Geometric shell growth and mode of life of a Devonian chonetoidean brachiopod

Morphological and statistical analysis of the chonetoid species Kentronetes variabilis from the Lower Devonian (Lochkovian) of the Argentine Precordillera demonstrate ontogenetic changes and allometric relationships between characters. A special study was made of spine distribution, morphology, and growth, compared to valve growth. The first, inner, developed spines (pairs 1–1'and 2–2') continued to grow after development of the following outer pairs. The spacing of spines, their diameter, and the density of growth rings vary from beak to posterolateral margins following a specific 2ⁿ geometric growth factor, compared to the regular, almost linear growth of the valves, attested by growth lines. The linear growth rate of outer spines (pairs 3–3'and 4–4') can be 6–8 times more rapid than that of the shell on the valve margin. Ontogenetic changes in spine morphology are interpreted as a response to changes in the mode of life.     

Two new Middle Devonian megaspores from Saudi Arabia

Two new megaspore species, Biharisporites jubahensis and Verrucisporites yabrinensis, are described from the Middle Devonian Jubah Formation of south central Saudi Arabia. Miospore-based biostratigraphy indicates that the age of the megaspore-bearing interval is no older than the early Givetian Geminospora lemurata Interval Zone, and not younger than Givetian. The presence of similar species in both Arctic Canada and Saudi Arabia suggests that the megaspore-producing plants achieved wide distribution in the Middle Devonian.     

Clustered trilobite assemblages in the Middle Devonian Hamilton Group

Clusters of well-preserved trilobites occur throughout lower and middle Paleozoic strata. This phenomenon, however, remains poorly understood. Several thin, regionally persistent beds in the Middle Devonian (Givetian) Hamilton Group of New York State display taphonomic and sedimentary features which indicate rapid burial by storm-generated clouds of fine-grained sediment. Furthermore, components in these units display little or no post-mortem disturbance and, therefore, reflect in faunal associations. These beds contain two types of clustered assemblages of the trilobites Phacops (Green), Greenops boothi (Green) and Dechenella rowi (Green). Body clusters comprise dense assemblages of three or more complete trilobite corpses. Moult clusters , in contrast, are composed of three more moult ensembles; i.e., articulated thoracopygidia in close proximity to corresponding cephala. Clusters are species-segregated and contain only a small portion of the total size range exhibited by trilobite species. Moreover, body clusters and moult clusters are morphometrically undifferentiated indicating that these two cluster modes represent related phenomena. Analogy with modern marine arthropods suggests that trilobites assembled into monospecific, age-segregated clusters and moulted prior to en masse copulation.     

The Trace Fossil Paleodictyon within The Cruziana Ichnofacies: First Record from The Devonian in Pennsylvania

Offshore marine deposits of hemipelagic dark-gray shales comprising the Middle Devonian Mahantango Formation have yielded the first evidence of the trace fossil Paleodictyon from Pennsylvania. Paleodictyon occurs in conjunction with a diversity of largely deposit-feeding trace fossils belonging to the Cruziana ichnofacies, and documents another example of a shallower-water occurrence of this ichnofossil in Paleozoic rocks.     

Substrate specific associations of epebionts on Middle Devonian brachiopods: Implications for paleoecology

A detailed study of over 2500 host brachiopods, from the Middle Devonian Hamilton Group of New York State, revealed distinct patterns of epibiont encrustation, that provide insight into taphonomy and paleoautecology of the host brachiopod shells and depositional environments. The concavo‐convex orthid, Tropidoleptus carinatus (Conrad), as well as strophomenid, and smooth athyrid brachiopods are among the most heavily encrusted. However, terebratulids of nearly identical size and shape are relatively clean of epibionts. This selective distribution strongly suggests that epibionts were discouraged from settling on punctate brachiopods. Brachiopods with small spines and frills were also nearly clean of epibionts, possibly because of entrapment of a mud layer, which made the outer layer of the host inhospitable for larval settling. Concavo‐convex taxa reveal high percent coverage and diversity of epibionts on the convex valve, which probably rested on the substrate during the life of brachiopod. This pattern is observed even on brachiopods that were buried with the convex valve downward. This implies complex post‐mortem histories involving multiple episodes of reorientation and colonization.     

A new homosporous,arborescent lycopsid from the Middle Devonian of Xinjiang,Northwest China

Abstract: A new arborescent lycopsid, Hoxtolgaya robusta gen. et sp. nov., is described from the Middle Devonian of Xinjiang, Northwest China. It has stems up to 90 mm wide with fusiform leaf bases and long linear microphylls. Sporophylls are not aggregated into strobili and are isomorphic, with sporangia homosporous and bearing Acinosporites‐type microspores. A syndrome of characters in Hoxtolgaya, including the arborescence and the homospory, implies that the arborescent habit is not necessarily correlated with the heterospory in the early evolution of arborescent lycopsids. Homosporous, arborescent lycopsids probably represent one of the transient forms between the Devonian herbaceous protolepidodendrids and the Devonian–Carboniferous heterosporous arborescent lycopsids.     

Biostratinomy and functional morphology of enrollment in two Middle Devonian trilobites

Although it is common knowledge that many trilobites enrolled, behavioral and functional aspects of enrollment are not at all well understood. Taphonomic details portrayed by enrolled trilobites in the Middle Devonian Hamilton Group (New York State) indicate that enrollment was a complex and morphologically constrained behavior. The trilobites Phacops rana (Green) and Greenops boothi (Green) are frequently enrolled in Hamilton strata; biostratinomic data indicate two very different enrollment postures. Interlocking morphologies (coaptative devices) and apodeme structure and disposition indicate that these postures reflect specific behaviors which involved interaction between tergal structures, inferred musculature, and the substratum. Phacops enrolled by burrowing forward and down into the sediment; dorsal muscles, attached to prominent articulating half-rings, imbricated the thorax such that each lappet overlapped the next posterior segment and locked into a posterior pleural facet. The pygidium was brought into place as the posterior segments of the thorax were placed into vincular notches along the lateral margin of the ventral cephalon. The pygidium locked with the cephalic vincular furrow to complete ‘perfect sphaeroidal’ closure. Greenops enrolled with the cephalon in an upright position at the sediment surface; a submarginal furrow on the ventral surface of the pygidium received the anterior rim of the cephalon. Relatively narrow articulating half-rings limited pleural rotation. Segments were loosely locked into narrow facets at the anterior margin of the next posterior lappet. In spite of rudimentary lappet and half-ring structures, Greenops displays an elaborate system of thoracopygidial muscles which involved dorsoventral and longitudinal attachments along the thorax and into the pygidium. Phacops, in contrast, displays very poorly developed apodemes which occur in the thorax only. Longitudinal muscle strength was likely less important during Phacops enrollment than is evident for the Greenops enrollment procedure. Conversely, Phacops clearly relied to a great degree upon competent closure devices which are poorly developed in Greenops. Biostratinomic data reveal different enrollment behaviors which reflect the function of different enrollment-related morphologies present in each species.     

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.     

Level-bottom brachiopod communities in the Middle Devonian of New York

Depositional environments of the Onondaga Limestone from central to southeastern New York are found to be normal, subtidal marine, due to the absence of characteristic supratidal or intertidal sedimentary features and the presence of a typical, diverse, marine level-bottom community framework. Post-mortem transport has not been extensive, as evidenced by low articulation ratios, lack of abraded valves, and complete ontogenetic gradations within species, which precludes large scale winnowing. Sedimentation rates appear to have been greatest in eastern New York where the Onondaga Limestone reaches a thickness almost three times that of the strata in central New York. Shaly beds in the central area represent periods of cessation of carbonate deposition rather than an influx of clastic material. Deposition terminated with the onset of deeper water characterized by a westerly advance of terrigenous sedimentation (the Marceilus Shale of the Hamilton Group). Nine brachiopod communities can be recognized in the Onondaga Limestone. There is a strong correlation between sediment-substrate and community type, reflecting the sedimentologic control of brachiopod community distribution. Sandy facies, cherty limestones and coral biostromes and bioherms are associated with inner-neritic deposition in Edgecliff time; argillaceous lime muds and lime sands are characteristic of mid-neritic deposition in Nedrow to Moorehouse time; and highly argillaceous lime muds are associated with outer-nerittc deposition in Seneca time.     

Shell repair and mode of life of Praenatica gregaria (Gastropoda) from the Devonian of Bohemia (Czech Republic)

Repaired shell injuries occuring in less than four per cent of examined specimens of the platyceratid gastropod Praenatica gregaria (Barrande, in Perner 1903) from the Lower Devonian (Pragian) of Bohemia probably result from non–lethal predatory attacks. The low frequency may reflect the resistance of the large, smooth, rapidly expanding shell to attack, but a high rate of fatal attacks could produce the same pattern. The habit of some platyceratid gastropods living on echinoderm calices may provide a refuge from benthic predators, but conclusive evidence for this mode of life in P. gregaria or the general assumption of platyceratid coprophagy is lacking.     

Integrated Middle Devonian chitinozoan and miospore zonation of the Amazonas Basin, northern Brazil

Middle Devonian chitinozoans were studied from 13 deep wells in the central part of the Amazonas Basin, northern Brazil, and compared with a newly established regional miospore zonation. Of the 23 chitinozoan species encountered, four are newly described: Ancyrochitina multibrachiata, Lagenochitina praeavelinoi, Ramochitina autasmirimense, and Ramochitina herculesi and seven species are retained in open nomenclature because of an insufficient number of well-preserved specimens. The stratigraphic intervals investigated herein correspond to three miospore interval range zones previously defined for the Amazonas Basin, namely, the Grandispora/Samarisporites spp. (GS); late Emsian-early Eifelian, Grandispora permulta (Per); late early Eifelian through the Eifelian-Givetian transition, and Geminospora lemurata-Chelinospora ex. gr. ligurata (LLi); early Givetian. A regional Middle Devonian chitinozoan biozonation, also consisting of three zones, is proposed for the investigated interval. These zones are the interval zone of Angochitina n. sp. A (early Eifelian), the interval zone of Alpenachitina eisenacki (late early to late Eifelian), and the total range zone of Ramochitina stiphrospinata (early Givetian).     

Signatures of hydrocarbon venting in a Middle Devonian Carbonate Mound (Hollard Mound) at the Hamar Laghdad (Antiatlas, Morocco)

Summary The Middle Devonian Hollard Mud Mound is situated in the eastern Hamar Laghdad, which is a small mountain range in the Tafilalt in SE Morocco. In contrast to the well known Lower Devonian Kess-Kess mounds, the Hollard Mound is of Middle Devonian age. The facies in the core of this mud mound differs from that of the other parts of the mound, and exhibits signatures of ancient hydrocarbon venting. The carbonate phases of the core facies are derived from the oxidation of vent fluids and consist of clotted micrite, a cryptocrystalline carbonate associated with spheres of uncertain origin, and a calcitic rim cement (rim cement B). These vent carbonates show δ¹³C values in the range of −11 to −20% PDB indicating that some of their carbon is derived from isotopically light hydrocarbons. Fossiliferous micrite has been affected by hydrocarbon venting in the proximity of the vent site, which is indicated by intermediate δ¹³C values between vent carbonates and not affected sediments. Bivalves occur in dense populations within the core facies. They form autochthonous shell accumulations and are almost exclusively articulated. it is likely that these bivalves were dependent on chemosynthesis similar to their counterparts at modern vents. The vent deposits also exhibit an unusual prasinophyte assemblage, which might have been linked to the specific nutrient availability at the vent site. The ancient vent site is characterized by an enhanced carbonate precipitation and rapid lithification. The latter is corroborated by the three-dimensional preservation of phytoplankton (prasinophytes and acritarchs) and the occurrence of stromatactoid pores. An early phase of carbonate corrosion predating the formation of vent carbonates affected the fossiliferous micrite of the core facies and is thought to be related to a phase of H₂S-rich venting.     

A new euphyllophyte Kunia venusta gen. et sp. nov. from the Middle Devonian of Yunnan,South China

Abstract Kunia venusta gen. et sp. nov. is reported from the late Middle Devonian (Givetian) Haikou Formation near Kunming City, Yunnan Province, China. This plant has three orders of naked axes that divide pseudomonopodially. The second‐ and third‐order axes occur in a helix. Fertile appendages are distantly spaced and helically inserted to the third‐order axis; they comprise equally dichotomous branches terminated by two clusters of paired and fusiform sporangia. Sterile appendages are dichotomous and distally recurved. A comparison is made with the basal euphyllophytes including the trimerophytes, cladoxylopsids, zygopterids, stauropterids, and some relevant genera of uncertain affinity. The new plant resembles them in dichotomous appendages with terminal elongated sporangia, but differs mainly in the three orders of pseudomonopodial axes bearing helical laterals. It is thus placed in the Euphyllophytina as incertae sedis. It is suggested that an evolutionary divergence in the branching pattern and appendage morphology might have occurred in the Middle Devonian euphyllophytes, that is, maintaining three dimensions versus yielding more or less planation.     

Fourier transform infrared characterization of the Middle Devonian non‐vascular plant Spongiophyton

Spectroscopy information (functional groups and semiquantitative data) by Fourier transform infrared (FTIR) spectroscopy of Spongiophyton Kräusel emend. Chaloner et al. is reported for the first time, in an attempt to identify spectroscopic patterns that would differentiate species of these taxa. A total of 33 specimens identified as S. lenticularis, S. nanum and S. minutissimum and preserved as compressions were analysed. They come from the same fossiliferous horizon of one outcrop situated in the uppermost Middle Devonian São Domingos Formation of the Paraná Basin in southern Brazil. The results reveal that functional groups in the 3000–2800 and 1800–700 cm^?1 wavenumber regions are present in all specimens. Infrared‐derived ratios (CH₂/CH₃, Al/Ox, Ox1/Ox2, C=O cont., C=C cont., Al2 and Ar/Al) and types of kerogen led to a comparison of the morphological characteristics of the studied material and indicate that compressions of S. lenticularis, S. nanum and S. minutissimum contain different organic compounds. S. nanum and S. minutissimum show higher oxidation and lower aromaticity of its organic matter than S. lenticularis. Additionally, S. lenticularis shows low A‐factor values, suggesting a composition similar to type III kerogen, while S. nanum and S. minutissimum showed intermediate to high A‐factor values, suggesting a composition similar to type II kerogen. In conclusion, the chemical‐analytical techniques and statistical analysis (using PCA technique) reveal a closer taxonomic relationship between S. nanum and S. minutissimum than either has with S. lenticularis, supporting their taxonomic separation, previously proposed in the literature on the basis of morphological features. Lastly, the chemical signatures support the hypothesis that links these enigmatic fossils to a type of lichen or other early inland plant.     

Autecology of Labrocuspis, a Middle Devonian omphalotrochid gastropod

Labrocuspis kobayashii , a Middle Devonian euomphalacean omphalotrochid gastropod from Japan has a wide callus pad on its base similar to that seen in the living trochid Umbonium. L. kobayashii could support the shell over the cephalopedal mass in a similar fashion to that seen in the Umbonium species. This reconstruction indicates that the labral projection in the outer lip was above the head, and the opisthocyrt basal lip and the sinus in the outer lip were loci of inhalant and exhalant currents, respectively. Therefore, the animal is assumed to have had only one ctenidium (the left). The presence of a wide callus pad is indicative of an infaunal life habit for the Umbonium species, Labrocuspis and some Paleozoic gastropods; clamping of the shell against the foot is also suggested for some omphalotrochid gastropods. These features indicate an additional adaptive strategy employed by euomphalaceans.     

A new Middle Devonian gasterocomid crinoid from central Turkey and revision of the Gasterocomidae

A juvenile cup, disarticulated arm plates, and columnals of the crinoid Arachnocrinus sarizensis n. sp. are described from the Eastern Taurus Mountains of central Turkey. This is the first Paleozoic crinoid based on a cup and arm plates reported from Turkey. Suggested revision of the Gasterocomidae includes transfer of Arachnocrinus and Ancyrocrinus to the family and rejection of Kopficrinus from the family. The stratigraphic range of Arachnocrinus is extended upward into the Middle Devonian from the Early Devonian. The Devonian of Turkey is of special interest because it includes Laurasian and Gondwanan components on different tectonic blocks. The paleogeographic range of Arachnocrinus is extended from the North American plate onto the Anatolian block, which would have been located on the southern edge of the Paleotethys Sea in the Middle Devonian in a shelf basin off the northern coast of Gondwana at approximately 42° south latitude.     

A reconsideration of Dimeripteris cornuta SCHWEITZER and CAI, a diminutive fossil plant from the Middle Devonian of Yunnan, China

Small plant fragments previously attributed to Dimeripteris cornuta SCHWEITZER and CAI and new specimens are described from the Middle Devonian (Givetian) Xichong Formation of Wuding, Yunnan Province, southwest China. A predominantly trifurcate branching pattern is recognized in lower orders of branching. Erect fusiform sporangia are borne in pairs terminally on a short pedicel on up to three times dichotomously-branching fertile unit. The beaked tips of the paired sporangia point outwards. This plant is distinguished from known plants by the trifurcate branching pattern and sporangia morphology, but is of problematic affinity. The plant is named Tauritheca cornuta (SCHWEITZER and CAI) WANG and BERRY nov. gen. and nov. comb.     

Miospore stratigraphy of Lower and early Middle Devonian deposits from Tidikelt, Central Sahara, Algeria

The Tidikelt region forms an outstanding area for subsurface Lower Devonian stratigraphy in the central Algerian Sahara. Sediments from five boreholes have revealed abundant and diverse assemblages of miospores, acritarchs, chitinozoa, scolecodont and microplant remains. The miospores are moderately well preserved. Three new miospore species (Dibolisporites saharansis nov. sp. Hassan Kermandji, Acinosporites conatus nov. sp. Hassan Kermandji and Scylaspora tidikeltense nov. sp. Hassan Kermandji) are described. Miospore assemblages vary through the regressive and transgressive sequences. Seven miospore assemblage biozones, including six new miospore assemblage biozones (Scylaspora tidikeltense-Perotrilites microbaculatus, Dictyotriletes emsiensis-Emphanisporites spinaeformis, Apiculiretusispora arenorugosa-Camptozonotriletes caperatus, Verrucosisporites polygonalis-Dictyotriletes subgranifer, Emphanisporites annulatus-Geminospora svalbardiae, Hystricosporites microancyreus-Grandispora protea, Calyptosporites velatus-Rhabdosporites langii) are proposed for the Lower and early Middle Devonian rocks of Tidikelt Plateau. The combined use of distinctive, wide distribution cosmopolitan and Gondwanan forms as biozonal and species characteristics permits accurate subdivision, dating and correlation of Tidikelt successions with other similar miospore zones of the Lower Devonian of Europe, Canada and other parts of Gondwana plate. The miospore data provide new explanations to stratigraphic relationships of regional rock units, sedimentary cycles and stratigraphic hiatus. The miospore biozones are proposed as a provincial biozonation, which may also be applied to other Palaeozoic rocks of similar miospore content.     

Sequence stratigraphy of the Early–Middle Devonian succession (Padeha Formation) in Tabas Block,East-Central Iran: Implication for mixed tidal flat deposits

The facies and sequence stratigraphic analyses of the Padeha Formation (Early–Middle Devonian) in the East-Central Iran are based on two measured stratigraphic sections in the Ozbak-Kuh (type section) and Deranjal (Dahaneh-Kalot section) Mountains in the northern Tabas Block, with a thickness of 492 and 320 m respectively. The field observations and laboratory studies were used to identify sixteen lithofacies and four architectural elements. The facies associations were classified into five categories including sandstone (St, Sp, Sr, Sh, Sl), fine grained (siltstone–claystone) or mud rock (Fl), interbedded sandstone–mud rock (Sr(Fl), Sr/Fl, Fl(Sr)), dolomite (Dl and Ds), and evaporate (El, Efl, Efm, Edl). Structural (wave and interference ripples, planar cross-bedding and herringbone, flaser-wavy and lenticular beddings, tepee and stromatolite structures, entrolothic foldings, shrinkage and syneresis cracks, rain drop imprints, and salt casts) and textural (mature–supermature quartzarenite) features, as well as predominant bimodal pattern of paleocurrents in most lithofacies, show that sediments of the Padeha Formation have been deposited in a tidal flat environment. The sandstone, interbedded sandstone–mud rock, and dolomite facies (SB and LA architectural elements) are related to subtidal and intertidal settings, and fine-grained sediments and evaporate facies (consisting of FF and E architectural elements) were deposited in the supratidal and sabkha settings. Detail sequence stratigraphic study led to identification of two 3rd order complete depositional sequences (DS₂ and DS₃) with SB₁ sequence boundaries and two incomplete sequences (DS₁ and DS₄). Supratidal and sabkha facies were formed during the fall of relative sea level and are related to the lowstand systems tract (LST). The intertidal facies were formed at a rise of relative sea level and are interpreted as transgressive (TST) and highstand (HST) systems tracts.     

Middle and Early Upper Devonian Miospore Zonations in Eastern Yunnan and the Significance of Their Stratigraphy

The Devonian marine and non marine deposits were widely distributed in East Yunnan, where rich marine faunas and floras were present. The Middle and early Upper Devonian miospores of Eastern Yunnan were also abundant and usually well- preserved. The present paper provides both analysis of qualitative and quantitative composition development of Middle and early Upper Devonian spore assemblage in Eastern Yunnan, and a palynological zonatlon containing four successive assemblage zones: 1.Zone of Calyptosporites velatus-Rhabdosporites langii (VL); 2. Zone of Archaeozonotriletes variabilis-Calyptosporites proteus (VP); 3. Zone of Geminospora lemurata-Cristatisporites triangulatus (LT); 4. Zone of Archaeoperisaccus ovalis-Lagenicula bullosum (OB). The Middle and early Upper Devonian spore assemblages of Eastern Yunnan may be compared with those in South China and West Qinling. The spore zonation possesses stratigraphical dating of the Devonian megafloras of the region, particularly those from the Middle and Upper Devonian. The proposed spore zonation is closely compared with that erected for the Middle and early Upper Devonian of Old Sandstone Continent and adjacent Region. Based on palynological data, the reconstruction of Paleogeography and Paleoecology are discussed.