Cheng Quan

Chang'an University · School of Earth Science and Resources

East Asian Paleogene climates have long been regarded as controlled by the planetary wind system, which might result in a climate pattern with three latitudinally distributed zones. Two humid zones located separately in the north and south were lithologically designated by coals and oil shales, while an arid zone in the middle was represented by re...

The timing of the development of the East Asian monsoon in the geologic past is critically important for paleoclimatological studies, yet few quantitative data are available. Based on palynomorphs from six formations, supplemented by leaf fossils from one of these formations in Fushun, northeastern China, we present a quantitative estimate of the e...

Proxy-based quantitative estimates of Eocene climates can be made from marine isotope records for ocean conditions or fossil plants for terrestrial environment. However, our understanding about Eocene terrestrial climates is derived mainly from North America and Europe, and little is known about East Asia. Previous qualitative paleoclimate studies...

Continental Paleogene climates have been well studied in Europe and North America, but very little is known from Asia because paleoclimatic results have only been reported from particular geological intervals. Here, based on 29 plant assemblages from 8 well age-controlled fossiliferous sites, we quantitatively reconstruct the climates through most...

Fossil genera with similar features to the winged fruits of the living Engelhardia Lesch. ex Blume (e.g., Palaeocarya G. Saporta) have been widely reported in Cenozoic fossil floras of the Northern Hemisphere. However, fossil winged fruits of Engelhardia with detailed anatomical structures have only been found in the upper Eocene of North America....

A detailed uplift history of the Tibetan Plateau is essential for disentangling the proposed geodynamical models and assessing its impacts on climate and biodiversity. However, when and how the plateau formed remains highly controversial. Here, we present unusual geochemical indicators of marine signatures in the Cenozoic terrestrial strata of the...

The 'man fern' (Cibotium, Cibotiaceae), a typical tree fern of tropical and subtropical climates, is today mainly distributed in the Hawaiian Islands, Mesoamerica, and tropical and subtropical regions of East and Southeast Asia. Reliable fossil records of this genus are rare. Only two cases of stem fossils have been reported from the Upper Cretaceo...

Ceratophyllum L. is a cosmopolitan genus of perennial aquatic herbs that occur in quiet freshwaters. Fossils of this genus have been widely reported from the Northern Hemisphere, most of them occurring in the temperate zone. Here, we describe two species of fossil fruits discovered from subtropical areas of China. The fossil fruit discovered in the...

Both Trapa L. and the extinct Hemitrapa Miki are aquatic plants in the family Lythraceae, with abundant fossil records in Eurasia and North America in the Cenozoic. However, documented materials are mainly based on fruit and pollen grains without reliable leaf fossils. Here, we report fossil leaves, fruit, and roots of Trapa and fruit of Hemitrapa...

The Neogene environment and paleovegetation of today’s semi-arid and arid Central Asia remain elusive. Little is known about the effect of paleoclimatic change on the distribution and ecological response mechanisms of aquatic plants, especially on the Tibetan Plateau. Here, we report a new species of Nelumbo Adanson, including leaves, receptacles,...

Quercus is the largest genus within the Fagaceae and has a rich fossil record. Most of the fossil material is attributed to the subgenus Quercus based on leaves, pollen or rarely acorns and fruits. Fossil records of Q. section Cyclobalanopsis characterized by ring-cupped acorns are relatively few and especially those described based on fruits are s...

The spatio-temporal vegetation patterns of East Asia in the Eocene remain uncertain. In this present paper, we conduct numerical analyses of plant functional types (PFTs) for 80 Eocene floras, subdivided into three-time intervals: the early, middle, and late Eocene. Based on PFT diversity spectra, five groups are identified from cluster analysis an...

Fagaceae have a long fossil history that can be dated back to the Late Cretaceous. They are the key element of several distinct biomes found throughout the Northern Hemisphere; therefore, it is a prime candidate for modern biogeographic analysis. Castanopsis (Fagaceae) species are the dominant trees of subtropical evergreen broad-leaved forests and...

Knowledge of paleolake evolution is highly important for understanding the past hydroclimate regime on the Tibetan Plateau and associated forcing mechanisms. However, the hydrological history of paleolakes on the central plateau, the core region of the plateau, remains largely inconclusive. Here we present new biomarker records from lacustrine depo...

Co-evolutionary relationships of plants and fungi are of great importance for the phylogeny of both groups. Nyssa was widely distributed in the northern hemisphere during the Cenozoic. Extant species of Nyssa exhibit a disjunct distribution between eastern North America, Central America and East Asia. Here, a new species, Nyssa nanningensis Xu & Ji...

Plant–arthropod interaction offers insight into the evolution of terrestrial ecosystems in the geological past. In this study, leaf fossils with arthropod traces, collected from the Miocene Huaitoutala flora, Qaidam Basin, northern Tibetan Plateau, were studied to provide direct evidence of plant–arthropod interactions on the Tibetan Plateau. The r...

Mummified fossil woods of Liquidambar from the Miocene of the Guiping Basin of Guangxi and the upper Pleistocene sediments of the Maoming Basin of Guangdong, South China are recognized as the new species Liquidambar guipingensis sp. nov. and the extant species Liquidambar formosana Hance, respectively. The fossil wood of L. guipingensis shows the g...

A new species, Syzygium guipingensis sp. nov. (Myrtaceae), is described based on mummified fossil wood from the Miocene Erzitang Formation of Guiping Basin, Guangxi, South China. This species represents the most ancient reliable fossil record of the genus Syzygium in eastern Asia, showing the greatest similarity to the extant species S. buxifolium...

A new species, Tetradium nanningense sp. nov. (Rutaceae), is described on the basis of well-preserved mummified wood from the upper Oligocene Yongning Formation of Nanning Basin, Guangxi Province, South China. This species represents the most ancient fossil evidence of the genus Tetradium in Asia, the region of its modern distribution. Its occurren...

Melia L. is a small genus of only 2–3 species, which is native to Indo-Malesia, India, Pakistan and southern parts of tropical Africa. Fossil records of Melia are known from the early Miocene to the Pleistocene. Here we describe some mummified fossil endocarps of Melia from the upper part of the Yongning Formation (late Oligocene) in Nanning Basin,...

The late Oligocene is an important deep-time analog for understanding future changes in the strength of the East Asian monsoon: it represents a climate warmer than today, yet follows the nascent uplift of the Tibetan–Himalayan orogeny during the Eocene Epoch. Here we quantify monsoon strength based on new oxygen isotope measurements on cellulose (δ...

Fokienia A. Henry & H. H. Thomas is a monotypic genus of the Cupressoideae Rich. ex Sweet (Cupressaceae), native to subtropical evergreen mesophytic forests in South China, northern Laos and Vietnam. The fossil record of Fokienia is very scanty, with only one known occurrence of foliage in the Oligocene of Longjing, Jilin, China. Here we report the...

The onset, characteristics, and drivers of paleo-monsoon conditions in East Asia remain a topic of heated debate. Records from the Eocene suggest pronounced rainfall seasonality consistent with monsoon rainfall across China, likely driven by migrations of the Inter-Tropical Convergence Zone. Model simulations indicate that modern-like monsoon circu...

Knowledge about within-species variation in stomatal frequency with varying elevation at very high elevations is rare, which is crucial for us to understand how alpine plants are adapted to the extreme environment. Here, we focus on the variation in stomatal frequency in Kobresia royleana (Nees) Boeckeler (Cyperaceae, Cyperales) along two altitudin...

The genus Elaeocarpus contains approximately 360 species and occurs in mesic forest communities from India, through to China, Southeast Asia, New Guinea, Australia, and New Caledonia. Elaeocarpus fossils are best known from the Eocene to the Miocene of Australia and the late Pliocene–early Pleistocene of India but have not been documented from East...

Pollen micromorphology provides one of the only morphological features characterising the phylogenetic lineages in oaks; therefore it is of great potential in understanding oak diversity in the geological past. In the present study, scanning electron microscopy (SEM) observations were made on the late Neogene pollen flora recovered from the well-kn...

New Dacrycarpus megafossils from southern China provide solid evidence for long-distance dispersals from Australia to Asia during the Miocene. This trans-equatorial migration route long post-dates the India-Asia suture and seems to have resulted from the collision of the Australian Plate and Asian Plate after the late Oligocene. Dacrycarpus is unli...

The collision between India and Eurasia in the mid‐Palaeogene facilitated terrestrial floristic exchange. However, due to the complexity of this geological event and scarcity of fossil record, the plant migration patterns between the two plates are still highly debated. In this study, we focus on the Oligocene floristic exchange between India and P...

In this paper, we describe a new species Magnolia nanningensis sp. nov., based on exceptionally well‐preserved mummified fossil woods from the late Oligocene of the Nanning Basin, Guangxi, South China. The features of these woods indicate a close affinity to the section Michelia of the subgenus Yulania belonging to the genus Magnolia sensu lato (Ma...

Podocarpium (Fabaceae), characterized by single seeded pods and paripinnate compound leaves, is well known as an extinct legume genus having extensive fossil occurrences from the Miocene of Central Europe and eastern Asia. However, the Palaeogene unambiguous megafossils are quite scarce, especially the Oligocene record from eastern Asia, which heav...

New genus and species Litseoxylon nanningensis gen. et sp. nov. is described on the basis of well-preserved mummified fossil wood from the upper Oligocene Yongning Formation of the Nanning Basin, Guangxi, South China. Litseoxylon gen. nov. is convincingly placed into the Litsea complex, the core group of the tribe Laureae of the Lauraceae family, a...

Fissistigma nanningense sp. nov. (Annonaceae) is described based on well-preserved mummified leaves from the Oligocene Yongning Formation of Nanning, Guangxi, southern China. They are characterized by retuse apex, a swollen petiole, eucamptodromous to simple brochidodromous secondary veins, percurrent tertiary veins, an obvious sub-vascular bundle...

Three new fossil species, two attributed to the genus Castanopsis (C. nanningensis and C. guangxiensis) and one to the organ genus Lithocarpoxylon (L. nanningensis) are described on the basis of well-preserved mummified wood from the upper Oligocene of Yongning Formation in the Nanning Basin, Guangxi Province, South China. The two species of Castan...

Stephania Loureiro is a large genus within Menispermaceae, with approximately 60 extant species naturally distributed in (sub)tropical areas in Asia, Africa, and a few in Oceania. This genus possesses highly characteristic endocarps that facilitate identification of extant and fossil specimens. Here, we report some well-preserved fossil fruits of S...

The genus Canarium contains approximately 78 species distributed in the Old World tropics and subtropics. Canarium is characterized by a distinctive drupaceous fruit with a trilocular endocarp derived from three fused pyrenes. Here, we describe new Canarium fossil fruits from the late Oligocene of the Yongning Formation, the Miocene of the Erzitang...

The genus Schima includes about 20 species and is distributed only in southern China and adjacent areas of Asia. The previous molecular phylogenetic analysis suggested Schima is in the tribe Gordoniae, along with Gordonia and Franklinia. However, because few fossils have been reported, the biogeographic origin of Schima is still poorly known. In th...

Extant species of Choerospondias (Anacardiaceae) are mainly distributed in the tropical and subtropical areas of East Asia. However, the fossil record indicates that the genus was more widespread in the past, with Cenozoic occurrences in both Europe and Asia. We investigated the pericarp anatomy of well-preserved mummified fossil fruits of Choerosp...

A new species of fossil conifer wood, Podocarpoxylon donghuaiense sp. nov., is described from the late Eocene of Nadu Formation in Baise Basin of the Guangxi Province, South China. This fossil wood is characterized by distinct growth rings, circular to oval tracheids in cross section, 1–2-seriate opposite pits on radial tracheid walls, uniseriate (...

A middle Pleistocene flora from a low latitude of southern China was paleoclimatically studied through a quantitative reconstruction. The fossiliferous layer is correlated to the fourth terrace of the local Youjiang River and its age is revised to be the middle Pleistocene based on the presence of Paleolithic stones in the fourth terrace. The Coexi...

A new species Camellia nanningensis was described on the basis of well-preserved mummified wood from the upper Oligocene Yongning Formation of Nanning Basin in Guangxi Province, South China. This represents the most ancient fossil wood assigned to Camellia, and the earliest fossil record of the family Theaceae in China. This fossil material shows t...

The Paleogene sedimentary records from southern China hold important clues to the impacts of the Cenozoic climate changes on low latitudes. However, although there are extensive Paleogene terrestrial archives and some contain abundant fossils in this region, few are accurately dated or have a temporal resolution adequate to decipher climate changes...

The Paleogene sedimentary records from southern China hold important clues to the impacts of the Cenozoic climate changes on low-latitudes. However, although there are extensive Paleogene terrestrial archives and some contain abundant fossils in this region, few are accurately dated and have a temporal resolution adequate to decipher climate change...

The occurrences of Staphylea L. (Staphyleaceae) fossils have been abundantly documented from the Cenozoic of Eurasia, but none has been confirmed from North America to date. In this study, we describe Staphylea levisemia sp. nov. on the basis of seed remains from the latest Miocene to earliest Pliocene of northeastern Tennessee, southeastern USA. T...

Cenozoic climatic and environmental changes in the arid Asian interior, and their possible relations with global climatic changes and the Tibetan Plateau uplift, have been intensively investigated and debated over past decades. Here we present 40-Myr (million years)-long n-alkane records from a continuous Cenozoic sediment sequence in the Dahonggou...

The modern European climatic regime is peculiar, due to its unitary winter but diverse summer climates and a pronounced Mediterranean climate in the south. However, little is known on its evolution in the deep time. Here we reconstruct the European summer climate conditions in the Tortonian (11.62-7.246 Ma) using plant fossil assemblages from 75 we...

A new cycad species, Cycas fushunensis sp. nov., is described from the Lutetian Jijuntun Formation at Fushun Coalmine, Liaoning Province, northeast China, based on a well-preserved partial frond containing about 15 leaflets. The fossil is characterized by a single strong vein per leaflet, decurrent leaflet base and haplocheilic stomata, suggesting...

Mammalian bipedalism has long been thought to have arisen in response to arid and open environments. Here we tested whether bipedalism coevolved with environmental changes using molecular and paleontological data from the rodent superfamily Dipodoidea and statistical methods for reconstructing ancestral characteristics and past climates. Our result...

In climatology, most variables are unequivocally defined and quantitatively described. These variables, such as temperature and precipitation, can also be quantified from geological proxies in paleoclimate studies. However, there are other climatic factors that, although strongly impacting ecosystems, have been ambiguously delineated and consequent...

Proxy-based quantitative estimates of Eocene climate conditions are abundantly available from marine isotope records and floral data. However, the available terrestrial data are mainly from North America and Europe, and only a few are known from East Asia. Previous qualitative studies on Chinese Eocene floras briefly illustrated the zonation of the...

The evolutionary history of Ginkgo is poorly understood for the Cenozoic Era because of the rarity of fossil reproductive organs. We here describe a new species, Ginkgo cranei sp. nov., on the basis of well-preserved ovulate organs and associated leaves from the Upper Paleocene Sentinel Butte Formation of North Dakota, USA. The ovulate organ is of...

Quantitative reconstructions of atmospheric CO2 by using terrestrial and marine records are critical for understanding the so-called “greenhouse” conditions in the Cretaceous, but data from terrestrial plants for several stages of this period remain quite limited. Using the stomatal index (SI) technique, here we estimate the Santonian (Late Cretace...

Three taxa of Late Cretaceous aquatic angiosperms, Queruexia angulata (Lesq.) Krysht, Cobbania corrugata (Lesq.) Stockey et al. and Nelumbites cf. extenuinervis Upchurch et al. from Jiayin of Heilongjiang, NE China, are described in detail. Among them, Cobbania and Nelumbites from the Upper Cretaceous in China are reported for the first time. The a...

Based on the gross morphology of leaves and fertile organs, including in situ spores, three species of Asplenium Linnaeus (Asplenium changcaium Sun et Liu, Asplenium popovii Samylina and Asplenium tiefanum Deng) have been recognized for the first time in the Lower Cretaceous Changcai Formation in the Helong area (Yanbian), eastern Jilin Province, C...

The living fossil Ginkgo, a relict of a once dominant gymnosperm, existed and flourished early in the Mesozoic, but only a vague outline of its evolutionary history in the Tertiary has been unveiled. Here we describe a new species, Ginkgo jiayinensis sp. nov., from the Wuyun Formation of Jiayin, China, which is another well-established Tertiary spe...

Using the stomatal index technique, the Campanian (Late Cretaceous) atmospheric carbon dioxide levels are estimated based on a sequence of fossil cuticles of Ginkgo adiantoides (Ung.) Heer. Ginkgo cuticles were sampled from 11 beds in the Taipinglinchang Formation at Jiayin, Heilongjiang Province, northeast China. In general, the regression functio...

Laic Crelaceous planlS were mainly colieckJ from the Upper Cretaceous Yong'aneun Formation and Talpingliochang FormalJoo In .lJaym area along Ihe HeilongJIang (Amur) River, Chioa. The comblrled megafloras conSISI of 34 species represenllng 26 gencra and Include horselalis (Cquisewm), fellls (Asple­ nium), ginkgoales (ClI1kgo), conifers (Cupresslf1o...