Shirleya grahamae gen. et sp. nov. (Lythraceae), Lagerstroemia-like fruits from the middle Miocene Yakima Canyon flora, central Washington State, USA

Shirleya grahamae Pigg & DeVore gen. et sp. nov. (Lythraceae) is established for silicified fruits from the middle Miocene Yakima Canyon of central Washington State, USA. The capsules are 10 mm long × 11.5-12.5 mm wide, enclosed in a persistent floral tube and contain 5-7 locules. They are loculicidally dehiscent, fracturing into fragments and leaving the central axis free. Placentation is axile. Five to seven mature seeds are tightly packed per locule, often with several smaller seeds. Seeds are winged, anatropous, and narrowly attached subapically to the central axis, curving basally and radially within the fruit. They are up to ～4.6 mm long × 1.9 mm wide, with a small, triangular embryo cavity and a prominent distal wing. The inflated wing is filled with a bilobed parenchymatous pad of tissue with a central cavity. Shirleya grahamae is assigned to the Lythraceae, and is most similar to Lagerstroemia, based on the synapomorphies of distally winged seeds and revolute cotyledons. Shirleya differs from Lagerstroemia in seed arrangement, and pericarp and wing anatomy. This study provides the first anatomical information about a Miocene Lagerstroemia-like fruit and documents further diversity of the Lythraceae in the Neogene of northwestern North America.     

Anatomically preserved Liquidambar (Altingiaceae) from the middle Miocene of Yakima Canyon, Washington state, USA, and its biogeographic implications

Liquidambar changii Pigg, Ickert-Bond & Wen sp. nov. (Altingiaceae) is established for anatomically preserved, middle Miocene infructescences from Yakima Canyon, Washington, USA. Specimens are spherical, ～2.5 cm in diameter, and have ～25-30 tightly packed, bilocular fruits per head. Fruits are 3.4-4.7 mm wide × 2.6-3.5 mm long and wedge shaped, fused at the base, and free distally. Each locule contains 1-2 mature, elongate seeds proximally and 5-9 aborted seeds of more irregular shape distally. Mature seeds are 1.5 mm long × 1.2 mm wide, elongate, and triangular transversely, with a slight flange. Seeds have a seed coat for which three zones can be well defined, a uniseriate outer palisade layer, a middle region of isodiametric cells comprising most of the integument, and a uniseriate inner layer of tangentially elongate cells lining the embryo cavity. Liquidambar changii is most similar to the eastern Asian L. acalycina H.-T. Chang on features of infructescence, fruit, and seed morphology and quite unlike the North American L. styraciflua L. and other species. Such a close relationship between these two species supports a Beringian biogeographic track between eastern Asia and western North America during the Miocene. Previous phylogenetic and allozyme analysis of modern Liquidambar demonstrates a close relationship between North American-western Asian taxa and suggests a North Atlantic biogeographic track in the middle Miocene. Together, these biogeographic tracks underscore the complexity of the biogeographic history of the Altingiaceae in the Northern Hemisphere throughout the Neogene.     

Anatomically preserved seeds of Nuphar (Nymphaeaceae) from the Early Eocene of Wutu, Shandong Province, China

Well-preserved seeds from the early Eocene of Wutu, Shandong, China are assigned to the genus Nuphar (Nymphaeaceae) based on morphology and anatomy. The seeds of Nuphar wutuensis sp. nov. are ellipsoidal to ovoid, 4-5 mm long with a clearly visible raphe ridge, and a truncate apex capped by a circular operculum ca. 1 mm in diameter bearing a central micropylar protrusion. These features, along with the testa composed of a uniseriate outer layer of equiaxial pentagonal to hexagonal surface cells and a middle layer 4-6 cells thick composed of thick-walled, periclinally elongate sclereids, correspond to the morphology and anatomy of extant Nuphar and distinguish this fossil species from all other extant and extinct genera of Nymphaeales. These seeds provide the oldest record for the genus in Asia and are supplemented by a similar well-preserved specimen from the Paleocene of North Dakota, USA. These data, together with the prior recognition of Brasenia (Cabombaceae) in the middle Eocene, indicate that the families Nymphaeaceae and Cabombaceae had differentiated by the early Tertiary.     

Clevelandodendron ohioensis,gen. et sp. nov., a slender upright lycopsid from the late Devonian Cleveland Shale of Ohio

Clevelandodendron ohioensis Chitaley & Pigg gen. et sp. nov. is an almost entire lycopsid plant known from a single compressed specimen from the Cleveland Shale member of the Upper Devonian Ohio Shale. This unique specimen is 125 cm long, consisting of an unbranched, slender, monopodial axis with a partially preserved plant base bearing thick appendages at one end, and a compact, terminal ovoid bisporangiate strobilus at the other. The stem is 2 cm wide for most of its length. Visible on the decorticated stem surface are helically arranged, elongate leaf traces and laterally compressed, slender leaves along the stem margin. The plant base bears 4-6 thick appendages. The terminal strobilus is compact, ovoid, 9 cm long and up to 6 cm wide, morphologically similar to those of some Lepidodendrales, and bears helically arranged sporophyll/sporangium complexes with narrow bases and distal laminae up to 18 mm long, turned upward. Megaspores are 320-360 μm, trilete and laevigate, lacking a gula; microspores are 30-42 μm, trilete, indistinctly punctate and possibly assignable to Calamospora or Punctatisporites. Clevelandodendron demonstrates that slender unbranched lycopsids with an isoetalean plant habit similar to the Carboniferous genera Chaloneria and Sporangiostrobus and Triassic Pleuromeia-like forms were present as early as the Late Devonian. The early occurrence of this unique habit suggests that diversification within the isoetalean clade sensu Rothwell and Erwin (including both Isoetales and Lepidodendrales) was well established prior to the Carboniferous.     

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.     

REPRODUCTIVE AND VEGETATIVE STRUCTURE OF NORDENSKIOLDIA (TROCHODENDRACEAE), A VESSELLESS DICOTYLEDON FROM THE EARLY TERTIARY OF THE NORTHERN HEMISPHERE

Nordenskioldia borealis Heer is described and its systematic position is reassessed based on examination of the type material and specimens recently collected from three Paleocene localities in North America (Almont, North Dakota; Melville, Montana; Monarch, Wyoming). The morphology of Nordenskioldia infructescences and fruits is clarified, and in particular, silicified specimens from Almont provide new details of fruit and seed anatomy. Fruits are schizocarpic, and individual fruitlets also dehisce to release flat reticulate seeds. These seeds occur in many Paleocene floras but have not been linked previously to Nordenskioldia. Anatomical details of infructescence axes are identical to those of distinctive long and short shoot systems that cooccur with Nordenskioldia, and neither the infructescence axes nor shoots have vessels in the secondary xylem. Comparison of the floras at Almont, Melville, and Monarch with those at other Paleogene localities in Asia, Europe, and North America provides association evidence supporting earlier conclusions that the Nordenskioldia plant bore simple, entire- to crenatemargined leaves with actinodromous venation. Such leaves have been previously assigned to extant genera such as Cercidiphyllum, Cocculus, and Populus but are treated here as Zizyphoides flabella (Newberry) comb. nov. Based on the combined morphological and anatomical details now available, the Nordenskioldia plant is assigned to the Trochodendrales as an extinct genus most closely related to extant Trochodendron.     

Beardia vancouverensis gen. et sp. nov. (Juglandaceae): permineralized fruits from the Eocene of British Columbia

Large numbers of permineralized juglandaceous fruits were identified in calcareous nodules from the Eocene Appian Way locality on Vancouver Island, British Columbia, Canada. The fruits, small dorsiventrally flattened nutlets, 4.5-7.0 mm long and 5.5-9.0 × 3-5 mm in diameter, were studied using cellulose acetate peels. They are wingless, ribbed, and have a lobed epicarp that surrounds the nutlet. Cells of the inner epicarp are thin-walled and traversed by a system of branching vascular strands. The stony nutlet wall is composed of fibers, with an outer layer of distinctive idioblasts. The fruits have a symmetry like that in Juglandaceae, subfamily Juglandoideae, tribe Platycaryeae, while the fibrous nut walls are like those of subfamily Engelhardioideae. This unique combination of characters indicates that these fruits represent a new genus and species of Juglandaceae: Beardia vancouverensis gen. et sp. nov. The excellent preservation of the Appian Way specimens has allowed a unique view of the internal fruit anatomy and external morphology. As the only wingless, flattened nuts known in the family, they further extend the range of morphological variation in fruits in the family. These fossils further support the hypothesis that North America was an important center of generic diversity for Juglandaceae during the early Tertiary.     

Phytogeographic implications of fossil endocarps of Menispermaceae from the Paleocene of Colombia

? Premise of the study: Fossil leaves of Menispermaceae were previously described from the Paleocene of Colombia. Because of strong homoplasy of leaf characters, the fossils could not be placed more specifically within recognized clades, and additional data were needed to specify intrafamilial and paleogeographic relationships during the Paleocene. ? Methods: Fossil endocarps of Menispermaceae were collected from the Cerrejón Formation, the recently discovered Bogotá flora, and Wyoming (～60 Ma). We surveyed the endocarp morphology of almost all extant genera, conducted character optimization, a molecular scaffold analysis, and critically reviewed the related fossil genera. ? Key results: Parallel syndromes of fruit characters have appeared in unrelated clades of the family according to current phylogenetic reconstructions. However, mapping selected endocarp characters across those clades that contain horseshoe-shaped endocarps facilitates identification and phylogenetic assessment of the fossils. Three fossil species are recognized. One of them belongs to the extant genus Stephania, which today grows only in Africa and Australasia. Palaeoluna gen. nov. is placed within the pantropical clade composed of extant Stephania, Cissampelos, and Cyclea; this morphogenus is also recognized from the Paleocene of Wyoming. Menispina gen. nov. shows similarity with several unrelated clades. ? Conclusions: The new fossils from Colombia reveal a complex paleobiogeographic history of the recognized clades within Menispermaceae, suggesting a more active exchange among neotropical, paleotropical, North American, and European paleoforests than previously recognized. In addition, the new fossils indicate that neotropical forests were an important biome for the radiation and dispersal of derived lineages in Menispermaceae after the Cretaceous-Paleogene boundary.     

Anatomical and developmental study of petrified Quercus (Fagaceae) fruits from the Middle Miocene, Yakima Canyon, Washington, USA

The first reported petrified acorns to show internal anatomical structure are here described from Middle Miocene (～15.6 million years old) chert of the Columbia River Basalt Group in Yakima Canyon, Washington. Quercus hiholensis Borgardt et Pigg sp. nov. is described from anatomical and morphological fruit features, as well as a little recognized anatomical feature, the umbilical complex. Acorns, each comprising a nut and its cupule, are up to 15.3 mm long and 18.8 mm wide with helically arranged, imbricate, tuberculate cupule scales. They show basal aborted ovules, short styles, broad stigmas, and lack grooves in their cotyledons. These characters and the developmental pattern seen in these fossil acorns demonstrate that Q. hiholensis conforms to genus Quercus (Fagaceae), subgenus Quercus, section Quercus (the white oaks). The correspondence of Q. hiholensis to the modern section Quercus reveals that the derived floral and fruit characters that distinguish section Quercus within the genus had evolved by the Middle Miocene.     

A new conifer species, Mesocyparis rosanovii sp. nov. (Cupressaceae, Coniferales), and Transberingian floristic connections

A new species, Mesocyparis rosanovii sp. nov. from the Lower Paleocene of the Amur Region, Russia, is characterized by a combination of advanced and primitive features: mixed opposite-alternate branching of vegetative shoots, leafy microsporophylls, as well as a relatively high number of sporangia per microsporophyll and seeds per cone scale. The Late Cretaceous-Early Paleogene genus Mesocyparis McIver et Basinger, which obviously embraces a natural group of closely related species, was a significant component of the vegetation in the middle and high latitudes of eastern Asia and western North America. The range of the genus testifies to terrestrial Transberingian connections between the continents during the Cretaceous warming phases.     

Seed germination ecophysiology of the Asian species Osmorhiza aristata (Apiaceae): comparison with its North American congeners and implications for evolution of types of dormancy

Osmorhiza aristata is an herbaceous perennial that grows primarily in Japan, through southern China, to the Himalayas. It closely resembles the eastern North American species O. claytonii and O. longistylis, and, together, the three species are an example of the well-known North American-Asian pattern of disjunction. Requirements for dormancy break and embryo growth were determined for seeds of O. aristata collected in Japan during the summers of 1998-2000. Embryos in fresh seeds were ca. 0.5 mm long, and they had to grow to 9 mm before the radicle emerged from the mericarp. Embryo growth and germination occurred during cold stratification at 5°C, the optimum temperature for germination. Gibberellic acid did not substitute for cold stratification. Thus, O. aristata seeds have deep complex morphophysiological dormancy (MPD). The type of MPD in O. aristata is similar to that in two western North American congeners but different from that in eastern North American congeners (nondeep complex MPD). Mapping the types of MPD onto a phylogeny of the genus suggests that nondeep complex MPD is derived from deep complex MPD. Although eastern North American-Asian disjuncts often exhibit morphological stasis, the taxa may differ greatly in physiological traits, such as seed dormancy.     

New Palaeomacromiid Dragonflies from the Upper Paleocene of Argentina

A new genus of palaeomacromiid dragonflies, Curviarculia , based on Curviarculia delicata sp. nov. and Curviarculia lamasi sp. nov., is erected from the upper Paleocene Maíz Gordo Formation of north–western Argentina. Phylogenetic relationships within Palaeomacromiidae are discussed, leading to a new family diagnosis.     

FOSSIL OPHIOGLOSSALES IN THE PALEOCENE OF WESTERN NORTH AMERICA

Vegetative and fertile frond segments of Botrychium have been recovered from Paleocene deposits of central Alberta, Canada. Specimens are preserved as coalified compressions that yield information about frond structure, sporangia, and spore ultrastructure. These fossils, described as Botrychium wightonii sp. nov., establish a megafossil record for the Ophioglossales, and demonstrate that modern-appearing species of the order were present in western North America by the earliest Tertiary. The largest vegetative fragments are up to 4.6 cm long and tripinnately compound, with opposite to subopposite branching. Ultimate segments are pinnatifid with dentate pinnules and open dichotomous venation. Fertile specimens are also tripinnately compound with a long rachis and subopposite to alternate pinnae. Sporangia are either submarginal and superficial, or marginal, and are all directed toward one surface of the pinnule. They are ovoid to subspheroidal and 0.8-2.0 mm in diameter. Some sporangia are apparently stalked, while others appear to be sessile. This variation results both from the ultimate frond segments being compressed in several different planes, and the fossils being exposed at different levels. Spores macerated from the sporangia are radial and trilete, and range 30–67 μm in diameter. Most are psilate, but some have a densely striate surface.     

MORPHOLOGICAL OBSERVATIONS ON THE EARLY IMBIBITION OF WATER BY SIDA SPINOSA (MALVACEAE) SEED

The chalazal area is confirmed as the site of initial water entry into prickly sida (Sida spinosa L.) seeds. Very early during imbibition of water, a kidney-shaped area of the seed coat separates from underlying cells forming a blister. This blister may also be induced in dry seeds (both afterripened and nonafterripened) when pressure is applied to the chalazal area. Blisters form more readily on afterripened seeds than on nonafterripened seeds, however, and the event is correlated with an increase in seed coat permeability to water. Immediately beneath the palisade layer of the blister lies a single layer of subpalisade cells. This layer is observed only in the region of blister formation. As the blister separates, the end walls of the subpalisade cells remain attached to the floor of the palisade layer. The subpalisade cells are thereby broken open, and their contents disgorged into the blister lumen. Evidence indicates that this separation of the palisade and subpalisade layers in the chalazal area initiates imbibition of water by prickly sida seeds.     

Vitis seeds (Vitaceae) from the late Neogene Gray Fossil Site, northeastern Tennessee, U.S.A.

This study focuses on morphometric and systematic analyses of the fossil Vitis seeds, recovered from the Gray Fossil Site (7-4.5 Ma, latest Miocene-earliest Pliocene), northeastern Tennessee, U.S.A. A multivariate analysis based on eleven measured characters from 76 complete fossil seeds recognizes three morphotaxa. Further comparisons with both selected modern and fossil vitaceous specimens confirm that these morphotaxa represent three new species, viz. Vitis grayensis sp. nov., Vitis lanatoides sp. nov., and Vitis latisulcata sp. nov. Furthermore, the close resemblance of the first two fossil grapes (V.grayensis and V.lanatoides) with two East Asian Vitis species provides further support concerning a strong eastern Asian aspect of the Gray fossil biota in the late Neogene southeastern North America, as previously evidenced by both animals (e.g. Pristinailurus bristoli [red panda]) and other plants (e.g. Sinomenium and Sargentodoxa).     

珍稀油料树种琴叶风吹楠果实和种子形态变异

以云南野生琴叶风吹楠(Horsfieldia pandurifolia)为材料,对39株树的果实和种子的形态进行变异分析。结果表明,果实和种子形态指标株间变异大,株内变异小。单株间果实表现出从卵圆形至长椭圆形的变异,果长4~7 cm,先端圆钝至锐尖,基部不同程度偏斜,与文献记录差异较大;单株间种子形态表现出从卵形、长卵形至细长形或近锥形的变异。单株平均种长变幅为25.05~35.66 mm,种宽为13.30~19.62 mm,长宽比为1.47~2.32,单因素方差分析表明此3个性状单株间差异极显著(p<0.01)。灰白色种皮具褐色斑块,种脐和胚位于种子基部,种子先端突尖或具短喙是稳定的性状,有重要的分类学价值。     

A NEW JURASSIC ISOETITES (ISOETALES) FROM THE WALLOWA TERRANE IN HELLS CANYON,OREGON AND IDAHO

A new species of the quillwort Isoetites, I. rolandii sp. nov. Ash and Pigg, is described from the Middle Jurassic Coon Hollow Formation in the Wallowa terrane in Hells Canyon, Oregon and Idaho. The new species is based on coalified impressions and mold-casts of an isoetaceous corm with narrow, elongate leaves, and isolated masses of leaves that have been studied using light microscopy. Plants are about 10 cm in maximum height and consist of cormose plant bases from 1.5 to 2.5 cm wide and 0.6 cm high bearing incomplete microphyllous leaves up to 8.4 cm long and 3.0 mm wide, that taper to about 2.0 mm wide distally. Associated casts of sporophyll bases and sporangia contain spherical structures about 440 μm in diameter that may represent megaspores. Based on the occurrence of complete plants, some specimens of I. rolandii are thought to have been preserved in situ. The new Isoetites provides the first floristic evidence for the occurrence of an aquatic or semiaquatic paleoenvironment in the Wallowa terrane. It is also the first documented occurrence of Isoetes-like plants in the Jurassic of western North America.     

Nowickea (Phytolaccaceae), a new genus with two new species from Mexico

A new genus of Phytolaccaceae,Nowickea, includes two new species,N. xolocotzii andN. glabra, each known only from its type locality in the states of Jalisco and Morelos, Mexico, respectively.Nowickea exhibits close relationship withPhytolacca by a robust herbaceous growth habit, 7- to 10-carpellate ovary, carpels with terminal, basally connate styles, basal placentation, and fleshy fruits. It is distinguished by green, subequal, herbaceous, and often lanceolate tepals 3 to 12 mm long, ovary and fruit elevated on a well developed gynophore, obovoid or obpyriform fruits, and narrowly ellipsoid seeds.     

Ariopsis macrosperma sp. nov. (Araceae) from the northern Western Ghats,India

Ariopsis macrosperma sp. nov. from Western Ghats of Maharashtra, India, is described and illustrated. It differs from the other two species in the genus, A. peltata and A. protanthera, in having a typical terrestrial habit, growing on the soil as undergrowth below the forest canopy, thick, leathery leaves and lower number of larger, ovoid and ribbed seeds.