The brachiopod fold: a neglected body plan hypothesis

Attention is drawn to Nielsen's radical body plan concept, here named the 'brachiopod fold hypothesis', under which brachiopods and phoronids are recognized to be transversely folded across the ontogenetic anterior–posterior axis so that, to make useful comparisons with other phyla, these organisms must be conceptually unfolded. Under the hypothesis brachiopod brachial and pedicle shell valves are respectively 'anterior' and 'posterior' rather than 'dorsal' and 'ventral' as traditionally described. The hypothesis makes sense of the symmetry axes of the brachiopod shell, is consistent with various indications from fossil and Recent brachiopods, and gives rise to predicted patterns of axis–determining gene expression that differ from those obtaining under the traditional view of the body plan, whilst the variety of folding movements in different lineages implies that superficially dissimilar morphogenetic folds may be fundamentally homologous. Convergent folding patterns are noted in some other organisms. A previous conjecture that inarticulate linguloid brachiopods were derived from halkieriid–like ancestors is elaborated with proposals that recognize possible functional continuities of coelomic and marginal sclerite functions, and it is noted that an ancestrally facultative fold could have become incorporated by genetic assimilation into the brachiopod developmental program. An experimental approach is outlined to test the possibility that some members of the 'small shelly fauna' may have been members of the halkieriid–like brachiopod stem lineage and it is also suggested that buoyancy modification may have been an important function of mineralization amongst Lower Cambrian floaters and swimmers, since negative buoyancy would facilitate access to the benthic niche

The Early Cambrian [Botomian] stem group brachiopod Mickwitzia from Northeast Greenland

The problematic brachiopod Mickwitzia Schmidt, 1888 is re−described based on new material of M. cf. occidens Walcott, 1908 from the Early Cambrian (Botomian) Bastion and Ella Island formations of Northeast Greenland. Etched material demonstrates that Mickwitzia has a lingulid−like juvenile (“larval”) shell with trails of nick−points, reflecting the movement of marginal setae. Juvenile and early mature ventral valves have a lingulid−like pseudointerarea with a pedicle groove. The shell of M. cf. occidens is only partially phosphatic, in particular around the juvenile–early mature shell in both valves. The phosphatic shell includes at least two types of cylindrical structures: (1) slender columns identical with the columns of acrotretoid brachiopods and (2) relatively thicker tubes which may be open to the exterior surface and have internal striations (on the ventral pseudointerarea). The striations are most likely imprints of microvilli and these tubes can be inferred to have contained setae. The thinner linguliform columns and thicker setigerous striated tubes are considered to be homologous with identical structures in the sellate and mitral sclerites of the problematic Micrina, which has been identified as a probable primitive stem group of the Brachiopoda. Mickwitzia represents a more derived member of the stem group Brachiopoda

Chemico-structure of the organophosphatic shells of siphonotretide brachiopods

The organophosphatic shell of siphonotretide brachiopods is stratiform with orthodoxly secreted primary and secondary layers. The dominant apatitic constituents of the secondary layer are prismatic laths and rods arranged in monolayers (occasionally in cross-bladed successions), normally recrystallized as platy laminae. Sporadically distributed, interlaminar, lenticular chambers, containing apatitic meshes of laths and aggregates of plates and spherulites, probably represent degraded, localized exudations of glycosaminoglycans (GAGs) with dispersed apatite. The shells of Helmersenia and Gorchakovia are perforated by canals with external depressions (antechambers) that possibly contained chitinous tubercles in vivo. The immature shell of Siphonotreta and most other siphonotretids is similarly perforated and pitted; but the mature part bears recumbent, rheomorphic, hollow spines that grew forward out of pits. Internally, spines pierce the shell as independent structures to terminate as pillars in GAGs chambers. Spines and pillars were probably secreted by collectives of specialized cells (acanthoblasts) within the mantle. The shell of the oldest siphonotretide, Schizambon, is imperforate but the ventral valve has a pedicle foramen that lies forward of the posterior margin of the juvenile valve. This relationship characterizes all siphonotretides, suggesting that the pedicle, in vivo, originated within the ventral outer epithelium and not from the posterior body wall as in lingulides

Adaptive strategies and environmental significance of lingulid brachiopods across the late Permian extinction

Linguliformbrachiopods are traditionally considered a conservative group which seems to pass through the late Permian extinction without any significant loss and even appear to thrive immediately after the extinction peak. In the Southern Alps, lingulids are very common in the post-extinction Mazzin Member (early Induan) of the Werfen Formation. Sparse occurrences are also known in the overlying Siusi and Gastropod Oolite members (late Induan and early Olenekian in age respectively). The recent discovery of well preserved specimens from a pre-extinction bed of the Bellerophon Formation (Changhsingian) has permitted a detailed comparative analysis, mostly based on the interior characters, preserved in the lingulid succession fromacross the extinction beds. The following effects on the lingulid populations have been analyzed: i) change in taxonomic assessment; ii) adaptive strategies during the surviving and recovery phases; and iii) environmental proxy connected with the killing mechanisms of the late Permian extinction. The pre-extinction individuals belong to Lingularia? cf. smirnovae Biernat and Emig, a species that is characterized by large-sized shells with a short lophophoral cavity. The post-extinction populations belong to different species and, probably, even to a different genus. The first post-extinction population (early Induan), with small-sized shells and long lophophoral cavity, has been referred to Lingularia yini (Peng and Shi). It records themost severe effects of the late Permian extinction on the marine ecosystems. The late Induan–Olenekian Lingularia borealis (Bittner), with large sized shells and long lophophoral cavity, appears during the first phase of the Triassic biotic recovery. Themain adaptive strategies of Lingularia yini, in comparisonwith the Permian species, include: i) shell miniaturization; ii) increasing of the lophophoral cavity surface (respiratory surface); and iii) increasing of shell width/ length ratio. Thesemodifications are interpreted as adaptations towards warming and hypoxia, two main killing mechanisms of themarine biota. The recovery species Lingularia borealis maintains a large lophophoral cavity, indicating an adaptation towards predominant low oxygenated bottom marine waters. The appearance and the great abundance of Lingularia yini in the Mazzin Member (early Induan) represent a proxy of dysaerobic conditions, which determined the appearance of the second phase of the Lilliput biota, characterized by the definitive disappearance of the rhynchonelliform brachiopods and calcareous algae in the Southern Alps

The new stem-group brachiopod Oymurania from the lower Cambrian of Siberia

A stem-group brachiopod, Oymurania gravestocki Ushatinskaya gen. et sp. nov. is described herein from the middle Atdabanian-lower Botoman Stages (-Cambrian Stage 3) of the Siberian Platform. The fossils were extracted from limestone beds of the Emyaksin, Perekhod, and Pestrotsvet formations as assemblages of disarticulated orthoconic to cyrtoconic porous shells in apatite preservation. The originally organophosphatic shells of Oymurania are externally similar to mitral sclerites (ventral valves) of the stem-group brachiopod Micrina, although no sellate-like sclerites, nor differentiated subapical area with apophyses were recognised in our material. The range of Oymurania shells with sub-central to posteromarginal apex is similar to that of ventral valves ofMickwitzia. Oymurania is also characterised by the system of radial and orthogonal canals open in pairs or triplets in small depressions or indentations of growth lamellae in the outer shell surface. The orthogonal (Micrina-Setatella type) and radial (horizontal setigerous tubes) canals are widespread among the early Cambrian stem-group brachiopods, such as Micrina, Mickwitzia, and Setatella. In addition to these canals, Oymurania exhibits a well-developed acrotretoid columnar microstructure, also known from Setatella. A broad subapical platform in cyrtoconic shells (presumably ventral valves) of Oymurania is interpreted homologous to the deltoid area in mitrals of Micrina and pseudointerarea/interarea in ventral valves of Setatella/paterinid brachiopods. Except with probable cell imprints and openings of orthogonal canals, no morphological differentiation was, however, reflected by the shell interior of Oymurania gravestocki. Being closely related to tannuolinids and mickwitziids, Oymurania complements the picture of diversification of the early Cambrian stem-group brachiopods that occurred in parallel with radiation of paterinids and other crow-group brachiopods on the Siberian Platform and worldwide

Soft-part preservation in a linguliform brachiopod from the lower Cambrian Wulongqing Formation [Guanshan fauna] of Yunnan, South China

Linguliform brachiopods were important components of early Cambrian benthic communities. However, exceptionally preserved soft parts in Cambrian linguliform brachiopods are extremely sparse, and the most important findings are from the early Cambrian Chengjiang Konservat Lagerstätte of Kunming, southern China. Here we describe the first record of preserved soft−part anatomy in a linguliform brachiopod from the early Cambrian Guanshan fauna (Wulongqing Formation, Palaeolenus Zone); a unit which is considerably younger than the Chengjiang fauna. The well preserved soft anatomy include linguliform pedicles, marginal setae and, in a few cases, an intact lophophore imprint. The pedicle has pronounced surface annulations, with its proximal−most part enclosing the apex of the ventral pseudointerarea; the pedicle is up to 51 mm long, corresponding to more than 4 times the sagittal length of the shell, and 12% of the maximum valve width. In details of their preservation, these new fossils exhibit striking similarities with the linguliforms from the older Chengjiang fauna, and all specimens are preserved in a compressed state as flattened impressions. The new linguliform has an elongate oval to subtriangular shell and an elongate triangular ventral pseudointerarea; the pedicle emerged from an apical foramen through a poorly preserved internal pedicle tube. The new linguliform is most similar to the mostly organic−shelled siphonotretoid−like brachiopod Acanthotretella spinosa, recently described from the classic middle Cambrian Burgess Shale Konservat Lagerstätte, British Columbia, Canada. The new species Acanthotretella decaius sp. nov. is described; it differs from A. spinosa in having a slightly thicker pedicle, and a larger and more rigid, probably partly mineralised shell, indicating that the mostly organic shell of A. spinosa may represent a secondary reduction of shell mineralisation. However, the spine−like setae of the new species are unfortunately poorly preserved only at the margin of the shell, but the new species is referred tentatively to the Superfamily Siphonotretoidea. The occurrence of A. decaius in the Guanshan fauna is the first lower Cambrian (Series 2, early Stage 4) record of both Acanthotretella and siphonotretoids, and it represents the first description of a lophophore and digestive tract from the siphonotretoid lineage

The problematic early Cambrian fossil Tumulduria incomperta represents the detached ventral interarea of a paterinid brachiopod

The organophosphatic early Cambrian (Terreneuvian, Cambrian Stage 2) fossil Tumulduria incomperta has been problematic ever since its original description in 1969. Comparison of abundant specimens from the Lower Cambrian of Siberia with co-occurring brachiopod valves show that T. incomperta represents the central portion of the ventral interarea of a paterinid brachiopod similar to Cryptotreta neguertchenensis, and that the domed central portion of typical Tumulduria specimens represents the ridge-like pseudodeltidium of the interarea