A new microphyllous plant from the Lower Devonian of Yunnan Province, China

Sterile and fertile shoots of a microphyllous vascular plant are described from the Lower Devonian Posongchong Formation (? upper Siegenian), Yunnan Province, southern China and placed in the new taxon Halleophyton zhichangense gen. et sp. nov. The sterile axes are characterized by conspicuous, contiguous swollen rhomboidal to hexagonal leaf bases and relatively broad spatulate laminae. The relief of the leaf bases is not so pronounced in the fertile areas, which bear numerous cauline sporangia with two equal valves. In vegetative characters the new fossils, although smaller and more fragmentary, resemble Drepanophycus gaspianus but in the absence of information on the fertile parts of the latter and because of the controversy surrounding the concept of the genus Drepanophycus, we decided to erect a new genus. Its relationship to the lycophytes sensu stricto is equivocal as exact details of the relationship of sporangium with leaf and sporangium vascularization remain unknown.     

Evolution and patterning of the ovule in seed plants

The ovule and its developmental successor, the seed, together represent a highly characteristic feature of seed plants that has strongly enhanced the reproductive and dispersal potential of this diverse group of taxa. Ovules encompass multiple tissues that perform various roles within a highly constrained space, requiring a complex cascade of genes that generate localized cell proliferation and programmed cell death during different developmental stages. Many heritable morphological differences among lineages reflect relative displacement of these tissues, but others, such as the second (outer) integuments of angiosperms and Gnetales, represent novel and apparently profound and independent innovations. Recent studies, mostly on model taxa, have considerably enhanced our understanding of gene expression in the ovule. However, understanding its evolutionary history requires a comparative and phylogenetic approach that is problematic when comparing extant angiosperms not only with phylogenetically distant extant gymnosperms but also with taxa known only from fossils. This paper reviews ovule characters across a phylogenetically broad range of seed plants in a dynamic developmental context. It discusses both well-established and recent theories of ovule and seed evolution and highlights potential gaps in comparative data that will usefully enhance our understanding of evolutionary transitions and developmental mechanisms.     

Latisemenia longshania,gen. et sp. nov., a new Late Devonian seed plant from China

The earliest known ovules in the Late Devonian (Famennian) are borne terminally on fertile branches and are typically enclosed in a cupule. Among these ovules are some that have terete integumentary lobes with little or no fusion. Here, we report a new taxon, Latisemenia longshania, from the Famennian of South China, which bears cupulate ovules that are terminal as well as opposite on the fertile axis. Each ovule has four broad integumentary lobes, which are extensively fused to each other and also to the nucellus. The cupule is uniovulate, and the five flattened cupule segments of each terminal ovule are elongate cuneate and shorter than the ovule. Associated but not attached pinnules are laminate and Sphenopteris-like, with an entire or lobate margin. Latisemenia is the earliest known plant with ovules borne on the side of the fertile axis and may foreshadow the diverse ovule arrangements found among younger seed plant lineages that emerge in the Carboniferous. Following the telome theory, Latisemenia demonstrates derived features in both ovules and cupules, and the shape and fusion of integumentary lobes suggest effective pollination and protection to the nucellus. Along with other recent discoveries from China, Latisemenia extends the palaeogeographic range of the earliest seed plants.     

Schischcatella (Fenestrata,Bryozoa) from the Devonian of the Rhenish Massif,Germany

Abstract: The Devonian fenestrate bryozoan, Schischcatella Waschurova, 1964 , possessed colonies in the form of low, erect bifoliate fronds that grew from an encrusting sheet‐like base with autozooecia arranged in biserial, bifurcating rows. This growth habit is unique in fenestrates, which normally had unilaminate arborescent colonies. Originally, Schischcatella was described from the Lower Devonian of Tajikistan. This article describes a new species, S. heinorum sp. nov., from the Middle Devonian of the Eifel (western Rhenish Massif, Germany) with additional material from the Lower Devonian of the Kellerwald (eastern Rhenish Massif, Germany). External and internal morphologies of this bryozoan have been studied using abundant material. The growth habit of Schischcatella suggests a completely different pattern of feeding currents than that in the normal fenestrate colony. The outflow of the filtered water occurred only on edges of colonies between rami. In the absence of chimneys (areas of vertical water expelling), such a functional morphology may have restricted extension of the colony in a distal direction. The evolution of Schischcatella is apparently an example of paedomorphosis, the genus evolved from an unknown semicosciniid species by the early ontogenetic interruption of colony development and further changes in the mode of growth.     

New cryptostomide bryozoans from the Upper Devonian of Transcaucasia and some aspects of the evolution of the order Cryptostomida

New bryozoans of the suborder Ptilodictyina of the order Cryptostomida are described from the Upper Famennian of Transcaucasia: those of the family Worthenoporidae, i.e., a genus Geranopora gen. nov. comprising three species G. fabulosa sp. nov. (type species), G. fida sp. nov., and G. finitima sp. nov., and those of a new family Mysticellidae fam. nov., i.e., a genus Mysticella gen. nov. comprising four species M. labyrinthica sp. nov. (type species), M. laudativa sp. nov., M. sacrosancta sp. nov., and M. ortiva sp. nov. Thus, a Late Devonian part of the evolutionary line of ptilodictyines is revealed that shows their radiation at the levels of families, genera, and species.     

Devonian floral assemblages and plant megafossils from the Iberian Peninsula: A review

This paper presents a review and critical analysis of the literature on Devonian floras of the Iberian Peninsula. Although the known outcrops of Devonian strata in the Iberian Peninsula are marine, in some cases, a few fragmentary remains of vascular plants are associated with faunal remains. Records include largely specimens from the Lower Devonian of Barrancos (Alentejo, Portugal) and the Upper Devonian of Sierra de Hornachos in Badajoz province, southwest Spain; the remainder consists of drifted plant fragments from scattered sites in the Iberian Peninsula ranging in age from Lochkovian to Upper Devonian-Earliest Carboniferous. The vegetation inferred for the Lower Devonian of the Iberian Peninsula is mainly based on palynological data and corresponds to herbaceous types characterised by bryophytes, rhyniophytes (Horneophyton, Cooksonia, Rhynia), trimerophytes (Psilophyton, Pertica and Hostinella), primitive lycophytes (Drepanophycus) and incertae sedis such as Nothia and Chaleuria, all flora that developed near the coast in low-lying and, at least periodically, wet areas. In the Middle Devonian, two vegetation strata can be recognised: herbaceous (Psilophyton) and semi-arboreal (Cladoxylales). Although three levels of vegetation in the Upper Devonian, have been described from outcrops worldwide, the scarce available data from the Iberian Peninsula only indicate an arborescent lycopsid vegetation and species with uncertain botanical affinity such as Sphenopteridium keilhauii Nathorst.     

Root evolution at the base of the lycophyte clade: insights from an Early Devonian lycophyte

Background and Aims The evolution of complex rooting systems during the Devonian had significant impacts on global terrestrial ecosystems and the evolution of plant body plans. However, detailed understanding of the pathways of root evolution and the architecture of early rooting systems is currently lacking. We describe the architecture and resolve the structural homology of the rooting system of an Early Devonian basal lycophyte. Insights gained from these fossils are used to address lycophyte root evolution and homology.Methods Plant fossils are preserved as carbonaceous compressions at Cottonwood Canyon (Wyoming), in the Lochkovian–Pragian (∼411 Ma; Early Devonian) Beartooth Butte Formation. We analysed 177 rock specimens and documented morphology, cuticular anatomy and structural relationships, as well as stratigraphic position and taphonomic conditions.Key Results The rooting system of the Cottonwood Canyon lycophyte is composed of modified stems that bear fine, dichotomously branching lateral roots. These modified stems, referred to as root-bearing axes, are produced at branching points of the above-ground shoot system. Root-bearing axes preserved in growth position exhibit evidence of positive gravitropism, whereas the lateral roots extend horizontally. Consistent recurrence of these features in successive populations of the plant preserved in situ demonstrates that they represent constitutive structural traits and not opportunistic responses of a flexible developmental programme.Conclusions This is the oldest direct evidence for a rooting system preserved in growth position. These rooting systems, which can be traced to a parent plant, include some of the earliest roots known to date and demonstrate that substantial plant–substrate interactions were under way by Early Devonian time. The morphological relationships between stems, root-bearing axes and roots corroborate evidence that positive gravitropism and root identity were evolutionarily uncoupled in lycophytes, and challenge the hypothesis that roots evolved from branches of the above-ground axial system, suggesting instead that lycophyte roots arose as a novel organ.     

A New Jawless Vertebrate from the Middle Devonian of Scotland

A new jawless vertebrate, Cornovichthys blaauweni gen. et sp. nov., is described from a single complete specimen from the Achanarras fish bed at Achanarras Quarry in Caithness, northern Scotland. The Achanarras fish bed consists of lacustrine laminites and was deposited in a major deep-water phase of the Orcadian lake during the deposition of the cyclic Caithness Flagstone Group. The Achanarras fish bed is of Middle Devonian (Eifelian) age. The new animal compares closely with Euphanerops, one ofthe Frasnian 'anaspids' (Hyperoartii) of the Escuminac Formation at Miguasha, Quebec, Canada.     

Depth-related changes in the arctic epibenthic megafaunal assemblages of Kangerdlugssuaq, East Greenland

The effects of depth-related differences in environmental factors (pressure, hydrography, habitat and disturbance) on the benthic megafauna at the mouth of Kangerdlugssuaq Fjord in Arctic Greenland were investigated using a towed camera platform and autonomous underwater vehicle photographs. Large reductions in faunal density (60,132 to 1881 individuals ha^-1) and increases in diversity (H'=0.93-2.54), through increases in richness (ES₍₂₂₀₎=7.6-18.8) and reductions in dominance (Berger-Parker index=0.77-0.38) were found between 270 and 720 m water depth. Assemblages were separated into distinct shallow, intermediate and deep faunas by multivariate community analysis. In the shallower sites there were high levels of iceberg disturbance, directly reducing diversity as well as creating a complex, patchy environment inhabited by high densities of mobile suspension-feeding epifauna. In the deeper areas there was small-scale disturbance from the deposition of drop stones, but at a relatively low frequency and magnitude, which allowed increased species diversity. Proportions of deposit-feeding epifaunal and infaunal taxa rose with depth, as indicated by an increase with depth in faunal traces. Decreases in faunal density probably resulted from decreased food supply with depth.     

A comparison between the Upper Triassic floras of China and the Rhaeto-Liassic floras of Europe and East Greenland

A survey is given of the geological and geographical distribution of the Upper Triassic floras of China, based chiefly on many articles published in Chinese during the last ten years. This is followed by a comparison between the Upper Triassic flora of China and the Rhaeto-Liassic floras of Europe (Sweden, Germany) and East Greenland. It is pointed out that important floral assemblages (for example that of the Liassic Thaumatopteris schenkii Assemblage-zone) occur considerably earlier in China (especially in S.W. China) than in Europe and East Greenland.     

A cyanolichen from the Lower Devonian Rhynie chert

The 400 million-year-old Rhynie chert has provided a wealth of information about various types of fungal interactions that existed in this Early Devonian paleoecosystem. In this paper we report the first unequivocal evidence of a lichen symbiosis from the Rhynie chert. Specimens of a new genus, Winfrenatia, consist of a thallus of superimposed layers of aseptate hyphae and, on the upper surface, numerous uniform depressions. Extending into the base of each depression are hyphae that form a three-dimensional netlike structure. Enclosed within each of the net spaces is a coccoid cyanobacterium, each cell of which is surrounded by a thick sheath. These photobiont cells divide in three planes, resulting in cell clusters of up to perhaps 64 individuals. The photobiont is parasitized by the fungus in the base of each net as new cyanobacterial cells are formed distally. Reproduction is by endospores and soredia. Affinities of the mycobiont appear closest to members of the Zygomycetes, while the photobiont is most similar to coccoid cyanobacteria of the Gloeocapsa and Chroococcidiopsis types. We speculate that this cyanobacterial symbiosis was well adapted to exploit and colonize new ecological niches, especially in the periodically desiccated environment postulated for the Rhynie chert paleoecosystem.     

Fossil plants with spores in the sporangia from the Upper Devonian (Frasnian) deposits of northern Timan

Fossil plants from the Upper Devonian (Frasnian) of northern Timan are studied. The sporangia contain well-preserved spores, which were studied in transmitted light using a scanning electron microscope. The genus Gutzeitia S. Snigirevsky, gen. nov. is established. Macroremains and in situ spores of G. timanica (Petros.) S. Snig., comb. nov. and ?Cephalopteris mirabilis (Nath.) Nath. are described. Microspores in Dimeripteris gracilis Schmalh. are identified, and microspores and megaspores in ?Cephalopteris mirabilis are studied for the first time.     

Diversity,structure and interactions of encrusting lithophyllic macrofaunal assemblages from Belgica Bank,East Greenland

Drop stones with lithophyllic macrofauna from Belgica Bank, Greenland Sea (79–81°N and 5–17°W) were collected during the Polarstern cruise in August 2000. The investigation focussed on species richness, diversity, abundance, and interactions of organisms colonizing rocks. On 101 stones collected by 10 dredges from nine sites, 113 taxa (58 genera, 38 families, 12 orders and 8 phyla) were recognized. Among the most abundant groups of organisms were foraminiferans, bryozoans and polychaetes, while the least abundant were anthozoans and ascidians. Abundance ranged from 1239 to 11,383 individuals m⁻² of rock surface area. Bryozoans were the most species-rich group, while anthozoans and ascidians were represented by the least number of species. Colonization occurred at various topographical levels and was classified into two categories. Stones were classified as primary space and were colonized by erect membranous, erect flexible, erect rigid, and flat encrusting organisms. The second level of colonization by epibionts occurred on erect forms of organisms (e.g. erect bryozoans) and was classified as a secondary space. Species composition and abundance between faunal assemblages of primary and secondary space differed greatly. Colonization occurred only on the top of stones and was considered a special adaptation to avoid burial by sedimentation and to enhance food supply. Only 3% of recruits were involved in any competitive interactions. Competition was considered to be of minor importance in structuring lithophyllic assemblages.     

A novel lycopsid from the Upper Devonian of Jiangsu, China

A new lycopsid, Monilistrobus yixingensis gen. et sp. nov., is described from the Wutung Formation (Famennian, Upper Devonian) of Jiangsu, China. The plant has many features typical of other Upper Devonian lycopsids, including dichotomous branching, helically arranged obovate expanded leaf bases, linear leaves with spiny appendages along the lateral margins, sporophylls widened proximally, and one elliptical sporangium attached to the adaxial surface. The most distinctive novel feature of the plant is that the modified sporophylls are arranged tightly into fertile zones or cone-like structures that are separated by lengths of axes with more lax sterile microphylls only: therefore the cone-like structures are strung along the branches like beads on a necklace.     

A new species of Laccognathus (Porolepiform Crossopterygii) from the Devonian of Latvia

A new species of the genus Laccognathus, L. grossi sp. nov., from the upper strata of the Gauja Formation of the Main Devonian Field (Lode locality, Latvia) is established. Morphological similarity and differences of the new species from the previously described L. panderi Gross from the lower strata of the Gauja and the Amata Formation of Latvia are discussed.     

An onychodont fish (Osteichthyes,Sarcopterygii) from the Early Devonian of China,and the evolution of the Onychodontiformes

Although the superbly preserved specimens of Onychodus jandemarrai have greatly advanced our understanding of the Onychodontiformes, a primitive sarcopterygian group with large parasymphysial tooth whorls, the scarcity of the otoccipital material in the group hampers further morphological comparisons between onychodonts and other sarcopterygian groups. Here we report a new onychodont Qingmenodus yui gen. et sp. nov. from the Early Devonian (Pragian) of South China that comprises well-ossified otoccipital and upper and lower jaw material. As one of the oldest known onychodonts, Qingmenodus shows for the first time the nearly complete structure of the otoccipital in onychodonts and provides an additional basis to address the phylogenetic position of the group. Its elongated otic shelf exhibits the posterior shift of the attachment for the basicranial muscle as in coelacanths and sheds light on the feeding mechanism of onychodonts. Qingmenodus displays a mosaic of primitive and derived onychodont features. The phylogenetic analysis places Qingmenodus immediately basal to the clade comprising Onychodus and Grossius.     

A new species of psammosteiform heterostracan (Agnatha) from the Givetian (Middle Devonian) of the Voronezh Region

A new pycnosteid species, Schizosteus shkurlatensis sp. nov. (Agnatha, Psammosteiformes), from the Staryi Oskol beds (Givetian, Middle Devonian) of the Voronezh Region is described. This is the first species of the genus Schizosteus Obruchev from the Central Devonian Field. The similarity between species of this genus and early species of Psammolepis is discussed.     

A new genus of lycopods from the Devonian of New York State

Drepanophycus colophyllus Grierson & Banks is transferred to llaskinsia Grierson & Banks on the basis of the morphology of its leaves and its anatomical structure. Slender, dichotomizing axes bear helically arranged leaves that are falcate, petiolate, simple, lanceolate and entire. A solid cylinder of metaxylem is surrounded by ridges of protoxylem. Pitting on tracheid walls is annular, helical, reticulate, scalariform to circular-bordered. This lycopod extends from early Givetian into early Frasnian time in the Catskill strata of eastern New York State. The effect of cleavage of fossiliferous rock on patterns observed on the exposed fossils and the bearing of recent research on the fossil history of lycopods are discussed.     

Molecular adaptation and the origin of land plants

The origin and diversification of land plants was one of the most important biological radiations. Land plants are crucial components of all modern terrestrial ecosystems. The first land plants had to adapt to a wide array of new environmental challenges including desiccation, varying temperatures, and increased UV radiation. There have been numerous studies of the morphological adaptations to life on land. However the molecular adaptations to life on land have only recently gained attention. These studies have greatly benefited from the recent advances in our understanding of the phylogenetic relationships between and among the charophycean algae and the basal land plant groups. In this review I summarize the current knowledge of a variety of physiological and biochemical adaptations to land including plant growth hormones, isoprene, phenolics, and heat shock proteins.