New Eocene fossil fruits and leaves of Menispermaceae from the central Tibetan Plateau and their biogeographic implications

Menispermaceae are a pantropical and temperate family with an extensive fossil record during the Paleogene period, especially in North America and Europe, but with much less evidence from Asia. The latest fossil evidence indicates a succession of tropical to subtropical flora on the central Tibetan Plateau during the Paleogene. However, the biogeographic histories of these floras are still unresolved. Here, we report on endocarps and leaves of Menispermaceae from the Middle Eocene of Jianglang village, Bangor County, central Tibetan Plateau. The endocarps belong to two genera: Stephania, which is characterized by a horseshoe-shaped endocarp and with one lateral crest ornamented by spiny to rectangular ribs, and a condyle area; and Cissampelos (s.l.), which has two characteristic lateral ridges and a conspicuous external condyle. Associated leaves belong to the genus Menispermites, and are characterized by actinodromous primary venation, brochidodromous secondary veins, entire margins, and the presence of marginal secondary veins. The biogeographic history of Menispermaceae is complex, but evidence from these new fossils indicates an early diversification of the group in Asia, probably in response to the warming climate during the Eocene. The Jianglang flora appears to be part of a boreotropical flora, connecting Asia with North American and European floras during the Middle Eocene. The modern distribution of menispermaceous taxa found in Jianglang, as well as other families represented in the Jianglang flora, show that a tropical to subtropical climate occurred during the Eocene in central Tibet.     

Fruits of Firmiana and Craigia (Malvaceae) from the Eocene of the Central Tibetan Plateau with emphasis on biogeographic history

The fossil record evidences an old origin and diversification of Malvaceae in the Northern Hemisphere. The central Tibetan Plateau was at a low elevation with a monsoon influence during the Eocene, allowing the development of a subtropical flora containing Malvaceae. The taxonomic study of fossils from the Eocene of what is now the Tibetan Plateau is still ongoing. Malvaceae fossils from the Eocene Jianglang flora, are attributed to sub-families Tilioideae and Sterculioideae, and are compared with modern species. A new specimen of Firmiana is described based on a fruit valve with a pinnate venation, the secondary veins starting at the ventral suture and reaching the midvein, and the seeds attached at the proximal part of the ventral suture. This specimen represents the earliest known occurrence of the genus. A new occurrence of Craigia is also reported based on detached membranous valves of a fruit capsule with a prominent fusiform locular area and radiating venation. Based on the fossil record of Firmiana and its modern distribution, we infer that the genus may have originated in East Asia and subsequently diversified in South China and Southeast Asia. The new occurrence of Craigia indicates that the genus was distributed in humid areas in South, Southwest and North China during the Eocene. Both fossil records evidence the important role that the Tibetan region played in the diversification of plants in East and Southeast Asia.     

Fossil fruits of Illigera (Hernandiaceae) from the Eocene of central Tibetan Plateau

Illigera (Hernandiaceae) is a liana genus distributed mainly in the tropical Asia and Africa. Previous fossil records suggested that Illigera was restricted in western North America during the Eocene. Recent paleobotanical investigation has unveiled a Paleogene flora that is totally different from today's vegetation in central Tibet. This provides novel insights into the paleoenvironmental change during the evolution of the Tibetan Plateau (TP). Here, we investigated 10 fruit impressions of Illigera from the early middle Eocene Niubao Formation in the Bangor Basin, central TP. The fossil winged fruits are characterized by their eroded fruit wings, well preserved fusiform locular areas with a median ridge bisecting the fruit, and short veins fanning radially outward. These features allow us to assign these fossils to Illigera eocenica, a species originally discovered in the Eocene of western North America. This is the second fossil occurrence of Illigera worldwide, and the first in Asia. Our finding suggests a warm and humid climate in the central TP during the early middle Eocene, and a close floristic link between Asia and North America during the Paleogene. We also propose a Northern Hemisphere origin and a Paleogene dispersal event from Northern Hemisphere to Africa for Illigera.     

Neogene Corylopsis seeds from eastern Tennessee

A new fossil species of Corylopsis (Hamamelidaceae), C. grisea Quirk & Hermsen sp. nov, based on seeds from the early Pliocene Gray Fossil Site (GFS), eastern Tennessee, USA, is described. The assignment of the seeds to Hamamelidaceae, subfamily Hamamelidoideae, is based on the overall size of the seeds, smooth testa, lack of a seed wing, and the presence of a terminal hilar scar. The assignment to the genus Corylopsis is based on seed size as well as the presence of a hilar facet, in addition to the hilar scar. Although Corylopsis persists only in East Asia today, its fossil record indicates that the genus was widespread across the Northern Hemisphere in the past. Prior to its discovery at GFS, Corylopsis was only known from the Paleogene in North America. The presence of C. grisea at GFS extends the fossil record of Corylopsis in North America to the Neogene and reinforces the interpretation of GFS as a forested refugium that provided a relatively moist, equable environment where subtropical to warm temperate plants could persist during a time of cooling and drying in the continental interior of North America. Its presence provides additional evidence for the biogeographic connection between the GFS paleoflora and the modern flora of eastern Asia.     

Fruits of icacinaceae (tribe iodeae) from the late paleocene of Western north america

The Icacinaceae occur pantropically today, but are well represented by fossil fruits of the warm Early Middle Eocene, when tropical plants that currently occupy low latitudes were more widely distributed in higher latitudes. Members of this family are first known in the Late Cretaceous; however, fossil fruits of tribe Iodeae are quite rare before the Eocene. In this paper we describe the first formally recognized Late Paleocene icacinaceous taxa from western North America. We name two new species of Icacinicarya based on anatomically preserved fruits and establish a new genus, Icacinicaryites, for impressions with a strong similarity to Icacinicarya that lack anatomical preservation. These new records from the Almont/Beicegel Creek flora in North Dakota and several localities in Wyoming, Colorado, and Montana complement records known from the Early Eocene of England and document an increased diversity of Iodeae and related forms in the Paleogene of western North America.     

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

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.     

山东植物区系的演变和来源

1　现代植物区系山东省位于我国东部、黄河下游 ,北濒渤海 ,东临黄海 ,地理范围介于北纬 34°2 3′～38°2 4′,东经 1 1 4°4 8′～ 1 2 2°4 3′之间。全省总面积为 1 5.72万 km2 ,占全国总面积的 1 .6%。属暖温带季风气候 ,沿海比较湿润 ,地带性植被主要是暖温带落叶阔叶林和松、栎类针阔叶混交林。山东省在中国植物区系的分区地位隶属于泛北极植物区、中国 -日本森林植物亚区、华北植物地区 [1 ]。据最近研究统计 ,现有野生维管植物 1 47科、61 4属 ,约 1 547种 (包括变种 ,下同 )。其中蕨类植物 2 4科 39属 1 0 5种 ,裸子植物 3科 3属 …     

Northern Hemisphere megafossil of Dacrycarpus (Podocarpaceae) from the Miocene of South China and its evolutionary and paleoecological implications

The modern genus Dacrycarpus (Endl.) de Laub. of the family Podocarpaceae, containing nine species, is mainly distributed in tropical mountain rainforests of the southwestern Pacific region, ranging from New Zealand to low‐latitude Asia. This genus has abundant fossil records in both hemispheres, but all the known megafossils were limited to Australasia and South America. Here we report on Dacrycarpus guipingensis sp. nov. from the Miocene Erzitang Formation of Guangxi, South China. This is the first megafossil of Dacrycarpus in the Northern Hemisphere. The new species is represented by mummified dimorphic foliage, ovuliferous shoots, and a male cone with in situ pollen. It resembles the extant Dacrycarpus imbricatus (Blume) de Laub., which is common in rainforests from southern China and northern Myanmar to Fiji. This paper presents the first data on the anatomical structure of seed cone and exine ultrastructure of Dacrycarpus in situ pollen grains from a fossil material by using computed tomography scanning and ultrathin sectioning. For comparative purpose, data on the pollen morphology and ultrastructure were obtained for modern D. imbricatus for the first time. The D. guipingensis fossils strongly suggest the Miocene arrival of Dacrycarpus in Asia from the Southern Hemisphere. Based on the modern ecological niche and related fossil elements, this fossil locality was probably covered by conifer–broad‐leaved mountain rainforests during the Miocene.     

Mummified Magnoliaceae woods from the upper Oligocene of South China,with biogeography,paleoecology, and wood trait evolution implications

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 (Magnoliaceae). Magnolia nanningensis is the first fossil record of the section Michelia from China, the modern diversity center of this group. These mummified woods provide fossil evidence supporting molecular dating that estimated an Oligocene age for divergence of the tropical evergreen section Michelia and the temperate deciduous section Yulania. Helical thickenings on vessel walls and a high degree of vessel grouping found in these fossil woods could be adaptive to temporary, possibly seasonal, droughts and, as suggested by other woods from the Nanning Basin, could be indicative of a monsoon‐influenced tropical climate in Guangxi during the late Oligocene. Helical thickenings have not been reported in magnoliaceous fossil woods prior to the Oligocene. The appearance of this trait was presumably a response to abrupt climate cooling near the Eocene–Oligocene boundary, followed by increase in climate seasonality. The associated increase of latitudinal zonation might be a possible trigger for divergence between the tropical evergreen sect. Michelia and the temperate deciduous sect. Yulania.     

A unique record of Cercis from the late early Miocene of interior Asia and its significance for paleoenvironments and paleophytogeography

The climatic impacts of the Tibetan Plateau since the Neogene and the phytogeographic pattern changes of formerly widely-distributed forest communities on the plateau remain poorly constrained. Today, Cercis L. (Fabaceae) is a well-known arborescent genus typically distributed in subtropical to warm temperate zones of the Northern Hemisphere, and Paleogene fossil occurrences from Eurasia and North America show a long history of the genus in mid-low latitudes of the Northern Hemisphere. Here, we describe a fossil species, Cercis zekuensis sp. nov. based on well-preserved fruits from the early Miocene of the northeastern Tibetan Plateau. Detailed morphological comparison (e.g., ventral margin with a veinless wing) of extant and fossil members of Cercis and other genera confirmed validity of the present taxonomic identity. Based on the comparison with extant relatives and their climate preferences, this unexpected occurrence of thermophilic Cercis in northeastern Tibetan Plateau indicates this area had higher temperature and precipitation in the Miocene than today. Integrated with inferred (paleo-)temperature lapse rates, this indicates a low paleoelevation of less than 2.4 km. In contrast with the present-day alpine climate here (~3.7 km), such a low elevation facilitated a more favorable habitat with comparatively high biodiversity and warm temperate forests at that time, as were evidenced by co-occurring megafossils. Moreover, the present existence of Cercis implies the genus was widespread in interior Asia during the early Neogene and shows its modern disjunction or diversification between eastern and central Asia was possibly shaped by the late Cenozoic regional tectonic uplift and consequential environmental deterioration.     

青藏高原杨属化石概述

杨属是北半球温带森林生态系统中一个重要的乔木属。文中概述了青藏高原古近纪和新近纪的杨属化石记录,并根据最新的地层学和年代学数据,对化石的时代进行厘定。化石记录显示杨属在青藏高原最早出现于始新世最晚期。在渐新世和中新世杨属在青藏高原的南部和北部都有较多的化石,但在高原中部尚未见记录。青藏高原具有杨属的化石植物群大多数都是温带落叶阔叶的河岸植被,进一步证实杨属在其演化历史早期就偏好温带近河岸的环境。青藏高原北部的早渐新世植物群中杨属多样性较高,并占据显著优势,表明这一地区在杨属早期分化中具有重要作用。     

Extinct genus Lagokarpos reveals a biogeographic connection between Tibet and other regions in the Northern Hemisphere during the Paleogene

The biodiversity history on the Qinghai–Tibetan Plateau (QTP) has remained unclear for a long time. Recent paleobotanical investigations provide opportunities for revealing the phytogeographic history on the QTP in deep time and for testing phylogeographic hypotheses. Lagokarpos is an extinct genus with unknown modern affinities. It is easily distinguished by its fruits, which have two long wings and an elliptical fruit body. Previously, Lagokarpos fossils have only been found from North America and Germany, ranging from the latest Paleogene to early middle Eocene. Recently, we found fossil fruits of this genus from the Niubao Formation, near Bangoin County in the central QTP. A new species, namely Lagokarpos tibetensis H. Tang, T. Su & Z. K. Zhou sp. nov., is described. This is the first fossil record of the genus in Asia. The occurrence of L. tibetensis indicates a close floristic linkage between the QTP and other floras in the Northern Hemisphere during the Paleogene. According to the floristic assemblages, we suggest the central QTP experienced a tropical or subtropical humid climate during that period.     

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.     

First fossil fruits of Elaeocarpus (Elaeocarpaceae) in East Asia: Implications for phytogeography and paleoecology

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 Asia before. Here we describe six new species of Elaeocarpus, E. nanningensis sp. nov. from the late Oligocene Yongning Formation of the Nanning Basin, E. presikkimensis sp. nov. from the Miocene Erzitang Formation of the Guiping Basin, E. prerugosus sp. nov., E. prelacunosus sp. nov., E. preserratus sp. nov., and E. preprunifolioides sp. nov. from the late Miocene Foluo Formation of the Nankang Basin in Guangxi, South China. This is the first reliable report for the genus occurring in East Asia, and the fossils indicate that Elaeocarpus had colonized this region by the late Oligocene and represented by a morphologically diverse group of species by the late Miocene. This sheds new insights into the timing and migration patterns of the genus in East Asia. Elaeocarpus is typically a rainforest genus occurring in mesic forests. Based on the habitat of their morphologically similar modern relatives we propose that these three sedimentary basins were warm and wet adjacent to mountainous regions with the evergreen or rain forests during the late Oligocene to Miocene.     

云南石林地区石灰岩山地种子植物区系成分的研究

云南石林是我国为数不多的世界地质公园。分析了云南石林地区石灰岩山地植物区系从科到种的 组成特征。调查统计分析的植物共有维管植物147科533属889种,其中种子植物占133科508属846种。 种子植物科级区系成分中,以亚热带科为主,暖温带科次之。属级区系成分中,热带分布属占优势,占总属数 的56．2％,温带分布属也占近三分之一,这表明该植物区系的过渡性即亚热带性质。在其种级区系组成中,中 国特有种有328种,占总种数的40％,其次是东亚分布种和热带亚洲分布种。石林石灰岩山地种子植物区系, 从科到种的组成都充分说明了该植物区系的亚热带性质及组成上的独特性。     

Extent and timing of floristic exchange between Australian and Asian rain forests

Aim We tested an entrenched concept – that the Australian rain forest flora is essentially a Gondwanan relict. We also assessed the role of regional‐level source–sink dynamics in the assembly of this flora. Location Eastern Australia. Methods To avoid potential biases inherent in selective studies undertaken to date, we used an analytical, whole‐of‐flora approach integrated with the fossil record. We identified disjunctions between woody Australian rain forest plant taxa and relatives on other land masses. To test the strength of the fossil evidence for the regional antiquity of this flora, we evaluated the proportion of these disjunct clades represented in the Australian fossil record, and to minimize the effects of biases in this record, we compared late Quaternary (i.e. late Pleistocene and Holocene, 126–0 ka), Pliocene and late Oligocene–early Miocene Australian pollen records interpreted as tropical rain forest. Using within‐species disjunctions as a proxy, we assessed the role of recent immigration from Asia into Australia. To assess the role of source–sink dynamics, we performed comparative analyses of disjunctions in major rain forest categories representing a north–south/climatic gradient. Results Southern Australian, cool temperate (microthermal) rain forests contain many floristic disjunctions with Gondwanan fragments and most of these clades have Gondwanan fossils. Disjunct clades in Australian mesothermal rain forest mostly occur in Asia/Malesia and a low proportion of these clades show pre‐Neogene records. Many clades in lowland tropical and ‘dry’ rain forest show disjunctions with Asia/Malesia and few have Australian fossil records. Rates of recent immigration from Asia/Malesia are high in these northern forests, and outweigh rates of recent emigration approximately nine‐fold. The late Quaternary fossil record has many more rain forest angiosperms than Oligocene–Miocene and Pliocene floras, consistent with extensive late Cenozoic immigration. Main conclusions The microthermal rain forests are largely Gondwanan relicts, but there is progressively greater, and more recent contribution from Asia/Malesia into more northern, and more lowland tropical rain forests. This variation reflects a strong gradient in geographic and ecological proximity between these forests and source floras in Asia/Malesia, and is consistent with a source–sink size model of immigration driven by late Cenozoic contractions and expansions of Australian rain forest.     

A new Oligocene Calocedrus from South China and its implications for transpacific floristic exchanges

? Premise of the Study: Calocedrus is among the genera with a typical eastern Asian-western North American disjunct distribution today. The origin of its modern distribution pattern can be better understood by examining its fossil record. ? Methods: The present article reports for the first time a new fossil species of this genus based on compressed material from the Oligocene Ningming Formation of Guangxi, South China, in its present major distribution area in eastern Asia. ? Key Results: Calocedrus huashanensis sp. nov. is most similar to the two extant eastern Asian species, C. macrolepis and C. formosana, in gross morphology of foliage shoots and bears a close resemblance to the latter in cuticle structure. It shows a general similarity to the North American fossil representatives of the genus in alternately branched foliage shoots but is clearly different from the European Paleogene species characterized by oppositely branched leafy shoots. ? Conclusions: This discovery provides new evidence for the floristic exchange of this genus between eastern Asia and North America before the Oligocene (most likely in the Eocene), presumably via the Bering land bridge. The flattened leafy shoots and dimorphic leaves with thin cuticle, open stomatal pits, and shallowly sunken guard cells of the present fossils suggest a rather humid climate during the Oligocene in the Ningming area, South China.     

An early Oligocene fossil acorn,associated leaves and pollen of the ring‐cupped oaks (Quercus subg. Cyclobalanopsis) from Maoming Basin,South China

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.     

Paleogene fossil fruits of Stephania (Menispermaceae) from North America and East Asia

Stephania Loureiro is a large genus within Menispermaceae, with approximately 60 extant species naturally distributed in tropical to subtropical areas in Asia and 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 Stephania from North America and East Asia. The specimens indicate the endocarps were bony or woody with an obovate to obovate-rotund outline and a horseshoe-shaped locule. The endocarp length varies from 4.7 to 8.3 mm, and width from 3.7 to 7.0 mm. The endocarp has a clear foramen in the central area and is surrounded by a keel with ribs running along the dorsal surface. Only one lateral crest develops on each side of the endocarp. Two new species are recognized: Stephania wilfii Han & Manchester sp. nov. from the Paleocene to Eocene of Wyoming (USA), and Stephania jacquesii Han & Manchester sp. nov. disjunct between the late Eocene of Oregon (USA) and the late Oligocene of Guangxi Province (China). In addition, on the basis of more detailed morphological comparative analyses, we transfer the fossils formerly treated as Diploclisia auriformis (Hollick) Manchester from the early Eocene of London Clay, and the middle Eocene of Alaska and Oregon to Stephania auriformis (Hollick) Han & Manchester comb. nov. These fossil materials indicate a broader biogeographic distribution for the ancestors of extant Stephania lineages. This finding enhances our knowledge of the taxonomic and morphological diversity of Stephania and provides new evidence concerning its phytogeographic history.