First fossil evidence of samaras of Ventilago Gaertn. (Rhamnaceae) from India and its implications

The Ventilago Gaertn. (Rhamnaceae) is widely distributed in pantropical areas of Africa, Asia, and Australia. However, fossil records of this taxon are sparse, which limits understanding of the evolution and biogeographic history of the genus. In the present study, we report and describe two new fossil species of Ventilago, V. siwalika sp. nov. from the Miocene sediments of Himachal Pradesh, western Himalaya, and V. pliocenica sp. nov. from the Pliocene sediments of Jharkhand, eastern India based on single-winged samaras. Ventilago pliocenica is characterized by a prominent midvein, obtuse to sub-round apex with mucronate tip, longitudinal secondary veins extending the full length of the fruit, and reticulate nature of higher-order veins, the presence of equatorial rim, the hypanthium, and short pedicel. On the other hand, V. siwalika is characterized by a prominent midvein, obtuse to sub-round apex with mucronate tip, longitudinal secondary veins extending the full length of the fruit, and reticulate nature of higher-order veins. Our discovery represents the first unambiguous fossil record of single-winged samara of Ventilago from India and provides valuable insights into the evolution of this genus. In this paper, we also review its biogeographic history and add new information to understand its hypothetical migration route. Present and earlier records of Ventilago also suggest that this genus was a common forest element during Neogene (Miocene time) in Asia.     

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.     

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.     

EOCENE RECORDS OF BEE FLIES (INSECTA, DIPTERA, BOMBYLIIDAE, COMPTOSIA): THEIR PALAEOBIOGEOGRAPHIC IMPLICATIONS AND REMARKS ON THE EVOLUTIONARY HISTORY OF BOMBYLIIDS

Abstract:  The first fossil records of Holarctic representatives of the genus Comptosia Macquart, 1840 from the middle Eocene Messel Pit, Germany, and the upper Eocene of Florissant, USA, are reported. The fossil from Messel, Comptosia pria sp. nov., is represented by a well-preserved wing, displaying characteristic wing venation, remains of the second wing and elements of the thorax. The Florissant fossil, C. miranda comb. nov., is preserved almost completely with both wings and most parts of the body visible. Extant members of the genus Comptosia are currently found only in Australia, and close relatives are known from southern South America. Previously this group of genera has been thought to be of Gondwanan origin, but these Eocene fossil representatives of Comptosia from the Holarctic region argue against this interpretation, and suggest that Comptosia and its relatives were more widespread in both Northern and Southern hemispheres during the Palaeogene. The fossil record of bee flies suggests a major radiation of bombyliids in the late Cretaceous or early Cenozoic. This assumption is supported by the comparatively young geological age of many bombyliid host taxa. A key factor driving the specialization of many bombyliid larvae on ground-dwelling hosts may have been avoidance of competition with tachinid flies and parasitic hymenopterans.     

Reproductive structures of Rhamnaceae from the Cerro del Pueblo (Late Cretaceous, Coahuila) and Coatzingo (Oligocene, Puebla) Formations, Mexico

Recently discovered fossil flowers from the Cretaceous Cerro del Pueblo and flowers and fruits from the Oligocene Coatzingo Formations are assigned to the Rhamnaceae. The Cretaceous flower, Coahuilanthus belindae Calvillo-Canadell and Cevallos-Ferriz, gen. et sp. nov., is actinomorphic with fused perianth parts forming a slightly campanulate to cupulate floral cup, with sepals slightly keeled and spatulate clawed petals. The Oligocene fossils include Nahinda axamilpensis Calvillo-Canadell and Cevallos-Ferriz, gen. et sp. nov. (characterized by its campanulate bisexual flower with stamens opposite, adnate to and enfolded by petals; and with the ovary ripening into a drupe), and a winged fruit assigned to Ventilago engoto Calvillo-Canadell and Cevallos-Ferriz, sp. nov. The flowers and drupe features indicate closer affinity to Zizipheae and/or Rhamneae, while the single samaroid fruit suggests the presence of Ventilagineae. However, the unique character combination in the fossil flowers precludes placing them in extant genera. Nevertheless, the history of the family is long and can be traced back to the Campanian. A detailed phylogenetic revision of the group that uses morphological characters from both extant and fossil plants is needed to better understand the significance of these records as well as other important fossils of the family.     

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.     

Miocene winged fruits of Loxopterygium (Anacardiaceae) from the Ecuadorian Andes

A new species of asymmetrically winged fruit is described from Miocene sediments of Andean Ecuador. The new fruit is readily placed in the genus Loxopterygium of the Anacardiaceae based on the size, position of the stigma, wing venation, and serration of the wing tip. The new fossil species is very similar to extant species of Loxopterygium now distributed in dry habitats of coastal Ecuador and Peru, as well as dry interior forests of Bolivia and northern Argentina. We use the fossil to calibrate a molecular-based phylogeny of some members of the Anacardiaceae, showing that dry forest habitats may have been present in South America for more than 10 million years.     

THE VENATION OF THE MATURE NYMPH WING PAD OF EPHEMEROPSIS TRISETALIS EICHWALD (EPHEMEROPTERA: HEXAGENITIDAE)

Abstract  The complete mature nymph wing pad with distinct venation has been collected from Jiuquan Basin, Northwest China, which is almost without precedent in the fossil history of insect. The venation with large radial and costal fields is considered as archaic feature which very probably represents the primitive stage in the evolution of Ephemeroptera. Furthermore, as compared with the modern adult wing, the stronger articular sclerites in the fossil nymphal wing pad of Ephemeropsis trisetalis probably shows that the reduction of the articulation in modern mayfly wing is a secondarily acquired character.     

三尾类蜉蝣成熟稚虫的翅芽脉相(蜉蝣目)(英语)

本文记述了三尾类蜉蝣Ephemeropsis trisetalis稚虫的脉相和翅关节区特征为化石昆虫保存史上罕见材料,脉相中显出颇大的径脉区和前缘区十分相似于某些原始的古生代蜉蝣,或许可以提示发达的径脉区和前缘脉区是蜉蝣翅进化史中的初期特征.再者,现代蜉蝣的翅关节区退化,一般由一组不发达,分化很弱的骨板和肩板组成;这个化石种的翅关节区具发达的骨板和肩板似指示现化蜉蝣翅关节区退化是后生获得的性状.     

First discovery of fossil Coloninae in Cretaceous Burmese amber (Coleoptera,Staphylinoidea, Leiodidae)

Coloninae is a tiny and distinctive subfamily of the highly diverse family Leiodidae (round fungus and small carrion beetles). Although only one leiodid fossil is known from the Mesozoic, its taxonomic position and classification are doubtful. We describe the first definitive Coloninae fossil, Colon burmiticum Yamamoto sp. nov., from Upper Cretaceous amber of Myanmar. Our finding sheds light on the origin and early evolutionary history of this peculiar genus and its subfamily.     

Moltenia rieki n. gen., n. sp. (Hymenoptera: Xyelidae?, a tentative sawfly from the Molteno Formation (Upper Triassic), South Africa

The fore wing of the new genus and speciesMoltenia rieki of the order Hymenoptera, tentatively belonging to the family Xyelidae, is described from the Molteno Formation (Upper Triassic) of South Africa. This fossil is a representative from the early evolutionary history of this order and the oldest Hymenoptera from central Gondwanaland. However, the plesiomorphic characters in its wing venation do not allow a clear assignment to a recent subgroup of the Hymenoptera, although generally it can be assumed that the Xyelidae had a much higher diversity during the Triassic than today.     

New fossil fruits of Carya (Juglandaceae) from the latest Miocene to earliest Pliocene in Tennessee,eastern United States

Fossil fruits including nuts and associated husk valves of a new species of Carya (Juglandaceae) are described from the latest Miocene to earliest Pliocene in northeastern Tennessee, eastern United States. The husk valves are elliptic, 1.2–4.5 mm thick, with a convex exterior face and a concave interior face; the nuts are globose to ovoid in shape, smooth and longitudinally ribbed on exterior surface, with a short protruding apex and a slightly 4-angled base; inner ribs, lacunae and primary septa are well-developed, while secondary septa are absent or weakly developed. The combination of these carpological characteristics clearly shows a close resemblance to the genus Carya in Juglandaceae. Detailed comparisons of carpological morphology and anatomy indicate that the present fossil taxon is different from both living and most other fossil species of the genus, and therefore warrants the designation of a new fossil species, Carya tennesseensis Huang et al., sp. nov. Carya tennesseensis displays a carpological similarity to C. ventricosa from the late Oligocene to early Pliocene in Europe, suggesting a potential species exchange of the genus between Europe and southeastern North America during the late Neogene. The new fossil species represents one of the few fruit fossil species of Carya from its modern distribution range in southeastern North America. It provides crucial information for better understanding the rapid diversification of the genus from the late Miocene to early Pliocene, and the origin and establishment of today's Carya biodiversity in this region.     

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.     

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.     

A new Old World vulture (Falconiformes: Accipitridae) from the Miocene of Gansu Province, northwest China

A large-sized and almost complete fossil vulture was discovered from the Late Miocene Liushu Formation of Linxia Basin in northwestern China. It is the best-preserved and the most complete fossil vulture yet discovered. The new genus and species Gansugyps linxiaensis is proposed and assigned to the family Accipitridae; morphology and limb proportions suggest it was chiefly an arboreal and soaring bird. It is more advanced than the other two known Miocene vultures from China. This new fossil increases our knowledge of the evolutionary history of vultures, and has implications for reconstructing the paleoecology of this region.     

The Gomphaeschninae (Odonata: Aeshnidae): new fossil genus, reconstructed phylogeny, and geographical history

ABSTRACT. Three new species of fossil Anisoptera (dragonflies) are reported from Tiffanian (Late Palaeocene) sediments of the Paskapoo Formation, near Red Deer, Alberta, Canada. The three species are all assigned to the new genus Alloaeschna , which is classified in the subfamily Gomphaeschninae.
Phylogenetic analysis of gomphaeschnine wing venation suggests that the subfamily is paraphyletic, with derived gomphaeschnines such as Boyeria and Linaeschna more closely related to the Brachytroninae and Aeshninae than are other gomphaeschnines. One of the most primitive gomphaeschnines, and the oldest fossil form, is the Jurassic Morbaeschna. Parallelism, convergence and reversal in the evolution of features of the wing venation is common.
The new genus Alloaeschna is at about the same evolutionary grade as the old-world genus Oligoaeschna and the new-world genus Gomphaeschna , and is one of the more primitive known members of the Gomphaeschninae. The new species are the first recorded Palaeocene gomphaeschnines and the oldest known gomphaeschnines from the Americas.
Several extant genera have fossil representatives on continents different from those on which they now survive, suggesting repeated crossing of land bridges and/or widespread ancestral species prior to the separation of the continents, and subsequent extinction leading to present relict distributions.     

A fossil fruit wing of Dipterocarpus from the middle Miocene of Fujian, China and its palaeoclimatic significance

Dipterocarpus zhengae sp. nov. is described from the middle Miocene Fotan Group of Zhangpu county, Fujian Province, Southeast China on the basis of a fruit wing. Three kinds of venation of the calyx longer lobes (enlarged sepals) occur in the winged fruits of extant Dipterocarpus. The fossil species is referred to the kind having three primary veins and within that category is most similar to extant Dipterocarpus gracilis Blume in the size of the longer lobe as well as its venation. The occurrence of the fruit wing of Dipterocarpus, together with palynological evidence, indicates unequivocally that a tropical climate and tropical rain forest occurred in Zhangpu during the middle Miocene, warmer and more humid than at the present day. The palaeobiogeography of Dipterocarpus is reviewed.     

Chaneya, a New Genus of Winged Fruit from the Tertiary of North America and Eastern Asia

A new genus is recognized on the basis of wind-dispersed fruits from the Eocene of western North America and Miocene of eastern Asia. The fruits consist of an accrescent hypogynous calyx of five obovate sepals and one or more globose fruit bodies. Although the fossils were formerly placed in the extant genera Porana (Convolvulaceae) and Astronium (Anacardiaceae), our investigation of numerous specimens from several floras in the western United States (e.g., Florissant, Green River, Clarno) and Canada (Whipsaw Creek, British Columbia) and the Yilan and Shanwang floras of China reveals unique characters that indicate that the fossils are a distinct genus, which we name Chaneya. Unlike Porana and Astronium, the fossil calyces have stomata that are longitudinally aligned, and early stages of fruit development show a gynoecium of five apocarpous carpels, of which only one or two usually enlarge at maturity. Precise systematic placement of the fossil genus is uncertain, but similarities to the extant Picrasma of the Simaroubaceae are suggestive of possible affinities. Two species are recognized: Chaneya tenuis (Lesq.) comb. nov., from the Eocene of western North America and northeastern China, and Chaneya kokangensis (Endo) comb. nov., from the Miocene of eastern Asia.     

First fossil record of the genus Nageia (Podocarpaceae) in south China and its phytogeographic implications

The first fossil record of the genus Nageia Gaertner from the Tertiary of southern China is reported. Nageia hainanensis sp. nov. is described from the Eocene Changchang Formation of Hainan Island. The extant species of Nageia are widely distributed in southeastern Asia, from northeastern India to southeastern China and southern Japan, and southward through Indochina to the Malay Archipelago and New Guinea. The fossil evidence of Nageia has been discovered from the Lower Cretaceous of the Far East of Russia and Japan. The distribution of both modern and fossil species of Nageia indicates that this genus could have originated in the northeastern part of Asia in the Early Cretaceous and spread to south China at least in the Eocene. Discovery of Nageia fossil record in south China provides important evidence for the geological and phytogeographic history of the genus.     

A new species of winged fruit from the Miocene of Ecuador: Tipuana ecuatoriana (Leguminosae)

Tipuana ecuatoriana sp. nov. (Leguminosae; Dalbergieae), is described from winged fruits collected in Miocene sediments of the Loja and Nabon Basins of Ecuador. Extensive comparisons with families that include genera with similar samaroid fruits or disperal units led to recognition of legume affinities for the fossil. Based upon the texture of the fruit, the position of the style remnant, and the venation of the fruit wing, the fossil conforms to the extant monotypic genus, Tipuana, native to subtropical Bolivia and northwest Argentina, but widely cultivated in the subtropics. Tipuana, and thus the Dalbergieae as presently recognized, were present in South America prior to the establishment of permanent land connection between North and South America.