A New Bivalved Arthropod from the Early Cambrian Sirius Passet Fauna, North Greenland

A new bivalved arthropod is described from the Lower Cambrian (?Upper Atdabanian) Buen Formation of North Greenland. Pauloterminus spinodorsalis gen. et sp. nov. possesses a bivalved carapace that covers the head, which has a single pair of antennae, and anteriormost thorax. No mouthparts are visible. The five‐segmented abdomen was limbless and terminated in a telson plus a pair of large, lobate uropods. A suite of at least six biramous thoracic limbs are present: the short endopods are made up of small, serial podomeres, while the exopods are lobate and may have functioned as gills as well as in swimming. Partially infilled guts are occasionally visible, suggesting that this animal may have been a sediment feeder. It is compared to other Cambrian bivalved arthropods, especially the waptiids Chuandianella ovata from the Lower Cambrian Chengjiang fauna (China) and Waptia fieldensis from the Middle Cambrian Burgess Shale (British Columbia). Of these three animals, the Greenland and Chinese taxa appear to be the most closely related. P. spinodorsalis possesses many typical arthropod features, but it also demonstrates more primitive characters that are more reminiscent of the lobopodians.     

Morphology and systematics of anthracomartidae (Arachnida: Trigonotarbida)

Abstract: X‐ray microtomography (XMT) was applied to three species of the extinct arachnid order Trigonotarbida, hosted in siderite nodules from the late Carboniferous British Middle Coal Measures. All three, Cryptomartus hindi ( Pocock, 1911 ), Cleptomartus plautus ( Petrunkevitch, 1949 ) and Maiocercus celticus ( Pocock, 1902 ), belong to the family Anthracomartidae. As well as providing interactive three‐dimensional visualisations of their likely appearance in life, XMT study has resolved new morphological detail, yielding key data about the likely stance and habitus of these early arachnids. Similarities in the form of the carapace, eyes and coxosternal region between anthracomartids and the Devonian genus Palaeocharinus Hirst, 1923 are highlighted. Analogies in limb disposition are drawn between anthracomartids and modern crab spiders (Araneae: Thomisidae), which exhibit sit‐and‐wait style predatory behaviour. Anthracomartids are relatively common fossils at many Coal Measures localities. Although they appear to exhibit limited morphological variation, numerous genera and species have been proposed in the literature, often distinguished from each other on rather trivial characters. From our reconstruction of well‐preserved examples, we have resolved a number of common features likely to be present in a typical anthracomartid. Comparisons to this model suggest that the characters on which many anthracomartid genera are distinguished may be artefacts of preservation. We hence treat Promygale Fri?, 1901 , Brachylycosa Fri?, 1904 , Perneria Fri?, 1904 , Coryphomartus Petrunkevitch, 1945 , Cryptomartus Petrunkevitch, 1945 , Pleomartus Petrunkevitch, 1945 , Cleptomartus Petrunkevitch, 1949 and Oomartus Petrunkevitch, 1953 as junior synonyms of the type genus of the family, Anthracomartus Karsch, 1882 . Only the genera Brachypyge Woodward, 1878 and Maiocercus Pocock, 1911 are additionally retained. Within Anthracomartus, Cleptomartus planus Petrunkevitch, 1949 , Cryptomartus meyeri Guthörl, 1964 , Cleptomartus hangardi Guthörl, 1965 and Cryptomartus rebskei Brauckmann, 1984 are treated as junior synonyms of A. hindi Pocock, 1911 (all syn. nov.). Cleptomartus plautus Petrunkevitch, 1949 and Anthracomartus denuiti Pruvost, 1922 are treated as junior synonyms of A. priesti Pocock, 1911 (both syn. nov.).     

Arthropod visual predators in the early pelagic ecosystem: evidence from the Burgess Shale and Chengjiang biotas

Exceptional fossil specimens with preserved soft parts from the Maotianshan Shale (ca 520 Myr ago) and the Burgess Shale (505 Myr ago) biotas indicate that the worldwide distributed bivalved arthropod Isoxys was probably a non-benthic visual predator. New lines of evidence come from the functional morphology of its powerful prehensile frontal appendages that, combined with large spherical eyes, are thought to have played a key role in the recognition and capture of swimming or epibenthic prey. The swimming and steering of this arthropod was achieved by the beating of multiple setose exopods and a flap-like telson. The appendage morphology of Isoxys indicates possible phylogenetical relationships with the megacheirans, a widespread group of assumed predator arthropods characterized by a pre-oral ‘great appendage’. Evidence from functional morphology and taphonomy suggests that Isoxys was able to migrate through the water column and was possibly exploiting hyperbenthic niches for food. Although certainly not unique, the case of Isoxys supports the idea that off-bottom animal interactions such as predation, associated with complex feeding strategies and behaviours (e.g. vertical migration and hunting) were established by the Early Cambrian. It also suggests that a prototype of a pelagic food chain had already started to build-up at least in the lower levels of the water column.     

Spinicaudatans and ostracods (Crustacea) from the Montceau Lagerstätte (Late Carboniferous, France): morphology and palaeoenvironmental significance

Spinicaudatans and ostracods form two components of the diverse arthropod fauna from the Montceau Lagerstätte (Stephanian, France). Spinicaudatans are represented by Montcestheria orri gen. and sp. nov. and Euestheria feysi sp. nov., and ostracods by a single species, Carbonita sp. aff. salteriana (Jones, 1862). Allied forms such as Montcestheria sp. aff. orri, Montcestheria sp. and Euestheria cebennensis (Grand'Eury, 1890), all from coeval localities in France, are also described. Montcestheria gen. nov. has carapace features, external (possibly sexual) dimorphism, preserved soft parts (e.g. appendages, gut) and resting eggs similar to those of Recent spinicaudatans, suggesting comparable lifestyles, reproductive strategies and feeding modes. Detailed anatomical comparisons are made with Cyzicus tetracerus from Recent ephemeral freshwater ponds. The ostracods belong to the Superfamily Carbonitoidea, which is a recurrent component of Carboniferous non‐marine biotas. Spinicaudatan‐rich assemblages typically occur in several Westphalian–Stephanian Lagerstätten (including Montceau) and localities from Europe and North America, where freshwater conditions prevailed, indicating that the group had already colonized continental waters by the Late Carboniferous. Similarities with the fauna from Recent temporary freshwater ponds (e.g. low diversity/high density spinicaudatan‐ostracod populations, synchronous spinicaudatan populations developing from resting eggs, high diversity/low density insects, amphibians) suggest the presence of temporary or ephemeral aquatic environments at Montceau as part of a complex limnic ecosystem. Flooding may have been the main driving force by which faunal and floral elements drifted away from their respective biotopes into the depositional areas, thus explaining the co‐occurrence of terrestrial (e.g. myriapods, scorpions, plants), amphibian and aquatic (e.g. conchostracans and syncarids from temporary and permanent settings, respectively) elements in fossil assemblages.     

A New Species of Scolecopteris from the Upper Carboniferous of Shanxi,China and Its Significance

A new species of the late Paleozoic fern Scolecopteris (Marattiales) is described and its relationship within the genus is discussed. Scolecopteris sinensis sp. nov., from Shanxi (coal balls of Taiyuan formation, upper Carboniferous), is similar to S. saharaensis, however some differences in characters are evidenced, such as pedicel and sporal ornamentation, The anatomy of the sporangium wails, pinule morphology, and general spore type support its association with the Minor group of Scolecopteris. The phylogenetic and ecological magnitude of Scolecopteris sinensis is also discussed.     

六足总纲系统发育研究进展与新分类系统

简要综述了昆虫分纲、分目的历史变化,包括昆虫分目多少的变化,昆虫是纲级还是总纲级阶元的变化,昆虫各目分类地位系统排列的变化以及六足总纲系统发育研究进展。根据近10年来形态特征与分子测序数据相结合的系统发育研究,整理出六足总纲与系统发育支序分析相一致的分类系统,对昆虫35目的分类运用了10个分类阶元。在此基础上,删减次要分类阶元,提出简明分类系统,既反映每个高级分类单元的单系性,明晰各目的共祖近度,又减少了分类阶元层次,方便各分类单元的识别与鉴定。六足总纲Hexapoda分为4纲:原尾纲Protura(包括蚖目Acerentomata、华蚖目Sinentomata、古蚖目Eosentomata),弹尾纲Collembola(包括弹尾目Collembola),双尾纲Diplura(包括双尾目Diplura),昆虫纲Insecta。昆虫纲分为单髁亚纲SubclassMonocondylia(包括石蛃目Archaeognatha)与双髁亚纲SubclassDicondylia。双髁亚纲分为衣鱼部DivisionZygentoma(包括衣鱼目Zygentoma)与有翅部DivisionPterygota。有翅部分为10个总目、27目。     

Nektaspid arthropods from the Lower Cambrian Emu Bay Shale Lagerstätte,South Australia,with a reassessment of lamellipedian relationships

Abstract: The lower Cambrian Emu Bay Shale on Kangaroo Island, South Australia, contains the only known Cambrian Burgess Shale‐type biota in Australia. Two new lamellipedian arthropods, Emucaris fava gen. et sp. nov. and Kangacaris zhangi gen. et sp. nov., from the Emu Bay Shale Lagerstätte are described as monotypic genera that are resolved cladistically as a monophyletic group that is sister to Naraoiidae + Liwiidae and classified within the Nektaspida as a new family Emucarididae. Shared derived characters of Emucarididae involve a bipartite, elongate hypostome and elongation of the pygidium relative to the cephalic shield and very short thorax. A monophyletic Liwiidae is composed of Liwia and the Ordovician Tariccoia + Soomaspis but excludes Buenaspis, and even the membership of Buenaspis in Nektaspida is contradicted amongst the shortest cladograms. New morphological interpretations favour affinities of Kwanyinaspis with Conciliterga rather than with Aglaspidida, and Phytophilaspis with Petalopleura.     

New Middle Cambrian bivalved arthropods from the Burgess Shale (British Columbia,Canada)

The morphology of two new bivalved arthropods, Loricicaris spinocaudatus gen. et sp. nov. and Nereocaris briggsi sp. nov. from the middle Cambrian (Series 3, Stage 5) Burgess Shale Formation (Collins Quarry locality on Mount Stephen, Yoho National Park, British Columbia, Canada), is described. The material was originally assigned to the genus Branchiocaris, but exhibits distinctive character combinations meriting its assignment to other taxa. Loricicaris spinocaudatus possesses an elongate and spinose abdomen comparable to the contemporaneous Perspicaris and Canadaspis, as well as chelate second head appendages and subtriangular exopods, comparable to Branchiocaris. Nereocaris briggsi possesses a laterally compressed carapace, elongate and delicate appendages and a medial eye located between a pair of lateral eyes on a rhomboidal eye stalk. Although undoubtedly congeneric with Nereocaris exilis from a slightly younger horizon of the Burgess Shale Formation, N. briggsi differs in overall proportions and segment number, warranting assignment to a new species. The newly described taxa were coded into an extensive cladistic analysis of 755 characters, and 312 extinct and extant panarthropods, including a variety of Cambrian bivalved arthropods from both the Burgess Shale and the Chengjiang Lagerstätten. Cambrian bivalved arthropods consistently resolved as a paraphyletic assemblage at the base of Arthropoda. Important innovations in arthropod history such as the specialization of the deutocerebral head appendages and a shift from a nekton‐benthic deposit feeding habit to a benthic scavenging/predatory habit, the symplesiomorphic feeding condition of Euarthropoda (crown‐group arthropods), were found to have occurred among basal bivalved arthropods.     

A new 'great-appendage' arthropod from the Lower Cambrian of China and homology of chelicerate chelicerae and raptorial antero-ventral appendages

The uniramous ‘great appendages’ of several arthropods from the Early to Middle Cambrian are a characteristic pair of pre‐oral limbs, which served for prey capture. It has been assumed that the morphological differences between the ‘great‐appendage’ arthropods indicate that raptorial antero‐ventral and anteriorly pointing appendages evolved more than once in arthropod phylogeny. One set of Cambrian ‘great‐appendage’ arthropods has, however, very similar short antero‐ventral appendages with a peduncle of two segments angled against each other (elbowed) and with stout distally or medio‐distally directed spines or long flexible flagellate spines on each of the four distal segments. Moreover, the head appendages of all these forms comprise the ‘great appendages’ and three pairs of biramous limbs. To this set of taxa we can add a new form from the Lower Cambrian Maotianshan Shale of southern China, Haikoucaris ercaiensis n. gen. and n. sp. It is known from three specimens, possibly being little abundant in the faunal community. It can be distinguished from all other taxa by the prominence of the proximal claw segment of its ‘great appendages’ and by only three distal spines (one on each of the distal segments). The similarity of the short, spiky ‘great appendages’ of Haikoucaris with the chelicera of the Chelicerata leads us to hypothesize that this particular type of ‘great appendages’ was the actual precursor of the chelicera. Homeobox gene and developmental data recently demonstrated the homology between the antenna of ateloceratans and the antennula of crustaceans on one side and the chelicera of chelicerates on the other. To this we add palaeontological evidence for the homology between the chelicerae of chelicerates and the ‘short great appendages’ of certain Cambrian arthropods, which leads us to hypothesize that the evolutionary path went from the ‘short great appendages’, by progressive compaction, toward the chelicera with only a two‐spined chela. The new form from China is regarded as the possible latest offshoot, whereas the other ‘great appendages’ arthropods with similar short grasping limbs were derivatives of the stem lineage of the crown‐group Chelicerata. Consequently, the chelicera with a chela with one fixed and one mobile finger is an autapomorphy of the crown group of Chelicerata, whereas a raptorial, but more limb‐like antenna, with more distal spine‐bearing segments, characterized the ground pattern of Chelicerata. Further taxa having ‘great appendages’, including the large Anomalocarididae, are also discussed in the light of their possible affinities to the Chelicerata and possible monophyly of all of these arthropods with raptorial anterior appendages.     

Microbial Methane Formation from Hard Coal and Timber in an Abandoned Coal Mine

About 7% of the global annual methane emissions originate from coal mining. Also, mine gas has come into focus of the power industry and is being used increasingly for heat and power production. In many coal deposits worldwide, stable carbon and hydrogen isotopic signatures of methane indicate a mixed thermogenic and biogenic origin. In this study, we have measured in an abandoned coal mine methane fluxes and isotopic signatures of methane and carbon dioxide, and collected samples for microbiological and phylogenetic investigations. Mine timber and hard coal showed an in-situ production of methane with isotopic signatures similar to those of the methane in the mine atmosphere. Enrichment cultures amended with mine timber or hard coal as sole carbon sources formed methane over a period of nine months. Predominantly, acetoclastic methanogenesis was stimulated in enrichments containing acetate or hydrogen/carbon dioxide. Molecular techniques revealed that the archaeal community in enrichment cultures and unamended samples was dominated by members of the Methanosarcinales. The combined geochemical and microbiological investigations identify microbial methanogenesis as a recent source of methane in abandoned coal mines.     

On the ground pattern of Annelida

Annelida, traditionally divided into Polychaeta and Clitellata, are characterized by serial division of their body into numerous similar structures, the segments. In addition, there is a non-segmental part at the front end, the prostomium, and one at the back, the pygidium. New segments develop in a prepygidial proliferation zone. Each segment contains four groups of chaetae made up of β-chitin, a pair of coelomic cavities separated by mesenteries, and septa. The nervous system is a rope-ladder-like ventral nerve cord with a dorsal brain in the prostomium. For the last stem species a trochophore larva and a benthic adult are commonly postulated. There are two conflicting hypotheses describing the systematization of Annelida: the first postulates a sister-group relationship of Polychaeta and Clitellata, the second sees Clitellata as a highly derived taxon forming a subordinate taxon within the polychaetes which, consequently, are regarded as paraphyletic. Depending on the hypothesis, different characters have to be postulated for the stem species of Annelida. Besides segmentation other characters such as nuchal organs, palps and antennae, body wall musculature, cuticle, parapodia as well as structure of the central nervous system and the foregut play an important role in this discussion. Here, the different characters and character states are critically reviewed and analyzed with respect to morphology and function. The consequences for systematization of their phylogenetic interpretation as autapomorphies, synapomorphies or plesiomorphies are outlined. The resulting hypotheses are compared with those relying on molecular data sets.     

Boron and Marine Life: A New Look at an Enigmatic Bioelement

On the occasion of the 200th anniversary of the discovery of boron, we review the oceanic biogeochemistry of boron as well as suitable analytical techniques for its determination. This overview includes aspects of biogeochemistry including geochemical stable isotope variations, uptake, transport, storage, nutritional value, toxicity, and distribution within biological materials, providing a framework for discussion of the role of boron in marine organisms, which remains largely enigmatic.     

Arthropod phylogeny inferred from partial 12SrRNA revisited: monophyly of the Tracheata depends on sequence alignment

A new hypothesis on arthropod evolution published by Ballard et al. (1992), based on partial 12SrRNA sequences, is re-analysed using the same data, but using different alignments. It is argued that there is no reason to reject monophyly of the Euarthropoda, Mandibulata and Tracheata.

Zusammenfassung

Die Phylogenie der Arthropoden, abgeleitet von Abschnitten der 12SrRNA, nochmals überdacht: Die Monophylie der Tracheaten hängt vom Sequenz-Alignment ab
Eine von Ballard et al. (1992) publizierte, auf partiellen 12SrRNA-Sequenzen beruhende Hypothese zur Stellung der Onychophora und Myriapoda innerhalb der Arthropoda, wird aufgegriffen und mit anderer Alinierung neu berechnet. Es wird gezeigt, da β es keinen Anlaß gibt, die Monophylie der Euarthropoda, Mandibulata und Tracheata zu bezweifeln.     

Arthropod phylogeny inferred from partial 12SrRNA revisited: monophyly of the Tracheata depends on sequence alignment

A new hypothesis on arthropod evolution published by Ballard et al. (1992), based on partial 12SrRNA sequences, is re-analysed using the same data, but using different alignments. It is argued that there is no reason to reject monophyly of the Euarthropoda, Mandibulata and Tracheata.     

新疆石炭系杜内-韦宪阶界线层的牙形类POLYGNATHUS COMMUNIS GANCAOHUENSIS (新亚种)POLYGNA-THUS COMMUNIS BRANSON ET MEHL 1934的种系发生

本文提出和描述了产自我国西北新疆南部喀拉塔克剖面的一个牙形类新亚种Polygnathus communis gan-caohuensis,并根据以前出版的信息,命名了另一个新亚种P0lygnathus communis carmanae.种系发生研究揭示了Polygnathus communis Branson et Mehl的亚种对于法门阶最上部至韦宪阶地层的划分是至关重要的.此种的cari-na→gancaohuensis谱系,连同发现于喀拉塔克剖面的菊石、有孔虫和腕足动物,被认为是划分杜内-韦宪阶界线的潜在的标志化石.因此,喀拉塔克剖面有可能成为全球杜内-韦宪阶界线的一个候选层型剖面.     

New species of spiriferids (Brachiopoda) from the Devonian and Carboniferous of Eastern Europe

In a revision of previously published materials, two new spiriferid species from the Late Tournaisian, Unispirifer subtornacensis sp. nov. and Mesochorispira ussuilensis sp. nov., and Atylephorus nalivkini sp. nov. from the Lytva Horizon of the Upper Devonian of Bashkiria are described. Based on the original collection, a new Serpukhovian subspecies, Podtsheremia duplicicosta triplicicosta, from the southern Ural Mountains and a new Late Carboniferous species, Purdonella kalashnikovi, from the polar Ural Mountains and adjacent islands are established.     

The Arthropod Alalcomenaeus cambricus Simonetta, from the Middle Cambrian Burgess Shale of British Columbia

More than 300 specimens of the previously rare arthropod Alalcomenaeus cambricus Simonetta have been collected from a new Burgess Shale locality in the Glossopleura Zone on Mount Stephen, British Columbia. This new material provides much more complete information on its morphology. The cephalon was covered by a shield. A pair of pedunculate eyes and three median eyes were followed by a large anterior appendage, the 'great appendage', bearing three long flagella. The two posterior head appendages, like those of the trunk, were biramous. They consisted of a segmented, inner branch, and a flap-like outer branch, fringed with long filaments. The trunk consisted of 11 somites, each protected by a tergite and bearing a pair of biramous limbs. The telson was paddle-like and fringed posteriorly with wide flat spines. Alalcomenaeus was probably a predator, moving mainly by swimming. It is now known to be one of the more abundant, widely distributed and longest ranging of Burgess Shale arthropod genera. Its affinities lie with the Arachnomorpha.     

The cranial morphology and relationships of the aberrant Carboniferous amphibian Spathicephalus mirus Watson

The skull of Spathicephalus mirus Watson, an amphibian from the Namurian, basal Upper Carboniferous, of Scotland is described. It shares with the Loxommatoidea a closed palate, palatal ornament and antorbital vacuities, and the family Spathicephalidae is recognized as the sister group of the Loxommatidae. A new diagnosis of the Loxommatoidea is presented together with one of the Spathicephalidae. An analysis of the functional morphology of the Spathicephalus skull suggests that it was incapable of rapid jaw closure required for catching fish. Instead it is proposed that Spathicephalus was a sluggish bottom-dwelling filter-feeder of small, soft invertebrates.     

New bryozoans from the Devonian-Carboniferous boundary beds of Transcaucasia

New species of Transcaucasian bryozoans are described: Primorela remota and Pseudoascopora subtilis (order Rhabdomesida) from the Upper Famennian and Anisotrypa hamata (order Trepostomida), Paranicklesopora stupenda, Euthyrhombopora barbara, Primorella testata, Ipmorella clara (order Rhabdomesida), and Taeniodictya vermiculata (order Cryptostomida) from the Tournaisian.