A new species of Leptocycas(Zamiaceae)from the Upper Triassic sediments of Liaoning Province,China

A new cycad,Leptocycas yangcaogouensis sp.nov.,was found in sediments from the Late Triassic in western Liaoning,China.The pinnately compound leaves(Pseudoctenis type)are screwed in a crown on the stem top.The leaflets are linear,with parallel veins and decurrent bases on the rachis.The leaf bases are persistent.The cataphylls intermix with the leaves.The female cone is ovoid in shape.The characteristics of the new plant are more similar to those of Leptocycas gracilis,a Triassic cycad from North America,but the new species differs from L.gracilis in the size of its stem(7-8 vs.3-5 cm in diameter,respectively),leaves(length × width 100 × 16 vs.30 × 7 cm,respectively)and leaf density along the stem(4-6 vs.1-2 bases/1 cm length,respectively).Both L.gracilis and L.yangcaogouensis,having leaves of the Pseudoctenis type,show a closer relationship to the extant Dioon of Zamiaceae.The present study provides evidence for the origin of the genus Dioon,which may have come from Leptocycas plants of the Triassic.It would be assumed that the extent cycads in Zamiaceae originate from the pteridosperms in the Late Paleozoic and have evolved through the stage of L.gracilis and L.yangcaogouensis in Late Triassic,and reaching the extant Dioon.     

KARYOTYPIC DATA ON NORTH AND CENTRAL AMERICAN ZAMIACEAE (CYCADALES) AND THEIR PHYLOGENETIC IMPLICATIONS

Chromosome numbers and karyotypes of species from four American Zamiaceae (Cycadales) are reported. Zamia shows interspecific and intraspecific chromosome variation, whereas Microcycas, Ceratozamia, and Dioon have constant karyotypes within each genus. In Zamia, all karyotypes have the same number of submetacentric and acrocentric chromosomes, but they differ in the number of metacentric and telocentric chromosomes. Centric fission of metacentric chromosomes is proposed to explain the karyotypic variation in this genus. Zamia shows karyological relationships with Microcycas and Ceratozamia, whereas Dioon appears very distinct from the other American cycad genera. Affinity among Zamia, Ceratozamia, and Microcycas karyotypes and distinctiveness of Dioon karyotypes are supported by comparative analysis of phenotypic characters in the four genera.     

Molecular evidence of cycad seed predation by immature Aulacoscelidinae (Coleoptera: Orsodacnidae)

Adult beetles in the small subfamily Aulacoscelidinae (superfamily Chrysomeloidea) are known to feed on the foliage and juices of New World cycads (Order Cycadales; family Zamiaceae), but the habits of larvae have long remained a mystery. We provide the first direct evidence that Aulacoscelidinae larvae feed on and develop within the megagametophyte of the Mesoamerican cycad, Dioon merolae (Zamiaceae). Phylogenetic analyses based on partial DNA sequences from 3 genes recover a cycad seed‐feeding larva proposed to belong to Aulacoscelidinae. These observations reveal a more intimate feeding relationship between Aulacoscelidinae and their New World cycad host plants than was previously recognized. Further, adult Aulacoscelidinae have long been noted to resemble Jurassic fossil chrysomeloids in the extinct subfamily Protoscelidinae. The molecular, morphological, ecological and fossil data reported herein are broadly compatible with an early association between Aulacoscelidinae and their gymnosperm hosts.     

LYSSOXYLON GRIGSBYI,A CYCAD TRUNK FROM THE UPPER TRIASSIC OF ARIZONA AND NEW MEXICO

Lyssoxylon grigsbyi Daugherty, a petrified stem with petiole bases, was originally described from the Upper Triassic Chinle Formation of Arizona and considered to be a member of the Williamsoniaceae. Investigation of additional material from a similar horizon in New Mexico, together with re-examination of preparations of the holotype, suggest that the plant, with its monoxylic stele, girdling leaf traces, and bicelled epidermal hairs is a true cycad. Cells of the New Mexico specimens contain structures interpreted as preserved nuclei.     

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.     

Onosma maculata sp. nov. (Boraginaceae) from Iran

Onosma maculata Ranjbar & Almasi, a new Boraginaceae species endemic to Iran, is described and illustrated. The new species belongs to Onosma sect. Onosma subsect. Onosma. It is closely related to O. nervosa Riedl, but is easily distinguished by its spotted stem and peduncle (vs no spots), leaves 14–23 cm long (vs leaves 4–12 cm long), pedicel 5–15 mm long (vs pedicel 3–5 mm long), higher density of hairs, and glabrous nutlet (vs villous). Moreover, meiotic chromosome number and behavior were studied in two populations of the new species and it was found to be diploid with 2n = 2x = 16.     

A new Neocalamites (Sphenophyta) with prickles and attached cones from the Upper Triassic of China

Remains of the extinct sphenophyte (horsetail) Neocalamites are most widespread in the Middle–Upper Triassic and are typically represented by stem and leaf fragments. Here we report on spectacular new finds of Neocalamites from the Late Triassic Yangcaogou Formation in Liaoning Province, China that include bedding surfaces dominated by nearly complete aerial stems with attached leaf whorls and rare bractless cones. They reveal a monopodial growth habit for the stems, which are covered with downward projecting prickles that probably provided protection against herbivores. These features provide the basis for a new proposed species, Neocalamites horridus. The nodes bear whorls of very long leaves mainly free to their bases, and one specimen bears an attached cone on a long peduncle. Identical dispersed cones have also been recovered. The leaves of adjacent monopodial stems most likely interlocked to support growth in large stands akin to the role now played by branches in large modern Equisetum species. The new Chinese Neocalamites is among the most confidently reconstructed species, and indicates a greater diversity of sphenophyte morphology during the Mesozoic than previously realized.     

Isolation,characterization and cross‐species amplification of microsatellite loci in the cycad genus Dioon (Zamiaceae). Potential utilization in population genetics studies of Dioon edule

Dioon edule (Zamiaceae) is an endemic Mexican cycad. Nineteen microsatellite loci were isolated from three enriched genomic libraries of D. edule var. angustifolium, D. tomasellii, and D. caputoi. Seven of these loci showed polymorphisms in D. edule. Levels of polymorphism were assessed using 16 individuals from each of seven populations throughout the range of this species. The number of alleles per locus ranged from two to five and the observed and expected heterozygosities ranged from 0.0 to 0.9821 and from 0.0088 to 0.6318, respectively. All loci show significant linkage disequilibrium. Three loci depart significantly from Hardy–Weinberg equilibrium.     

Pseudodioon akyoli gen. et sp. nov., an extinct member of Cycadales from the Turkish Miocene

Fossil cycadalean leaves recorded from the Miocene plant assemblage of Soma, western Turkey, are described and assigned to an extinct genus and species, Pseudodioon akyoli. Leaf macromorphology suggests affinity with members of the Zamiaceae (subfam. Encephalartoideae), particularly with modern Dioon. Micromorphological features on the other hand indicate affinity with modern Cycas (Cycadaceae). Ordinary cells on the adaxial epidermis are isodiametric and are not differentiated into thick- and thin-walled cells. This is similar to Encephalartoideae-like fossils reported from the Cenozoic of the Northern Hemisphere, and even from the Mesozoic. Shared macromorphological traits of P. akyoli and other coeval Encephalartoideae-like fossil cycadalean leaves from Europe suggest that an extinct group of cycads inhabited southern Europe from the western part of Turkey, through Greece and France to Switzerland in the north during the Oligo-Miocene.     

A new scanilepiform from the Lower Triassic of northern Gansu Province,China, and phylogenetic relationships of non‐teleostean Actinopterygii

A new scanilepiform, Beishanichthys brevicaudalis gen. et sp. nov. , is named and described based on fossils from the Lower Triassic lake deposits exposed in Beishan area, Gansu Province, China. The discovery documents a new record of this group, which is significantly older than other known scanilepiforms from China, and is slightly younger than Evenkia from the Lowest Triassic of Central Siberia. Although the Beishan beds were previously interpreted as Late Permian in age, based on megaplant fossils, this new discovery supports the reinterpretation of the deposits as Early Triassic in age, based on vertebrate fossils from the same locality and horizon. Phylogenetic analysis was conducted to resolve the relationships of Scanilepiformes with other actinopterygian clades, and the inter‐relationships within Scanilepiformes. Contrary to previous thought that scanilepiforms are closely related to the Amiidae, the phylogenetic results of this study recognize the Scanilepiformes as stem‐group neopterygians. Relationships of the Scanilepiformes and Australosomus with other neopterygians remain unresolved. With a characteristic long‐based dorsal fin, scanilepiforms represent a small group that emerged in Early Triassic freshwater environments, inhabited Eurasia and North America during the Middle–Late Triassic, briefly invaded the marine environment by the Late Triassic in Europe, and became extinct at the end of Triassic. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 595–612.     

Cataphylls of the Middle Triassic cycad Antarcticycas schopfii and new insights into cycad evolution

Cataphylls associated with the Middle Triassic stem genus Antarcticycas are described, and their impact on understanding cycad evolution is discussed. The cataphylls of Antarcticycas are triangular in outline and flattened adaxially with lateral flanges. The outer surfaces are covered with a ramentum of filamentous hairs, the epidermis is a single cell layer thick, and the ground tissue is parenchymatous with mucilage canals and sclereids. Vascular bundles form a distinct inverted omega-shaped pattern characteristic of the Cycadales observed in petioles of extant species. The structures in Antarcticycas are interpreted as cataphylls based on overall morphology, presence of straight vascular strands in the cortex of the associated stem, and lack of fascicular cambia in the vascular bundles. Because much of the overall diversity of Cycadales is represented by fossils, integrating fossil taxa into explicit phylogenetic hypotheses is important for understanding cycad evolution. Therefore, character and minimum age mapping were performed on a phylogeny of extant and fossil taxa including Antarcticycas. The results suggest that major extant lineages of Cycadales had diverged by the Permian to Triassic and that certain synapomorphies for Cycadales had evolved by the Permian. Evidence of insect feeding on Antarcticycas suggests that associations between cycads and insects are ancient.     

Memecylon trunciflorum sp. nov. (Melastomataceae–Olisbeoideae) from the Udzungwa Mountains,southern Tanzania

Described and illustrated is Memecylon trunciflorum R. D. Stone, an evidently localized endemic of the Udzungwa Mountains in southern Tanzania. The new species was previously confused with the vegetatively similar but distantly related M. erythranthum Gilg and M. semseii A. Fern. & R. Fern., from which it is distinguished by its anther connectives bearing a dorsal oil‐gland and by its ellipsoid to obovoid fruits (vs anther connective gland absent and fruits globose in M. erythranthum and M. semseii). The new species is placed in M. sect. Magnifoliata R. D. Stone together with M. magnifoliatum A. Fern. & R. Fern., from which it differs by its smaller leaves mostly 9.5–15.0 × 3.5–6.0 cm (vs 18–35 × 8–13 cm), transverse veins 8–18 pairs (vs 25–28 pairs), short‐pedunculate inflorescences with secondary axes well developed (vs peduncles and secondary axes absent), white flowers (vs bluish purple), and smaller fruits mostly 11.5–14.5 × 9–11 mm on longer fruiting pedicels 8.0–13.5 mm (vs fruits 17–20 × 12–14 mm on pedicels 5.0–7.5 mm). Despite its local endemism, Memecylon trunciflorum has been assessed as ‘Least Concern’ according to IUCN criteria, although this assessment is dependent on the continued safeguarding of the Udzungwa Mountains National Park.     

Heterozygote excess in ancient populations of the critically endangered Dioon caputoi (Zamiaceae,Cycadales) from central Mexico

Dioon caputoi is a long‐lived cycad known from only four populations that range in size from 50 to 120, mostly adult individuals. Dioon caputoi has the most narrow geographical range of all Dioon spp. (less than 10 km), existing completely within the boundaries of the Tehuacán–Cuicatlán Biosphere Reserve, Mexico. Negative inbreeding values were found in all four populations (F_IT = ?0.242) and within subpopulations (F_IS = ?0.379). Only c. 10% of the total genetic variation was partitioned among populations (F_ST = 0.099). We also found that most mean values of genetic variation (A = 1.91 ± 0.12; P = 78.9 ± 10.2; H_E = 0.35 ± 0.01) are within the range reported for other Dioon species with larger populations and with wider geographical ranges. These results support recent findings that rare plant species maintain high levels of genetic diversity. The heterozygote excess found at all loci is discussed in detail from a neutral evolutionary perspective, leaving arguments as working hypotheses for further research. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 436–447.     

COMPARATIVE POLLEN MORPHOLOGY AND TAXONOMIC AFFINITIES IN CYCADALES

Scanning and transmission electron microscopy of pollen grains of 29 species, representing the ten extant genera of Cycadales, has provided valuable insight into their relationships. Pollen grains of these taxa are boat-shaped, monosulcate, and bilaterally symmetrical. They range from narrowly to widely elliptical or subcircular when viewed distally, and have an exine surface of psilate, foveolate, or fossulate. Pollen wall ultrastructure of Cycadales is typically tectate with alveolate∗∗∗spongy exine. The nexine is laminated in all genera. Nexine 1 (footlayer) is present in most species as a thin and often discontinuous layer. There is consistent variation in thickness of the sporoderm layers among the genera but relative uniformity within them. Pollen characteristics are well correlated with macro- and micromorphological features, chromosome numbers, geographical distribution, and postulated pollination mode. A close affinity between Encephalartos, Lepidozamia, and Macrozamia is recognized. Pollen characteristics of the genus Bowenia show some similarity with those of the latter group. Except for two species of Macrozamia which are narrowly elliptic, all of the genera have widely elliptic pollen and share a psilate exine surface and the thinnest sexine with nearly identical arrangement of alveoli. Pollen grains of the species in the genus Dioon exhibit a unique morphology but are more similar to Stangeria than they are to those of taxa in Zamiaceae. The circular outline of the grains and the foveolate exine surface are characters shared by these two genera, but several morphological features distinguish Dioon from Stangeria. Ceratozamia and Zamia share a widely elliptic shape, foveolate exine surface and nearly identical sexine, as well as morphological features and chromosome numbers. They differ from Microcycas in sexine thickness, gross morphology and chromosome numbers. The pollen grains of Cycas circinalis and C. revoluta differ in size and structure of the sexine from all other genera and from each other, substantiating their distinct subgeneric delimitations.     

Phylogeny of the cycads based on multiple single-copy nuclear genes: congruence of concatenated parsimony,likelihood and species tree inference methods

Background and aims

Despite a recent new classification, a stable phylogeny for the cycads has been elusive, particularly regarding resolution of Bowenia, Stangeria and Dioon. In this study, five single-copy nuclear genes (SCNGs) are applied to the phylogeny of the order Cycadales. The specific aim is to evaluate several gene tree–species tree reconciliation approaches for developing an accurate phylogeny of the order, to contrast them with concatenated parsimony analysis and to resolve the erstwhile problematic phylogenetic position of these three genera.

Methods

DNA sequences of five SCNGs were obtained for 20 cycad species representing all ten genera of Cycadales. These were analysed with parsimony, maximum likelihood (ML) and three Bayesian methods of gene tree–species tree reconciliation, using Cycas as the outgroup. A calibrated date estimation was developed with Bayesian methods, and biogeographic analysis was also conducted.

Key Results

Concatenated parsimony, ML and three species tree inference methods resolve exactly the same tree topology with high support at most nodes. Dioon and Bowenia are the first and second branches of Cycadales after Cycas, respectively, followed by an encephalartoid clade (Macrozamia–Lepidozamia–Encephalartos), which is sister to a zamioid clade, of which Ceratozamia is the first branch, and in which Stangeria is sister to Microcycas and Zamia.

Conclusions

A single, well-supported phylogenetic hypothesis of the generic relationships of the Cycadales is presented. However, massive extinction events inferred from the fossil record that eliminated broader ancestral distributions within Zamiaceae compromise accurate optimization of ancestral biogeographical areas for that hypothesis. While major lineages of Cycadales are ancient, crown ages of all modern genera are no older than 12 million years, supporting a recent hypothesis of mostly Miocene radiations. This phylogeny can contribute to an accurate infrafamilial classification of Zamiaceae.     

A PHYLOGENETIC ANALYSIS OF DIOON (ZAMIACEAE)

A phylogenetic analysis of all the intrageneric taxa of the genus Dioon Lindley (Zamiaceae) was undertaken by using chloroplast DNA restriction fragment length polymorphism analysis. Wagner parsimony analysis on a 187 character matrix yielded two equally parsimonious trees, differing only for the position of D. caputoi. The consensus tree has two well-defined major clades. The first is composed of D. mejiae, D. rzedowskii, and D. spinulosum; the second is composed of D. califanoi, D. caputoi, D. edule var. angustifolium, D. edule var. edule, D. holmgrenii, D. merolae, D. purpusii, D. tomasellii var. sonorense, and D. tomasellii var. tomasellii. A phenetic analysis of the same data showed results broadly congruent with the cladistic analysis. This resulting phylogeny is partially congruent with morphological data and is also compatible with the biogeography of the genus. Modem species of Dioon may have evolved as a consequence of a very fast succession of vicariance events that mainly occurred during the early Cenozoic. The short time between each of these events may not have allowed the accumulation of a large number of morphological synapomorphies for the groups of species.     

A surface microrelief on the leaves of Glossophyllum florinii (?Ginkgoales) from the Upper Triassic of Lunz,Austria

The cuticles of many extant seed plants display distinct surface microreliefs, which represent adaptations to certain habitat conditions or mechanical defences against herbivores and phytopathogenic microorganisms. Although microreliefs have variously been noted in fossil cuticles, hypotheses relating to the effectiveness of these structures in fossil plants have not been advanced to date. A surface microrelief composed of longitudinally orientated idiocuticular striae occurs on the leaves of the enigmatic Carnian (Late Triassic) gymnosperm (?ginkgophyte) Glossophyllum florinii Kräusel from the Northern Calcareous Alps of lower Austria. Most striae originate from the tips of the papillae on the stomatal subsidiary cells. The G. florinii surface microrelief may have (1) reduced leaf wettability, (2) produced or enhanced the self‐cleaning effect of the leaf, (3) prevented the formation of a water film on the leaf surface, and/or (4) mechanically stabilized the leaf. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 87–95.