Appomattoxia ancistrophora gen. et sp. nov., a new Early Cretaceous plant with similarities to Circaeaster and extant Magnoliidae

A new genus and species of fossil angiosperm (Appomattoxia ancistrophora) is established based on well-preserved fruiting units and associated pollen from the Early Cretaceous (Early or Middle Albian) Puddledock locality in the Potomac Group sequence of Virginia, eastern North America. Fruiting units are small, unilocular, and with a single, pendulous, orthotropous seed. The fruit surface is characterized by densely spaced unicellular spines with hooklike tips, which probably functioned in biotic dispersal. Pollen grains adhering to the stigmatic area of many specimens are monocolpate and tectate with granular to columellate infratectal structure, and are similar to dispersed grains assigned to Tucanopollis and Transitoripollis. Comparison of fossil Appomattoxia ancistrophora with extant plants reveals an unusual combination of characters that includes similarities with some magnoliid taxa, particularly Piperales (Piperaceae, Saururaceae) and Laurales (Chloranthaceae), as well as the monotypic ranunculid family Circaeasteraceae. Appomattoxia ancistrophora differs from extant Piperales in having a pendulous rather than erect ovule, and differs from extant Circaeaster in details of the fruit wall, as well as the presence of monosulcate rather than tricolpate pollen.     

POLLEN MORPHOLOGY AND PLANT TAXONOMY OF WHITE OAKS IN EASTERN NORTH AMERICA

An evaluation of possible approaches to fossil oak pollen identification utilized scanning electron microscopy to examine exine-surface features of 171 collections, representing 16 Quercus subgenus Lepidobalanus species and varieties of eastern North America. Twenty qualitative pollen morphological characters were defined and tabulated for each of 217 pollen grains. The data were subjected to cluster analysis and cluster diagrams were compared with published white oak taxonomy. Pollen morphology and plant taxonomy compared well in series of the subgenus Lepidobalanus due primarily to consistency of character presence and absence within species and varieties. Pollen morphology of white oaks appears to reflect plant systematics above the species level. Use of routine SEM analysis to identify series of white oaks among fossil pollen grains likely will yield valid results.     

New insights into the pollen morphology of the genus Gunnera (Gunneraceae)

Pollen grains from ten species of Gunnera, chosen to represent the six different subgenera in the genus, were examined using light and scanning electron microscopy. The aim of the study was to explore characters that have the potential to define different types of pollen within Gunnera and to study the evolution of these characters in light of the phylogeny of the genus. According to our results, there are three main types of pollen in the examined species of Gunnera. Type 1, unique for the South American species G. herteri (subgenus Ostenigunnera), is characterised by an imperfect reticulum with sinuous undulating-creasted muri. A reticulum with equidimensional polygonal lumina is typical for the plesiomorphic type of pollen (type 2) present in subgenera Gunnera, Misandra and Panke. Lastly, pollen grains of subgenera Pseudogunnera and Milligania are characterised by a reticulum with lumina of variable shape and size (type 3). In G. macrophylla (subgenus Pseudogunnera), the lumina in the apocolpia are of a different shape and size from the lumina in the mesocolpia (type 3a), while in G. dentata, G. monoica and G. cordifolia (subgenus Milligania), the lumina are identical in the apocolpium and the mesocolpium (type 3b).

The identification of pollen types will possibly allow the interpretation of the different specimens of Tricolpites reticulatus, the fossil species believed to be allied to the extant Gunnera.

In addition to the revision on the pollen of Gunnera, a brief comparison between the pollen of this genus and its sister group Myrothamnus is reported.     

Origin and divergence of Afro-Indian Picrodendraceae: linking pollen morphology,dispersal modes,fossil records,molecular dating and paleogeography

The pantropical Picrodendraceae produce mostly spheroidal to slightly oblate, echinate pollen grains equipped with narrow circular to elliptic pori that can be hard to identify to family level in both extant and fossil material using light microscopy only. Fossil pollen of the family have been described from the Paleogene of America, Antarctica, Australia, New Zealand, and Europe, but until now none have been reported from Afro-India. Extant pollen described here include representatives from all recent Picrodendraceae genera naturally occurring in Africa and/or Madagascar and south India and selected closely related tropical American taxa. Our analyses, using combined light microscopy and scanning electron microscopy, show that pollen of the Afro-Indian genera encompass three morphological types: Type 1, comprising only Hyaenanche; Type 2, including Aristogeitonia, Mischodon, Oldfieldia and Voatamalo; Type 3, comprising the remaining two genera, Androstachys and Stachyandra. Based on the pollen morphology presented here it is evident that some previous light microscopic accounts of spherical and echinate fossil pollen affiliated with Arecaceae, Asteraceae, Malvaceae, and Myristicaceae from the African continent could belong to Picrodendraceae. The pollen morphology of Picrodendraceae, fossil pollen records, a dated intra-familial phylogeny, seed dispersal modes, and the regional Late Cretaceous to early Cenozoic paleogeography, together suggest the family originated in the Americas and dispersed from southern America across Antarctica and into Australasia. A second dispersal route is believed to have occurred from the Americas into continental Africa via the North Atlantic Land Bridge and Europe.     

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.     

Normapolles flowers of fagalean affinity from the Late Cretaceous of Portugal

 Exquisitely preserved, charcoalified fossil flowers with in situ pollen of the Normapolles-type from the Late Cretaceous of Portugal are described and a new genus and species of Fagales, Normanthus miraensis, are established. Floral organization and structure of floral organs were studied with scanning electron microscopy and microtome sections. Flowers are actinomorphic, epigynous, and pentamerous; the perianth is simple; stamens alternate with tepals; pollen is oblate and vestibulate; the exine is thick and the tectum is scabrate-microgranulate; the gynoecium is bicarpellate and unilocular; the fruits are probably one-seeded. Comparisons with extant taxa demonstrate that N. miraensis shares many similarities with Fagales and in particular with Betulaceae. However, it is not identical with any extant taxon and cannot be included in any extant family. The combination of characters found in the fossil flowers is congruent with wind-pollination syndromes present in many extant angiosperms and clearly indicates wind-pollination of N. miraensis. Received June 13, 2000 Accepted September 28, 2000     

Elatides sandaolingensis n. sp. (Cupressaceae sensu lato) — A new fossil conifer with cones from the Middle Jurassic of Xinjiang,northwestern China

Here we describe a new conifer fossil, Elatides sandaolingensis Z.X. Wang and B.N. Sun n. sp., from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China. The materials consist of compressions represented by well-preserved leafy shoots, pollen cones, and seed cone. Leaves are characterized by long triangular shapes, with straight apex and entire margins, and two stomatal bands on the abaxial surface. Pollen cones are terminally disposed on the ultimate leafy shoots, borne singly or in clusters. Pollen sacs are long-oval shaped, with three pollen sacs fused together. Pollen grains are spherical and have small germinal papilla and few wrinkles. The seed cone is oblong, with more than 35 helically arranged bract-scale complexes, which are characterized by long-oval shape and triangular apex. Compared with the extant nine genera of Taxodiaceous Cupressaceae in the morphology of seed cone and pollen cones, the present fossil consistently shares many characteristics with the extant genus Cunninghamia, but differs in other aspects. After being compared with the reported fossil records of the fossil plants, the current species is found to be different from any known species; thus, the present fossil is referred to as a new species of Elatides. From the similarity between the present fossil and Cunninghamia, it can be inferred that there may be a genetic relationship between these two genera. Additionally, the new species has thin cuticles and slightly sunken stomata, which can provide evidence indicating that the climate of the Turpan-Hami Basin in Middle Jurassic might have been warm and humid. By studying the geological history and geographical distribution of Elatides, it can be inferred that this genus may have originated in Switzerland, and it was migrated from Xinjiang to North and South China through the Ural Mountains.     

FIRST FOSSIL EVIDENCE FOR THE PRIMITIVE ANGIOSPERM FAMILY LACTORIDACEAE

Pollen of the primitive angiosperm family Lactoridaceae has been recovered from Turonian-Campanian (Upper Cretaceous) sediments from eleven boreholes off the southwest coast of southern Africa. This is the first report of the Lactoridaceae in the fossil record. The one extant species of the Lactoridaceae is confined to the Juan Fernandez Islands located off the coast of Chile. The occurrence of lactoridaceous pollen in Cretaceous deposits of southern Africa suggest that this primitive angiosperm family may have been more widespread in the Southern hemisphere during the Cretaceous.     

Upatoia barnardii gen. et sp. nov., an araucarian pollen cone with in situ pollen from the Late Cretaceous (Santonian) of Georgia,USA

Upatoia barnardii gen. et sp. nov., a conifer pollen cone from the Late Cretaceous (Santonian) Eutaw Formation of Upatoi Creek, Georgia, USA, is known from lignified and fusainised mesofossils that preserve its three-dimensional structure. The cone consists of numerous helically arranged microsporophylls, each composed of a thin stalk and distal lamina. Three elongate pollen sacs are attached to the base of the lamina. Pollen grains isolated from the pollen sacs are relatively large (52 – 75 μm), spheroidal to ellipsoidal in outline, lack sacci, and have a thickened equatorial exine that is often strongly folded. Pollen of Upatoia barnardii indicates a close relationship to extant Araucariaceae. Microsporophylls of U. barnardii confirm suggestions from previous studies of fossil material that some Mesozoic Araucariaceae had only three pollen sacs per microsporophyll, in contrast to extant species that often have more than ten pollen sacs per microsporophyll.     

A scolopocryptopid centipede (Chilopoda: Scolopendromorpha) from Mexican amber: synchrotron microtomography and phylogenetic placement using a combined morphological and molecular data set

The first scolopocryptopid centipede known from the fossil record is a specimen of the subfamily Scolopocryptopinae in Miocene amber from Chiapas, southern Mexico. It is described here as Scolopocryptops simojovelensis sp. nov. , displaying a distinct combination of morphological characters compared to extant congeners. Anatomical details of the fossil specimen were acquired by non‐invasive 3D synchrotron microtomography using X‐ray phase contrast. The phylogenetic position of the new species is inferred based on a combination of morphological data with sequences for six genes (nuclear 18S and 28S rRNA, nuclear protein‐coding histone H3, and mitochondrial 12S rRNA, 16S rRNA, and protein‐coding cytochrome c oxidase subunit I) for extant Scolopendromorpha. The data set includes eight extant species of Scolopocryptops and Dinocryptops from North America, east Asia, and the Pacific, rooted with novel sequence data for other blind scolopendromorphs. The molecular and combined data sets, analysed in a parsimony/direct optimization framework, identified a stable pattern of two main clades within Scolopocryptopinae. North American and Asian species of Scolopocryptops are united as a clade supported by both morphological and molecular characters. Its sister group is a Neotropical clade in which the type species of Dinocryptops is nested within a paraphyletic assemblage of Scolopocryptops species; Dinocryptops is placed in synonymy with Scolopocryptops. The strength of support for the relationships of extant taxa from the molecular data allow the Chiapas fossil to be assigned with precision, despite ambiguity in the morphological data; the fossil is resolved as sister species to the extant Laurasian clade. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 768–786.     

Establishing a fossil record for the perianthless Piperales: Saururus tuckerae sp. nov. (Saururaceae) from the Middle Eocene Princeton Chert

Investigations of small permineralized flowers from the Middle Eocene Princeton Chert, British Columbia, Canada have revealed that they represent an extinct species of Saururus. Over 100 flowers and one partial inflorescence were studied, and numerous minute perianthless flowers are borne in an indeterminate raceme. Each flower is subtended by a bract, and flowers and bracts are borne at the end of a common stalk. Five stamens are basally adnate to the carpels. Pollen is frequently found in situ in the anthers. Examined under SEM and TEM, pollen grains are minute (6-11 μm), monosulcate, boat-shaped-elliptic, with punctate sculpturing and a granulate aperture membrane. The gynoecium is composed of four basally connate, lobed carpels with recurved styles and a single ovule per carpel. Flower structure and pollen are indicative of Saururaceae (Piperales), and in phylogenetic analyses using morphological characters, the fossils are sister to extant Saururus. The fossil flowers are described here as Saururus tuckerae sp. nov. These fossil specimens add to the otherwise sparse fossil record of Piperales, represent the oldest fossils of Saururaceae as well as the first North American fossil specimens of this family, and provide the first evidence of saururaceous pollen in the fossil record.     

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.     

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.     

Pollen morphology of Chamaebuxus (DC.) Schb., Chodatia Paiva and Rhinotropis (Blake) Paiva (Polygala L., Polygalaceae)

Polygala L. is a large and highly diverse genus with complex taxonomy, but pollen morphological information for this taxon is scarce. In the present study, pollen characters have been used to assess the taxonomic delimitation and phylogenetic relationships of three newly established subgenera of Polygala: Chamaebuxus, Chodatia and Rhinotropis (sensu Paiva). The pollen morphology of 22 species has been examined using light microscopy and scanning electron microscopy of acetolysed material. The pollen of 15 of the species is examined for the first time. The pollen grains are isopolar, radially symmetrical, tectate and, typically, polyzonocolporate with numerous colpi running parallel to the polar axis, and an endocingulum around the equator. Two pollen types can be distinguished: Type I, which includes species belonging to Rhinotropis, and Type II, which includes species from Chamaebuxus and Chodatia. The two pollen types are described and the pollen of the three studied subgenera is illustrated. Despite the low infrageneric morphological diversity observed within the genus Polygala, quantitative characters of pollen grains support the current classification of the subgenera Chamaebuxus, Chodatia and Rhinotropis, and reveal a closer relationship between the first two taxa. Pollen characters are shown to be a useful and informative tool for assessing taxonomic position and phylogenetic relationships within Polygalaceae, especially at higher taxonomic levels.     

The oogonial aperture of Oedogonium decipiens var. decipiens (Oedogoniales, Chlorophyta) and its systematic significance

Pollen morphology of ten Brazilian species within the South American clade (ca. 20 species) of Symplocos section Barberina (Symplocaceae) was analyzed with scanning electron microscopy to assess their reported androdioecious breeding system. All species exhibited pollen dimorphism. Pollen from male individuals is well developed and 3-colporate, whereas that from morphologically hermaphroditic individuals is malformed, often completely fragmented, lacks cytoplasm, and has no germination pores. Our results suggest that the morphologically hermaphroditic species of S. section Barberina with malformed pollen are cryptically dioecious.     

POLLEN MORPHOLOGY AND DETAILED STRUCTURE OF FAMILY COMPOSITAE,TRIBE CICHORIEAE. I. SUBTRIBE STEPHANOMERIINAE

A survey of pollen morphology in 20 species representing the 11 genera of the North American subtribe Stephanomeriinae by light, scanning electron, and transmission electron microscopy revealed 10 of the 11 genera to have echinate, tricolporate pollen grains, Lygodesmia being the only genus with echinolophate pollen. Sectioned exines of most of the species examined are similar, being composed of ektexine and endexine. The ektexine surface is composed of spines which typically have globose perforate bases. A cavus occurs as a separation between the basis (foot layer) and the columellae in all of the genera examined except Chaetadelpha. Pollen of the two species of Glyptopleura were found to be strikingly different in exomorphology. Pollen of the putatively self-fertile G. marginata has much shorter spines than the closely related G. setulosa. Atrichoseris, Anisocoma, Calycoseris, Glyptopleura, Pinaropappus, Prenanthella, and most species of Malacothrix have pollen which lack paraporal ridges. The remaining genera, Chaetadelpha, Lygodesmia, Rafinesquia, and Stephanomeria have well-developed ridges of fused spine bases around the apertures. Pollen characters, particularly those of the aperture region, have been found to be systematically useful in the subtribe, therefore acetolyzed material gives more useful information than untreated pollen.     

The metareticulate pollen morphology of Alternanthera Forssk. (Gomphrenoideae,Amaranthaceae) and its taxonomic implications

Alternanthera (Amaranthaceae) is a diverse genus largely restricted to the American Tropics that belongs to the alternantheroid clade containing C₄ and C₃–C₄ intermediate species. This research focuses on the study of pollen characters by studying 13 species, representatives of the two major clades and subclades of Alternanthera. General palynological comparisons were conducted with light microscopy (LM), scanning electron microscopy (SEM) and with confocal laser scanning microscopy (CLSM) for exine ultrastructure. Twenty-five characters were measured and described for Alternanthera and among these, 14 pollen characters were used to discriminate pollen groups using cluster analysis and canonical analysis of principal coordinates (CAP). Pollen form and ornamentation, pores number, spines length, number of ektexinous bodies and nanospines on the ektexinous bodies on pore membranes, arrangement of nanopores and spines on structural elements, and metareticula form were taxonomically important and therefore used to construct the first palynological key to the alternantheroid clade species. Our study indicates that the seemingly subtle morphological variation of pollen is useful for recognising three main pollen types within Alternanthera. The much needed palynological terminology for describing the mesoporium in the metareticulate pollen of Amaranthaceae is provided.     

POLLEN MORPHOLOGY OF THE CHLORAEINAE (ORCHIDACEAE: DIURIDEAE) AND RELATED SUBTRIBES

Pollen organization and morphology of the South American Chloraeinae (Orchidaceae) was examined by scanning electron microscopy and compared with that of the remainder of the otherwise Australasian Diurideae. All five genera of the Chloraeinae, Bipinnula, Chloraea, Codonorchis, Gavilea, and Geoblasta, and at least one genus from each of the other subtribes were sampled. The Australasian Diurideae are diverse in pollen organization and morphology. The two genera of the Acianthinae, Corybas and Acianthus, have very different pollen and their classification is questioned. Monad pollen organization of Pterostylis (Pterostylidinae) is reinterpreted as primitive and not secondarily derived. Pollen of the Chloraeinae is uniform in exine morphology and organization. Most species sampled have reticulate pollen which tends to be foveolate distally. The basic pollen unit of all Chloraeinae is the tetrad, except Codonorchis which possesses monads. Pollen morphology and organization of the Chloraeinae is most similar to the Caladeniinae, which supports the contention that the Chloraeinae including Codonorchis should be retained in the Diurideae.     

Panbiogeography of Nothofagus (Nothofagaceae): analysis of the main species massings

Aim The aim of this paper is to analyse the biogeography of Nothofagus and its subgenera in the light of molecular phylogenies and revisions of fossil taxa. Location Cooler parts of the South Pacific: Australia, Tasmania, New Zealand, montane New Guinea and New Caledonia, and southern South America. Methods Panbiogeographical analysis is used. This involves comparative study of the geographic distributions of the Nothofagus taxa and other organisms in the region, and correlation of the main patterns with historical geology. Results The four subgenera of Nothofagus have their main massings of extant species in the same localities as the main massings of all (fossil plus extant) species. These main massings are vicariant, with subgen. Lophozonia most diverse in southern South America (north of Chiloé I.), subgen. Fuscospora in New Zealand, subgen. Nothofagus in southern South America (south of Valdivia), and subgen. Brassospora in New Guinea and New Caledonia. The main massings of subgen. Brassospora and of the clade subgen. Brassospora/subgen. Nothofagus (New Guinea–New Caledonia–southern South America) conform to standard biogeographical patterns. Main conclusions The vicariant main massings of the four subgenera are compatible with largely allopatric differentiation and no substantial dispersal since at least the Upper Cretaceous (Upper Campanian), by which time the fossil record shows that the four subgenera had evolved. The New Guinea–New Caledonia distribution of subgenus Brassospora is equivalent to its total main massing through geological time and is explained by different respective relationships of different component terranes of the two countries. Global vicariance at family level suggests that Nothofagaceae/Nothofagus evolved largely as the South Pacific/Antarctic vicariant in the breakup of a world‐wide Fagales ancestor.