Architecture and function of the lophophore in the problematic brachiopod Heliomedusa orienta (Early Cambrian, South China)

The detailed structure of the lophophore is a key diagnostic character in the definition of higher brachiopod taxa. The problematic Heliomedusa orienta Sun and Hou, from the Lower Cambrian Chengjiang Lagerstätte of Yunnan, southwestern China, has a well-preserved lophophore, which is unlike that of any known extant or extinct brachiopods. Based on a comparative study of lophophore disposition in H. orienta and the extant discinid Pelagodiscus atlanticus, the in- and excurrent pattern and shell orientation of H. orienta are described and discussed. Reconstructions of lophophore shape and function are based on numerous specimens and comparison with P. atlanticus. The lophophore is composed of a pair of lophophoral arms that freely arch posteriorly rather than coiling anteriorly as commonly seen in fossil and recent lingulids. The lophophore is attached to the dorsal lobe of the mantle; it has neither calcareous nor chitinous supporting structures, and is disposed symmetrically on either side of the valve midline. The mouth can be inferred to be located at the base of the two brachial tubes, slightly posterior to the anterodorsal projection of the body wall. The lophophoral arms bear laterofrontal tentacles with a double row of cilia along their lateral edge, as in extant lingulid brachiopods. The main brachial axes are also ciliated, which presumably facilitated transport of mucous-bound nutrient particles to the mouth. The unique organization of the lophophore in Heliomedusa is not like any known fossil and living brachiopods. This clearly demonstrates that H. orienta is not a member of any crown group. It is here considered as a member of the brachiopod stem group, which challenges recent interpretations of a close discinid affinity.     

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.     

弱硅化腕足类化石酸处理方法的改进

本文介绍了一种获取灰岩中弱硅化腕足类化石的酸处理方法。以四川省布拖县浪珠乡万吨山剖面上奥陶统铁足菲克组近顶部的生屑灰岩为实验对象, 通过准备样品、悬吊样品、配酸解液、酸解样品等实验流程, 配合遮盖液(如二乙酸纤维素丙酮溶液)的使用, 可获得保存精美的、弱硅化的腕足类实体化石, 且该化石保存有主突起、铰齿和匙形台等重要的细小结构。相较于传统方法, 该方法可获取硅化程度更低的壳相化石。该方法为灰岩中其他多门类弱硅化保存的化石之酸解分析提供了新思路和参考。     

Brachiopod shell thickness links environment and evolution

While it is well established that the shapes and sizes of shells are strongly phylogenetically controlled, little is known about the phylogenetic constraints on shell thickness. Yet, shell thickness is likely to be sensitive to environmental fluctuations and has the potential to illuminate environmental perturbations through deep time. Here we systematically quantify the thickness of the anterior brachiopod shell which protects the filtration chamber and is thus considered functionally homologous across higher taxa of brachiopods. Our data come from 66 genera and 10 different orders and shows well-defined upper and lower boundaries of anterior shell thickness. For Ordovician and Silurian brachiopods we find significant order-level differences and a trend of increasing shell thickness with water depth. Modern (Cenozoic) brachiopods, by comparison, fall into the lower half of observed shell thicknesses. Among Ordovician–Silurian brachiopods, older stocks commonly have thicker shells, and thick-shelled taxa contributed more prominently to the Great Ordovician Biodiversification but suffered more severely during the Late Ordovician Mass Extinction. Our data highlight a significant reduction in maximum and minimum shell thickness following the Late Ordovician mass extinction. This points towards stronger selection pressure for energy-efficient shell secretion during times of crisis.     

广西马平灰岩腕足动物群的时代问题--兼评"Fauna of the Maping Limestone of Kwangsi and Kweichow"

ｒａｂａｕ（１９３６）记述的马平动物群中所有的腕足类化石标本并不是产于真正的马平（灰岩）组,而是产自有关地区下石炭统最上部的罗城组和上石炭统下部的黄龙组下部,时代为Ｓｅｒｐｏｕｋｈｏｖｉａｎ－Ｂａｓｈｋｉｌｉａｎ期,而不是Ｕｒａｌｉａｎ期     

Morpho-anatomical differences of the Early Cambrian Chengjiang and Recent lingulids and their implications

Since the publication of Darwin's theory of evolution in 1859, lingulids have probably been the most widely quoted examples of arrested evolution. This, to some degree, may be because few anatomical features are impressed either on or inside valves so these may not adequately reflect the extent of change incurred during lingulid evolution. Two lingulid brachiopods, Lingulella chengjiangensis and Lingulellotreta malongensis, from the Early Cambrian Chengjiang Lagerstätte (Yunnan, South China) show preservation of a series of soft parts, notably the lophophores, setae mantle canals and perfectly impressed visceral region, which are vital to understanding the evolution and lifestyle of brachiopods. Analysis of the valve interiors favours the claim that an epifaunal mode of life could be a plesiomorphic state in contrast to the infaunal one of modern lingulids. Based on these fossils, the three‐pseudosiphon formation of setae found in Recent lingulids is proposed to be an apomorphy derived as an adaptation to an infaunal lifestyle. Comparison between the interior of the fossils with that of modern lingulids does little to support the widespread notion that the morphology of this lineage has remained remarkably constant since at least the early Palaeozoic.     

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.     

A preliminary phylogenetic study of late Palaeozoic spiriferoid brachiopods using cladistic and Bayesian approaches

The brachiopod Superfamily Spiriferoidea diversified greatly and was widely distributed in the late Palaeozoic (Carboniferous–Permian), and yet its phylogeny has been seldom investigated with analytical methods. This is reflected in the current flux of very different classification schemes for this superfamily. This paper provides the first attempt to investigate the phylogenetic relationships of spiriferoid brachiopods through both cladistic and Bayesian analyses involving 24 discrete and continuous characters. The continuous characters, from morphometric data, have been separately discretized using the gap weighting method, and the ‘as such’ option in TNT. Our results highlight the potential significance of continuous characters in reconstructing and elucidating phylogenies, as much as qualitative characters. Building on the outcomes of the analyses, we also briefly evaluate existing classification schemes of Spiriferoidea. We found that none of the existing classifications fully reflect the phylogeny properly; major families within the superfamily, such as Spiriferidae, Choristitidae, and Trigonotretidae, turned out to be polyphyletic. Although this study is considered preliminary, due to the selection of and restriction to certain taxa, combined with the use of a relatively small number of characters, it nevertheless demonstrates that potentially the true phylogenetic relationships of spiriferoid taxa sharply contrast with any of the existing classification schemes. This highlights the need to develop an alternative scheme that takes into account a more comprehensive range of phylogenetic variables.     

Shell Structure And Inferred Growth, Functions And Affinities Of The Sclerites Of The Problematic Micrina

The stratiform laminae of Micrina sclerites originally consisted of rheomorphic successions of monolayers of micrometric–sized, apatitic tablets, presumably interleaved with chitin and glycosaminoglycans (GAGs). Paired laminae enclose slot–like chambers swelling into lobes distally that originally contained GAGs and deposits of spherulitic and prismatic apatite. The laminae are pervaded by apatitic tubes, apparently secreted by microvillous setoblasts and containing, at the surface, chitinous setae. Internal markings suggest that the triangular (sellate) sclerite supported a pair of muscles and the planospiral (mitral) sclerite, a medial muscle and gonadal sacs flanked by a pair of crescentic muscle bases. Both sclerites were secreted by a mantle with a circumferential fold. The sellate and mitral sclerites are homologized with the anterior and posterior shells of Halkieria and could have become the dorsal and ventral valves of the ancestral brachiopod by a sequence of transformations. These include: the folding of the halkieriid body axis; accelerated mixoperipheral growth of the anterior (dorsal) shell to enclose, with the posterior (ventral) shell, a mantle cavity lined with modified ciliated epithelium of the foot; reduction of sclerite–secreting epithelium to the locus of the brachiopod pedicle epithelium; and the anterior (dorsal) spread of gonadal lamellae.     

The development and shell microstructure of the pseudodeltidium and interarea in thecideide brachiopods

Abstract: This study examines the development of the delthyrium, pseudodeltidium and interarea, their growth during the early juvenile stages of ontogeny and the extrapolation of morphology from adult shells to the probable juvenile state. Examination of the development and shell microstructure of the cardinalia of early juvenile thecideide brachiopod ventral valves from Jurassic, Cretaceous and Holocene specimens suggests that the delthyrium develops early in ontogeny and that the initial development of the pseudodeltidium precedes that of the interarea. Also, until the interarea is formed, the postero‐lateral flanks of the ventral umbo are palintropic. The development of the interarea can be seen to be a consequence of the lateral extension of the early juvenile hinge line. Initially, the pseudodeltidium consists solely of a thin plate of primary shell material. Comparison of the morphology of the pseudodeltidium of early juveniles with that of adults suggests that the initially curved lateral profile of the pseudodeltidium is retained, or even accentuated, in the ontogeny of lacazellines, but in all thecidellinines, with the exception of Pachymoorellina and Minutella, following the appearance of the interarea, the pseudodeltidium becomes flattened and often appears continuous with the interarea. However, we do not support any proposal that suggests that, in thecideides, only those forms in which the delthyrium is closed by a dorsally convex plate should be considered to possess a pseudodeltidium sensu stricto, mainly because of physiological differences and the prospect of possible taxonomic confusion in the future. Instead, we propose the term planodeltidium for a flat pseudodeltidium, typically developed in the thecidellinines, and rugideltidium for a convex pseudodeltidium, typically developed in the lacazellines, but also in Pachymoorellina and Minutella. Despite the presence of a rugideltidium, we believe the affinities of Minutella are more strongly with the thecidellinines and have included it in the new subfamily Minutellinae of the family Thecidellinidae.