Cenozoic plant diversity of Yunnan： A review

Yunnan in southwestern China is renowned for its high plant diversity.To understand how this modern botanical richness formed,it is critical to investigate the past biodiversity throughout the geological time.In this review,we present a summary on plant diversity,floristics and climates in the Cenozoic of Yunnan and document their changes,by compiling published palaeobotanical sources.Our review demonstrates that thus far a total of 386 fossil species of ferns,gymnosperms and angiosperms belonging to 170 genera within 66 families have been reported from the Cenozoic,particularly the Neogene,of Yunnan.Angiosperms display the highest richness represented by 353 species grouped into 155 genera within 60 families,with Fagaceae,Fabaceae,Lauraceae and Juglandaceae being the most diversified.Most of the families and genera recorded as fossils still occur in Yunnan,but seven genera have disappeared,including Berryophyllum,Cedrelospermum,Cedrus,Palaeocarya,Podocarpium,Sequoia and Wataria.The regional extinction of these genera is commonly referred to an aridification of the dry season associated with Asian monsoon development.Floristic analyses indicate that in the late Miocene,Yunnan had three floristic regions:a northern subtropical floristic region in the northeast,a subtropical floristic region in the east,and a tropical floristic region in the southwest.In the late Pliocene,Yunnan saw two kinds of floristic regions:a subalpine floristic region in the northwest,and two subtropical floristic regions separately in the southwest and the eastern center.These floristic concepts are verified by results from our areal type analyses which suggest that in the Miocene southwestern Yunnan supported the most Pantropic elements,while in the Pliocene southwestern Yunnan had abundant Tropical Asia(Indo e Malaysia)type and East Asia and North America disjunct type that were absent from northwestern Yunnan.From the late Miocene to late Pliocene through to the present,floristic composition and vegetation types changed markedly,presumably in response to altitude changes and coeval global cooling.An integration of palaeoclimate data suggests that during the Neogene Yunnan was warmer and wetter than today.Moreover,northern Yunnan witnessed a pronounced temperature decline,while southern Yunnan experienced only moderate temperature changes.Summer precipitation was consistently higher than winter precipitation,suggesting a rainfall seasonality.This summary on palaeoclimates helps us to understand under what conditions plant diversity occurred and evolved in Yunnan throughout the Cenozoic.

Wu Zhengyi and his contributions to plant taxonomy and phytogeography

Wu Zhengyi,a master in science and a giant in botany,was born in Jiujiang,Jiangxi province,China 100 years ago and passed away on June 20,2013 in Kunming,China.He was an Academician of the Chinese Academy of Sciences,Professor and Director Emeritus of Kunming Institute of Botany,Chinese Academy of Sciences。

A few words on our commemoration of 80 years at KIB

Eighty years ago, a botanical institute called"the Yunnan Provincial Institute of Agricultural and Forestry Botany"was jointly founded by the Jingsheng Biology Institute and the Yunnan State Department of Education in Kunming. At that time, the Heilong Dragon Park became the institute's home (Fig. 1) and the research conducted there mostly focused on taxonomy and economical botany. When the institute came under the auspices of the Chinese Academy of Science, it still had fewer than 20 staff members.

Sharing earth with all life

The Anthropocene is marked by twin crises:climate change and loss of biodiversity.Climate change has dominated the headlines,reflecting,in part,the greater complexity that characterizes the biodiversity crisis(Corlett,2020).Since the Convention on Biological Diversity(CBD)was adopted in 1993,the conservation of biological diversity has become a global concern.If we want a sustainable future,we must protect biodiversity.However,biodiversity loss remains one of the most challenges today.

Paleogene integrative stratigraphy,biotas,and paleogeographical evolution of the Qinghai-Tibetan Plateau and its surrounding areas

The Paleogene is a crucial period when terrestrial and marine ecosystems recovered from major disruptions and gradually approached their modern states.In the Qinghai-Tibetan Plateau and its surrounding regions,the Paleogene also represents a significant phase of tectonic evolution in the Qinghai-Tibetan Plateau-Himalaya orogeny,reorganization of Asian climates,and evolution of biodiversity.Due to limitations in research conditions and understanding,there are still many controversies regarding stratigraphic divisions in the Qinghai-Tibetan Plateau and its surrounding regions.In recent years,extensive studies on sedimentary petrology,magnetostratigraphy,and isotope dating have been conducted in the region.Numerous fossils have been discovered and reported,contributing to a more systematic understanding of biostratigraphy.These studies have laid a solid foundation for the comprehensive investigation of the stratigraphy,biotas and paleogeographic evolution of the Qinghai-Tibetan Plateau and its surrounding regions during the Paleogene.In this paper,we integrate recent research on fossils,isotopic dating,magnetostratigraphy,and geochemistry to refine the stratigraphic divisions and correlation framework of different tectonic units in the region,building upon previous studies.Since the Second Tibetan Plateau Scientific Expedition and Research,the knowledge of Paleogene floras has gradually expanded.This paper discusses the biostratigraphic significance of extinct and newly appeared taxa based on the latest dating results of these plant species.The new understanding of fossil species such as the“Eucalyptus”and Arecaceae establishes connections between the Paleogene flora of the Qinghai-Tibetan region and the biotas of Gondwana,specifically Oceania and South America.The evolutionary history of key taxa near the Yarlung Zangbo suture zone indicates that the collision between the Indian and Eurasian plates occurred approximately 65-54 Ma.Paleoelevation reconstructions,based on plant fossils,suggest that the Hengduan Mountain had already formed their current topographic pattern prior to the Early Oligocene.The warm and humid lowlands adjacent to the main suture zones in the Paleogene Qinghai-Tibetan Plateau served as the primary pathway for biota exchanges.The relatively low elevation of the Himalaya during the Paleogene did not effectively block the moisture from the Indian Ocean.

Vegetation changes across the Eocene-Oligocene transition:Global signals vs.regional development

The Eocene-Oligocene transition(EOT)marked a rapid global cooling event,often considered as the beginning of the modern icehouse world.Influenced by various factors,including tectonic activity and paleogeographic settings,the terrestrial records indicate a diverse response of fauna and vegetation to this global event.We examined nine macrofossil assemblages from seven fossil localities on the southeastern margin of the Tibetan Plateau and from the mid-latitudinal Europe ranging from the latest Bartonian and Priabonian(37.71-33.9 Ma)to the Rupelian(33.9-27.82 Ma).Our aims were to trace and compare the vegetation history of both regions in the late Eocene and early Oligocene.The results show that both regions experienced changes in vegetation composition in response to climate change,characterized by a decrease in the percentages of broad-leaved evergreen elements and distinctive changes in general vegetation types.A general change in the overall vegetation type from subtropical broad-leaved evergreen forests in the late Eocene to temperate broad-leaved mixed deciduous evergreen forests,or mixed mesophytic forests,in the early Oligocene is recognized in both regions.The results indicate a clear change in leaf architecture,leaf margin states,and secondary venation types in the mid-latitudinal Europe,while the results from the southeastern margin of the Tibetan Plateau show a distinct reduction in leaf size.Our data suggest that both global and regional factors played key roles in shaping the vegetation in the two regions.

新生代青藏高原生长对东亚水循环及生态系统的影响

新生代青藏高原生长和全球温度变化驱动东亚气候和生态系统发生了剧烈变化.本文综述了前人的相关研究成果,基于前期的数值模拟工作,进一步探讨了青藏高原地形地貌演变对东亚水循环及生态系统的影响.目前,关于青藏高原新生代以来地形地貌的演化还存在争议,但近年来不同学科的证据一致认为,青藏高原生长过程具有区域差异性.最新的古气候数值模拟表明,青藏高原北部抬升显著改变了亚洲气候系统,促使东亚降水显著增加,尤其是中国南方冬季降水增加更为明显,地表径流也相应增加,对东亚水系格局产生了重要影响;土壤含水量也随之增加,冬季深层土壤含水量增加尤为显著,导致中国东部植被从干旱、半干旱转变为湿润、半湿润植被类型,造就了现今东亚植被和生物多样性格局.目前,关于青藏高原生长对东亚降水的影响,科学界已有深入认识,但是还有若干关键问题亟待解决,包括:对青藏高原地形地貌演化的进一步限定、对气候转型期高精度的古气候数值模拟,以及进一步解析新生代东亚水循环和生态系统的耦合关系.理解这些关键问题对预测未来全球气候急剧变化背景下的生态系统响应及其演变趋势具有重要的参考意义.

Cenozoic plants from Tibet:An extraordinary decade of discovery,understanding and implications

Plant fossils play an important role in understanding landscape evolution across the Tibetan Region,as well as plant diversity across wider eastern Asia.Within the last decade or so,paleobotanical investigations within the Tibet Region have led to a paradigm shift in our understanding of how the present plateau formed and how this affected the regional climate and biota.This is because:(1)Numerous new taxa have been reported.Of all the Cenozoic records of new plant fossil species reported from the Tibet(Xizang)Autonomous Region 45 out of 63(70%)were documented after 2010.Among these,many represent the earliest records from Asia,or in some cases worldwide,at the genus or family level.(2)These fossils show that during the Paleogene,the region now occupied by the Tibetan Plateau was a globally significant floristic exchange hub.Based on paleobiogeographic studies,grounded by fossil evidence,there are four models of regional floristic migration and exchange,i.e.,into Tibet,out of Tibet,out of India and into/out of Africa.(3)Plant fossils evidence the asynchronous formation histories for different parts of the Tibetan Plateau.During most of the Paleogene,there was a wide east-west trending valley with a subtropical climate in central Tibet bounded by high(>4 km)mountain systems,but that by the early Oligocene the modern high plateau had begun to form by the rise of the valley floor.Paleoelevation reconstructions using radiometrically-dated plant fossil assemblages in southeastern Tibet show that by the earliest Oligocene southeastern Tibet(including the Hengduan Mountains)had reached its present elevation.(4)The coevolution between vegetation,landform and paleoenvironment is evidenced by fossil records from what is now the central Tibetan Plateau.From the Paleocene to Pliocene,plant diversity transformed from that of tropical,to subtropical forests,through warm to cool temperate woodland and eventually to deciduous shrubland in response to landscape evolution from a seasonally humid lowland valley,to a high and dry plateau.(5)Advanced multidisciplinary technologies and novel ideas applied to paleobotanical material and paleoenvironmental reconstructions,e.g.,fluorescence microscopy and paleoclimatic models,have been essential for interpreting Cenozoic floras on the Tibetan Region.However,despite significant progress investigating Cenozoic floras of the Tibetan Region,fossil records across this large region remain sparse,and for a better understanding of regional ecosystem dynamics and management more paleobotanical discoveries and multidisciplinary studies are required.

滇东南普阳盆地含煤地层孢粉组合特征及其年代

云南地处青藏高原东南缘,生物多样性丰富,季风气候特征明显。新生代剧烈的构造活动形成了众多山间盆地,其间保存了大量精美的植物化石,是探讨新生代以来植被、植物多样性和地球环境演变的理想地区。但是长期以来,由于地层年代学证据的缺乏,这些新生代沉积盆地的地质年代还存在很大争议。本研究通过对滇东南地区富宁县普阳盆地的含煤地层开展深入的孢粉学研究,探讨盆地含煤地层年代及其古气候演化过程。剖面下部煤层孢粉组合以杉粉属(Taxodiaceaepollenites)为主,指示以杉科为主的湿润沼泽森林,冬春季相对湿润;剖面上部孢粉组合指示以常绿栎类为主的亚热带常绿-落叶阔叶混交林,生长山核桃粉属(Caryapollenites)、冬青粉属(Ilexpollenites)、胡桃粉属(Juglanspollenites)、枫香粉属(Liquidambarpollenites)等亚热带常见树种,气候温暖湿润,季节分明。结合新发现的哺乳动物化石证据和周边地区不同地质时代的孢粉组合,普阳盆地含煤地层的沉积时代应为晚始新世;同时,孢粉组合也表明滇东南地区植被现代化面貌至少在晚始新世就已经开始出现。

Tibetan Plateau: An evolutionary junction for the history of modern biodiversity

Holding particular biological resources,the Tibetan Plateau is a unique geologic-geographic-biotic interactively unite and hence play an important role in the global biodiversity domain.The Tibetan Plateau has undergone vigorous environmental changes since the Cenozoic,and played roles switching from"a paradise of tropical animals and plants"to"the cradle of Ice Age mammalian fauna".Recent significant paleontological discoveries have refined a big picture of the evolutionary history of biodiversity on that plateau against the backdrop of major environmental changes,and paved the way for the assessment of its far-reaching impact upon the biota around the plateau and even in more remote regions.Here,based on the newly reported fossils from the Tibetan Plateau which include diverse animals and plants,we present a general review of the changing biodiversity on the Tibetan Plateau and its influence in a global scale.We define the Tibetan Plateau as a junction station of the history of modern biodiversity,whose performance can be categorized in the following three patterns:(1)Local origination of endemism;(2)Local origination and"Out of Tibet";(3)Intercontinental dispersal via Tibet.The first pattern is exemplified by the snow carps,the major component of the freshwater fish fauna on the plateau,whose temporal distribution pattern of the fossil schizothoracines approximately mirrors the spatial distribution pattern of their living counterparts.Through ascent with modification,their history reflects the biological responses to the stepwise uplift of the Tibetan Plateau.The second pattern is represented by the dispersal history of some mammals since the Pliocene and some plants.The ancestors of some Ice Age mammals,e.g.,the wholly rhino,Arctic fox,and argali sheep first originated and evolved in the uplifted and frozen Tibet during the Pliocene,and then migrated toward the Arctic regions or even the North American continent at beginning of the Ice Age;the ancestor of pantherines(big cats)first rose in Tibetan Plateau during the Pliocene,followed by the disperse of its descendants to other parts of Asia,Africa,North and South America to play as top predators of the local ecosystems.The early members of some plants,e.g.,Elaeagnaceae appeared in Tibet during the Late Eocene and then dispersed and were widely distributed to other regions.The last pattern is typified by the history of the tree of heaven(Ailanthus)and climbing perch.Ailanthus originated in the Indian subcontinent,then colonized into Tibet after the Indian-Asian plate collision,and dispersed therefrom to East Asia,Europe and even North America.The climbing perches among freshwater fishes probably rose in Southeast Asia during the Middle Eocene,dispersed to Tibet and then migrated into Africa via the docked India.These cases highlight the role of Tibet,which was involved in the continental collision,in the intercontinental biotic interchanges.The three evolutionary patterns above reflect both the history of biodiversity on the plateau and the biological and environmental effects of tectonic uplift.

Pliocene flora and paleoenvironment of Zanda Basin, Tibet, China

This paper describes a plant megafossil assemblage from the Pliocene strata of Xiangzi, Zanda Basin in the western Qinghai-Tibet Plateau. Twenty-one species belonging to 12 genera and 10 families were identified. Studies show that the Pliocene vegetation in Zanda Basin was mostly deciduous shrub composed of Cotoneaster, Spiraea, Caragana, Hippophae,Rhododendron, Potentilla fruticosa, etc. Leaf sizes of these taxa were generally small. Paleoclimate reconstruction using Coexistence Analysis and CLAMP showed that this area had higher temperature and precipitation in the Pliocene than today, and distinct seasonal precipitation variability was established. The reconstructed paleoelevation of Zanda Basin in the Pliocene was similar to modern times. In the context of central Asian aridification, the gradual drought in the area beginning in the late Cenozoic caused vegetation to transition from shrub to desert, and the flora composition also changed.

Fossil seeds of Euryale(Nymphaeaceae) indicate a lake or swamp environment in the late Miocene Zhaotong Basin of southwestern China

Euryale （Nymphaeaceae） is a monotypic genus distributed in eastern Asia, but fossils in its native distribu- tion are sparse as compared to Europe where the genus has disappeared. Here, we describe a new fossil species, Euryale yunnanensis sp. nov., from the late Miocene Zhaotong Basin in southwestern China on the basis of seed remains. Char- acteristics including an ellipsoidal to almost spherical shape, a smooth surface, a germination cap that covers the micro- pyle, the separation of the hilum and germination cap （mi- cropyle）, elliptic testa surface cells with undulate margins and thickened cell walls collectively indicate a close affinity with the genus Euryale in Nymphaeaceae. Comparisons of seed morphology and anatomy demonstrate that the newly described fossil species differs from modem and other fossil species of Euryale and thus verify the assignment to a new fossil species. As the modem Euryale is an aquatic plant, preferably living in swamps and lakes with shallow and stable water, we hypothesize that a lake and/or swamp environment with shallow water was present near Shui- tangba in the Zhaotong Basin, where the Shuitangba homi- noid lived during the late Miocene. This inference is generally consistent with paleoenvironmental data extracted from avian, fish, frog, turtle and crocodile fossils, as well as from aquatic pollen. The fossil history suggests a compara- tively recent （Miocene） emergence of Euryale, which is roughly in agreement with the divergence time inferred from the molecular information.

Fossil leaves of Berhamniphyllum(Rhamnaceae) from Markam,Tibet and their biogeographic implications

A new occurrence of buckthorn fossil leaves is reported from the upper Eocene strata of Markam Basin,southeastern Tibet,China.The leaf margin is either entire or slightly sinuous.Secondary veins are regularly spaced,forming eucamptodromous venation.These secondaries exist as straight lines from midvein to near margin and then arch abruptly upward and enter into a margin vein.The tertiary veins are densely spaced and parallel,and are percurrent to secondary veins.This leaf architecture conforms with Berhamniphyllum Jones and Dilcher,an extinct fossil genus reported from America.Our fossils are characterized by their dense secondaries,with secondary veins on the upper half portion of the blade accounting for over 40%of all secondaries.A new species,Berhamniphyllum junrongiae Z.K.Zhou,T.X.Wang et J.Huang sp.nov.,is proposed.Further analysis shows that confident assignment among Rhamnidium,Berchemia,and Karwinskia cannot be made based on leaf characters alone.Berhamniphyllum might represent an extinct common ancestor of these genera.In this study,several fossil Berchemia from Yunnan and Shandong are emended and reassigned to Berhamniphyllum.A new complex,namely the Berchemia Complex,is proposed based on morphology,molecular evidence,and the fossil record.This complex contains the fossil leaves of Rhamnidium,Karwinskia,Berchemia,and Berhamniphyllum.The historical biogeography of the Berchemia Complex is also discussed in this paper.This complex might have originated in the late Cretaceous in Colombia,South America,and dispersed to North America via Central America during the Eocene.Subsequently,the complex moved from North America to East Asia via the Bering Land Bridge no later than the late Eocene.Besides,the complex migrated from North America to Europe via the North Atlantic Land Bridge and then migrated further to Africa.In East Asia,it first appeared in Markam on the Qinghai-Tibetan Plateau,and then dispersed to other regions of Asia.

The Paleogene to Neogene climate evolution and driving factors on the Qinghai-Tibetan Plateau

The growth of the Qinghai-Tibetan Plateau(QTP)during the Cenozoic drove dramatic climate and environmental change in this region.However,there has been limited comprehensive research into evolution of climate during this interval.Here we present a quantitative reconstruction using Bioclimatic Analysis(BA)and Joint Probability Density Functions(JPDFs)based on data available for 48 fossil floras,including macrofossils and palynological fossils collected in the QTP area from the Paleogene to Neogene(66–2.58 Ma).Both methods indicate that there was an overall decline in temperature and precipitation.Paleoclimatic simulations using Hadley Centre Coupled Model version3(HadCM3)show that the most prominent climate change was very likely driven by QTP orographic evolution from the late Eocene,which was accompanied by a shift in temperature from a latitudinal distribution to a topographically controlled pattern.In addition,with the growth of the QTP,temperature and precipitation decreased gradually in the northeastern part of the plateau.Different sources of evidence,including plant fossil records,climate simulations and other proxies,indicate that the topographic evolution of the QTP and other geological events,in conjunction with global cooling,may have been the main factors driving climate change in this region.This research can provide insights into Cenozoic environmental change and ecosystem evolution.

The Tibetan Plateau is a natural laboratory for studying organic evolution and environmental change

Today’s fauna and flora are a continuation from their geological past. In order to better understand how patterns of biodiversity form and organic evolution takes place, it is necessary to study these flora and fauna over time.

Late Eocene sclerophyllous oak from Markam Basin, Tibet, and its biogeographic implications

Sclerophyllous evergreen broad-leaved forests,mainly made up of sclerophyllous oak,Quercus section Heterobalanus(Øerst.)Menitsky,Fagaceae,represent the most typical forest type in the Hengduan Mountains.Their distribution pattern is closely related to the growth and formation of the Qinghai-Tibetan Plateau(QTP).The oldest fossil record of Quercus sect.Heterobalanus so far discovered is from the middle Miocene of the Gazhacun Formation in Namling County,southern Tibet.However,our recent discovery of leaf fossils from the upper Eocene of Lawula Formation in Markam Basin,southeastern Tibet,illustrates that their origin is nearly 20 Myr older than previously assumed.By integrating the results from geometric morphometrics,geographical range expansion,and ecological niche shifts of this section in what is now the QTP and the Hengduan Mountains,we infer that the leaves of Quercus sect.Heterobalanus were already adapted to cool and dry conditions in some local regions no later than in the late Eocene.Then,with the growth of the QTP and late Cenozoic global cooling,the expansion of cooler and drier habitats benefited the spread and development of this section and their leaves exhibited morphological stasis through stabilizing selection.Based on published fossil records and recent discoveries,we argue that Quercus sect.Heterobalanus appeared in the subtropical evergreen and deciduous broad-leaved mixed forests of the southeastern margin of what is now the QTP no later than in the late Eocene.Some taxa spread westwards along the Gangdese Mountains and later the Himalaya,and others spread eastwards and southeastwards,gradually becoming a dominant group of species in the Hengduan Mountains.This dispersal route is contrary to the previous“northwards hypothesis”of this section,and further supports the hypothesis of an East Asian origin for Quercus section Ilex Loudon.

Evolution of stomatal and trichome density of the Quercus delavayi complex since the late Miocene

A fossil oak species,Quercus tenuipilosa Q.Hu et Z.K.Zhou,is reported from the upper Pliocene Ciying Formation in Kunming,Yunnan Province,southwestern China.The establishment of this species is based on detailed morphologic and cuticular investigations.The fossil leaves are elliptic,with serrate margins on the apical half.The primary venation is pinnate,and the major secondary venation is craspedodromous.The tertiary veins are opposite or alternate-opposite percurrent with two branches.The stomata are anomocytic,occurring only on the abaxial epidermis.The trichome bases are unicellular or multicellular.The new fossil species shows the closest affinity with theextant Q.delavayi and the late Miocene Q.praedelavayi Y.W.Xing et Z.K.Zhou from the Xiaolongtan Formation of the Yunnan Province.All three species share similar leaf morphology,but differ with respect to trichome base and stomatal densities.Q.tenuipilosa.Q.praedelavayi,and Q.delavayi can be considered to constitute the Q.delavayi complex.Since the late Miocene,a gradual reduction in trichome base density has occurred in this complex.This trend is the opposite of that of precipitation,indicating that increased trichome density is not an adaptation to dry environments.The stomatal density(SD)of the Q.delavayi complex was the highest during the late Miocene,declined in the late Pliocene,and then increased during the present epoch.These values show an inverse relationship with atmospheric CO_2 concentrations,suggesting that the SD of the Q.delavayi complex may be a useful proxy for reconstruction of paleo-CO_2 concentrations.

Words from the Editors

We are pleased to present the first issue of Plant Diversity,a new international Plant Science journal focusing on evolutionary processes that create and maintain plant diversity.We believe that a journal devoted to understanding how plant diversity evolves and is maintained is highly needed.This understanding is impossible to achieve without integrated knowledge of the evolutionary processes

A tribute to my supervisor Professor Zhengyi Wu

I was saddened and quite shocked when I received the phone called informing me that Professor Zhengyi Wu had died. It was the evening of June 20th, 2013. I had just got off a plane from Tibet and turned on my mobile phone. Prof. Wu had been unwell for neady a year. I knew that this would happen one day, but refused to believe it would really come to pass. A few days ago, some students and I were on a field trip in Tibet. Sitting in our jeep, I watched the mountains and roadside as we drove, and tried to identify the plants along the route. The mountain roads in Tibet seem to always make my students sleepy. I woke them up.