Anatomical reinvestigation ofArchaeopteris macilenta from the Upper Devonian (Frasnian) of South China

Archaeopteris macilenta is one of the most widespread plants in the Late Devonian.Based on fossils from the Frasnian Huangjiadeng Formation,Yichang District of Hubei Province,for the first time we study in detail the anatomy of this progymnosperm plant in South China.Ultimate axes are protostelic with three xylem sympodia and lack secondary tissue.Penultimate axes are eustelic,bearing eight sympodia and a thin band of secondary xylem.Radially symmetrical sympodia of mesarch primary xylem produce traces of appendages in a spiral arrangement.Archaeopteris macilenta and A.halliana (A.roemeriana) are dominant in the Frasnian and Famennian,respectively.Comparisons with these two species from other tectonic plates indicate consistent stelar architectures.Global spread,continuous occurrence,and identical anatomy during the Late Devonian indicate that Archaeopteris survived the Frasnian-Famennian extinction event.In this time,endemic genera and cosmopolitan taxa,including Archaeopteris,suggest the palaeogeographic isolation of South China and certain associations with other plates.     

ACTINOXYLON BANKSII GEN. ET SP. NOV.: A PROGYMNOSPERM FROM THE MIDDLE DEVONIAN OF NEW YORK

A new genus from a Middle Devonian locality near Cairo, N. Y., is described. Actinoxylon gen. nov. is based upon pyritic petrifactions. Three orders of branching are present: penultimate branch, ultimate branch, and leaf. The penultimate branch bears spirally arranged ultimate branches and leaves, the leaves apparently replacing the branches in the spiral. The ultimate branches bear opposite to subopposite and decussate leaves. The leaves are non-planated, unwebbed structures which show at least three dichotomies. Each segment of the leaf is terete as are all other axes. Internally the penultimate branch has a six-lobed actinostele with mesarch protoxylem areas, one or two per lobe. Secondary xylem is visible in the oldest parts of several specimens. The xylem has helical-reticulate, reticulate, scalariform and circular-pitted elements. The presumptive areas of phloem are occupied by cells with dark contents. The cortex is composed of a parenchymatous inner region and a sclerenchymatous outer region. The ultimate branch traces are at first three-lobed protosteles, later becoming four-lobed. Several ultimate branch traces also possess secondary xylem while within the cortex of the penultimate branch. The leaf traces are terete strands. Below each forking of a leaf segment there is a corresponding forking of the vascular strand. Actinoxylon is compared with the progymnosperms Actinopodium, Svalbardia, Archaeopteris, Siderella, and Tetraxylopteris. The anatomy of the penultimate branch of Actinoxylon is similar to that of Actinopodium, Archaeopteris macilenta, and Siderella. The ultimate branch traces of Archaeopteris and Actinoxylon are similar. The ultimate branch stele and pattern of trace formation in Actinoxylon is similar to the stelar configuration and trace formation in the r + 2 axes of Tetraxylopteris schmidtii. The unwebbed leaves are similar to those of Archaeopteris fissilis, Svalbardia, and the terminal units of the Aneurophytales.     

ARCHAEOPTERIS MACILENTA,ANATOMY AND MORPHOLOGY OF ITS FROND

The stem, rachides, and pinnae of Archaeopteris macilenta, formerly considered to be a fern of Devonian age, comprise a branch system in which the ultimate divisions heretofore referred to as pinnules are the leaves. The primary vascular system of the “frond” is a lobed siphonostele from which leaf traces arise in a spiral sequence. The anatomy of the “rachis” and of the “pinnae” is shown to be similar to that of the stem, Callixylon, which bore these “fronds.” Branching, epidermal pattern and stomates are described for the spirally arranged leaves (fertile pinnules). Attachment and dehiscence of sporangia as well as their stomates are reported. Archaeopteris is retained in the Class Progymnospermopsida which includes plants with gymnospermous anatomy and pteridophytic reproduction. It is suggested that Actinopodium, Svalbardia and Siderella are related closely to Archaeopteris and that this group of genera shows evolutionary stages in webbing of leaves and planation of branch systems. The opportunities for ontogenetic studies of the arborescent genus Archaeopteris are pointed out.     

Microconchid tubeworms across the upper Frasnian – lower Famennian interval in the Central Devonian Field,Russia

Abstract: Microconchid tubeworms (Tentaculita) encrusting brachiopod shells have been investigated from the upper Frasnian – lower Famennian (Upper Devonian) deposits of the Central Devonian Field, Russia. The condition of microconchids and associated encrusting taxa is reported for the first time from the early Famennian recovery interval (crepida Chron) following the Frasnian–Famennian mass extinction. Two species, one new (Palaeoconchus variabilis sp. nov.) and the second one in open nomenclature (Palaeoconchus sp.), are described. Compared to lower Famennian specimens, they seem to be preferentially grouped on the anterior parts of the brachiopod host shells, which are interpreted as the most suitable sites away from the sea‐bottom and sediment. During the late Frasnian (Late rhenana Chron), microconchids, outnumbered by cornulitids and as abundant as foraminifers, were also associated with trepostome bryozoans, tabulates, rugose corals and various problematic encrusters. During the early Famennian recovery interval encompassing the crepida Chron, microconchids greatly outnumbered all associated encrusters, including the previously dominant cornulitids, while foraminifers, tabulates and rugose corals vanished. Early Famennian microconchids, represented by the single, albeit very abundant, species Palaeoconchus variabilis sp. nov., were opportunists that rapidly colonised the environment during the ongoing transgression following the regression‐driven biotic crisis in the area of the Central Devonian Field. In comparison to their late Frasnian predecessors and even other Middle Devonian specimens, no size reduction (the so‐called Lilliput effect) of early Famennian microconchid tubes was observed. It is probable that microconchids either rapidly attained their ‘normal’ sizes or they did not suffer any dwarfism following the Frasnian–Famennian event.     

An anatomically preserved lycopsid from the Upper Devonian of South China

An anatomically preserved lycopsid, Lobodendron fanwanensegen. et sp. nov. Liu, Wang, Xue & Meng, is described from the Upper Devonian (Famennian) Wutong Formation of Changxing County, Zhejiang Province, China. The fossil plant bears slender, dichotomously branched axes. The vascular strand consists of solid terete primary xylem and lobed secondary xylem, which implies the result from the activity of possibly discontinuous cambium. The new plant has character combinations that do not conform to any branches in the canopy of the tree-lycopsids known previously, but resemble those of the basal part of some pseudoherbaceous lycopsids. This new plant may exemplify a Late Devonian lycopsid with a pseudoherbaceous growth habit.     

ON THE ANATOMY AND MORPHOLOGY OF LATERAL BRANCH SYSTEMS OF ARCHAEOPTERIS

A large collection of specimens, consisting primarily of Archaeopteris macilenta, forms the basis for an anatomical and morphological study of the lateral branch systems. Emphasis is placed on the main axes of the branch systems, which are found to be characterized by a variable number of orthostichies of lateral appendages, ranging from 7 to probably 11. The number of orthostichies seems to correspond to the number of ribs of the stele. There may be an equal number of rows of leaves on either side of the plane in which the two rows of lateral branches lie, or one side may have one or two more leaf orthostichies than the other. In all specimens for which both part and counterpart were available, however, there is a greater density of leaves on one side (hypothesized to be the adaxial) than on the other, caused primarily by an apparent abortion of leaf primordia early in development on the side of least density. Leaves and branches occur in the same ontogenetic spiral and in one specimen comprise a ‘phyllotaxis’ of 2/9 and contact parastichies of 5 + 9. On both the main and lateral axes, leaves are characterized by long, decurren bases which essentially ensheath the axrs on which thay are borne. Phyllotaxis and the correlated stelar form apparently vary considerably within Archaeopteris. It is suggested that Archaeopteris was more closely related to the coniferophytes than to any other gymnosperms.     

华南泥盆系法门阶浅海相四射珊瑚的组合序列

晚泥盆世弗拉期末F-F的生物灭绝事件,导致珊瑚以及许多浅海底栖生物灭绝,几乎整个法门期珊瑚群都处于残存阶段,华南地区目前只在湖南的个别地方发现少量的Smithiphyllum。直到法门期末珊瑚和其它一些造礁底栖生物开始复苏,出现了不少与典型泥盆纪珊瑚存在很大差别的新分子。华南泥盆纪最晚期(Strunian)的珊瑚可划分成上、下两个组合:上部Cystophrentis组合;下部Eocaninophyllum组合。泥盆—石炭纪之交的另一次生物灭绝事件,使新生的泥盆纪最晚期的珊瑚又遭灭绝。     

Late Devonian global ostracod palaeobiogeography

A global Late Devonian ostracod database is constructed, incorporating new materials from South China and Northwest China. Four palaeobiogeographical units (Cathaysia, North America, Europe and peri‐Gondwana) are recognized during the Frasnian and five palaeobiogeographical units (Cathaysia, North America, Europe, Siberia and Australia) in the Famennian. Three controlling factors (climatic zonation, geographical isolation and global sea‐level changes) are identified to have played roles in shaping the palaeogeographical regionalization of ostracod faunas in the Late Devonian. The ostracod palaeobiogeography in the Frasnian was mainly influenced by climatic zonation, while rapid changes in tectonic configuration in the Famennian drastically altered the global palaeobiogeography of ostracods. The palaeobiogeographical regionalization of ostracod faunas suggests that Laurussia and Gondwana continued to draw near during the Late Devonian, with the first collision occurring in Southern Central Europe in the Famennian. The South China plate drifted northward to the Kazakhstan plate away from the Australian plate, which gradually became isolated during the Famennian Stage.     

The fate of the homoctenids (Tentaculitoidea) during the Frasnian–Famennian mass extinction (Late Devonian)

The homoctenids (Tentaculitoidea) are small, conical‐shelled marine animals that are among the most abundant and widespread of all Late Devonian fossils. They were a principal casualty of the Frasnian–Famennian (F‐F, Late Devonian) mass extinction, and thus provide an insight into the extinction dynamics. Despite their abundance during the Late Devonian, they have been largely neglected by extinction studies. A number of Frasnian–Famennian boundary sections have been studied, in Poland, Germany, France, and the USA. These sections have yielded homoctenids, which allow precise recognition of the timing of the mass extinction. It is clear that the homoctenids almost disappear from the fossil record during the latest Frasnian ‘Upper Kellwasser Event’. The coincident extinction of this pelagic group, and the widespread development of intense marine anoxia within the water column, provides a causal link between anoxia and the F‐F extinction. Most notable is the sudden demise of a group, which had been present in rock‐forming densities, during this anoxic event. One new species, belonging to Homoctenus is described, but is not formally named here.     

The primary vascular system of Callixylon

The primary vascular system of Callixylon (main stem axes of Archaeopteris) is composed of a discontinous cylinder of longitudinal bundles from some of which traces diverge radially in a regular, helical pattern. Tangential divisions and fusions of primary xylem strands occur in approximately equal frequency and, thus, the number of bundles in the system remains constant in the specimen studied. Following divergence, traces increase in size and become ribbed distally. The available evidence suggests that all of the traces observed in this study are branch traces. Branch bases, embedded in secondary xylem, have the anatomical characteristics of penultimate axes of lateral branch systems of Archaeopteris. It is suggested that some regions of main axes of Archaeopteris might have borne lateral branch systems only, lacking entirely simple leaves. Lateral branch systems, therefore, might have been more similar to compound leaves than previously supposed.     

Sphenophyllum Brongniart (Sphenopsida) from the Upper Devonian of South China

Sphenophyllum Brongniart constitutes a common and important element in the Carboniferous and Permian floras around the world, but its records in the Devonian Period are limited and remain to be investigated with respect to their morphology, anatomy, and distribution. Diversity of Sphenophyllum during Late Devonian in South China was significant with the presence of the well-known species including S. lungtanense Gothan and Sze, S. pseudotenerrimum Sze, and S. changxingense Huang et al. In this article, we describe a new species of the genus, S. fanwanense n. sp., from the Fanwan section of Changxing County, Zhejiang Province, characterized by having at least two orders of axes, with six spoon-shaped leaves attached at the node. A comprehensive review of previous records of Sphenophyllum from the Upper Devonian of South China shows that at least four species (S. fanwanense, S. lungtanense, S. pseudotenerrimum and S. changxingense), each with distinct leaf morphology, occur in South China in the Late Devonian, mostly from the Wutong Formation (Famennian). Given the lack of concrete evidence for anisophylly or heterophylly among the Devonian Sphenophyllum species (and more broadly, the Devonian sphenopsids), we suggest that the development of these features later in the Carboniferous and Permian periods was likely related to an increase in complexity of forest communities through time.     

Progressive drowning of carbonate platform in the Moravo-Silesian Basin (Czech Republic) before the Frasnian/Famennian event: facies,compositional variations and gamma-ray spectrometry

The Moravo-Silesian Basin (MSB; eastern Czech Republic and southern Poland) hosted an extensive shallow-water carbonate platform in the Middle Devonian to Frasnian interval. The platform drowned in a stepwise fashion from the Palmatolepis hassi to the Pa. linguiformis zone. Three types of drowning successions were revealed from conodont biostratigraphy, facies, microfacies and gamma-ray spectrometry data: (A) drowning to periplatform turbidite setting; (B) drowning to (hemi)pelagic seamount setting and (C) drowning associated with the stratigraphical gap. In the lower Pa. hassi zone, rapid subsidence caused the platform to drown locally along the N–S to NW–SE trending faults (type A drowning). In the upper Pa. rhenana to the Pa. linguiformis zone, the drowning accelerated in the western part of the MSB due to locally higher subsidence rates combined with the Late Frasnian biotic crisis (type B). In the southern part of the basin, the platform emerged shortly before the Frasnian/Famennian (F/F) boundary and drowned in the Early to Late Famennian (type C). The primary cause of drowning was differential subsidence at the Laurussian passive margin. Eustatic sea-level fluctuations, if any, contributed only to a minor extent to the Late Frasnian drowning, but were effective in type C drowning during the Famennian. The drowning boundaries are associated with increased contents of K and Th, reflecting the deceleration of carbonate production. Uranium contents display isolated peaks that roughly correlate with the drowning boundaries or the stratigraphic gaps associated with the F/F boundary. The uranium contents are considered to reflect local depositional conditions and are not suitable for stratigraphic correlation. On the other hand, from the K and Th contents, we can infer Late Frasnian sea-level fluctuations with duration on the order of 1 Myr. These cyclic variations in K and Th contents proved to be useful in platform-to-basin stratigraphic correlation.     

Correct author citation of the Late Devonian plant <Emphasis Type="Italic">Archaeopteris</Emphasis> (Progymnospermopsida)

Archaeopteris as a representative fossil plant genus in the class Progymnospermopsida has been extensively recorded in the Late Devonian of the world. Although its phylogenetic, paleoecological, and biostratigraphic significances are well known, the author citation of the generic name Archaeopteris remains problematic. Here, it is shown that Dionys Stur was the first to treat Archaeopteris as an independent genus.Therefore, the generic name should be cited as Archaeopteris (Dawson) Stur, while Archaeopteris Dawson and Archaeopteris (Dawson) Lesquereux are incorrect.     

First <Emphasis Type="Italic">Sycidium</Emphasis> (Charophyta) from the Upper Devonian of Thuringia (Germany)

The well-preserved silicified fragment of a utricular envelope assigned to Sycidium aff. clathratum Peck, discovered in the earliest Famennian “incisus bed” of the Bohlen section near Saalfeld in Thuringia (Central Germany), is the first evidence of charophytes present in the West European Late Devonian. The occurrence of 16 longitudinal ridges is shared with the basal Mississippian S. clathratum. The specimen shows delicate structures of the external layer including ovoid elements at its basal end which are interpreted as casts of rudimentary bract cells and/or bracteoles, rarely observed in the fossil record of charophytes. S. aff. clathratum is among the rare Sycidiaceae that survived the terminal Frasnian Kellwasser Event. This single specimen was probably attached to a piece of wood and thereby transported into fully marine basinal environments that characterize the Famennian of the Bohlen section.     

New Late Devonian strophomenids (Brachiopoda) from Mongolia

Two new strophomenid species, Leptagonia barunkhuraica sp. nov. (Famennian) and Floweria mongolica sp. nov. (Frasnian), from the Upper Devonian of the Barunkhurai Depression of southern Mongolia are described.     

Devonian Vertebrates From Colombia

Vertebrate remains are reported from the Emsian–Eifelian Floresta Formation and the Late Devonian (?Frasnian) Cuche Formation of north‐eastern Colombia. The material from the Floresta Formation is associated with a marine invertebrate fauna and includes an arthrodire and probably a rhenanid. Several vertebrate‐bearing localities are recorded from the Cuche Formation; vertebrates occur with plant remains and lingulid fragments. They include an acanthodian (Cheiracanthoides? sp.), a chondrichthyan (Antarctilamna? sp.), placoderms (Bothriolepis sp., Asterolepis? sp. and an undetermined groenlandaspidid or primitive brachythoracid arthrodire), a stegotrachelid actinopterygianand three sarcopterygians (a cosmine‐covered form tentatively referred to an osteolepidid, the porolepiform Holoptychiussp., and the rhizodontid Strepsodus? sp.). This assemblage suggests a Late Frasnian age and is surprisingly similar to Late Devonian vertebrate assemblages found in similar facies of Europe and North America, notwithstanding the presence of the Gondwanan chondrichthyan Antarctilamna?. key words: Vertebrata, Devonian, Colombia, South America, biostratigraphy, palaeobiogeography.     

Evolution of Devonian biodiversity of radiolarians with two porous spheres and one main spine

Features and major distinctions in the appearance and distribution of radiolarians with two porous spheres and one main spine in the Devonian basins are considered. Four solitary population waves and expansion scenario of radiolarians of this morphotype have been recognized in the Devonian. A new species, Radiobisphaera rozanovi sp. nov., from the Upper Emsian, Upper Eifelian, and Middle Frasnian of the southern Ural Mountains, from the Upper Eifelian of the Prague Basin (Barrandian), from the Middle-Upper Frasnian of the Rudnyi Altai, and from the Lower Famennian of the Timan-Pechora Basin and the Polar Ural Mountains is described.     

The history of Devonian-Carboniferous reef communities: Extinctions,effects, recovery

Summary Analysis of the taxonomic composition, diversity and guild structure of five “typical” reef and mud mound communities ranging in age from Late Devonian-Early Carboniferous indicates that each of these aspects of community organization changed dramatically in relation to three extinction events. These events include a major or mass extinction at the end of the Frasnian; reef communities were also effected by less drastic end-Givetian and mid-late Famennian extinctions of reef-building higher taxa. Peak Paleozoic generic diversities for reef-building stromatoporoids and rugose corals occurred in the Eifelian-Givetian; reef-building calcareous algal taxa were longranging with peak diversity in the Devonian. These three higher taxa dominated all reef-building guilds (Constructor, Binder, Baffler) in the Frasnian and formed fossil reef communities with balanced guild structures. The extinction of nearly all reef-building stromatoporoids and rugose corals at the end of the Frasnian and the survival of nearly all calcareous algac produced mid-late Famennian reef communities dominated by the Binder Guild. Despite the survival of most calcareous algae and tabulate corals, the mid-late Famennian extinction of all remaining Paleozoic stromatoporoids and nearly all shelf-dwelling Rugosa brought the already diminished Devonian reef-building to a halt. These Devonian extinctions differ from mass extinctions by the absence of a statistically significant drop in taxonomic diversity and by their successional and cumulative effects on reef communities. Tournaisian mud mounds contain communities markedly different from the frame-building communities in Late Devonian and Visean reefs. Mound-building biotas consist of an unusual association dominated by erect, weakly skeletonized members of the Baffler Guild (chiefly fenestrate Bryozoa; Pelmatozoa) and laterally expanded, mud-binding algae/stromatolites and reptant Bryozoa. The initial recovery to reefs with skeletal frameworks in the Visean was largely due to the re-appearance of new species of abundant colonial rugose corals (Constructor Guild) and fenestrate Bryozoa. This Frasnian-Visean evolution in the taxonomic composition and structure of the reef-building guilds is also expressed by abrupt changes in biofacies and petrology of the reef limestones they produced. Thus, “typical” Frasnian reef limestones with balanced guild structures are framestones-boundstones-bafflestones, Famennian reefs are predominantly boundstones, Tournaisian mud mounds are bafflestones and Visean reefs are bafflestones-framestones.     

Famennian conodonts from the Hongguleleng Formation at the Bulongguoer stratotype section,western Junggar,Northwest China

The Hongguleleng Formation, the highest and most important Devonian marine carbonate horizon in western Junggar, contains an endemic shallow-water Icriodus–Polygnathus conodont fauna with rare palmatolepids and other genera. The conodont faunas from the Bulongguoer and the Wulankeshun sections are similar, with high abundance (about 40%) of endemic taxa (13 taxa), indicating isolation of the Junggar Basin during early Famennian. Non-endemic species in the faunas suggest that the Lower Member of the Hongguleleng Formation is assignable to the Pa. rhomboidea Zone to the Pa. marginifera marginifera Zone of early Famennian, not including the Frasnian–Famennian boundary. The Upper Member may be of late Famennian–early Tournaisian in age on the basis of our preliminary faunal analysis.     

New discoveries of tetrapods (ichthyostegid‐like and whatcheeriid‐like) in the Famennian (Late Devonian) localities of Strud and Becco (Belgium)

The origin of tetrapods is one of the key events in vertebrate history. The oldest tetrapod body fossils are Late Devonian (Frasnian–Famennian) in age, most of them consisting of rare isolated bone elements. Here we describe tetrapod remains from two Famennian localities from Belgium: Strud, in the Province of Namur, and Becco, in the Province of Liège. The newly collected material consists of an isolated complete postorbital, fragments of two maxillae, and one putative partial cleithrum, all from Strud, and an almost complete maxilla from Becco. The two incomplete maxillae and cleithrum from Strud, together with the lower jaw previously recorded from this site, closely resemble the genus Ichthyostega, initially described from East Greenland. The postorbital from Strud and the maxilla from Becco do not resemble the genus Ichthyostega. They show several derived anatomical characters allowing their tentative assignment to a whatcheeriid‐grade group. The new tetrapod records show that there are at least two tetrapod taxa in Belgium and almost certainly two different tetrapod taxa at Strud. This locality joins the group of Devonian tetrapod‐bearing localities yielding more than one tetrapod taxon, confirming that environments favourable to early tetrapod life were often colonized by several tetrapod taxa.