A new genus of male cones of voltzialean affinity,Uralostrobus voltzioides nov. gen., nov. sp., from the Lower Permian of the Urals (Russia)

A new genus and species of male cones of coniferophyte from the Lower Permian (Artinskian and Kungurian) deposits of the Urals, Russia is described: Uralostrobus voltzioides Naugolnykh nov. gen., nov. sp. The cone shows characters typical of some representatives of conifers belonging to the order Voltziales: more or less isometrical bracts of rhombic shape, prolonged sporangia with attenuate apices, and bisaccate pollen of Illinites-type. General information on the associated female seed scales and vegetative leafy shoots is given as well.     

CHILBINIA GEN. NOV., AN ARCHAIC SEED FERN IN THE LATE TRIASSIC CHINLE FORMATION OF ARIZONA, USA

Abstract:  A seed-bearing pinnate leaf found in the Chinle Formation of Late Triassic age in Arizona appears to be an archaic seed fern that more closely resembles typical Carboniferous–Permian seed fern leaves than any younger form. The fossil, which is described here as Chilbinia lichii gen. et sp. nov., incorporates characters of several Palaeozoic taxa. These characters include pinnules that have reticulate venation resembling that in some species of Lonchopteris , cordate bases similar to those found in many species of Sphenopteris , and, except for a significant difference in size, linear ovate seeds that resemble the radially symmetrical seeds found attached to a species of Alethopteris in the Permian of China. The presence of this fossil in strata of Late Triassic age is further evidence that plant extinctions after the Permian–Triassic ecological crisis followed a stepwise pattern and apparently continued for at least 20–25 myr after the end of the Permian in western parts of the Triassic paratropical belt.     

Phylogeny and Divergence Times of Gymnosperms Inferred from Single-Copy Nuclear Genes

Phylogenetic reconstruction is fundamental to study evolutionary biology and historical biogeography. However, there was not a molecular phylogeny of gymnosperms represented by extensive sampling at the genus level, and most published phylogenies of this group were constructed based on cytoplasmic DNA markers and/or the multi-copy nuclear ribosomal DNA. In this study, we use LFY and NLY, two single-copy nuclear genes that originated from an ancient gene duplication in the ancestor of seed plants, to reconstruct the phylogeny and estimate divergence times of gymnosperms based on a complete sampling of extant genera. The results indicate that the combined LFY and NLY coding sequences can resolve interfamilial relationships of gymnosperms and intergeneric relationships of most families. Moreover, the addition of intron sequences can improve the resolution in Podocarpaceae but not in cycads, although divergence times of the cycad genera are similar to or longer than those of the Podocarpaceae genera. Our study strongly supports cycads as the basal-most lineage of gymnosperms rather than sister to Ginkgoaceae, and a sister relationship between Podocarpaceae and Araucariaceae and between Cephalotaxaceae-Taxaceae and Cupressaceae. In addition, intergeneric relationships of some families that were controversial, and the relationships between Taxaceae and Cephalotaxaceae and between conifers and Gnetales are discussed based on the nuclear gene evidence. The molecular dating analysis suggests that drastic extinctions occurred in the early evolution of gymnosperms, and extant coniferous genera in the Northern Hemisphere are older than those in the Southern Hemisphere on average. This study provides an evolutionary framework for future studies on gymnosperms.     

Morphogenesis of the reproductive shoots of Welwitschia mirabilis and Ephedra distachya (Gnetales), and its evolutionary implications

For decades, Gnetales appeared to be closely related to angiosperms, the two groups together forming the anthophyte clade. At present, molecular studies negate such a relationship and give strong support for a systematic position of Gnetales within or near conifers. However, previous interpretations of the male sporangiophores of Gnetales as pinnate with terminal synangia conflict with a close relationship between Gnetales and conifers. Therefore, we investigated the morphogenesis of the male reproductive structures of Welwitschia mirabilis and Ephedra distachya by SEM and light microscopy. The occurrence of reduced apices to both halves of the antherophores of W. mirabilis gives strong support for the assumption that the male ‘flowers’ of W. mirabilis represent reduced compound cones. We assume that each half of the antherophore represents a lateral male cone that has lost its subtending bract. Although both halves of the antherophores of Ephedra distachya lack apical meristems, the histological pattern of the developing antherophores supports interpreting them as reduced lateral male cones as well. Therefore, the male sporangiophores of Gnetales represent simple organs with terminal synangia. Although extant conifers do not exhibit terminal synangia, similar sporangiophores are reported for some Cordaitales, the hypothetical sister group of conifers. Moreover, several Paleozoic conifers exhibit male cones with terminal sporangia or synangia. Therefore, we propose that conifers, Cordaitales and Gnetales originated from a common ancestor that displayed simple sporangiophores with a terminal cluster of sporangia.     

MORE EVIDENCE FOR CONIFER DIVERSITY IN THE UPPER TRIASSIC OF NORTH CAROLINA

An ovulate strobilus from the Upper Triassic Deep River Basin, North Carolina, has helically arranged, loosely aggregated, elongated, spatulate bracts with axillary ovule-bearing appendages with about 8–10 ovules attached in two lateral rows, with outwardly directed micropyles. The axillary ovuliferous appendage is homologous with the voltzialean fertile dwarf shoot, but probably not directly evolved from it. More credible is a suggested origin from a completely fertile axillary appendage such as that of the Lower Permian Trichopitys. The occurrence of this cone, Metridiostrobus palissyaeoides, gen. and sp. nov., along with Compsostrobus neotericus and Voltzia andrewsii, reflects considerable diversity among conifer ovulate cones during the Upper Triassic.     

A new Early Cretaceous relative of Gnetales: Siphonospermum simplex gen. et sp. nov. from the Yixian Formation of Northeast China

Background

Knowledge on fossil and evolutionary history of the Gnetales has expanded rapidly; Ephedra and ephedroids as well as the Gnetum-Welwitschia clade are now well documented in the Early Cretaceous. However, hypotheses on evolutionary relationships among living and fossil species are hampered by restricted knowledge of morphological variation in living groups and recent studies indicate that gnetalean diversity and character evolution may be more complex than previously assumed and involve additional extinct groups (Bennettitales, Erdtmanithecales and unassigned fossil taxa).

Results

Here we describe a new fossil related to Gnetales, Siphonospermum simplex from the Early Cretaceous Yixian Formation, an impression/compression of a reproductive shoot. The slender main axis bears one pair of opposite and linear leaves with primary parallel venation. The reproductive units are ovoid, without supporting bracts and borne on one median and two lateral branches. The most conspicuous feature of the fossil is the long, thread-like micropylar tube formed by the integument. Each ovule is surrounded by two different layers representing one or two seed envelopes; an inner sclerenchymatous layer and an outer probably parenchymatous layer.

Conclusions

The vegetative and reproductive features of Siphonospermum simplex exclude a relationship to any other group than the Gnetales. A combination of opposite phyllotaxis, linear leaves and ovules surrounded by seed envelope(s) and with a long exposed micropylar tube are known only for extant and extinct Gnetales. Siphonospermum simplex constitutes a new lineage within the Gnetales. Its morphology cannot be directly linked to any previously known plant, but the organization of the reproductive units indicates that it belongs to the Gnetum-Welwitschia clade. Based on the absence of cone bracts and the inferred histology of the seed envelope(s) it could be related to Gnetum, however, there are also affinities with the ephedran lineage, some of which are likely plesiomorphic features, others perhaps not. Phylogeny and character evolution in the Bennettitales, Erdtmanithecales and Gnetales are currently only partly understood and under debate; the exact systematic position of Siphonospermum simplex, i.e., its position within the Gnetales, cannot be resolved with certainty.     

Implications of fossil conifers for the phylogenetic relationships of living families

Fossils have played a central role in our understanding of the evolution of conifers. Interpretation of the seed cone as a compound strobilus and the homologies of the ovuliferous scales of modern conifers with the axillary dwarf shoot of Pennsylvanian forms are based on fossils. Similarly, early evolutionary trends involving the reduction, fusion, and planation of the fertile and sterile elements of the axillary dwarf shoot, leading to structures characteristic of modern families, are documented in Late Permian and Triassic conifers. However, a phylogeny elucidating the derivation of modern families from fossil forms based on shared derived features has been elusive. The present cladistic treatment using 11 characters of ovulate cones and one of pollen grains suggests three phylogenetic groups of Late Paleozoic conifers, represented loosely by the Emporicaceae, Utrechtiaceae, and Majonicaceae of Mapes and Rothwell. The Taxaceae appears to have diverged from ancestors within the Utrechtiaceae, whereas the other modern families owe their origins to the Majonicaceae. The origin of the Taxodiaceae appears to have been biphyletic.Taxodium, Cupressus andSciadopitys are strongly linked toDolmitia of the Majonicaceae, butCryptomeria, Cunninghamia andAraucaria are grouped together and diverge basal to the former taxa.Pinus branches from a position basal to the known genera of the Majonicaceae and all modern families except the Taxaceae.Podocarpus also diverges basal toMajonica but may share an ancestor with this genus;Cepahalotaxus diverges basal to theDolmitiaPseudovoltzia subclade but distal toMajonica. Similarly, the Cheirolepidiaceae originated from basal members of the Majonicaceae and shows no close phylogenetic relationship with any modern family. Except for a strong linkage betweenCycadocarpidium and theAraucariaCunninghamia subclade, genera of the Voltziaceae appear to have branched more or less independently from within the Majonicaceae and show no strong affinity with modern conifers. Thus differences between modern conifer families are due mainly to their divergence from different Paleozoic ancestors.     

Turonian Pinaceae of the Raritan Formation, New Jersey

 Fossil vegetative and reproductive structures from deposits of the Raritan Formation in New Jersey (Turonian, Upper Cretaceous, ∼90 MYBP) include ferns, gymnosperms, and angiosperms. Gymnosperms collected from this formation have been known since the beginning of the 20th century. Fossil leaves, wood and seed cones have been are identified as belonging to the Cupressaceae, Pinaceae, and Taxodiaceae. In the present contribution, we describe a series of fossil dwarf shoots, leaves and a pollen cone belonging to the family Pinaceae. Fossils are charcoalified with preserved three-dimensional structure and excellent cellular detail. The dwarf shoots are assigned to a new species Prepinus crossmanensis sp. nov. and to the previously described Pinus quinquefolia Jeffrey. The new species Prepinus crossmanensis differs in size, shape, presence of hypodermis, sclerenchyma and stomata in the cataphylls, and number and shape of needle leaves from previously known species. Also, isolated leaves were found that were assigned to the new species, Prepinus raritanensis sp. nov. The new species is differentiated by the size and shape of stomata, the presence of number of layers of the hypodermis; and the cell-shape and number of layers of the mesophyll and transfusion tissue. A previously undescribed male cone, Amboystrobus cretacicum gen. and sp. nov., has an axis with spirally attached microsporophylls, each bearing two abaxial ovoid microsporangia. The pollen grains are monosulcate and bisaccate (eusaccate), with an elliptical corpus, granulate exine sculpture, and honeycomb-like wall structure. Received March 21, 2000 Accepted November 13, 2000     

Fossils and seed plant phylogeny reanalyzed

In a cladistic analysis of Recent seed plants, Loconte and Stevenson (1990) obtained results that conflict with our 1986 analysis of both extant and fossil groups and argued that fossil data had led us to incorrect conclusions. To explore this result and the general influence of fossils on phylogeny reconstruction, we assembled new “Recent” and “Complete” (extant plus fossil) data sets incorporating new data, advances in treatment of characters, and those changes of Loconte and Stevenson that we consider valid. Our Recent analysis yields only one most parsimonious tree, that of Loconte and Stevenson, in which conifers are linked with Gnetales and angiosperms (anthophytes), rather than with Ginkgo, as in our earlier Recent and Complete analyses. However, the shortest trees derived from our Complete analysis show five arrangements of extant groups, including that of Loconte and Stevenson and our previous arrangements, suggesting that the result obtained from extant taxa alone may be misleading. This increased ambiguity occurs because features that appear to unite extant conifers and anthophytes are seen as convergences when fossil taxa are interpolated between them. All trees found in the Complete analysis lead to inferences on character evolution that conflict with those that would be drawn from Recent taxa alone (e.g., origin of anthophytes from plants with a “seed fern” morphology). These results imply that conclusions on many aspects of seed plant phylogeny are premature; new evidence, which is most likely to come from the fossil record, is needed to resolve the uncertainties.     

Post-Variscan late Palaeozoic Northern Hemisphere gymnosperms: the onset to the Mesozoic

During the Carboniferous gymnosperms became highly diverse. However, several groups became diminished toward the end of the Carboniferous and Permian, and were extinct by the end of the Permian. Relatively few groups managed to survive the Permian—Triassic transition. These remaining forms represent the gymnosperms that developed during the Mesozoic. This paper, that focuses on Euramerican floras, addresses specific problems related to the study of the gymnosperms of the uppermost Carboniferous and Permian and deals with the major floral changes in the latest Carboniferous and Permian. The major late Palaeozoic Northern Hemisphere gymnosperms are briefly discussed and special attention is given to the peltasperms and conifers, two groups that extended from the late Palaeozoic into the Mesozoic.