A bryozoan fauna from the Santa Lucia Formation (Lower–Middle Devonian) of Cantabrian Mountains,NW Spain

The Lower to Middle Devonian Santa Lucia Formation of NW Spain contains a rich and well-preserved bryozoan fauna. An assemblage containing 14 species is described from two localities in Cantabrian Mountains, Abelgas, and Paradilla. One new genus with one new species is described: Isostylus abelgasensis n. sp. n. gen. Another two new species have been found: Microcampylus minor n. sp. and Acanthoclema parvula n. sp. Additionally, the following species were identified: Cyclotrypa communis (Ulrich 1890), Fistuliphragma gracilis Ernst 2008a, Leioclema attenuatum Duncan 1939, Hemitrypa cf. tenella Barrande in Počta 1894, Fenestella aff. parallela Hall 1881, Anastomopora adnata (Hall 1883), Semicoscinium rhombicum Ulrich 1890, Quadrisemicoscinium discretum (Prantl 1932). Three species are described in open nomenclature: Trepostomata sp. indet.1 and 2, and Semicoscinium sp. The described bryozoan fauna shows connections to the Lower Devonian of Bohemia, and to the Middle Devonian of Rhenish Massif and North America.     

Species–area relationships in the Hantam-Tanqua-Roggeveld,Succulent Karoo,South Africa

The Hantam-Tanqua-Roggeveld subregion is part of the Succulent Karoo hotspot of biodiversity which stretches along the southwestern side of South Africa and Namibia. Forty Whittaker plots were surveyed in the spring of 2005, in eight vegetation associations, to gather diversity data for the Hantam, Tanqua Karoo and Roggeveld areas. Seven plot sizes were used to construct species–area curves using three different models namely: the untransformed linear function, the power function and the exponential function. In general, the power and exponential functions produced a more significant fit to the data than the untransformed linear function. Linear regressions using environmental parameters indicated that altitude, mean annual precipitation and mean annual temperature were significant predictors of species richness at the 1, 10, 100 and 1000 m² scales. To illustrate the variation in species–area curves and species richness across the landscape, a transect through the study area is discussed. The transect stretches eastwards from the Tanqua Karoo across the escarpment into the Roggeveld and crosses five different vegetation associations. Differences between associations were found in species richness in the 1000 m² plots. Each association also produced species–area curves with their own characteristics. Slope values for the samples within an association did not differ significantly, although the intercept value often did. Comparisons between associations along the transect revealed significant differences in the slope value between the associations, except for the Dicerothamnus rhinocerotis Mountain Renosterveld which did not differ significantly from the associations bordering it on either side.     

Isotopic evidence of environmental changes during the Devonian–Carboniferous transition in South China and its implications for the biotic crisis

The Devonian–Carboniferous (D–C) transition coincides with the Hangenberg Crisis, carbon isotope anomalies, and the enhanced preservation of organic matter associated with marine redox fluctuations. The proposed driving factors for the biotic extinction include variations in the eustatic sea level, paleoclimate fluctuation, climatic conditions, redox conditions, and the configuration of ocean basins. To investigate this phenomenon and obtain information on the paleo-ocean environment of different depositional facies, we studied a shallow-water carbonate section developed in the periplatform slope facies on the southern margin of South China, which includes a well-preserved succession spanning the D–C boundary. The integrated chemostratigraphic trends reveal distinct excursions in the isotopic compositions of bulk nitrogen, carbonate carbon, organic carbon, and total sulfur. A distinct negative δ¹⁵N excursion (~−3.1‰) is recorded throughout the Middle Si. praesulcata Zone and the Upper Si. praesulcata Zone, when the Hangenberg mass extinction occurred. We attribute the nitrogen cycle anomaly to enhanced microbial nitrogen fixation, which was likely a consequence of intensified seawater anoxia associated with increased denitrification, as well as upwelling of anoxic ammonium-bearing waters. Negative excursions in the δ¹³C_carb and δ¹³C_org values were identified in the Middle Si. praesulcata Zone and likely resulted from intense deep ocean upwelling that amplified nutrient fluxes and delivered ¹³C-depleted anoxic water masses. Decreased δ³⁴S values during the Middle Si. praesulcata Zone suggests an increasing contribution of water-column sulfate reduction under euxinic conditions. Contributions of organic matter produced by anaerobic metabolisms to the deposition of shallow carbonate in the Upper Si. praesulcata Zone is recorded by the nadir of δ¹³C_org values associated with maximal △¹³C. The integrated δ¹⁵N-δ¹³C-δ³⁴S data suggest that significant ocean-redox variation was recorded in South China during the D–C transition; and that this prominent fluctuation was likely associated with intense upwelling of deep anoxic waters. The temporal synchrony between the development of euxinia/anoxia and the Hangenberg Event indicates that the redox oscillation was a key factor triggering manifestations of the biodiversity crisis.     

Late Devonian–Early Carboniferous palynology of the CSDP-2 Borehole in the southern Yellow Sea,China

Late Devonian–Early Carboniferous spores recovered from the depths of 2140 m to 2031.3 m in the CSDP-2 Borehole in the southern Yellow Sea are systematically documented to characterize palynological associations for the stratigraphic correlation. The taxonomic study identifies 96 species belonging to 48 genera, including a re-examination of 33 previously described and indeterminate species. Three palynological associations are established for the Wutong Formation in the CSDP-2 Borehole. In ascending order, these are the late Famennian Aneurospora asthenolabrata–Geminospora lemurata (AL) Association from the proposed Guanshan Member, the latest Famennian Cymbosporites circinatus–Asperispora acuta (CA) Association from the lower part of the proposed Leigutai Member, and the Tournaisian Auroraspora macra–Lophozonotriletes involutus (MI) Association of the upper part of the latter member. The AL and CA associations are correlated with the palynological assemblages in the Guanshan Member, and those in the lower to middle parts of the Leigutai Member of the Wutong Formation that outcrops in the suburbs of Nanjing and southern Jiangsu. The MI Association is similar to those derived from the uppermost beds of the subsurface Wutong Formation and is dated as early Tournaisian.     

The arthropodBundenbachiellus giganteus from the Lower Devonian Hunsrück Slate, Germany

The large Hunsrück Slate arthropodBundenbachiellus giganteus (Broili, 1929) has been interpreted as a myriapod, a possible arthropleurid, and most recently as a distorted specimen of the arachnomorph arthropodCheloniellon. The Single specimen lacks the head region and is characterised by a narrow trunk bearing a series of biramous appendages with a segmented endopod and filamentous exopod. A reinvestigation demonstrates that it is not referable toCheloniellon but represents a distinct taxon of whichEschenbachiellus wuttkensis Briggs &;Bartels, 2001, also based on a Single Hunsrück Slate specimen, is a junior synonym. Minor features ofStürmer &;Bergström’s (1978) reconstruction ofCheloniellon, which were based on the holotype ofBundenbachiellus giganteus, are revised.     

Conodont biostratigraphy of Upper Devonian–Lower Carboniferous deposits in eastern Alborz (Mighan section), North Iran

The Devonian/Carboniferous (D/C) transition is characterized by a major transgressive/regressive cycle which led to a widespread ocean anoxia known as the Hangenberg Black Shale Event (HBSE), as well to a major sea-level fall (Hangenberg Sandstone Event, HSSE), recognized around the world. Both events are known as the Hangenberg Crisis. In order to examine the D/C transition in shallow water environment, the Mighan section in eastern Alborz was studied in terms of conodont biostratigraphy and stable isotope geochemistry. Twenty-five conodont species belonging to seven genera were identified and 5 conodont zones discriminated; namely, the Bispathodus aculeatus aculeatus Zone, Bispathodus costatus Zone, Bispathodus ultimus Zone, Siphonodella praesulcata Zone, costatus-kockeli Interregnum, and the sulcata Zone. Below the Devonian–Carboniferous boundary (DCB), the Hangenberg Black Shale and Hangenberg Sandstone equivalents were recognized, representing the Hangenberg Crisis that highly affected trilobite, ammonoid, brachiopod and conodont faunas at Mighan and worldwide. The kockeli Zone of the latest Famennian is missing at Mighan due to the lack of conodonts, probably related with the major environmental changes linked with the Hangenberg Crisis recognizable worldwide. Carbon isotopes measured of micrites from Mighan indicate a proximal depositional environment of a shallow shelf with terrestrial input and the oxygen isotope values from conodont apatite suggest warm seawater temperatures of tropical and subtropical setting in the study area.     

Nahecaris stuertzi,a phyllocarid crustacean from the Lower Devonian Hunsrück Slate

Restudy ofNahecaris based on new stereo radiographs and re-examination of actual specimens has added the following new information.Nahecaris ranges in shape from long and slender to short and stout, the carapace from deep and rounded to long and narrow. Carapace ornament consists either of thin ridges parallel to the ventral border or inclined rows of irregular lines or cusps, but these characters cannot be consistently correlated with other variables to permit definition of separate species. The carapace is fused to the head, and possibly to the anterior end to the thorax in the area in front of the median dorsal plate. A post-mandibular pyritised band may represent the carapace adductor muscle, which inserts on the mesolateral carinae. The mandible resembles that ofCeratiocaris, while the maxillae are like those of extant leptostracans. The 8 thoracopods are biramous: they have an exopod with finger-like lobes and a stenopodous endopod. Posteriorly the endopods reduce markedly, forming a conical, anteriorly open feeding basket. The predominance of dorso-ventrally flattened specimens, and the wide ventral gape of the carapace valves suggests those valves were spread apart in life.Nahecaris may have pressed itself to the substrate by flexing the carapace rim outward and incurving the ventral region of the carapace. The stout second antennae may have stabilised and propelled the animal over the substrate.Nahecaris is thought to have been benthic and to have fed, likeCeratiocaris, by lifting the substrate into suspension, as well as by picking up large fragments directly with the mandibles.     

Amplectosporangium—A New Genus of Plant from the Lower Devonian of Sichuan,China

A new genus and species Amplectosporangium jiangyouense, is described from the Pingyipu Formation (Siegenian) of the Lower Devonian in Jiangyou district, Sichuan, China. The plant consists of naked dichotomizing erect axes with terminal fertile branch-systems. Sporangia are oval-shaped each with a short stalk and are arranged along the inner side of dichotomized axes in single row. By comparison with other known Devonian plants the genus is considered to represent a new morphological type of early land plants. Based on the theory of the telomic origin of the ovular integument, the plant may represent an early stage in the evolution of integument.     

Discovery of radiolaria and conodonts in the Carboniferous–Permian of San Salvador Patlanoaya (Puebla,Mexico); biostratigraphic implications

New identifications of radiolaria and conodonts allow clarifying the biostratigraphy of the Pennsylvanian and Permian deposits in San Salvador Patlanoaya (Puebla State, Mexico). The radiolaria are sometimes relatively common in the series, but weakly diversified and endemic. Among the conodonts, Streptognathodus bellus is characteristic of the Late ‘Virgilian’ sensu Baars, probably coeval with the Early Wolfcampian sensu Thompson or Wilde, the Bursumian of Ross & Ross or the Orenburgian–Asselian stage or substage of Russia (sensu Davydov). San Salvador Patlanoaya is a key-section for the Pennsylvanian–Permian boundary in Mexico.     

Hsüa robusta,a new land plant from the Lower Devonian of Yunnan,China

A new psilophytic plant, Hsüa robusta, is found in the Xujiachong Formation (Emsian) of the Lower Devonian from the Qüjing (= Kütsing) district of Yunnan, China. This plant is tentatively referred to the Cooksoniaceae of Rhyniales. Hsüa gen. nov. Type species: Hsüa robusta (Li et Cai) C. S. Li. Diagnosis: Plants erect and then creeping. Main axes dividing pseudomonopodially and bearing dichotomous lateral branches which somewhat differentiate into vegeta, tire and fertile ones, with dichotomous root-like and rhizophore-like appendages. Sporangia terminal, round to reniform or wide reniform, dehiscing along distal margin into two equal halves. Spores homosporous, trilete. Stomata anomocytic. Protostele centrarch. Hsüia robusta (Li et Cai) C. S. Li, comb. nov. Cooksonia zhanyiensis Li et Cai, Acta Geologica Sinica, 52 (1) 1978, p. 10, pl. II, fig. 6.——Taeniocrada robusta Li et Cai,ib. p. 10, pl. II, fig. 7—14. Diagnosis: Characters same as in generic diagnosis. Main axes 6—10 mm wide and at least 24 cm long, with vascular strands 1.2—2.4 mm acr oss. Fertile branches 3—4 times equally or unequally dichotomous, 10—1.5 mm in width and up to 11 cm in length, possessing a vascular bundle of 0.5 mm in its greatest diameter. Branches circinately coiled in apical regions. Axial tubercles, root-like and rhizophore— like appendages arising from the main axes usually anterior to the lateral branches. Axial tubercle round with a diameter of 2.2—2.4 mm, having a vascular bundle about l mm across. Rootlike branches 3 times bifurcate, 1—0.3 mm wide and up to 1.5 cm long, with a vascular bundle about 0.1 mm across. Rhizophore-like appendages forked, 3—1.7 mm in width, possessing a vascular bundle of 0.7 mm in its greatest diameter. Root-like protuberances sometimes arising from rhilzophore-like branches. Epidermal cells of axes generally elongate, measuring 60—290μby 25—60 μ. Stomata mainly fusiform, 90—110 μ long and 50—60μ wide, consisting of a pair of guard cells enclosing a pore 6—15μ in length and 1—3μ in width. Cuticle of guard cells quite thick. Stomatal density about 5 per mm². Sporangia 0.8—4.2 mm high, 1.0—8.2 mm across, usually having a dehiscent distal border which measures 50—100μ broad. Demarcation between sporangium and its stalk quite clear. Epidermal cells of basal part of sporangial walls elongate, about 100 μ long and 30μ wide, but those of distal part isodiametrally polygonal, about 50μ in diameter. Stomata, radially arranged scattering over sporangial walls, generally round about 50μ in diameter and 50 per sporangium. Spores round, 18—36μ (average 27μ) indiameter, and smooth. Tracheids of protoxylem about 10μ across; those of metaxylem about 30μ across, with scalariform thickening. This plant is similar to Renalia hueberi Gensel in general morphology, but differs from the latter in possessing root-like and rhizophore-like branches. The generic name is derived from Prof. Hsü Jen. This paper is a thesis for M. Sc.     

Lower–middle Frasnian organic carbon isotope record of conodonts in East European Platform

The early–middle Frasnian boundary interval of the northern part of the East European Platform (northwest of Russia) corresponding to the transitans-punctata isotope event is revealed by biostratigraphically constrained conodont carbon stable isotope data (δ¹³C_con). The dynamics of δ¹³C_con demonstrate a three-fold pattern with positive peaks at the onset of the main phase of the transitans-punctata isotope event (upper part of Montagne Noire 4 conodont Zone, MN4; up to -22.5‰ VPDB), and during the late part of the event (lower and middle parts of MN6 Zone; up to -24.0‰ VPDB and -22.1‰ VPDB). The stratigraphic level near the MN5 and MN6 boundary is marked by a prominent negative excursion in δ¹³C_con (down to -31.8‰ VPDB) that resembles the negative excursion in the terminal phase of the transitans-punctata isotope event worldwide. The δ¹³C_con variations in different taxa are generally consistent but demonstrate some differences in values and amplitudes. It is assumed that variations in the carbon isotopic compositions of conodonts were mainly controlled by changes in the isotope composition of the planktons as the main food source for conodonts.     

Trigonotarbid arachnids from the Lower Devonian (Siegenian) of Bürdenbach (Lahrbach Valley,Westerwald area,Rhenish Slate Mountains,Germany)

An association of trigonotarbid arachnids from Siegenian strata of Bürdenbach in the Westerwald is described. The Bürdenbach trigonotarbids are numerically dominated by a new genus and species, Spinocharinus steinmeyeri n. gen. n. sp., which is characterized by a rounded carapace with a unique pattern of thorn-like tubercles. A comparison with previously described Devonian trigonotarbids suggests that the new taxon is most probably assignable to the family Palaeocharinidae, which is well represented by the genus Palaeocharinus from the Scottish Rhynie cherts. This would constitute the first record of Palaeocharinidae from the Rhenish Devonian and contrasts sharply with previously described Emsian associations from the Rhenish Slate Mountains, which are dominated by archaeomartids. A rare archaeomartid, Archaeomartus cf. levis, occurs in association with the putative palaeocharinids.     

Late Devonian and early Carboniferous microfloras from the Hakkarı Province of southeastern Turkey

Late Devonian and early Carboniferous miospore and microphytoplankton assemblages are described for the first time from southeastern Turkey. The preliminary data show that assemblages recorded from the upper part of the Yiginli Formation are late Famennian in age and can be correlated with the VH, LL, LE Miospore biozones of western Europe, whilst assemblages from the overlying Köprülü Formation are considered to be middle to late Tournaisian in age and are tentatively assigned to the PC Miospore biozone of western Europe. Regional palynological correlations with other sections in North Africa and the Middle East are proposed. A new species, Verruciretusispora loboziakii is described.     

A tubicolous animal from the Hunsrück Slate (Lower Devonian,Southern Germany)

A small tube-dwelling organism is interpreted as living in the sediment. This life habit is inferred from the difference between lined burrows and tubes and between soft and rigid tubes. Lamellar extensions of the body-wall may have provided irrigation for the vermiform part of the body. The gaping aperture lacks an operculum and may have been open permanently; the apertural rim was strengthened by spicules. The specimen lacks segmentation and is thus not an annelid worm. Considering the theoretical suitability of the crinoid “forest” in the Hunsrück sea for such worms, the absence of Annelida is curious.     

A pyritized lepidocoleid machaeridian (Annelida) from the Lower Devonian Hunsrück Slate,Germany

A machaeridian, Lepidocoleus hohensteini sp. nov., is described from the Hunsrück Slate (Lower Emsian) of Germany. The available material includes a unique example preserving evidence of the soft tissues, only the second machaeridian specimen to do so and the first lepidocoleid. This specimen shows that the plates are attached to alternate segments in the trunk. The morphology is consistent with an annelid affinity of the Lepidocoleidae and confirms the unity of the Machaeridia. This discovery adds an important group to the known diversity of this famous late Palaeozoic marine Konservat-Lagerstätte.     

U–Pb zircon age,sedimentary facies,and sequence stratigraphy of the Devonian–Carboniferous boundary,Daposhang Section,Guizhou, China

The Devonian–Carboniferous (D/C) boundary marks a major extinction event and an important evolutionary turnover in Paleozoic biotas, but understanding D/C boundary events is hampered by a lack of accurate and consistent ages. As a result, although a global stratotype section and point (GSSP) for the D/C boundary was defined in the section La Serre trench E’ (Montagne Noire, France) and accepted and ratified by the IUGS, problems arose concerning both the lithology of the La Serre section and the identification of S. sulcata. Due to these problems, in the very recent years, the ICS decided to reconsider the GSSP. The Daposhang section in Muhua, Guizhou, China, is one of the most important reference sections for the D/C boundary: it is a conformable succession across the D/C boundary that contains conodont faunas characteristic of restricted subtidal facies. A biostratigraphically well constrained tuff near the D/C boundary in the Daposhang section provides an excellent opportunity to date this important event in Paleozoic history. Sensitive high-mass resolution ion microprobe (SHRIMP) U–Pb analyses were conducted on single zircons extracted from the tuff (between beds 0 and 1, which may be temporally equivalent to the Hangenberg black shale) as well as an analysis of sedimentary facies and sequence stratigraphy across the D/C boundary in the Daposhang section. Bed 0 is below the D/C boundary, in the Upper praesulcata conodont zone, and bed 1 is directly above the D/C boundary, in the sulcata conodont zone. Zircon analyses yielded 20 concordant dates that form a cluster with a ²⁰⁶Pb/²³⁸U concordia age of 359.6 ± 1.9 Ma. An abundance of bioclastic packstone and wackestone and a lack of obvious high-energy sedimentary structures or evidence of reworking in layers spanning the D/C boundary suggest conformable deposition in a quiet, restricted subtidal paleoenvironment. Fifty-nine fifth-order cycles combine to depict a shallowing-upward succession (beds 029-01; highstand systems tract of a third-order sequence representing the terminal Devonian). The highstand systems tract is overlain by a deepening-upward succession (beds 0-29; maximum flooding surface and transgressive systems tract of a third-order sequence representing predominantly the basal Carboniferous) that begins at the D/C boundary. Fischer plot analysis of high-frequency cycles at the D/C boundary confirms this well developed sea-level pattern, which, together with the tuff and black shale depositional events, has potential for global correlation. On the basis of this new single-zircon SHRIMP age and previously published work on the biostratigraphy and sequence stratigraphy of the D/C transition interval, the deposition of tuff took place at approximately 359.6 Ma, and the age of the D/C boundary at Daposhang, Guizhou, China, is estimated at 359.58 Ma.     

Occurrence of the Lower Cambrian anemone-style trace fossils in the Zabuk Formation (Mardin–Derik,SE Turkey)

The Zabuk Formation of the Derik Group exposed over much of south-eastern Turkey presents a succession composed of shallow marine and fluvial siliciclastic sedimentary rocks. The Lower Cambrian assemblages containing abundant anemone-style trace fossils are known from most major palaeocontinents such as Laurentia, Baltica, and Gondwana. These have possible affinities with semi-infaunal dwelling anemones on siliciclastic mid-latitude shelves of West Gondwana in early Fortunian deposits. Among them is Bergaueria, characterized by plug-shaped burrows as exemplified by Bergaueria perata, that is, a characteristic trace fossil of the Cambrian globally.     

Miospore assemblages from the Silurian–Devonian boundary,in borehole A1-61, Ghadamis Basin,Libya

Well preserved and diversified miospore assemblages have been recorded from a relatively continuous sequence in borehole A1-61 which spans the Silurian–Devonian boundary in the northwestern part of the Ghadamis Basin, Libya. The sequence is represented by early Devonian Lochkovian beds of the Tadrart Formation that transgress onto the Silurian Ludlow-Pridoli beds of the upper part of the ‘Alternances Argilo-gréseuses’ Formation. The present work demonstrates a succession of miospore assemblages from closely sampled layers that have been stratigraphically dated as Ludlow–middle Pridoli and early Lochkovian by chitinozoans and acritarchs. Over 80 species of cryptospores and trilete spores have been identified. Modified detailed morphological interpretations are given. The miospore assemblages are correlated with miospore zonation schemes established for the type sequences of the Welsh Borderland, and those previously described from Libya. Early occurrences of some species as Streelispora newportensis on the western Gondwana plate, are put forward by comparison with the Old Red Sandstone continent. Phytogeographic and palaeobotanic implications based on these observations are discussed.     

Blackened bioclasts and bituminous impregnations in the Koněprusy Limestone (Lower Devonian), the Barrandian area, Czech Republic: implications for basin analysis

Carbonate reef talus facies of the Koněprusy Limestone (Pragian, Lower Devonian, Barrandian) locally exhibit widespread impregnation by organic matter resulting in a partial to complete blackening of the limestones. Two contrasting types of impregnation are recognized: blackening of individual carbonate fossils and bioclastic layers within the limestone originated very early during diagenesis. The blackening is due to finely dispersed organic matter and possibly some iron sulphides and clay minerals that selectively adhered to the outer layers of corals, bryozoans, and crinoid fragments, leaving other fossils unaltered. These darkened fossils are similar to black pebbles—i.e., reworked, dark to black limestone clasts and bioclasts that are known to occur exclusively in shallow-water zones of both ancient and modern carbonates. The alteration of fossil fragments may have taken place in very shallow-water environments, possibly those of saline and reducing back-reef lagoons or supratidal-intertidal zones, with organic matter being derived from decayed algae and microbes, or early vascular terrestrial plant material. Following the coloration, the blackened fossils were removed from their original position by waves or storms and transported into relatively deeper-water reef slope settings to form graded, “salt-and-pepper”-colored bioclastic beds. The presence of blackened fossils in the carbonate succession may point to episodic emergence and indicates a vanished vegetated siliciclastic hinterland that may once have existed to the west or south from the present-day erosive edge of the Barrandian Devonian strata. Subvertical veins cutting the Koněprusy Limestone and filled with black solid bitumen and blackened calcite resulted from a subsequent but substantially later diagenetic event, which is a testament of aqueous and petroleum fluid migration through the succession during deeper burial. Microthermometric characteristics of the aqueous inclusions embedded in vein calcite indicate that the veins were precipitated by brines of low to moderate salinity (0.5–9.5 wt% NaCl equiv.) with temperatures in the range of 87–116°C. The bitumen in the veins is epi-impsonite (R_r?=?0.70–1.90%), which is interpreted as degraded petroleum residuum that experienced thermal alteration at around 120°C. The AFT modeling combined with fluid inclusion microthermometry and wider geological considerations indicate that the veins originated during the Variscan orogeny, most probably upon deep burial of the Lower Paleozoic strata in Carboniferous time.