Anatomical knowledge of early chondrichthyans and estimates of their phylogeny are improving, but many taxa are still known only from microremains. The nearly cosmopolitan and regionally abundant Devonian genus Phoebodus has long been known solely from isolated teeth and fin spines. Here, we report the first skeletal remains of Phoebodus from the Famennian (Late Devonian) of the Maïder region of Morocco, revealing an anguilliform body, specialized braincase, hyoid arch, elongate jaws and rostrum, complementing its characteristic dentition and ctenacanth fin spines preceding both dorsal fins. Several of these features corroborate a likely close relationship with the Carboniferous species Thrinacodus gracia , and phylogenetic analysis places both taxa securely as members of the elasmobranch stem lineage. Identified as such, phoebodont teeth provide a plausible marker for range extension of the elasmobranchs into the Middle Devonian, thus providing a new minimum date for the origin of the chondrichthyan crown-group. Among pre-Carboniferous jawed vertebrates, the anguilliform body shape of Phoebodus is unprecedented, and its specialized anatomy is, in several respects, most easily compared with the modern frilled shark Chlamydoselachus . These results add greatly to the morphological, and by implication ecological, disparity of the earliest elasmobranchs.
A comprehensive review and phylogenetic analysis of genera and species presently assigned to the rhynchonelliform superfamily Nisusioidea and family Nisusiidae suggests that this short‐lived but important group of brachiopods first appeared in peri‐Gondwana during the second half of the Cambrian Series 2, before going extinct by the end of Drumian times. Nisusiides achieved their maximum morphological disparity and geographical distribution during the Wuliuan Age, and Laurentia was probably the major centre of their dispersal. A new phylogenetic analysis suggests an early separation of the lineages of spinose and non‐spinose nisusiids. The non‐spinose nisusiids probably evolved in Laurentia by the end of Cambrian Series 4. The new nisusiid genus Bellistrophia is described. The new species Nisusia multicostata represents the first documented rhynchonelliform (kutorginide) brachiopod from the Miaolingian (Drumian) of the Alborz Mountains, Iran.
A Llandovery (mid-upper Aeronian) brachiopod fauna is described for the first time from the Niur Formation of Central Iran. It is dominated by two succeeding rhynchonellide species Stegocornu procerum Dürkoop, 1970 and Stegocornu denisae sp. nov. In addition, there are three common and four rarer brachiopod species, including Dalejina? rashidii sp. nov., Isorthis (Ovalella) inflata sp. nov. and Striispirifer? ocissimus sp. nov. The Stegocornu Association gives a distinct biogeographic signature to the mid to late Llandovery rhynchonellide-dominated shallow-water brachiopod faunas of Central Iran, Kope-Dagh and Afghanistan. Its proliferation in temperate latitude peri-Gondwana was one of the earliest signs of biogeographical differentiation of the brachiopod faunas in the early Silurian. The affinities of Stegocornu and Xerxespirifer are discussed.
Abstract. Bulk-carbonate carbon isotope ratios are a widely applied proxy for investigating the ancient biogeochemical carbon cycle. Temporal carbon isotope trends serve as a prime stratigraphic tool, with the inherent assumption that bulk micritic carbonate rock is a faithful geochemical recorder of the isotopic composition of seawater dissolved inorganic carbon. However, bulk-carbonate rock is also prone to incorporate diagenetic signals. The aim of the present study is to disentangle primary trends from diagenetic signals in carbon isotope records which traverse the Permian-Triassic boundary in the marine carbonate-bearing sequences of Iran and South China. By pooling newly produced and published carbon isotope data, we confirm that a global first-order trend towards depleted values exists. However, a large amount of scatter is superimposed on this geochemical record. In addition, we observe a temporal trend in the amplitude of this residual δ 13 C variability, which is reproducible for the two studied regions. We suggest that (sub-)sea-floor microbial communities and their control on calcite nucleation and ambient porewater dissolved inorganic carbon δ 13 C pose a viable mechanism to induce bulk-rock δ 13 C variability. Numerical model calculations highlight that early diagenetic carbonate rock stabilization and linked carbon isotope alteration can be controlled by organic matter supply and subsequent microbial remineralization. A major biotic decline among Late Permian bottom-dwelling organisms facilitated a spatial increase in heterogeneous organic carbon accumulation. Combined with low marine sulfate, this resulted in varying degrees of carbon isotope overprinting. A simulated time series suggests that a 50 % increase in the spatial scatter of organic carbon relative to the average, in addition to an imposed increase in the likelihood of sampling cements formed by microbial calcite nucleation to 1 out of 10 samples, is sufficient to induce the observed signal of carbon isotope variability. These findings put constraints on the application of Permian-Triassic carbon isotope chemostratigraphy based on whole-rock samples, which appears less refined than classical biozonation dating schemes. On the other hand, this signal of increased carbon isotope variability concurrent with the largest mass extinction of the Phanerozoic may proPublished by Copernicus Publications on behalf of the European Geosciences Union.
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