FRUITS AND SEEDS OF TRIBE GORDONIEAE (THEACEAE) FROM THE EOCENE OF NORTH AMERICA

A new genus and species Gordoniopsis polysperma and two new species of Gordonia (Gordonieae, Camellioideae, Theaceae) are described based on fossil fruit and seed remains. These specimens are part of a large flora consisting of various plant organs from the middle Eocene Claiborne Formation in western Kentucky and Tennessee. Gordoniopsis is a five-valved loculicidally dehiscent capsule similar to capsules of Gordonia but differing in having unwinged seeds and a greater number of seeds per locule. The two Gordonia species are among the earliest unequivocal records of the genus and two of only four fossil Gordonia species known with in situ seeds. Two extinct genera, Gordoniopsis and Andrewsiocarpon, and the extant genus Gordonia in the tribe Gordonieae are known from the middle Eocene Claiborne flora, suggesting an early radiation within the tribe. Based on a survey of Recent fruits and seeds we concur with Keng's proposal to merge Laplacea with Gordonia.     

FOSSIL FLORAL EVIDENCE OF MALPIGHIACEAE AND AN EARLY PLANT-POLLINATOR RELATIONSHIP

Fossil flowers with affinities to Malpighiaceae have been discovered in the Middle Eocene Claiborne formation of northwestern Tennessee. The new taxon Eoglandulosa warmanensis gen. et sp. nov. Taylor and Crepet, has paired glands on the five sepals, clawed petals and tricolporate pollen with reticulate ornamentation and an unusual infratectal wall structure of anastomosing elements. The fossil is similar in wall structure to some extant species of Malpighiaceae. Glandular floral morphology in extant species is associated with specific anthophorid bee pollinators and the fossil evidence suggests that such specific plant-pollinator relationships existed during the Eocene. This fossil species also suggests that by the Eocene, South American floral elements had migrated to North America via island pathways, and that the Mississippi embayment was nearly frost-free.     

The mangrove palm Nypa in the geologic past of the New World

The paleontological history of Nypa, known today as the mangrovepalm, is traced through geological time back to the Late Cretaceous. Emphasis is laid on the New World occurrences, especially in NorthAmerica, where Nypa is known from fossil fruits and pollen. In SouthAmerica, the stratigraphic range of this palm extends from theMaastrichtian to the late Eocene, whereas in North America, Nypa isrestricted to only the Eocene. Nypa occurs as pollen all along theAmerican Gulf Coast from the early Eocene (Ypresian) to the late Eocene(Priabonian), while fruit records come from the early and middle Eoceneof Maryland and Texas, respectively. The floristics of these Eocenemangroves, including possible mangrove associates, and the developmentof mangrove vegetation in the neotropics through the Tertiary andQuaternary, are discussed. New paleobotanical evidence from a middleEocene faunal and floral assemblage in Texas, the Casa Blanca flora (LaredoFormation, Claiborne Group), which contains fossil Nypa pollen andfruits, is described in detail.     

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.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF SOUTHEASTERN NORTH AMERICA: PHILODENDRON LEAF REMAINS

Large leaves, new to the fossil record, from the Claiborne Formation in western Tennessee have been collected and analyzed. Careful analyses of venation and cuticular anatomy indicate that these fossil leaves contain specific characters found in modern species of the genus Philodendron subgenus Meconostigma. Features of venation were taxonomically more useful than other features. Reports of fossils of the Araceae are scarce and those few fossils assigned to the family frequently have been inadequately studied. This report establishes a reliable and carefully documented occurrence of the Araceae in the fossil record, and provides information concerning the differentiation of the family in time and the distributions of the genus Philodendron.     

EUPHORBIOID INFLORESCENCES FROM THE MIDDLE EOCENE CLAIBORNE FORMATION

Inflorescences from the Claiborne Formation of western Tennessee are remarkably similar to those of the tribe Hippomaneae, subfamily Euphorbioideae, of the Euphorbiaceae. The fossil inflorescences are spikes of bract-subtended cymules of at least three florets each. Florets are composed of at least three stamens. Palynological features of the fossils are also shared by the Hippomaneae. Fossil pollen is tricolporate, prolate (26.9 × 20.6 μm; P/E = 1.3), with lalongate pores. Exine structure is tectate columellate with a perforate tectum. The exine is reticulate and the muri are conspicuously striate. These specimens represent the first fossil floral evidence of the Euphorbiaceae. It is surprising that inflorescences of the Hippomaneae so modern in aspect existed in the Middle Eocene, since the tribe is universally considered to be one of the most advanced in the family.     

EXTINCT TRANSITIONAL FAGACEAE FROM THE OLIGOCENE AND THEIR PHYLOGENETIC IMPLICATIONS

Fruits, catkins, and associated leaves of at least two extinct trigonobalanoid taxa have been discovered at an Oligocene fossil plant locality rich in fagaceous remains. These fossils exhibit a mosaic of fruit and pollen characters found in the two extant subfamilies Castaneoideae and Fagoideae of Fagaceae. Comparison with cladograms based on modern taxa suggests that these extinct taxa were similar to the ancestors of subfamily Fagoideae and may have been intermediate between Fagus and the modern trigonobalanoid genera. Pollen types isolated from the fossil staminate catkins provide unique character states that are transitional between modern pollen types in Fagaceae and are important in understanding the evolution of exine micromorphology within the family. This analysis provides a striking example of the use of character data from fossils to determine character-state adjacency prior to polarization of characters using outgroup comparison. Because of the mosaic nature of their character complexes, these fossils support monophyly in both the family Fagaceae and the subfamily Fagoideae. In addition, the occurrence of trigonobalanoid fossils in the Oligocene of North America has interesting biogeographic implications and provides insights into the nature of North American Fagaceae during the Tertiary.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: A MIMOSOID INFLORESCENCE

Six examples of a spicate inflorescence from the Middle Eocene Claiborne Formation in western Tennessee have been investigated. Individual flowers are small, alternately arranged, nearly sessile, and perfect. The style protrudes 2 mm beyond the floral envelope and terminates in a slightly swollen, rounded stigma. Both the style and the stigma are hairy. Ten stamens are exserted and extend 5 mm beyond the floral envelope. Anthers are small, versatile, and dehisce longitudinally. Pollen grains are tricolporate, tectate, and are in permanent tetrahedral tetrads 32 μm in diameter. Comparison of the fossil inflorescences with those of extant families having multiple pollen grain configurations suggests that the fossil inflorescences are most closely allied to the subfamily Mimosoideae of the Leguminosae. These are the first structurally documented remains of the Mimosoideae from the Middle Eocene.     

An ancient example of Platycladus (Cupressceae) from the early Miocene of northern China: origin and biogeographical implications

A number of cupressaceous fossil leaves are described in this paper that come from the lower Miocene of Inner Mongolia, northern China. These new examples of fossil leaves are compressed, have their branchlets arranged in a plane, either spreading or ascending, and are flat. These specimens also possess scale-like leaves that are dimorphic, preserved facial leaves that are rhomboidal and have obtuse apices, and lateral leaves that are boat-shaped and have slightly incurved apices. The epidermal cells of these fossils are quadrangular, rectangular, or narrowly rectangular, while the leaves are amphistomatic, and stomata are elliptical or oblong, and monocyclic. In comparison with both living and related fossil taxa, we identify the new fossils as Platycladus preorientalis sp. nov. Taken in combination with a molecular phylogenetic analysis, the new fossils suggest that the genus Platycladus originated in the Oligocene in mid and high Asian latitudes. Thus, the ancient Platycladus lineage probably migrated from Europe to Asia following the closure of the Turgai strait during the Oligocene.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: A CATKIN WITH JUGLANDACEOUS AFFINITIES

Three specimens of one type of fossil catkin from the Middle Eocene of Tennessee are excellently preserved and have been investigated morphologically. The flowers on these catkins are subtended by elongate, three-lobed bracts, are exclusively staminate, and have three conspicuous, obovate, perianth parts that bear large peltate scales. The stamens are well preserved and contain triporate pollen grains that are equivalent to the dispersed pollen genus Momipites. Floral morphology, cuticular features, and pollen indicate close affinities with the extant genera Engelhardia, Oreomunnea, and Alfaroa of the Juglandaceae; but because the fossil catkins are distinct and are a dispersed plant organ, they are placed in a new form genus: Eokachyra. These fossil flowers represent a rare opportunity to correlate the micro- and macrofossil record and to compare the relative rates of evolution of these features. The fossil catkins also demonstrate that much structural information may be gained from the study of fossil angiosperm flowers. The similarities between the staminate flowers of the fossil catkins and the staminate flowers of Engelhardia, Oreomunnea, and Alfaroa confirm the idea that this complex has had a long evolutionary history and suggest that the pollination system of certain extant genera was well developed during Middle Eocene times.     

福建漳浦中中新世几种藓类植物研究*

亚洲新生代藓类植物化石非常稀少,漳浦琥珀是目前已知的我国低纬度地区唯一含苔藓植物化石的新生代琥珀矿床。漳浦琥珀源自中中新世气候适宜期(~14-17 Ma)的龙脑香树脂,温暖潮湿的热带季雨林气候和广泛分布的龙脑香科植物,为藓类植物的生长、多样性演化和三维立体保存提供了条件。本研究共报道了漳浦县中中新世藓类植物化石3个属种,包括:白发藓属Leucobryum Hampe、牛舌藓属Anomodon Hook.et Taylor和扁枝藓属Homalia(Brid.)B.S.G.。首次利用三维X射线断层扫描技术(Micro-CT)研究了白发藓属化石,发现其与现生种短枝白发藓Leucobryum humillimum Cardot非常相似,因此将其命名为短枝白发藓(相似种)Leucobryum cf.humillimum。此外,当前牛舌藓属Anomodon和扁枝藓属Homalia化石是东亚地区的首例记录。漳浦琥珀中的藓类植物化石揭示了中中新世时期苔藓植物的多样性,丰富了东亚低纬度地区新生代苔藓植物的化石记录。     

BOTANICAL AFFINITIES OF SOME TRIPROJECTACITES FOSSIL POLLEN

During Late Cretaceous to Oligocene times, fossil pollen of the Triprojectacites group (also known as Aquilapolles or triprojectates), comprised a temporally and environmentally distinctive element of palynofloras in eastern Asia and western North America. Several species of this group serve as biostratigraphic index fossils for this interval. Using electron microscope and numerical analyses, primarily of North American triprojectate pollen, it is possible to recognize the presence of three distinct subgroups. One group, corresponding to the fossil genus Mancicorpus, has no morphologically close modern representative. Characters resembling those in Santalaceae (Santalales, Rosidae) occur in the second group, which is represented by a previously underscribed fossil triprojectate genus. The third group contains retipilate, isopolar pollen and strioreticulate, isopolar pollen. Forms exhibiting the latter morphology are commonly assigned to the genus Integricorpus, while the retipilate morphology characterizes another underscribed genus. This third group may have some phylogenetic connection to Apiaceae (Apiales, Asteridae). Pollen of some other extant families exhibits triprojectate features, although no close fossil representative can be presently identified. The triprojectate morphology is thus interpreted in modern and fossil forms as resulting from convergence rather than close phylogenetic relationships at the group level.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: STIPULATE LEAVES OF THE RUBIACEAE INCLUDING A PROBABLE POLYPLOID POPULATION

A comparative study was made of the gross morphology, fine venation and cuticular features of Leitneria fioridana Chapman, the single living representative of the order Leitneriales and Leitneria eocenica (Berry) Brown, presumbaly a related fossil species. In addition to the type material, newly collected fossil specimens were investigated from clay pits in the Middle Eocene, Claiborne Formation, of western Tennessee and Kentucky. Foliate stipules attached to the petioles of several specimens suggest the assignment of this fossil leaf type to the genus Leitneria is incorrect. The nature of the gross morphology, fine venation and cuticular features confirms the misidentification. Previously, various specimens of this fossil leaf type have been placed in eight species of seven genera in seven families of six angiosperm orders, none of which are correct systematically. The gross morphology, venation and cuticular characters of the fossil leaf are distributed among a few extant South American genera of arborescent Rubiaceae. The fossil is an extinct rubiaceous leaf type which cannot be placed within a single modern subfamily, tribe or genus of the family. The organ genus, Paleorubiaceophyllum is proposed for these leaves. Three varieties of a single fossil species, P. eocenicum, are recognized. One variety with epidermal cells nearly twice the size of the others may represent a polyploid population.     

New fossil leaves and fruits of Lauraceae from the Middle Miocene of Fujian,southeastern China differentiated using a cluster analysis

The fossil record of Lauraceae can be traced back to the Early Cretaceous of eastern Asia based on fossil flowers. Here, we refer a number of new occurrences of leaf and fruit fossils of Lauraceae from the Middle Miocene of Zhangpu, Fujian, China, to seven species. These data provide evidence supporting the fact that a diverse subtropical, or tropical, Lauraceae-dominated evergreen forest surrounded this region 15 million years ago (Mya). The Lauraceae fossils presented in this paper provide evidence for the evolution of this group as well as new materials that enable the study of the Fujian Province Neogene flora. The fossils described in this paper fill in the gaps in studies about Lauraceae pollen in the Middle Miocene from Fotan, Fujian, China. In addition, these fossils also enrich the Middle Miocene fossil records of Lauraceae in eastern Asia, especially improving the study of the macrostructures and reproductive organs of fossil Lauraceae from southern China. The similarity between fossil and modern fruits shows that during the Middle Miocene the fruit morphological of Lauraceae have changed very little. We also identify families where the fossils we report belong to their closest relatives and can be used to reconstruct the paleoenvironment of Fujian in the Middle Miocene.     

ADDITIONS TO THE MIOCENE SUCKER CREEK FLORA OF OREGON AND IDAHO

Study of plant microfossils and additional macrofossils collected from sediments in the vicinity of Sucker Creek along the Oregon-Idaho boundary has provided additional data on the composition of this fossil flora. A total of seven taxa new to the flora have been recognized. These include colonies of Botryococcus; pollen of Podocarpus, Pachysandra, and the Onagraceae; and leaves of Cephalotaxus californica and Vaccinium sophoroides. The occurrence of Podocarpus in this mid-Miocene flora is particularly significant in view of the limited number of records of this genus in the Tertiary of North America. In addition to these new records, the presence of three additional taxa—Castanea, Nyssa, and Ilex—previously recognized on the basis of leaf remains, is further substantiated by the occurrence of fossil pollen in the sediments.     

Floral data from the mid-Cenozoic of central Pakistan

Floral remains are generally rare in the fossil record of Pakistan. We present here new discoveries of mid-Cenozoic wood and pollen of Oligocene deposits from central Pakistan. The palynological sample derives from claystone stratigraphically positioned above the wood fossils, but pollen and spores as well as the wood are all of Oligocene age. The three fossil wood samples derive from the top of the lower unit of the Chitarwata Formation, and the palynological sample comes from claystone situated in the middle unit of the formation. The wood samples are described and found to represent two species of the morphotaxon Terminalioxylon (Combretaceae): T. burmense Mädel-Angeliewa and Müller-Stoll [Madel-Angeliewa, E., Müller-Stoll, W.R., 1973. Kritische Studien über fossile Combretaceen-Hölzer: über Hölzer von Typus Terminalioxylon G. Schönfeld mit einer Revision der bisher zu Evodioxylon Chiarugi gestellten Arten. Palaeontographica 142B, 117–136.] and T. sulaimanense sp. nov. These fossils are close to modern species of Terminalia, which occur in moist deciduous or semi-evergreen tropical forests. The palynological assemblage is composed of a mixture of pollen and spore types from different origins. There is a dominance of hygrophilous ferns, pines, Amaranthaceae–Chenopodiaceae–Caryophylaceae, but also the occurrence of Palmae, which are typical of tropical rainforests. This assemblage suggests that the depositional system is set in a context of nearby mountains with a minimum altitude of 2000 to 2500 m and characterized by differentiated forest belts above the tropical lowland vegetation of lower elevations. Wood, pollen and spores support the view of a fluvial environment surrounded by a tropical forested habitat. The separate stratigraphic position of the pollen assemblage above the fossil wood could explain its slightly different, more varied, palaeoenvironmental signal.     

The Fossil Record of the Epacridaceae

Fossil pollen and macrofossils of Epacridaceae are uncommonand are mainly known from Tasmania and other parts of south-easternAustralia. Most epacrids have generalized ericalean pollen althoughthe pollen of some genera is distinctive. Ericalean pollen isknown from the late Cretaceous. The first occurrence ofParipollisorchesis pollen, which is consistent with some extantEpacrisspecies,probably means that Epacridaceae, and possibly the tribe Epacrideae,had differentiated by the Middle Eocene. The fossil record at present provides minimum ages of the firstoccurrences of major subfamilial taxa. Macrofossils of subfamilyRicheoideae and of several morphotypes of the tribe Epacrideaeare known from the Early Oligocene. Tribe Cosmelieae pollenand macrofossils are known from the Early Pleistocene, and areprobablySprengelia. The oldest Australasian fossils of tribeStyphelieae are leaves in latest Oligocene–Early Mioceneparts of the Latrobe Valley coal. Endocarps identified as Epacridaceaefrom the Eocene of England need further investigation. PollenofMonotoca, or a close relative, is known from the mid-Miocene.PossibleTrochocarpaleaves occur in Late Oligocene/Early Miocenesediments, and fossil leaves indistinguishable from the extantTasmanian rainforest species,T. gunniiandT. cunninghamii, areknown from the Early Pleistocene in Tasmania. Epacridaceae; macrofossils; microfossils; Cretaceous; Cainozoic     

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.     

The first named Ediacaran body fossil, Aspidella Terranovica

Aspidella terranovica Billings, 1872 was first described from the late Neoproterozoic Fermeuse Formation (St. John's Group) on the Avalon Peninsula of eastern Newfoundland, approximately 1km stratigraphically above the famous Ediacaran biota at Mistaken Point, and several kilometres below the base of the Cambrian. Aspidella has been reinterpreted perhaps more than any other Precambrian taxon, and has variously been regarded as a fossil mollusc or ‘medusoid’, a gas escape structure, a concretion, or a mechanical suction mark. Our studies indicate that Aspidella includes a wide variety of preservational morphs varying from negative hyporeliefs with a raised rim and ridges radiating from a slit (Aspidella‐type preservation), to flat discs with a central boss and sharp outer ring (Spriggia preservation), to positive hyporeliefs with concentric ornamentation (Ediacaria preservation). Specimens occur in a continuum of sizes, with preservational styles dependent on the size of the specimen and the grain size of the host lithology; the elongation of specimens is tectonic. Aspidella is confirmed as a body fossil from observations of complex radial and concentric ornamentation, mutually deformed borders in clusters of specimens, and occurrence on the same bedding planes as certain distinctive Ediacaran taxa. Aspidella is indistinguishable from, and has priority over, several of the most common genera of late Neoproterozoic discoidal body fossils worldwide. Similar fossils from Australia are interpreted as holdfasts of frond‐like organisms. The density of specimens in the Aspidella beds suggests levels of benthic biomass in the Neoproterozoic that could rival those of modern marine communities. The serial growth forms, PalaeopascichnusIntrites, Neonereites renariusYelovichnus, associated with Aspidella, are interpreted as body fossils of unknown affinities rather than trace fossils. A new, trilobed, Ediacaran body fossil, Triforillonia costellae gen. et sp. nov., is described from the Aspidella beds of the Fermeuse Formation.     

EOCENE FLORAL EVIDENCE OF LAURACEAE: CORROBORATION OF THE NORTH AMERICAN MEGAFOSSIL RECORD

Numerous megafossils of Lauraceae have been reported from the early Tertiary of North America, but the subfamilial affinities are usually not well understood due to the great morphological variability found in extant taxa. The flowers of Androglandula tennessensis gen. et sp. nov. Taylor, from the Middle Eocene Claiborne Formation, are six-parted, pedicellate, bracteate, and have stamens with paired basal staminal glands. The flowers have ethereal oil cells and paracytic stomates throughout. The fossil species has affinities with the subtribe Cinnamomineae, and this supports suggestions that the Middle Eocene climate of the southeastern U.S. was subtropical. The existence of this fossil, and reports of the subtribe from the Eocene of Europe, indicate a South American-North American-European-southeast Asian paleodistribution suggesting that extinction in North America and Europe was the cause of the tribe's current disjunct distribution.