Tonganoxichnus a new insect trace from the Upper Carboniferous of eastern Kansas

Upper Carboniferous tidal rhythmites of the Tonganoxie Sandstone Member (Stranger Formation) at Buildex Quarry, eastern Kansas, USA, host a relatively diverse arthropod-dominated ichnofauna. Bilaterally symmetrical traces displaying unique anterior and posterior sets of morphological features are well represented within the assemblage. A new ichnogenus, Tonganoxichnus, is proposed for these traces. T. buildexensis, the type ichnospecies, has an anterior region characterized by the presence of a frontal pair of maxillary palp impressions, followed by a head impression and three pairs of conspicuous thoracic appendage imprints symmetrically opposite along a median axis. The posterior region commonly exhibits numerous delicate chevron-like markings, recording the abdominal appendages, and a thin, straight, terminal extension. T. buildexensis is interpreted as a resting trace. A second ichnospecies, T. ottawensis, is characterized by a fan-like arrangement of mostly bifid scratch marks at the anterior area that records the head- and thoracic-appendage backstrokes against the substrate. The posterior area shows chevron-like markings or small subcircular impressions that record the abdominal appendages of the animal, also ending in a thin, straight, terminal extension. Specimens display lateral repetition, and are commonly grouped into twos or threes with a fix point at the posteriormost tail-like structure. T. ottawensis is interpreted as a jumping structure, probably in connection with feeding purposes. The two ichnospecies occur in close association, and share sufficient morphologic features to support the same type of arthropod producer. T. buildexensis closely mimics the ventral anatomy of the tracemaker, whereas T. ottawensis records the jumping abilities of the animal providing significant ethologic and paleoecologic information. The presence of well-differentiated cephalic, thoracic, and abdominal features, particularly in T. buildexensis, resembles the diagnostic tagmosis and segmentation of insects. Detailed analysis of trace morphology and comparison with described Paleozoic insect fossils and extant related forms suggest a monuran as the most likely tracemaker.     

The origin and paleoecologic significance of the trace fossil Asteriacites in the Pennsylvanian of Kansas and Missouri

The trace fossil Asteriacites, recorded in Cambrian to Recent shallow- and deep-marine facies, is traditionally interpreted as the resting trace of asterozoans. Well-preserved specimens of A. lumbricalis are abundant in Pennsylvanian (Upper Carboniferous) shallow- and marginal-marine siliciclastic deposits of eastern Kansas and western Missouri. Detailed morphologic analysis of these specimens suggests that they record the activities of mobile epifaunal ophiuroids. Evidence of a brittle star (ophiuroid) producer rather than sea star (asteroid) is provided by (1) trace-fossil morphologic features reflecting the anatomy of the producer (e.g., well-differentiated central structure, slender vermiform arms) and ophiuroid burrowing technique (e.g., proximal arm expansion, arm branching), and (2) mode of occurrence (e.g., gregarious behavior, horizontal and vertical repetition). Vertical and horizontal repetition produces complex aggregates of A. lumbricalis that are interpreted either as escape structures (fugichnia) or as feeding structures, respectively. Ophiura texturata is proposed as a modern analogue for the A. lumbricalis producer, based on inferred life habit and feeding behavior. Asteriacites lumbricalis is present in two different intertidal trace-fossil assemblages. The first assemblage is characterized by high diversity and records tidal flats developed outside of embayments under normal marine conditions. The second assemblage consists of A. lumbricalis together with a few other ichnotaxa and represents a depauperate association that developed in restricted tidal flats within an embayment or estuarine setting. This challenges the conventional view of Asteriacites as a normal-marine salinity indicator. Some echinoderms, and particularly asterozoans, penetrate and inhabit modern environments of depressed salinity. The presence of Asteriacites in Pennsylvanian marginal-marine facies of Kansas and Missouri provides evidence that ophiuroids had adapted to brackish-water conditions by the late Paleozoic.     

贵州罗甸纳水上石炭统(宾夕法尼亚亚系)地层的再研究

本文详细描述了贵州罗甸纳水上石炭统(宾夕法尼亚亚系)剖面的生物地层和年代地层,其牙形刺序列自上而下可详细划分为：Streptognathodus isolatus, S. wabaunsensis, S. tenuialveus, S. firmus, Idiognathodus nashuiensis , Streptognathodus simulator, S. guizhouensis , S. gracilis-S, excelsus , S. cancellosus , S. clavatulus , S. nodocarinatus , Idiognathodus podolskensis , Mesogondolella clarki -Idiognathodus robustus , Diplognathodus ophanus-D, ellesmerensis, Idiognathoides ouachitensis, Streptognathodus expansus, Idiognathoides sulcatus parva, Idiognathodus primulus-Neognathodus bassleri, Idiognathodus primulus-Neognathodus symmetricus, Neognathodus symmetricus, Idiognathoides corrugatus-I, pacificus, I. sinuatus, I. sulcatus sulcatus, Declinognathodus noduli ferus 和 Gnathodus bilineatus bollandensis 等带。 Declinognathodus noduliferus 和 Streptognathodus isolatus 的首次出现分别代表上石炭统(宾夕法尼亚亚系)和二叠系的开始。根据牙形刺和有孔虫的序列,罗甸纳水剖面的上石炭统(宾夕法尼亚亚系)地层自下而上可划分为罗苏阶(Luosuan)、滑石板阶(Huashibanian)、达拉阶(Dalaan)和马平阶(Mapingian),并可与俄罗斯的巴什基尔阶(Bashkirian)、莫斯科阶(Moscovian)、卡西莫夫阶(Kasimovian)和格舍尔阶(Gzhelian),北美的莫罗阶(Morrowan)、阿托克阶(Atokan)、得梅因阶(Desmoinesian)、密苏里阶(Missourian)和弗吉尔阶(Virgilian)进行对比。另外,本文也详细讨论了剖面中的石炭系中间界线及石炭-二叠系界线。     

Large Pelycosaur Footprints from the Lower Pennsylvanian of Alabama,USA

Relatively large tetrapod footprints from Lower Pennsylvanian strata of the Pottsville Formation in the Warrior basin coal field of Alabama were previously assigned to the ichnogenus Attenosaurus. However, three morphotypes are present, and we assign these tracks to Attenosaurus, a new ichnogenus (Alabamasauripus) and Dimetropus. Most of these tracks apparently were made by large pelycosaurs, for which there is no bone record older than Late Pennsylvanian.     

Reappraisal of Oklahomaichnus,a supposed amphibian trackway from the Pennsylvanian of Oklahoma,USA

Oklahomaichnus millsii Sarjeant, from the Pennsylvanian (Missourian) Nellie Bly Formation of Oklahoma, is not the trackway of a lepospondyl amphibian, as originally proposed. Instead, it is the undertrack of an arthropod.     

Late Pennsylvanian to Middle Permian revised algal and foraminiferan biostratigraphy and palaeobiogeography of the Lycian Nappes (SW Turkey): Palaeogeographic implications

Shallow-marine limestones associated to a Palaeotethyan seamount in the Teke Dere unit of the Tavas Nappe (Lycian Nappes, SW Turkey) are essentially latest Moscovian-Kasimovian in age. The wide range of microfauna and -flora of the series show biogeographic affinities comparable to those from the northern Palaeotethyan borders (especially to assemblages from the Carnic Alps, Urals, Donbass and Darvaz). These biogeographic affinities seem to persist until the end of the Early Permian (Artinskian). The Middle Permian fauna is represented by the typical warm, tropical assemblages known at the same time in the Palaeotethys (NW Caucasus, Darvaz, south China, Primorie and Japan), and in the Neotethys (Transcaucasia, central Iran, southern Afghanistan and Sibumasu). The new Kasimovian algae and incertae sedis Novantiellopsis elliottii n. gen. n. sp., Uvanellopsis fluegelii n. gen. n. sp., Tubiphytes rauzerae n. sp. and Asselodiscus davydovi n. sp. are described.     

The trace fossil Nummipera eocenica from the Tatra Mountains,Poland: morphology and palaeoenvironmental implications

Jach, R., Machaniec, E. & Uchman, A. 2011: The trace fossil Nummipera eocenica from the Tatra Mountains, Poland: morphology and palaeoenvironmental implications. Lethaia, Vol. 45, pp. 342–355. The tubular trace fossil Nummipera eocenica Hölder 1989 occurs in a single stratigraphical horizon in Eocene nummulitic limestones of the Tatra Mountains, Poland. The wall of N. eocenica is built of Discocyclina and Nummulities (larger foraminifera) tests, very rarely of the Ditrupa (Polychaeta) tube fragments, bivalve shell fragments, echinoid spines and coralline algae. Morphotype are distinguished on the basis of wall composition and structure. Morphotype A is dominated by fusiform Discocyclina tests, which were preferentially selected by the trace makers for construction of a well‐constructed and resistant wall. Morphotype B contains more robust tests of Nummulites, while morphotype C is dominated by saddle‐shaped tests of Discocyclina. Nummipera eocenica was produced during a period of seafloor stabilization caused by a deepening. The succession of the morphotypes B, A reflects diminishing energy and increasing water depth. Probably morphotype C represents even lower energy environment than morphotype A. The trace fossil is interpreted as a domichnion, which wall was constructed for protection. The trace maker can be considered between polychaetes and crustaceans; however, comparisons to the closest recent analogues, the polychaete Diopatra cuprea or alpheid shrimps, are not satisfactory. □Bartonian, burrow, Carpathians, large foraminifera, trace fossils.     

Saccoglossus testa from the Mazon Creek fauna (Pennsylvanian of Illinois) and the evolution of acorn worms (Enteropneusta: Hemichordata)

The limited fossil record of enteropneust hemichordates (acorn worms) and the few external features that distinguish the four families have provided a challenge to our understanding of the evolution of the group and their various feeding adaptations. The middle Pennsylvanian Saccoglossus testa sp. nov. from the Mazon Creek, Westfalian D Carbonate Formation, Francis Creek Shale of northern Illinois provides evidence for the exploitation of surface sediments. Saccoglossus testa has a long proboscis characteristic of the extant genus Saccoglossus, a specialist in surface deposit feeding. The collar is as long as it is wide. The anterior trunk lacks a distinctively wide branchial region. These three features distinguish it from its sympatric enteropneust species Mazoglossus ramsdelli Bardack that has a proboscis characteristic of an infaunal deposit feeder. It is the seventh known species of fossil enteropneust, including a resting trace of a Lower Triassic fossil that has collar lips that characterize the extant deep‐sea family Torquaratoridae, and which represents a second parallel evolution of surface deposit feeding. An analysis of the seven fossils shows that the earliest Enteropneusta had a relatively simple harrimaniid‐like body plan, and that the spengelid, the torquaratorid and lastly the most complex ptychoderid body plan appeared in that chronological order.     

The vacuolar-type ATPase from insect plasma membrane: immunocytochemical localization in insect sensilla

Summary This comparative immunocytochemical investigation provides evidence that the electrogenic potassium pump of insect sensilla is a vacuolar-type proton ATPase energizing potassium-proton antiport, as was shown recently for the electrogenic potassium pump in the larval midgut of the sphinx moth Manduca sexta. Antennal sensilla of the saturniid moth Antheraea pernyi were probed with antibodies to the midgut vacuolar-type ATPase. The monoclonal antibodies recognized their epitopes in the native and SDS-denatured state, and bound specifically to the subunit with the relative molecular mass (M_r) of 67000 (antibody 86-3) or to the subunits of M_r 28000 and 16000 (antibody 47-5). Both antibodies labelled the apical region of the auxiliary cells, as was demonstrated by immunofluorescence microscopy. Immunogold-electron microscopy localized the binding sites of the 47-5 antibody in the highly folded apical plasma membranes of the auxiliary cells. Labelling was selective and was detected in all types of examined sensilla (S. trichodea, S. styloconica, S. coeloconica). These findings are in agreement with the current view that an electrogenic potassium pump is situated in the apical plasma membrane of the auxiliary cells and that the pump is involved in driving the receptor current. They support the hypothesis that a proton-motive force generated by a vacuolar-type ATPase provides an alternative to the classical Na⁺/K⁺-ATPase to energize animal plasma membranes.     

Early Pennsylvanian (Langsettian) fish assemblages from the Joggins Formation,Canada, and their implications for palaeoecology and palaeogeography

A review of all available specimens of fossil fishes from the classic Pennsylvanian Joggins locality of Nova Scotia, Canada, reveals the existence of a diverse community of chondrichthyans (xenacanthids, ctenacanthids and the enigmatic Ageleodus), acanthodians (gyracanthids), sarcopterygians (rhizodontids, megalichthyids, rhizodopsids and dipnoans) and actinopterygians (haplolepids). Reassessment of supposed endemic species (Ctenoptychius cristatus, Sagenodus plicatus, Gyracanthus duplicatus) indicates they are invalid, and overall, the assemblage comprises cosmopolitan taxa that were widespread around the coasts of tropical Pangaea. Strontium isotope analysis of fish remains and a critical study of their facies context suggest that these fish communities occupied bodies of water with salinities across the marine–freshwater spectrum. This preponderance of euryhaline forms implies a community structure quite distinct from that of today and might represent a transitory phase prior to the establishment of fully freshwater fish communities. Interpretation of fish ecology provides further evidence that the Joggins Formation was deposited in a paralic setting, and the recognition of one previously undetected brackish incursion strengthens the link between sedimentary cycles at Joggins and Milankovitch‐induced glacio‐eustatic change. Furthermore, interregional correlation of these marine transgressions supports palynostratigraphical arguments for an early Langsettian age for the Joggins Formation. This places tighter constraints on the age of the earliest known crown amniote, Hylonomus lyelli, an important calibration point used in phylogenomic studies.     

The youngest record of Carcharopsis (Chondrichthyes) from the Pennsylvanian of Xinjiang Uyghur Autonomous Region,China

A chondrichthyan tooth having a serrated edge, from the Late Pennsylvanian of Xinjiang Uyghur Autonomous Region, China, was identified as Edestus sp. [Cheng, Z., Lucas, S.G., Zidek, J., 1996. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1996 (11), 701-707]. Because of the lack of bilateral symmetry, the tooth does not belong to Edestus or any other known edestoid. It is too incomplete for a definite identification, but its features are consistent with Carcharopsis, which also has serrated teeth, and not with any other described late Paleozoic chondrichthyan genus. If it is Carcharopsis, it is the latest occurrence of the genus, which was previously known only from the Mississippian and the Early Pennsylvanian.     

Influence of transportation and time-averaging in fossil assemblages from the Pennsylvanian of Oklahoma

The relative effects of shell destruction, transportation, and time-averaging have been assessed in seventeen soft-bottom, benthic fossil collections from the Pennsylvanian Wewoka Formation in Hughes County, Oklahoma. Invertebrates were identified to species level, and each was subject to a taphonomic evaluation to determine whether it was in place. Species that were not deemed parautochthonous (living at the location of collection shortly before final burial) were eliminated from the faunal lists to reduce the effects of time-averaging and transportation in the fossil assemblages. Both reciprocal averaging and non-metric multidimensional scaling ordinations revealed two biofacies. These were distinguished using both presence/absence data and numerical counts of species despite an overwhelming number of taxa occurring in both biofacies. Comparison between ordinations based on entire faunal lists with those based on parautochthonous faunal lists indicates that the two biofacies faithfully reproduce differences in the living distributions of member taxa but that the two biofacies underwent different degrees of time-averaging and transportation. Time-averaging leads to accumulation of taxa over time, thereby making presence/absence composition of collections more similar. Transportation, on the other hand, homogenizes originally patchy population distributions and makes relative abundances more uniform. Based on differences in abundances and presence/absence patterns, the stratigraphically lower biofacies underwent more transportation and more time-averaging than the upper biofacies.     

Pennsylvanian (Late Carboniferous) calcareous microfossils from Cedro Peak (New Mexico,USA). Part 1: Algae and Microproblematica

At Cedro Peak (Manzanita Mountains, central New Mexico), the Pennsylvanian succession is divided into four formations: Sandia, Gray Mesa, Atrasado, and lower part of Bursum. The sampled limestones of the Gray Mesa and Atravasado formations yielded three age distinctive fossil assemblages: (a) latest Atokan/early Desmoinesian; (b) late early Desmoinesian, and (c) middle Virgilian. The calcareous algae, incertae sedis algae and cyanobacterial-foraminiferal consortia are described here. The following new taxa are named: Epimastoporaceae n. fam.; Anthracoporellopsis novamexicana n. sp.; Tubiphytidae n. fam.; and Latitubiphytes n. gen.     

The trace fossil Thalassinoides from the Upper Ordovician of Tuva

Traces of Thalassinoides (the tunnels of unknown burrowing organisms) are described from carbonates of the Khondelensky layers of the Upper Ordovician of Tuva. Hitherto, this fossil was unknown in the Ordovician of the USSR. They demonstrate great similarity with Thalassinoides from coeval deposits of the Great Basin, USA. The traces are assumed to have been made in terrigenous-carbonate sediments deposited on the areas of a gently sloping shelf in quiet water, below wave base. □ Upper Ordovician, terrigenous-carbonate sediments, burrows.     

A New Ichnospecies of Arthrophycus from the Late Carboniferous (Pennsylvanian) of Michigan,USA

A unique ichnofossil assemblage from Pennsylvanian-age sandstones near Eaton Rapids, Michigan, USA, comprises straight-to-curved traces preserved in convex hyporelief, with the transverse ridges and median grove associated with the ichnogenus Arthrophycus. The Michigan traces show some branching or pseudo-branching (also known from other Arthrophycus specimens) and are among the smallest structures (millimeters in diameter) attributed to this ichnogenus. The orientation of the Michigan Arthrophycus burrows is distinct from other ichnospecies of this taxon in the pronounced co-planar alignment of the burrows, as opposed to the multiplanar, interweaving, “bundled” nature typical of the ichnogenus. On this basis we assign the Michigan specimens to a new ichnospecies of Arthrophycus. The “paralleling behavior” of the new taxon may reflect a strategy of the tracemaker to avoid previous burrows and reflect differences in resource availability or current energy. This new taxon supports previous records of the occurrence of this ichnogenus in Upper Carboniferous strata.     

The first agetopanorpine mecopteroid insect from Middle Permian of China (Insecta: Mecoptera: Permochoristidae)

Sinoagetopanorpa permiana n. gen., n. sp., first Chinese Permian Agetopanorpinae, is described and compared to the other genera of this group.     

Pennsylvanian (Late Carboniferous) calcareous microfossils from Cedro Peak (New Mexico,USA). Part 2: Smaller foraminifers and fusulinids

The Pennsylvanian stratigraphic section of the Manzanita Mountains (central New Mexico, USA) is restudied at Cedro Peak. This Pennsylvanian succession is divided into the Sandia Formation, Gray Mesa Fm, Atrasado Fm, and lower part of Bursum Fm. The sampled limestones of the Gray Mesa and Atrasado formations yielded three age-distinctive fossil assemblages: they are of (a) latest Atokan/early Desmoinesian, (b) late early Desmoinesian, and (c) middle Virgilian. The following new taxa of foraminifers are proposed: Endoteboidea n. superfam.; Spireitlinidae n. fam.; Millerellinae n. subfam.; Pseudonovella marshalli n. sp.; Pseudonovella ohioicus nom. nov.; Pseudoacutella n. gen.; Pseudoacutella hoarei nom. nov.; Plectofusulina manzanensis n. sp. Translated or emended names are: Staffelloidea, Ozawainelloidea, Ozawainellidae, and Profusulinellidae.     

Microproblematic endophytes and epiphytes of fern pinnules from the Upper Pennsylvanian of France

Ten different types of microproblematic structures occur on the surfaces, and in epidermal and hypodermal cells of fern pinnules preserved in the Late Pennsylvanian Grand-Croix cherts from France. These include thick-walled spheres, some with a prominent exit site, aggregations of variously sized spheres that may be surrounded by a membrane or sheath, a cylindrical structure composed of polygonal compartments, and branched hyphae/filaments with intercalary swellings protruding from a small opening in the surface (probably a stomatal pore). Some of the structures likely were produced by some fungal group, and may represent (resting) spores, microsclerotia, and conidiophores. The affinities of the others, and sometimes even their biological nature, remain difficult to determine. Nevertheless, documenting these structures is important because it provides the first indication for vascular plant leaves that were colonized by different types of organisms in a late Paleozoic ecosystem.     

Fish and tetrapod communities across a marine to brackish salinity gradient in the Pennsylvanian (early Moscovian) Minto Formation of New Brunswick,Canada, and their palaeoecological and palaeogeographical implications

Euryhaline adaptations in Pennsylvanian vertebrates allowed them to inhabit the marine to freshwater spectrum. This is illustrated by new assemblages of fish and tetrapods from the early Moscovian Minto Formation of New Brunswick, Canada. Fish include chondrichthyans (xenacanthids and the enigmatic Ageleodus), acanthodians (gyracanthids and acanthodiforms), sarcopterygians (rhizodontids, megalichthyids and dipnoans), and actinopterygians (eurynotiforms). Tetrapods include small‐ to medium‐sized, and largely aquatic, stem tetrapods (colosteids) and anthracosaurs (embolomeres). A key finding is that the parautochthonous fossil assemblages are preserved across a salinity gradient, with diversity (measured by the Simpson Index) declining from open marine environments, through brackish embayments, and reaching a nadir in tidal estuaries. Chondrichthyans dominate the entire salinity spectrum (65% of fossils), a distribution that demonstrates a euryhaline mode of life, and one large predatory chondrichthyan, Orthacanthus, may have practised filial cannibalism in coastal nurseries because its heteropolar coprolites contain juvenile xenacanthid teeth. In contrast, other fish communities were more common in open marine settings while tetrapods were more common in coastal brackish waters. While all these faunas were also likely to have been euryhaline, their osmoregulation was, perhaps, less versatile. The demonstration of widespread euryhalinity among fish and aquatic tetrapods explains why Pennsylvanian faunas generally show a cosmopolitan biogeography because taxa were able to disperse via seaways. It also resolves the paradox of enriched strontium isotopic signatures observed in these faunas because organisms would have been, at times, exposed to continental water bodies as well. Therefore, our new findings contribute to the long‐running debate about the ecology of Pennsylvanian fishes and tetrapods.