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.     

Fossil history of the Menispermaceae (Ranunculales)

The Menispermaceae family (Ranunculales) includes more than 500 extant species. Climbers represent the majority of this family, and these plants are now distributed in all of the tropical and subtropical regions of Americas, Asia and Africa. Their endocarps show a characteristic shape that is easily identified in the fossil floras. Numerous fossil leaves are also mentioned in the literature, but the methods for identifying these fossils are often outdated and need revision. Four tribes of Menispermaceae are present in the fossil record, including Fibraureae, Menispermeae, Pachygoneae and Tinosporeae. Only the Anomospermeae are potentially lacking. The Menispermaceae appear in the Palaeocene, and are common in the Eocene, of Europe and in North America as represented by endocarps and locule-casts. The species of this age show morphological diversification, corresponding with the current familial diversity. A few specimens from the Cretaceous period could be part of this Angiosperm family. Fossils after the Eocene are scarcer, despite several relatively rich outcrops. The maximal expansion of the Menispermaceae during the Palaeocene/Eocene limit correlates with the thermal optimum of the early Tertiary and also parallels the maximal expansion of tropical and subtropical floras. The relative richness of the Eocene era in Menispermaceae could be a consequence of more intense sampling in some regions. The bias in tribe representation can be explained by evolution, ecology or taphonomic hypotheses.     

First macrofossil record of Icacinaceae in East Asia (early Oligocene,Wenshan Basin) and its ecological implications

Icacinaceae are well represented in the modern tropical flora of East Asia, but this family has no confirmed macrofossils from this region. Most of the unambiguous fossils (e.g., endocarps) are from the Paleogene of North America and Europe, where the family is no longer present. Here we report a fossil endocarp of the liana genus Iodes from the Oligocene Wenshan flora, southwestern China. The fossil is relatively large (ca. 20 mm length, 11 mm width) and documents a vascular bundle inside the endocarp wall, a pattern of ridges enclosing few areoles, and an asymmetrical apex and rounded base. On the basis of these characteristics, we described a new species, Iodes elliptica, which represents the first Icacinaceae fruit fossil record from Asia. This fossil, consistent with recent reports of Iodes pollen from the Eocene of Hainan, indicates a long-standing presence of the genus in SE Asia, dating back to the Paleogene. Based on the climatic data of modern Iodes, and other fossil occurrences from Wenshan, we hypothesize that the climate in the region was subtropical during the Oligocene, supporting a rainforest, with an overall mixed regional flora of subtropical and tropical elements.     

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。     

Fossil moonseeds from the Paleogene of West Gondwana (Patagonia,Argentina)

Premise of the Study

The fossil record is critical for testing biogeographic hypotheses. Menispermaceae (moonseeds) are a widespread family with a rich fossil record and alternative hypotheses related to their origin and diversification. The family is well‐represented in Cenozoic deposits of the northern hemisphere, but the record in the southern hemisphere is sparse. Filling in the southern record of moonseeds will improve our ability to evaluate alternative biogeographic hypotheses.

Methods

Fossils were collected from the Salamanca (early Paleocene, Danian) and the Huitrera (early Eocene, Ypresian) formations in Chubut Province, Argentina. We photographed them using light microscopy, epifluorescence, and scanning electron microscopy and compared the fossils with similar extant and fossil Menispermaceae using herbarium specimens and published literature.

Key Results

We describe fossil leaves and endocarps attributed to Menispermaceae from Argentinean Patagonia. The leaves are identified to the family, and the endocarps are further identified to the tribe Cissampelideae. The Salamancan endocarp is assigned to the extant genus Stephania. These fossils significantly expand the known range of Menispermaceae in South America, and they include the oldest (ca. 64 Ma) unequivocal evidence of the family worldwide.

Conclusions

Our findings highlight the importance of West Gondwana in the evolution of Menispermaceae during the Paleogene. Currently, the fossil record does not discern between a Laurasian or Gondwanan origin; however, it does demonstrate that Menispermaceae grew well outside the tropics by the early Paleocene. The endocarps’ affinity with Cissampelideae suggests that diversification of the family was well underway by the earliest Paleocene.     

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.     

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.     

黑龙江依兰始新世植物群的研究

本文报道了黑龙江依兰煤矿早第三系始新统达连河组的上、下两个层位所产的植物化石的区系组成及植物群特征：一个系产于煤层下部的砂页岩中的植物群A,另一个是产于煤层上部油页岩中的植物群B。植物组合共计35科49属52种,其中有蕨类植物2种,裸子植物10种,被子植物40种,其中包括1个新种。对其进行区系成分和叶相分析表明,A段植物群的古植被为北亚热带常绿阔叶和落叶阔叶混生林;B段植物群的古植被为暖温带落叶阔叶林。通过与邻近地区的国内外相应植物群比较及植物群的属种地史分布分析,确定A段植物群的时代为早始新世,B段植物群的时代很可能为晚始新世。两个层位间植物群区系成分的变化,表明始新世我国东北地区发生了较明显的气温下降过程,即古气候发生了由亚热带向暖温带的变化。     

A Study on the Eocene Flora in Yilan County,Heilongjiang, China

This paper reports the fossil flora in the Yilan Coal Mine, Heilongjiang Province.The Paleogene flora of the Dalianhe formation includes two groups according to their geology column:one (flora A) is from sand-shale above the lower coal seams and the other (flora B) is from oil shale above the upper coal beds. The flora contains two species of pteridophytes, 10 species of gymnosperms and over 40 species of angiosperms including 1 new species and 10 uncertain species, assigned to 49 genera and 35 families. The analysis of the floristic composition and their foliar physiognomy showes that the flora A consists of elements in evergreen broad-leaved and deciduous broad-leaved forests indicating its subtropical feature, whereas the flora B consists of elements in deciduous broad-leaved forests indicating a warm-temperate feature. Compared to the Paleogene floras of Northeastern Asia and North America, the flora A is assigned to the early Eocene, and the flora B to the late Eocene. These results seem to indicate that in the Eocene, paleoclimatic decline took place in Northeast China.     

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.     

Cyclocarya cf. paliurus (Batal.) Iljinskaja (Juglandaceae) from the Hunchun Formation (Eocene), Jilin Province, China

The leaflet architecture of Cyclocarya cf. paliurus (Batal.) Iljinskaja from the Hunchun Formation (Middle Eocene) shows similarity to that of modern C. paliurus (Batal.) Iljinskaja and the specimen is the oldest fossil record in Europe and Asia. The distributions of C. cf. paliurus and other fossil records,such as Glyptostrobus, Metasequoia, Nyssa, and Liquidambar, in Hunchun flora show that it would have been a warmer-temperature to subtropical climate in Hunchun District during the Eocene period.     

Cyclocarya cf. paliurus （Batal.） Iljinskaja （Juglandaceae） from the Hunchun Formation （Eocene）, Jilin Province, China

The leaflet architecture of Cyclocarya cf. paliurus （Batal.） Iljinskaja from the Hunchun Formation （Middle Eocene） shows similarity to that of modem C. paliurus （Batal.） Iljinskaja and the specimen is the oldest fossil record in Europe and Asia. The distributions of C. cf. paliurus and other fossil records, such as Glyptostrobus, Metasequoia, Nyssa, and Liquidambar, in Hunchun flora show that it would have been a warmer-temperature to subtropical climate in Hunchun District during the Eocene period.     

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.     

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 and foliage of Pueraria (Leguminosae, Papilionoideae) from the Neogene of Eurasia and their biogeographic implications

? Premise of the study: Pueraria (Leguminosae, Papilionoideae) is native in East Asia, South Asia, Southeast Asia, and Oceania and is well known as a rampant invasive weed in the southeastern United States (P. montana; better known as kudzu), but relatively little is known about its early evolution and biogeographic origin. ? Methods: On the basis of comparative analyses of the fruit and leaflet architecture of closely related extant and fossil taxa, we studied the fossil history and biogeography of Pueraria. ? Key results: Fossil Pueraria is recognized on the basis of distinctive fruit and foliage from the Mio-Pliocene of middle latitudes in China, Japan, Abkhazia, and Croatia. Recognition of P. miothunbergiana from the Mio-Pliocene of China and Japan is reinforced by a trifoliolate leaf as well as isolated lateral and terminal leaflets. Pueraria shanwangensis sp. nov. represents the first recognition of fossil Pueraria fruits. This fruit species co-occurs with P. miothunbergiana in the Middle Miocene Shanwang flora and possibly represents the same population. Pueraria maxima (Unger) comb. nov., previously named as Dolichites maximus or Desmodium maximum, is recognized on the basis of leaflets from the Miocene of Croatia and Abkhazia. Other prior fossil reports of Pueraria and Dolichites are reevaluated. ? Conclusions: Pueraria had begun to diversify by at least the Middle Miocene and had spread into the Mio-Pliocene subtropical and temperate floras of the Balkan Peninsula, the Caucasus, and eastern Asia, which suggests the present diversity of this genus in tropical Asia and Oceania might have originated from the mid-latitudes of Eurasia.     

云南剑川盆地上始新统双河组双壳类新材料及其地质意义*

青藏高原东南缘一直被认为是高原隆升造成物质逃逸的主要场所,很好地记录了反映高原隆升的沉积、构造和生物演化事件。剑川盆地是青藏高原东南缘的一个拉分盆地。作为剑川盆地最主要的沉积地层,双河组内仅有少量双壳类化石的报道,缺少详细的化石对比研究。本文系统描述双河组最新发现的双壳类共4属9种,分别为Unio,Acuticosta,Cuneopsis和Anodonta 4属,Acuticosta huoerguosica,Acuticosta sp.,Acuticosta subimmutata,Acuticostacf.chinensis,Acuticostacf.cuspidata,Acuticostacf.semiretiaria,Unio cf.subparallelus,Cuneopsis cf.subceltiformis和Anodonta cf.subpiscinalis 9种,均属于古异齿目珠蚌科。其中Acuticosta和Cuneopsis为现代东亚的特征属,Acuticosta chinensis和Cuneopsis subceltiformis为长江以南的现生种,而Acuticosta chinensis以往只在新疆有过化石记录。结合双河组的侵入脉体的年龄和介形虫标准化石的研究确定双河组的时代为晚始新世,故本次研究不仅为相关双壳类的产出时代可以延伸至晚始新世提供了有力的证据和新材料,而且反映出剑川盆地双河组在晚始新世为亚热带的淡水三角洲前缘-前三角洲的沉积环境。     

Palaeoskapha sichuanensis gen. et sp. nov. (Menispermaceae) from the Eocene Relu Formation in western Sichuan, West China

Palaeoskapha sichuanensis gen. et sp. nov. of Menispermaceae is described here for the first time based on a well preserved fossil fruit. The specimen was found in the Relu Formation of western Sichuan, West China. The specimen, straight, boat-shaped endocarp with large ventral condyle, clearly belongs to the tribe Tinosporeae. The wide aperture of the double condyle, combined with a whole shape not deeply invaginated, indicates a genus different from what was already known to science for this tribe. This fossil widens the distribution of the tribe during Eocene from North America and Europe to Asia, where it was formerly unknown.     

Fossil legumes from the Middle Eocene (46.0 Ma) Mahenge Flora of Singida, Tanzania

Middle Eocene age caesalpinioid and mimosoid legume leaves are reported from the Mahenge site in north-central Tanzania. The Mahenge flora complements a sparse Paleogene tropical African fossil plant record, which until now consisted of a single macrobotanical assemblage, limited palynological studies in West Africa and Egypt, and fossil wood studies primarily from poorly dated deposits. Mahenge leaf macrofossils have the potential to add significantly to what is known of the evolutionary history of extant African plant groups and to expand our currently limited knowledge of African Paleogene environments. The site is associated with a kimberlite eruption and demonstrates the potential value of kimberlite-associated lake deposits as much-needed resources for African Paleogene floras. In this report we document a relatively diverse component of the flora consisting of the leaves of at least five species of Leguminosae. A new species of the extant genus Acacia (Mimosoideae), described herein, is represented by a bipinnate leaf. Another taxon is described as a new species of the extant genus Aphanocalyx (Caesalpinioideae), and a third leaf type may be related to the extant genus Cynometra (Caesalpinioideae). Two additional leaf types are less well understood: one appears to be referable to the Caesalpinioideae and subfamily affinities of the other taxon are unknown.     

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 …