Early Oligocene Calocedrus (Cupressaceae) from the Maoming Basin,South China,and its paleogeographic and paleoclimatic implications

The genus Calocedrus Kurz is characterized by flattened branches covered with decussate and dimorphic leaves. In this study, we describe fossil foliage of Calocedrus discovered in the Shangcun Formation (early Oligocene) of the Maoming Basin, South China. The fossils have wedge‐shaped branchlet segments and scale‐like strongly dimorphic leaves, similar to the fossil species Calocedrus lantenoisi (Laurent) Tao. There have been no detailed studies of the morphological features and cuticle structures of C. lantenoisi. Therefore, a newly emended diagnosis of the species C. lantenoisi is given based on a detailed study of leaf morphology and cuticular characters exhibited by the Maoming fossils. This is one of the earliest fossil records of Calocedrus in the world, providing additional evidence for the early biogeographic history of this genus and supporting the inference that eastern Asian Calocedrus is primitive among all the living species. The extant species of Calocedrus are mainly distributed in mountainous regions. On the basis of the “nearest living relative” analysis, we propose that the Maoming Basin was adjacent to a mountainous region during the early Oligocene.     

Diverse epiphyllous fungi on Cunninghamia leaves from the Oligocene of South China and their paleoecological and paleoclimatic implications

The unique co-occurrence of thyriothecia belonging to three fossil genera of epiphyllous fungi, Stomiopeltites Alvin & Muir (Micropeltidaceae), Callimothallus Dilcher, and Trichothyrites Rosendahl (Microthyriaceae), are reported on the leaves of the same host plant, Cunninghamia shangcunica Kodrul, Gordenko & Sokolova from the Oligocene Shangcun Formation of the Maoming Basin, South China. In China, Stomiopeltites is identified for the first time, Callimothallus is known from the Oligocene and Miocene of Guangxi and Zhejiang provinces, and Trichothyrites previously has been found only in the Eocene palynological assemblages of the Maoming Basin. The presence of abundant and diverse epiphyllous micromycetes, together with the taxonomic composition of the Shangcun megaflora and pollen assemblage, as well as quantitative climatic estimates obtained using Climate Leaf Analysis Multivariate Program, confirm the existence of a warm and humid climate in this region during the late early Oligocene. The geographic and stratigraphic distributions, comparisons with extant analogues, as well as ecological and paleoclimatic implications of the fossil fungi are discussed.     

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.     

Fossil pinnae,sporangia, and spores of Osmunda from the Eocene of South China and their implications for biogeography and paleoecology

The genus Osmunda L. contains approximately 10 extant species widely distributed in tropical and temperate regions, with the greatest concentration of species in East and Southeast Asia. Osmunda is characterized by dimorphic or commonly hemidimorphic fronds with dimorphic pinnae. Its geological history has been traced back to the Triassic. Most records of the genus are based on rhizomes and rarely on pinnae bearing sporangia and spores. Here, we describe fossil pinnae, sporangia, and spores of Osmunda lignitum (Giebel) Stur recovered from the middle Eocene of the Changchang Formation in the Changchang Basin, Hainan Island and the Youganwo Formation in the Maoming Basin, Guangdong, South China. The fossils closely resemble the extant Osmunda banksiifolia (C. Presl) Kuhn of the subgenus Plenasium on the basis of their morphological and anatomical structures. The present occurrence of O. lignitum indicates subg. Plenasium flourished and extended from the high latitude regions such as Northeast China to the low latitude areas of South China during the middle Eocene. Large numbers of specimens described here also indicate that Osmunda was the dominant understory fern element beneath mixed evergreen broad-leaf angiosperm and gymnosperm forests living in a warm and humid environment.     

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.     

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.     

<Emphasis Type="Italic">Camellia nanningensis</Emphasis> sp. nov.: the earliest fossil wood record of the genus <Emphasis Type="Italic">Camellia</Emphasis> (Theaceae) from East Asia

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 that Camellia occurred in China as early as the late Oligocene, suggesting more ancient radiation of this genus than estimated by molecular dating.     

Mummified fossil of Keteleeria from the Late Pleistocene of Maoming Basin,South China,and its phytogeographical and paleoecological implications

Keteleeria is a small genus of Pinaceae now mainly restricted to eastern Asia. Although this genus has been documented with a wide distribution in the geologic record of Europe, North America, and Asia, its history in low‐latitude areas (including South China) has remained obscure. In this paper, a fossil wood of Keteleeria sp. is described from the Late Pleistocene (29–27 ka BP) of the Maoming Basin, South China. This wood is the most ancient megafossil evidence of Keteleeria within the modern distribution area of this genus. The fossil records of Keteleeria suggests that this thermophyllous genus migrated into South China by the Middle Pleistocene escaping from glacial cooling and became widespread over this region in the Late Pleistocene beginning from the interglacial stage preceding the Last Glacial Maximum. The analysis of growth rings in the fossil wood and its comparison with those of modern Keteleeria davidiana (Bertrand) Beissner indicates that in the Late Pleistocene of Maoming Basin (29–27 ka BP) there was a humid climate with less pronounced seasonality of precipitation than that seen in the subtropical monsoonal climate of modern northeastern Vietnam. Apparently, the Maoming Basin was influenced by interglacial regime with summer–monsoon circulation. The previously proposed method to distinguish between evergreen and deciduous conifers based on growth ring anatomy, is not reliable because of the wide variance and ambiguity in its results.     

Diversity of Menispermaceae from the Paleocene and Eocene of South China

We present here the earliest known Asian fossil records of the Menispermaceae based on fossil fruits from Paleocene and Eocene localities in South China. A new genus and species, Paleoorbicarpum parvum sp. nov., and two new species of Stephania Loureiro, S. ornamenta sp. nov. and S. geniculata sp. nov., are recognized from Paleocene deposits of the Sanshui Basin, Guangdong, and a new occurrence of the widespread Eocene species Stephania auriformis (Hollick) Han & Manchester is recognized from the Maoming Basin, Guangdong. The Paleocene Stephania specimens described here represent the earliest fossil endocarp record of the Menispermaceae in eastern Asia. This discovery shows that the moonseed family had arrived in tropical and humid South China by at least the middle Paleocene, which provides important evidence for the origin and phytogeographic history of the family.     

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.     

华南地区新生代木乃伊化木化石研究进展*

中国木化石资源虽然很丰富, 但研究程度相比叶片化石要低得多, 且主要集中在中国北部和西南地区, 研究领域大多局限于分类学方面, 而木化石分析与古气候、古生态等领域的交叉和系统研究还很少。大多数情况下木化石的保存方式都为石化或矿化保存, 而木乃伊化保存的木化石则非常罕见。木乃伊化保存的化石在植物分类、系统演化及古气候研究等方面具有独特而重要的价值。近年来, 本团队在华南地区发现大量木乃伊化保存的新生代木化石, 它们分别产自广西南宁盆地晚渐新世邕宁组、广西桂平盆地中新世二子塘组以及广东茂名盆地晚更新世地层中。本文总结了这些木乃伊化保存的木化石的研究进展, 包括多样性组成、古植被、古气候及古生态等特征, 并对木兰属、蒲桃属和枫香属的植物地理史进行了讨论。综合分析植物群组成特征和木材生长轮以及木化石中具有气候指示意义的解剖结构特征, 推测广西南宁盆地晚渐新世时期生长着常绿(少数落叶)阔叶林, 为温暖湿润的亚热带季节性气候; 广西桂平盆地中新世时期生长着常绿—落叶、阔叶—针叶混交林 (阔叶为主), 为温暖湿润的亚热带气候; 广东茂名盆地晚更新世生长着常绿—落叶、阔叶—针叶混交林, 为温暖湿润的亚热带至热带季风气候。这是中国低纬度地区首次系统研究木乃伊化保存的木化石, 为新生代植物群的系统演化和古环境分析等提供了重要的木化石依据。     

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.     

A new species of Liquidambar (Altingiaceae) from the late Eocene of South China

Journal of Plant Research - A new fossil leaf species, Liquidambar bella (Altingiaceae), is described from the lower part of the Eocene Huangniuling Formation, Maoming Basin, South China....     

Late Eocene leaves of Nageia (section Dammaroideae) from Maoming Basin,South China and their implications on phytogeography

A new species of Nageia, Nageia maomingensis sp. nov., is described from the Late Eocene of Maoming Basin, Guangdong Province, South China. According to its cuticular characteristics, the present fossil species is assigned to N. sect. Dammaroideae, showing the most resemblance to the extant species N. motleyi (Parl.) de Laub. At present, this section is mainly distributed in the Pacific islands region from South Asia to New Guinea. The discovery of N. maomingensis demonstrates that plants of this section inhabited South China at least by the Eocene. The distribution and living habitat of extant N. sect. Dammaroideae imply that South China was warmer during the Late Eocene than today, and may represent one of the centers for early diversification of Nageia and its section Dammaroideae during the Eocene. In addition, we postulate that plants of sect. Dammaroideae migrated southward in response to climate cooling after the Eocene.     

Liverwort fossils from the Late Triassic of Baiyin City,Gansu Province,and their geological significance

Although liverworts are widely distributed around the world with a large number of extant species, reliable fossil records are relatively rare. Here, we report a new species, Ricciopsis baojishanensis Han and Yan, n. sp. (Ricciaceae) and an unnamed species, Hepaticites sp. from the Late Triassic Nanying’er Formation in Baojishan Basin, Baiyin City, Gansu Province, Northwest China. The generic designation is based on detailed comparison of the gross morphology with related fossil and extant species. The new species is characterized by its rosette-forming thallus, dichotomous branching, ribbon-like segments and entire margins. The current fossils represent the first record of liverwort from the Late Triassic in Baojishan Basin, Gansu Province. Based on the different fossil records of the Ricciaceae, we suggest that these taxa were widely distributed during Late Triassic to Oligocene worldwide, mainly in warm temperate and tropical environments, similar with their current distribution. The discovery of the present fossils indicates that the climate of Baojishan Basin in Late Triassic is warmer and more humid than that of today.     

Paliurus Fruits from the Oligocene of South China and Their Phytogeographic Implications

Paliurus favonii Unger is recognized and described based on fruits from the Oligocene Ningming flora of Guangxi, South China. Characteristics of the present specimens include circular winged fruits that are 10.0–11.5 mm in diameter with a central endocarp at 3.0 to 4.0 mm in diameter. The specimens fall into the morphological range of the fossil species P. favonii, which has been observed in other Cenozoic sites in the Northern Hemisphere. The present discovery represents the lowest latitude distribution of P. favonii in the world, and we are presenting the first P. favonii fossil described with detailed cuticular characteristics from China. Further, this finding demonstrates that the genus existed in the Oligocene Ningming region, South China, and provides new information for understanding the fossil history. The dispersal mode for winged fossils demonstrates that wind dispersal is well-represented in the Oligocene Ningming flora.     

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.     

Fossil involucres of Ostrya (Betulaceae) from the early Oligocene of Yunnan and their biogeographic implications

A new fossil occurrence of Ostrya (Betulaceae) is reported based on 14 involucre impressions from the lower Oligocene of Löhe Basin, Yunnan Province, Southwest China. They are characterized by their bladder-like shape with longitudinal veins and perpendicular or branched intercostal veins that form a reticulate venation. The discovery of these fossil involucres represents the earliest unequivocal fossil record of Ostrya in East Asia and the record at the lowest latitude. Its fossil history suggests that the modern distribution pattern of Ostrya might have been established since the early Oligocene, and that this genus has inhabited low altitude areas since then.     

麂属(Muntiacus)的分类及其系统发育研究

麂属(Muntiacus)是一群中小型草食有蹄类,为东洋界的特有属,是典型的热带、亚热带类群。在我国,麂属的分布仅限于长江流域及其以南各地,为我国南方的重要毛皮兽。 以往对麂属的研究多限于一般的形态分类,对其系统发育尚未深入研究。在种级分类上至今还存在一些混乱现象。本文试图用分支系统学(Cladistics)的方法,对麂属的种级分化进行系统整理,并对其系统发育作一初步探讨。     

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.