Should it prove possible to link pollen types in this family to specific desert vegetation, it would be feasible to trace vegetation successions in the geological past. Nevertheless, the morphological similarity of pollen grains in the Chenopodiaceae rarely permits identification at the generic level. Although some pollen classifications of Chenopodiaceae have been proposed, none of them tried to link pollen types to specific desert vegetation types in order to explore their ecological significance. Based on the pollen morphological characters of 13 genera and 24 species within the Chenopodiaceae of eastern central Asia, we provide a new pollen classification of this family with six pollen types and link them to those plant communities dominated by Chenopodiaceae, for example, temperate dwarf semi‐arboreal desert (Haloxylon type), temperate succulent halophytic dwarf semi‐shrubby desert (Suaeda, Kalidium, and Atriplex types), temperate annual graminoid desert (Kalidium type), temperate semi‐shrubby and dwarf semi‐shrubby desert (Kalidium, Iljini, and Haloxylon types), and alpine cushion dwarf semi‐shrubby desert (Krascheninnikovia type). These findings represent a new approach for detecting specific desert vegetation types and deciphering ecosystem evolution in eastern central Asia.
• More Malpighiales: Woods of Achariaceae and/or Salicaceae from the Deccan Intertrappean beds, India

  Rashmi Srivastava, Regis B. Miller, and Pieter Baas

  J Syst Evol. 2019, 57(2): 200-208.
  https://doi.org/10.1111/jse.12455

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  The families Achariaceae and Salicaceae (Malpighiales) are characterized by wood anatomical ranges that partly overlap. Formerly these families were treated together in the polyphyletic Flacourtiaceae and a much more narrowly circumscribed Salicaceae. Here we attribute two recently collected fossil woods from the Deccan Intertrappean Beds to the clade that contain these two families, i.e., the Parietal Clade of the Malpighiales. The new genus Elioxylon shares features with several extant genera of Achariaceae and Salicaceae, but does not completely match with any of them. A new record of Hydnocarpoxylon indicum Bande & Khatri is a good match for extant Hydnocarpus Gaertn. (Achariaceae). Elioxylon and Hydnocarpoxylon share an absence of parenchyma, the presence of septate fibres and 1–3 seriate heterocellular rays with long uniseriate margins consistent with Achariaceae and Salicaceae. Elioxylon has mixed simple and scalariform perforations, whereas Hydnocarpxylon has exclusively scalariform perforations. Other Deccan fossils formerly attributed to “Flacourtiaceae” in the literature are critically discussed and mostly excluded from Achariaceae and Salicaceae. Elioxylon and Hydnocarpoxylon from the Maastrichtian ‐ Danian of India are the oldest fossil records of the Parietal Clade of the Malpighiales. With their occurrence on the Indian plate during its northward journey from Gondwana to Laurasia, these fossils provide further support for an ‘out‐of‐India’ hypothesis for Achariaceae and/or Salicaceae. “Baileyan trends” in vessel perforation plate and vessel grouping evolution are apparent in the phylogeny of the Parietal Clade.