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–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.     

云南中新世杉科化石木研究

利用光学显微镜和扫描电镜首次对云南省楚雄州晚中新世石灰坝组石鼓村层的钙化木材进行了解剖学研究.鉴定出两种类型的木材:柳杉型落羽杉型木(Taxodioxylon cryptomeripsoides Schonfeld)和杉木型落羽杉型木(Taxodioxylon cunninghamioides Watari).二者分别与现代柳杉属和杉木属具有最接近的亲缘关系.根据这两种杉科化石木现存最近亲缘种的生态环境,并综合其他资料,推测该地区在晚中新世为温暖湿润的亚热带气候环境.     

Occurrence of Christella (Thelypteridaceae) in Southwest China and its indications of the paleoenvironment of the Qinghai–Tibetan Plateau and adjacent areas

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.     

红杉属植物在中国云南中新世首次发现

化石标本采自中国云南省中部楚雄市吕合煤矿的石鼓村层 ,该层为灰色至浅灰色细砂岩至粗砂岩、粉砂质粘土层和褐煤层 ,地质时代属中新世 ① 。石鼓村层的孢粉组合以被子植物占绝对优势(90 % ) ,其中热带、亚热带植物成分类型较多 ,如金缕梅科、棕榈科等 ,温带植物成分类型较少 ,如榆科、桦木科等 ;裸子植物花粉占整个孢粉组合的 9% ,主要是松科和杉科花粉 ;蕨类植物孢子仅占整个孢粉组合的 1% ,以水龙骨单缝孢为主。本次发现的标本小枝长约 2 .8ｃｍ ,宽约 1.6ｃｍ ,小枝基部稍膨大。叶螺旋状排列 ,基部扭转 ,在同一平面排成 2列 ,以 30～ 70…     

Late Miocene Palaeocarya (Engelhardieae: Juglandaceae) from Southwest China and its biogeographic implications

Fossil fruits of Palaeocarya (Juglandaceae) are described from late Miocene sediments of southeastern Yunnan, China. The fruits present a tri-lobed wing consisting of an intact oblong-ovate middle lobe and two lateral lobes. The lobes are apically obovate, and have pinnate venation. The middle lobe is thicker at the base and gradually tapers to the apex. The nutlet, located at the base of the winged fruit, is round and hispid, and is subdivided by a septum into two compartments. Based on extensive morphological comparisons to previously documented fossil fruits, we found that the fossil fruits align most closely with members of the genusPalaeocarya, but have a unique combination of characters. Thus, we describe the fossils as a new species, Palaeocarya hispida sp. nov. This species represents an important range expansion for low-latitude occurrences of Palaeocarya in the late Miocene and therefore substantially improves our understanding of the biogeographic history of the genus. We propose that the wide distribution of Palaeocarya and relatively narrow distributions of close relatives, Engelhardia, Alfaropsis, and Oreomunnea, might be associated with a stepwise cooling and a major ice sheet expansion in the Antarctic and Arctic from the late middle Miocene to early Pliocene. In particular, the climatic oscillations during the Quaternary, such as the last glacial maximum, might have led to a decrease in the geographic distribution of Engelhardieae.     

Early Oligocene fruits and leaves of Burretiodendron (Malvaceae s.l.) from South China

The genus Burretiodendron Rehder is currently endemic to an area near the China–Vietnam border and the limestone mountains of Thailand and Myanmar. The fossil records of this genus were previously found only from the Miocene of Yunnan, Southwest China, and the Oligocene of Guangxi, South China. Here, we describe fossil fruits and associated leaves of Burretiodendron, which were discovered in the lower Oligocene Shangcun Formation of the Maoming Basin, Guangdong, South China. Morphological comparison with extant and fossil Burretiodendron taxa indicates that fruit fossils belong to the species Burretiodendron parvifructum J. Lebreton Anberrée & Z. K. Zhou. This is one of the earliest fossil records of the genus, providing additional evidence for the early biogeographic history of this genus and supporting the inference that the genus originated in South China. According to the habitat conditions of modern species, we speculate that there were limestone mountains around the Maoming Basin in the early Oligocene.     

A Miocene leaf fossil record of Rosa (R. fortuita n. sp.) from its modern diversity center in SW China

Roses (Rosa, Rosaceae) are arguably the most admired ornamental plants in the world. Southwestern China is the center of diversity for many extant native species of Rosa and fossils found in this region are critical for understanding the evolution of this genus. Herein, we report a leaf fossil record with good preservation from the late Miocene of Yunnan Province. The opposite and odd-pinnate leaf is composed of seven elliptical leaflets, with close, crenulate, and regular marginal teeth. The stipules are lanceolate and adnate to the petiole. Additionally, the secondary veins are semicraspedodromous, showing the same venation pattern as most living Rosa species in southwestern China. On the basis of the extensive morphological comparisons, we propose a new species, R. fortuita T. Su et Z.K. Zhou n. sp. This is the first confirmed Rosa leaf fossil record in China, and the discovery of R. fortuita n. sp. indicates that Rosa existed in southwestern China by the late Miocene. It suggests that Rosa was distributed in subtropical or temperate forests and shared a similar ecological niche as Rosa in Europe during the Oligocene and Miocene. The modern diversification of Rosa in southwestern China is thought to have been closely associated with the continuous uplift of the Qinghai-Tibet Plateau since the late Miocene, creating complex topography and a variety of climate conditions.     

First fossils of Zygogynum from the Middle Miocene of Central Yunnan,Southwest China,and their palaeobiogeographic significance

The collision of Gondwana and Laurasia effected plant exchanges between the two continents and thus affected the evolution of the flora of Yunnan, Southwest China. Consequently, the plant diversity in Yunnan has been enriched such that it has attracted the attention of botanical researchers. In this paper, we describe winteraceous fossil leaves collected from the Middle Miocene in Central Yunnan. Compared with the leaves of today’s plants, they share cuticular features and leaf morphological characters with Zygogynum Baillon (Winteraceae); the fossil plant is designated as a new species, Zygogynum poratus n. sp. Liang and Zhou. Based on its historical and recent distribution, the genus probably had migrated into Yunnan from Gondwana during or before the Middle Miocene, spreading beyond its current geographic range.     

Pliocene white pine (Pinus subgenus Strobus) needles from western Yunnan,southwestern China

Abstract

The fossil records of Pinus L. are abundant since the Late Cretaceous, especially for the subgenus Pinus L. (the hard pines). However, those of the subgenus Strobus (D. Don) Lemmon (the white or soft pines) are not well documented. In this study, we describe a new species of white pines, Pinus plioarmandii sp. nov., from the Pliocene of western Yunnan, southwestern China. This species mostly resembles extant P. armandii Franch. in gross morphology and cuticular structure of needles. Molecular dating showed that the Asian white pines split into two lineages around the Late Miocene, and the Pinus armandii clade diverged at the Early Pliocene. The present fossil occurrence indicates that Pinus armandii might originate in southwest China and probably in the western Yunnan. The glacial events during the Pleistocene might be the major factors for the retreat of the white pines from western Yunnan, and the rapid uplift of the Yunnan Plateau and deep incision of the river valleys since the Pliocene had posed a significant geographic barrier for their reconnection in the subsequent warm climate condition.     

古猿闭壳龟的新材料及该属的地史分布

云南禄丰古猿产地发现的古猿闭壳龟(Cuora pitheca Yeh,1981),是闭壳龟属的首次化石记录。它的发现,把该属的历史从现代一下追溯至中新世晚期。本文除补充记述古猿闭壳龟的新材料外,并结合近年国内外的新发现,综述了该属龟类的地史分布。     

云南开远小龙潭动物群的性质及时代的进一步探讨

本文在前人工作的基础上,结合新材料,对小龙潭动物群作了新的补充与论述.小龙潭动物群在组合面貌上与西瓦立克的 Chinji, Nagri 层动物群相近,动物群的时代不早于 Chinji 层而早于云南禄丰石灰坝动物群,为晚中新世.     

Dental evolution of the orang-utan in China

The main focus of this paper is to report the spatial and temporal distributions of fossil and subfossil orang-utans in south and southwest China from the late Miocene through Pliocene and Pleistocene. We will describe fossil samples and their odontometrics in order to estimate their body size through time. Furthermore we will also discuss their taxonomic status and evolutionary changes. Then, we will explore the possible reasons why and how these living fossils retreated to their present habitats.     

The microsoroid ferns: Inferring the relationships of a highly diverse lineage of Paleotropical epiphytic ferns (Polypodiaceae, Polypodiopsida)

The relationships of the microsoroid ferns were studied using a DNA sequence-based phylogenetic approach. Nucleotide sequences for up to four chloroplast genome regions were assembled for 107 samples from 87 species. Microsoroids s.l. include six lineages of which two are species rich. The results indicate that several genera are not monophyletic (e.g. Microsorum), several controversial genera are confirmed to be monophyletic (e.g. Leptochilus), and some genera new to science should be recognized (M. membranaceum clade). Unique insights were gained into the biogeographic history of this highly diverse epiphytic vascular plant lineage that is widespread in continental Asia to Australasia. Evidence was found for splits into lineages diversifying in parallel in continental Asia and Malesia. No evidence was recovered for an African radiation because all African microsoroid species either also are found in Asia or have sister species in continental Asia. In contrast, evidence for independent radiations were discovered for the Australasian region.     

福建中中新世紫萁属(紫萁科)的一个新种*

丰富的化石记录显示中生代和新生代紫萁属在北半球广泛分布.本文描述了发现于福建中中新世佛昙群的紫萁属一新种:漳浦紫萁(Osmunda zhangpuensis Z.X.Wang and B.N.Sun,sp.nov.).化石小羽片保存完好,呈线状披针形,边缘具齿,羽状脉序.在该属现生种中,当前化石种与现生种Osmunda banksiifolia在形态上最相似.当前化石的发现表明,早在中中新世紫萁属就已生长在福建省.漳浦紫萁和之前报道过的附生真菌化石以及龙脑香科、桑科、豆科、荨麻科、胡桐科、大戟科、樟科等植物的存在,表明福建在中中新世时期的气候温暖湿润.     

The diversity and paleoenvironmental significance of Calophyllum (Clusiaceae) from the Miocene of southeastern China

Abstract

Three species within the genus Calophyllum collected from middle Miocene Fotan Group sediments in Zhangpu County, Fujian, southeastern China are described in this paper. These fossils include Calophyllum zhangpuensis sp. nov., Calophyllum striatum, and Calophyllum suraikholaensis. The new fossil species C. zhangpuensis sp. nov. is oval, possesses entire leaves with closely spaced parallel secondary veins and has a round, or slightly retuse, apex. These specimens represent the first known fossil records of this relative wide leaf-type form of Calophyllum from China and have a length:width (L:W) ratio less than 3:1. In combination with the known modern geographic distribution and habitats of this wide leaf-type Calophyllum and other plants, data suggest that the middle Miocene Fotan flora is indicative of a warm climate. Thus, based on available fossil data, we speculate that this genus probably originated in India during the Paleocene before spreading from India to Bangladesh and into China, Sumatra, Malaysia, Indonesia, and Java during the Neogene, leading to its modern distribution. At least, the 3 fossil species in this region can explain floristic exchange between India, Fujian, and South China, which is consistent with previous studies; the occurrence of these 3 species indicates that Calophyllum began to diversity in China no later than the Miocene.     

First middle Miocene sivaladapid primate from Thailand

Sivaladapids are a group of Asian adapiform primates that were previously documented from deposits dating to the middle Eocene through the late Miocene in Pakistan, India, Myanmar, Thailand, and China. The group is notable for the persistence of three genera, Sivaladapis, Indraloris and Sinoadapis, into the late Miocene. In Thailand, sivaladapids were previously documented only from late Eocene deposits of the Krabi mine. Here, we describe the first Southeast Asian Miocene sivaladapid, Siamoadapis maemohensis gen. et sp. nov. from a 13.3 to 13.1 Ma lignite layer from the Mae Moh coal mine, Thailand. It differs from other Miocene sivaladapids by its distinctly smaller size and in features of the dentition. This discovery enhances the paleoecological diversity of the middle Miocene primate fauna of Thailand, which now includes sivaladapids, a loris, tarsiids, and hominoids. In this respect, the fossil primate community from the middle Miocene of Thailand is similar in its composition to roughly contemporaneous assemblages from southern China, India, and Pakistan. However, the Thai fossils represent a distinct genus, suggesting a different biogeographic province with distinctive paleoenvironments.     

Terpenoid Compositions and Botanical Origins of Late Cretaceous and Miocene Amber from China

The terpenoid compositions of the Late Cretaceous Xixia amber from Central China and the middle Miocene Zhangpu amber from Southeast China were analyzed by gas chromatography-mass spectrometry (GC-MS) to elucidate their botanical origins. The Xixia amber is characterized by sesquiterpenoids, abietane and phyllocladane type diterpenoids, but lacks phenolic abietanes and labdane derivatives. The molecular compositions indicate that the Xixia amber is most likely contributed by the conifer family Araucariaceae, which is today distributed primarily in the Southern Hemisphere, but widely occurred in the Northern Hemisphere during the Mesozoic according to paleobotanical evidence. The middle Miocene Zhangpu amber is characterized by amyrin and amyrone-based triterpenoids and cadalene-based sesquiterpenoids. It is considered derived from the tropical angiosperm family Dipterocarpaceae based on these compounds and the co-occurring fossil winged fruits of the family in Zhangpu. This provides new evidence for the occurrence of a dipterocarp forest in the middle Miocene of Southeast China. It is the first detailed biomarker study for amber from East Asia.     

Samaras of Ventilago (Rhamnaceae) from the upper Miocene of Lincang,Yunnan, China and their phytogeographic implications

The genus Ventilago Gaertn. usually occurs in hot, dry, and low-altitude tropical environments. The fossil record of this genus is meager, so its origin and geographic history are still poorly understood. We describe a fossil winged fruit species-Ventilago lincangensis K. N. Liu & S. P. Xie sp. nov. from the upper Miocene Bangmai Formation of Lincang city, Yunnan Province, China. The fossil conforms to the modern genus Ventilago in architecture of the prominent fruit wing including a prominent midvein, a set of minor longitudinally secondary veins extending the full length of the fruit and reticulate higher order venation. The elliptical fruit body at the base of the wing shows an equatorial rim and has a short pedicel. This is the first report of Ventilago fruits in China, and is significant for the investigation of the origin and evolutionary history of this genus.