Cenozoic xeromorphic vegetation in China and its spatial and temporal development in connection with global changes

Cenozoic xeromorphic vegetation in China and its spatial and temporal development in connection with global changes

Available online at www.sciencedirect.com ScienceDirect Palaeoworld 22 (2013) 86–92 Cenozoic xeromorphic vegetation in China and its spatial and tem...

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Available online at www.sciencedirect.com

ScienceDirect Palaeoworld 22 (2013) 86–92

Cenozoic xeromorphic vegetation in China and its spatial and temporal development in connection with global changes Wei-Ming Wang ∗ , Jun-Wu Shu Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China Received 12 May 2013; received in revised form 11 July 2013; accepted 13 August 2013 Available online 27 August 2013

Abstract Remarkable angiosperm evolution and significant vegetation changes took place during the Cenozoic on earth. Specifically, Neogene is a crucial period in forming the framework of the current vegetation and in restructing earth ecosystem after the Eocene–Oligocene “greenhouse-icehouse” climate transition. Under the background of global changes, regional vegetation in China was synchronously developed. Some local changes had made extensive influences, such as the topographic changes, especially the uplift of the Qinghai-Tibet Plateau, and the changes in seasonal atmospheric circulation and precipitation, i.e., the initiation and evolvement of Asian monsoon system, along with other geographic changes in land and sea. The expansion of xeromorphic vegetation, represented by forest steppe, steppe and desert, played an ever-increasing role, which was closely related with the development of angiospermous xerophytes. The distributions of some representative angiospermous pollen types with their parent plants of xerophilous origination in the Cenozoic are summarized in the paper. Evidence shows that the origination and evolution of the angiospermous xerophytes underwent a series of developments, which were prompted largely by environmental variations. Pollen studies from some representative sites in China show that spatial and temporal development of Cenozoic xeromorphic vegetation is largely consistent with the global changes. © 2013 Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS. All rights reserved. Keywords: Xerophytes; Vegetation; Global changes; Cenozoic; China

1. Introduction The Cenozoic witnesses a series of extraordinary changes in land and sea, resulting in severe climate variations. The global warm climates had already begun a long and irregular slide into the glaciations, which characterized the late Cenozoic (Frakes, 1979). The Cenozoic climate was generally warm and stable in its early stage, and became deteriorated from the middle stage, with a worldwide glacier developed in the late stage (Berger, 1982; Shackleton et al., 1984; Barrett, 2003). The Eocene–Oligocene boundary, at about 33.5 Ma ago, marks the transition from “greenhouse-” to “icehouse-world” (Abelson et al., 2008). This changing tendency of the Cenozoic climate did not go straightforward, but was punctuated by periods of warmth, which were especially distinct during 19.5–16.5 Ma and 4.6–3.8 Ma (Kennett and von der Borch, 1985). Zachos et al.



Corresponding author. Tel.: +86 25 83282222. E-mail address: [email protected] (W.-M. Wang).

(2001) summarized the Cenozoic deep sea ␦18 O temperature curve, changes in ice-sheets at Polar Regions, major global climatic, tectonic and biological events in their paper, which is widely recognized in the academic circle. One of the major influences on the extant vegetation in China is the Asian monsoon system. The present East China is part of the Asian monsoon region. Geographically, taking the Daxing’anling, Yinshan, and Helanshan Mts. and the east border of the Qinghai-Tibet Plateau as an approximate boundary (Fig. 1), its southeast part is affected mainly by the Pacific Ocean monsoon, and its southwest part by the Indian Ocean monsoon, occupying about 47.6% of the whole Chinese territory (Zhang, 1989). There are many discussions on the initiation and development of the Asian monsoon in the geological past (Wang, 1984, 2009; Zhang, 1984, 1989; Liu, 1999; Sun and Wang, 2005; Wang et al., 2009), which is believed to be closely related with the intensified Cenozoic environment changes. Topographically, the formation of three macro-landform complexes in China, including some major mountain systems, numerous intermontane plateaus, basins, and plains, also played a great role.

1871-174X/$ – see front matter © 2013 Elsevier B.V. and Nanjing Institute of Geology and Palaeontology, CAS. All rights reserved. http://dx.doi.org/10.1016/j.palwor.2013.08.001

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Fig. 1. Geographic map of China showing major topographic units and monsoon influenced regions.

In the Cenozoic of China, the temperate and warm temperate xeromorphic vegetation, such as forest steppe, steppe and desert played an increasing role. This paper aims at the major changing process of the Cenozoic xeromorphic vegetation in China under the background of the global changes. To better understand the vegetation changes, we also summarize the distribution of some representative angiospermous pollen types with their parent plants of xerophilous origination in the Cenozoic of China. 2. Evolution of main angiospermous xerophytes in the Cenozoic of China The angiospermous xerophytes are well represented in the Cenozoic of China. Their families or genera have varying biostratigraphic significances in different time ranges (Song et al., 2004; Wang et al., 2006). The origination and evolution of xerophytes underwent a series of development, and were prompted largely by environmental variations, resulting in the formation of xeromorphic vegetation. Some of the xerophytes are especially representative, such as Chenopodiaceae, Nitrariaceae, Asteraceae, Poaceae, and Caryophyllaceae. Chenopodiaceae was first recorded from the Cenomanian Qingshankou Formation in the Songliao Basin (Gao et al., 1999). It is widely distributed in the Cenozoic of China, especially in Northwest China. Pollen records from the Qaidam Basin show

two distinct flourishing stages of the plant in the Oligocene and the Middle Miocene respectively (Zhu et al., 1985; Fig. 2). Currently, there are about 42 genera and 190 species of Chenopodiaceae, which are distributed mainly in deserts, and coastal and saline habitats in China. Nitraria is a genus of halophilic in the family Nitrariaceae (previously assigned to the family Zygophyllaceae). The sometimes thorny shrubs have alternate and entire fleshy leaves, and are distributed mainly in Northwest China. Nitraria first occurred in the Late Paleocene upper part of the Qijiachuan Formation in Xining area, and fully developed in the Late Eocene–Early Oligocene as revealed by the pollen data from the Qaidam Basin (Zhu et al., 1985; Fig. 3). There are four fossil pollen genera, related to Astereae, Tubuliforae, Artemisia, and Cichorieae respectively in Asteraceae. All of them were first recorded in the Qaidam Basin, with Astereae from the Eocene Lulehe Formation, Tubuliforae from the Eocene lower part of the Lower Ganchaigou Formation, and Artemisia and Cichorieae from the Oligocene Upper Ganchaigou Formation, respectively (Zhu et al., 1985). Although the origin of Asteraceae might be dated back to the Paleogene, it did not thrive until the Middle Miocene, and only became widely distributed in the Pliocene (Wang, 2004). Most members of Asteraceae are herbaceous, but a significant number of them are also shrubs, vines, and trees. This family has a worldwide distribution and is most common in the

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Fig. 2. Percentage and number of species of Chenopodiaceae in Cenozoic pollen assemblages of the Qaidam Basin (data after Zhu et al., 1985).

arid and semi-arid regions of subtropical and lower temperate latitudes. Poaceae was first recorded from the Eocene First Member of the Kongdian Formation in the coastal region of the Bohai Sea (IPEDPMPI and NIGPAS, 1978), and the Eocene lower part of the Lower Ganchaigou Formation in the Qaidam Basin (Zhu et al., 1985) respectively. It was generally insignificant in the Middle Cenozoic, and became widely distributed from the late Cenozoic, especially in the Holocene. The grass family is now one of the most widely distributed and abundant groups of plants on earth. Though Caryophyllaceae was first documented from the Eocene Lower Ganchaigou Formation in the Qaidam Basin (Zhu et al., 1985), its main developing stage was in the Quaternary. As a cosmopolitan family with mostly herbaceous plants, Caryophyllaceae is best represented in temperate climates, with a few species growing on tropical mountains. To evaluate the evolution and development of the angiospermous xerophytes in the Cenozoic of China, we give the first occurrence of some angiospermous pollen that show affinities to the modern terrestrial herbs and shrubs in the Appendix. The early angiospermous pollen Clavatipollenites Couper, 1958 and

Asteropollis Hedlund et Norris, 1968 show affinity with Chloranthaceae. Modern plants in Chloranthaceae are mostly herbs or shrubs, distributed mainly in the tropical and subtropical areas (How, 1982). We consequently propose that parts of the early angiosperms are possibly inherent with xerophilous attribute. Xerophytes that occurred during the Late Cretaceous–Paleogene include Dipsacaceae, Chenopodiaceae, Apiaceae, Centrolepidaceae, Zygophyllaceae, and Astereae (Asteraceae). They are mostly considered as thermophilic plants with arid adaptation because of the overall tropical and subtropical conditions at that time (Song et al., 1983). More plant types occurred during the Eocene–Oligocene, which is proposed as a major stage for the transition of the angiospermous xerophytes from the former thermophilic type to the cold resistant one. There are Tubuliforae, Artemisia and Cichorieae (Asteraceae), Lamiaceae, Poaceae, Polygonaceae, Amaranthaceae, Caryophyllaceae, Ericaceae, Ranunculaceae, Cyperaceae, and Geraniaceae in the recognized plants. The angiospermous xerophytes became more cold resistant as a result of the climate deterioration since the terminal Eocene. However, most of the plants were still relatively rare in the floras, except for Nitraria, which was fully developed in

Fig. 3. Percentage and number of species of Pokrovskaja (Nitraria) in Cenozoic pollen assemblages of the Qaidam Basin (data after Zhu et al., 1985).

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Northwest China, while Chenopodiaceae also became important at the late stage. Though the distribution of the angiospermous xerophytes got two temporary retreats in the Miocene Climate Optimum and the mid-Pliocene warm period respectively, the terrestrial angiospermous herbs became increasingly important all through the late Cenozoic. Chenopodiaceae was flourishing in the Middle–Late Miocene, while Asteraceae and Poaceae also developed gradually. The angiospermous xerophytes had their full development and diversification in the Pliocene, and went on further in the Quaternary. It is notable that the main angiospermous xerophytes in the Neogene are mainly those originated earlier, although there are also some limited new types, such as Portulaceae and Acanthaceae, etc. 3. Cenozoic xeromorphic vegetation in China and its spatial and temporal development The origination and development of the angiospermous xerophytes resulted in the formation of xeromorphic vegetation, such as forest steppe, steppe and desert. The early temperate xeromorphic vegetation was first recognized from the latest Eocene Qaidam Basin, indicated by steppe and shrub. Evidence from some related pollen floras shows that the original steppe was rather monotonous in components with an isolated distribution (Wang and Zhang, 1990). Since its appearance, the xeromorphic vegetation had gone through a series of expansion and concentration, which mostly displays a linkage with the global changes. Pollen record from the Linxia Basin, the northeast boundary of the Qinghai-Tibet Plateau (Fig. 1), shows a forest recovery at about 21.8 Ma (Shi et al., 1998). During the Miocene Climate Optimum, steppe was much concentrated or even disappeared in many places (Song et al., 2008). Since the Middle Miocene (ca. 14 Ma), forest steppe and steppe have gradually expanded and fully developed with its distribution extending from Northwest China to Northeast and North China (Wang, 1990, 1996, 1999; Wang and Zhang, 1990), especially in the Pliocene. At the same time, original semi-desert and desert began to form in the Northwest, and vegetation in the Qinghai-Tibet area gradually showed differentiation compared with those in other regions because of the constant uplift of the plateau. In the Quaternary, the former Tertiary steppe was partly replaced by desert in the Northwest, but fully developed in the Songliao Plain, Inner Mongolia Plateau, Loess Plateau, and North China Plain. Alpine frigid desert and steppe were formed on the Qinghai-Tibet Plateau. The distribution of the vegetation in China gradually took the present shape during this stage. The major development stages of Cenozoic vegetation in China are summarized in Table 1. A generally changing tendency of Neogene vegetation in China is also revealed by the Integrated Plant Record (IPR) vegetation analysis. Based on the semi-quantitative reconstruction method, we found there was an increase in the broad-leaved deciduous components in the northern areas during the Neogene. At the same time, an increase of sclerophyllous and herbaceous components in western, central, and northern China occurred. However, there is no noticeable change in the Neogene vegetation of southern China.

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The Pliocene is characterized by an increasing contrast in vegetation between southern and northern China (Jacques et al., 2013). 4. Cenozoic vegetation in China and its relation with global changes Accumulated evidence from land and sea has already shown an overall deterioration of the Cenozoic climate (Zachos et al., 2001). Despite some punctuated periods of warmth (Kennett and von der Borch, 1985), the global warm climates had already begun a long and irregular slide into the glaciations, which characterized the late Cenozoic (Frakes, 1979). During the Eocene–Oligocene “greenhouse-icehouse” climate transition, the angiospermous xerophytes experienced a major transition from the original thermophilic type to the later cold resistant one, which directly led to the formation of temperate vegetation. Meanwhile, great topographic changes in China were also accompanied with vegetation variations. At the beginning of the Paleocene, the geography of China was relatively even. Both geographic and climatic patterns were mainly a continuation of those in the Cretaceous. The temperature on earth was commonly high, and the northern boundary of tropical and subtropical zones was located at much higher latitude than today (Leopold, 1969). The Chinese continent was situated mostly within the tropical and subtropical zones except part of warm temperate area in North China (Wu, 1980), and an arid subtropical climate prevailed over all of southern and northwestern China (Wang, 1984). The mountains formed in the Paleozoic and the Mesozoic were leveled after a long period of denudation (Wu, 1980). Affected by the Himalayan orogeny, the Tethys was retreated from the west part of the Chinese Continent at the terminal Eocene. Under the huge northward drifting forces of the South Continent, the crust in the Middle Asian Plain and the former leveled mountains began to rise, and the present geographic features gradually took shape. With the great tectonic movements and climate changes, seasonal atmospheric circulation and precipitation, i.e., Asian monsoon system, simultaneously occurred. Meteorologically, monsoon is defined as a wind system caused by thermal differences between continent and ocean in the winter and summer. Its intensity and the scope primarily depend on the relative sizes between continent and ocean, latitudinal location of the continent, and the latitudinal thermal differentiation of the Earth (Gao, 1984). Additionally, the thermal differentiation between plateau and the free atmosphere at the same altitude in the surrounding region is the same as that between continent and ocean, leading to the specific distribution of air pressure and the field of air flow (Ding, 1992). Strengthening of the Asian monsoon has been linked to the uplift of the Qinghai-Tibet Plateau after the collision of India and Asia in the Eocene (e.g., Garzanti et al., 1987; Aitchison et al., 2008), resulting in an enlarged ratio between Eurasia continent and the Pacific Ocean. When we discuss the development of monsoon in the geological past, it is not necessary to be completely comparable with that of today. The Early Miocene pollen floras exhibit a broad influence of the increasing summer monsoon in many places

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Table 1 Major development stages of Cenozoic vegetation in China. Main stages

Distribution characteristics of vegetation

Stage 4 (since ca. 2.6 Ma)

Steppe replaced by desert in Northwest China, fully developed in the Songliao Plain, Inner Mongolia Plateau and Loess Plateau; alpine frigid desert and steppe formed in the Qinghai-Tibetan Plateau; temperate mixed coniferous-deciduous broad-leaved forests, and cold temperate coniferous forests developed in Northeast China Main development period for steppe, gradually expanding from Northwest China to Northeast and North China; original desert formed in Northwest China; more developed warm temperate to temperate deciduous forest in Northeast and North China, partly coniferous forest in Northern-Northeast China; vegetation differentiated in Southwest China First occurrence of original steppe in Northwest China; temperate vegetation spreading in North China, with frequent occurrence of conifers showing some fluctuations Vegetation largely along the latitude for distribution, with tropical, subtropical-warm temperate forests fully developed, and some sparse woods existing in the subtropical arid regions

Stage 3 (ca. 14–2.6 Ma)

Stage 2 (ca. 33.9–14 Ma) Stage 1 (ca. 65.5–33.9 Ma)

of China, which is consistent with the Qinghai-Tibet Plateau uplift acceleration and a subsequent Miocene Climatic Optimum (e.g., CETQXP, 1983; Pan et al., 1998; Zachos et al., 2001; Böhme, 2003; Zheng and Yao, 2006), indicating that a strong summer monsoon prevailed in the Early Miocene, which reached to the maximum during the Miocene Climate Optimum. A further development of the xeromorphic vegetation in the late Cenozoic is closely related with global cooling and an intensified winter monsoon in East Asia. According to the meteorological study, the strength of summer monsoon in the Middle and higher latitude regions during the strong winter monsoon years will be less active, and the precipitation decreases following a strong winter monsoon (Sun and Chen, 2000). This is also proved by the IPR vegetation analysis. An increase of sclerophyllous and herbaceous components in the Neogene of western, central, and northern China is recognized to be indicative of aridification. The aridification of northern China is due to a strengthening of the winter monsoon. Because there is no major change in the vegetation of southern China (Wang, 2006), the weakening of East Asian summer monsoon in the lower latitude is improbable. The Pliocene cooling is responsible for colder winters in Siberia, and the winter high pressure over Siberia becomes higher. As a result, the winter monsoon winds are stronger. The evolution of the summer and winter monsoons is thus not coupled (Jacques et al., 2013).

The Early Miocene vegetation in China shows an initiative control of the summer monsoon, which reached the maximum in the Miocene Climate Optimum. A prevailing winter monsoon with minor oscillations followed after the Middle Miocene at about 14 Ma years ago. The spatial and temporal development of Cenozoic vegetation in China is well evidenced to be closely connected with global changes, along with some local topographic variations, especially the uplift of the Qinghai-Tibet Plateau, and seasonal atmospheric circulation and precipitation, i.e., the initiation and evolvement of monsoon climate. Acknowledgements We thank Dr. Sangheon Yi and an anonymous reviewer for their helpful comments. This study is funded by the CAS Strategic Priority Research Program (XDA05120101) and the Pilot Project of Knowledge Innovation, CAS (KZCX2-YW105). Appendix A. List of some angiospermous pollen with affinities to modern terrestrial herbs and shrubs in the geological past of China (in chronological order with age of first occurrence in brackets) (Song et al., 1999, 2004; Wang et al., 2006).

5. Conclusions After the Eocene–Oligocene “greenhouse-icehouse” climate transition, the overall tropical and subtropical zones in China had been gradually replaced by the warm temperate and temperate zones in the middle and higher latitudes, which was accompanied by a major transition of the angiospermous xerophytes from the thermophilic types to the cold resistant ones, resulting in the formation of the temperate xeromorphic vegetation. The earliest temperate xeromorphic vegetation was recognizable from the latest Eocene. Since the Middle Miocene (ca. 14 Ma), forest steppe and steppe gradually expanded to become fully developed with its distribution extending from the Northwest China to the Northeast and the North China. The xeromorphic vegetation got further developed with extraordinary vegetation differentiation occurred in the Pliocene and Quaternary.

Chloranthaceae Clavatipollenites Couper, 1958 (Early Cretaceous) Asteropollis Hedlund et Norris, 1968 (Early Cretaceous) Chloranthacearumpollenites Nagy, 1969 (Miocene) Dipsacaceae Scabiosapollis Song et Zheng, 1980 (Late Cretaceous) Jianghanpollis Wang et Zhao, 1979 (Late Cretaceous) Morinoipollenites Wang et Zhao, 1979 (Late Cretaceous) Chenopodiaceae Chenopodipollis Krutzsch, 1966 (Late Cretaceous)

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Apiaceae Nanlingpollis Sun et He, 1980 (Late Cretaceous) Hydrocotaepites Zheng, 1985 (Eocene) Umbelliferaepites Biswas, 1962 (Pliocene) Centrolepidaceae Centrolepidacidites Wu et Yu, 1981 (Paleocene) Nitrariaceae

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Geraniaceae Geraniapollis Song et Zhu, 1985 (Oligocene) Portulaceae Rugaepollis Engelhardt, 1966 (Miocene) Acanthaceae Acanthus L., 1753 (Pliocene) Rostellularia Rchb., 1837 (Pliocene)

Pokrovskaja Boitzova, 1979 emend. Zhu, 1999 (Paleocene) References Asteraceae Echitricolporites Van der Hammen, 1956 ex Germeraad Hopping et Muller, 1968 (Paleocene) Tubulifloridites Cookson, 1947 ex Potonié, 1960 (Eocene) Artemisiaepollenites Nagy, 1969 (Oligocene) Cichoreacidites Sah, 1967 (Oligocene) Lamiaceae Labitricolpites Ke et Shi, 1978 (Eocene) Poaceae Graminidites Cookson, 1947 ex Potonié, 1960 (Eocene) Polygonaceae Persicarioipollis Krutzsch, 1962 (Eocene) Amaranthaceae Vaclavipollis Krutzsch, 1966 (Eocene) Caryophyllaceae Caryophyllidites Couper, 1960 emend. Krutzsch, 1966 (Eocene) Ericaceae Ericipites Wodeh., 1933 emend. Krutzsch, 1970 (Eocene) Ranunculaceae Ranunculacidites Sah, 1967 (Eocene) Cyperaceae Cyperaceaepollis Krutzsch, 1970 (Oligocene)

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