Chemico-structural phylogeny of the discinoid brachiopod shell

Stratiform shells of living discinids are composed of membranous laminae and variously aggregated, protein-coated granules of apatitic francolite supported by proteinaceous and chitinous nets in glycosaminoglycans (GAGs) to form laminae in rhythmic sets. The succession is like that of living lingulids but differs significantly in the structure of the periostracum, the nature of baculate sets and in its organic composition. In particular, discinids have a higher level of amino acids although with relatively lower acidic and higher basic concentrations; and their overall lower organic content is owing to lower levels of hydrophilic components, like GAGs and chitin. The organic constituents are not completely degraded during fossilization; but data are presently too meagre to distinguish between linguloid and discinoid ancestries. Many differences among three of the four described extant genera emanate from transformations with a long geological history. Pelagodiscus is characterized by regular, concentric rheomorphic folding (fila) of the flexible periostracum and the plastic primary layer and by sporadically developed hemispherical imprints of periostracal vesicles. Both features are more strikingly developed in Palaeozoic discinids. In the oldest discinid, the Ordovician Schizotreta, and the younger Orbiculoidea and related genera, vesicles were persistent, hexagonal close-packed arrays fading out over fila. They must have differed in composition, however, as the larger vesicles of Schizotreta were simple (possibly mucinous), whereas the smaller vesicles of Orbiculoidea and younger genera were composites of thickly coated spheroids, possibly of lipoproteins, which survive as disaggregated relicts in Pelagodiscus. Baculate sets within the secondary layer are also less well developed in living discinids, being incipient in Pelagodiscus and restricted to the dorsal valve of Discinisca. The trellised rods (baculi) with proteinaceous cores are composed of pinacoids or prisms of apatite, depending on whether they are supported by chitinous nets or proteinaceous strands in GAGs. This differentiation occurred in Schizotreta but in that stock (and Trematis) the baculate set is symmetrical with baculi subtended between compact laminae, whereas in younger and post-Palaeozoic species the outer bounding lamina(e) of the set is normally membranous and/or stratified. The most striking synapomorphy of living discinids is the intravesicular secretion of organsiliceous tablets with a crystalline habit within the larval outer epithelium and their exocytosis as a close- or open-packed, transient, biomineral cover for larvae. Canals, on the other hand, are homologous with those pervading lingulid shells. Both systems interconnect with chitinous and proteinaceous sets and have probably always served as vertical struts in an organic scaffolding supporting the stratiform successions. A phylogenetic analysis based mainly on shell structure confirms the discinoids as the sister group of the linguloids but, contrary to current taxonomic practice, also supports the inclusion of acrotretoids within a ''discinoid'' clade as a sister group to the discinids.     

Mineral phase in linguloid brachiopod shell: Lingula adamsi

The linguloid brachiopod shell family has been the focus of several studies because of the similarity in the composition of the mineral phase of these shells to that of human bone. However, ultrastructural features of Lingula shells have not yet been fully demonstrated at high magnification using Transmission Electron Microscopy (TEM) and Electron Diffraction. Ultrastructural characterization of the mineral phase in Lingula shells will improve our understanding of the biomineralization processes and mineral/organic interaction in more complex systems such as in bone or in other human mineralized tissues. In this study, the mineral phase of Lingula adamsi was characterized using a combination of ultrastructural and crystallographic techniques. The results showed that L. adamsi shells consist of apatite crystals of varying size, shape, and orientation in different areas of the shell. The c-axis of apatite was parallel to the shell surface and crystals were organized in different laminae. Compared to trabecular bovine bone, L. adamsi shells demonstrated a higher crystallinity and a lower amount of carbonate and organic compounds. This study therefore demonstrated how dissimilar organic matrix between L. adamsi shell and trabecular bone can modify the ultrastructural characteristics of apatite crystals in these two biomineralized tissues.     

Macromolecules in brachiopod shells: characterization and diagenesis

An immunological investigation was conducted of soluble intra-crystalline macromolecules isolated from living and fossil brachiopod shells, which had previously been used for an immunologically based study of phylogeny (serotaxonomy). The soluble intra-crystalline macromolecules comprised 0.03% by weight of the extant shell material. Bulk analysis and gel electrophoresis indicated that the organic material is predominantly glycoprotein, and contains up to 30% by weight carbohydrate. Treatment of the macromolecules with periodate and proteinase K revealed that antibodies were raised predominantly against the carbohydrate moieties. Using a specially adapted dot blot immunobinding assay (DIBA) the decay in immunological signal over geological time was determined. Pleistocene shells have lost between 99 and 99.9% of immunological reactivity, and original antigenic determinants form a declining proportion of total organic matter. It is suggested that condensation reactions between amino acids and sugars account for the rapid destruction of determinants; this has important implications for the direction of future studies on fossil macromolecules. □ Serotaxonomy, biomolecular palaeontology, glycoproteins, melanoidins, brachiopods.     

Comparative hydrodynamic stability of brachiopod shells on current-scoured arenaceous substrates

Selected brachiopod specimens were placed on a well-sorted medium grained sand (0.5 mm) in a recirculating flume and subjected to a constant unidirectional current of 26–27 cm/sec. The specimens were placed in each of six feasible life-orientations. In three orientations, the valves were reclining with either the lateral, anterior or posterior profile upcurrent. In three orientations, the commissural plane was perpendicular to the substrate with either the pedicle or brachial valve upcurrent, or a lateral margin upcurrent. Destabilization indices were calculated for each specimen in each orientation based on the elapsed time until the specimen reoriented. Nonstrophic biconvex brachiopods are most stable with the valves reclining on the substrate. Among orientations with the valves erect, the pedicle valve upcurrent and the brachial valve upcurrent are the most and least stable orientations, respectively. Shell stability increased among nonstrophic specimens that were more equi-biconvex. Biconvex strophic brachiopods are also more stable if the valves were equi-biconvex, cither compressed (biplanate) or inflated, provided that the specimen was propped up on its hingeline with a lateral margin upcurrent. Catacline interareas afforded greater stability than procline, apsacline or orthocline interareas when the valves were perpendicular to the substrate. Accentuated plication reduced shell stability. Alate biconvex forms were often more stable if the valves were vertical to the substrate rather than reclining. Alate biconvex geometries are more stable than nonstrophic and strophic biconvex forms if the commissural plane is perpendicular to the substrate and parallel to the current. Broad catacline interareas increased stability in all orientations. Plano-convex and dorsi-biconvex alate forms are usually less stable than equi-biconvex or ventri-biconvex alate geometries. Concavo-convex geometries are stable in all orientations except with the valves nearly vertical to the substrate and parallel to the current. Spines greatly retard sediment-scour and maintain concavo-convex specimens in orientations with the valves elevated above the substrates. D Brachiopoda, shell shape, hydrodynamic stability.     

Koskinoid perforations in brachiopod shells: function and mode of formation

Clusters of tiny perforations called 'koskinoid' have been interpreted as having accommodated byssus-like fibrils for attachment in Devonian Uncites and the Orthotetacea, which range from the Silurian through the Permian. The mode of formation of the tiny holes has inspired three hypotheses: (1) The fibrils protruded at the commissure and then were surrounded by growing shell. (2) The fibrils protruded from a juvenile foramen at the beak, which then was covered by shell that caused them to separate and migrate. (3) The fibrils penetrated their own shells like pedicles of living brachiopods are known to penetrate calcite. The first two hypotheses assume that the fibrils were produced by a rudimentary pedicle within the shell; the third postulates that the fibrils are diverticula from the outer epithelium of the mantle. The facts that the clusters are not confined to the beak region, but occur scattered over the ventral valve, and that they also penetrate the interara, favor the third theory. All Orthotetacea except Thecospira have the koskinoid perforations; all Davidsoniacea except Morinorhynchus lack them. This suggests that the two genera should trade places in the classification.     

Epithelial moulds on the shells of the early Palaeozoic brachiopod Lingulella

A fine hexagonal network of microscopic ridges has been discovered on the internal shell surface of the cosmopolitan Lower Palaeozoic inarticulate genus Lingulella. The micro-ornamentation has been recognised on all well-preserved specimens examined, from Cambrian and Ordovician successions in Ireland, North America, Gt. Britain, Spitzbergen, and China. Examination of the mantle of the related Recent genus Lingula revealed that the outer epithelial cells are arranged in a hexagonal close-packed pattern comparable in size and form to the micro-ornamentation in the fossil shells. This fact, along with the discovery of a polygonal pattern on some organic layers in the Lingula shell, suggests that the micro-ornamentation in fossil Lingulella is the mould of the outer epithelium responsible for shell secretion. No trace of epithelial moulds has yet been found in specimens of Lingula from younger Palaeozoic successions. This difference may prove to be a useful diagnostic feature in distinguishing linguloid inarticulates from one another. Epithelial moulds , Lingulella, Palaeoglossa, Lingula, Cambrian, Ordovician, Recent.     

An intracrystalline chromoprotein from red brachiopod shells: implications for the process of biomineralization.

1. The red colour of some terebratulid brachiopod shells is caused by a small chromoprotein that occurs within the calcium carbonate matrix of the shell. 2. This carotenoid-protein complex was isolated from within the calcite shell of three different brachiopod genera and may therefore be involved in the process of biomineralization. 3. The apparent molecular weight of this protein, as judged by SDS-PAGE, is 6.5 kDa. 4. The partial N-terminal amino acid sequence of the protein is virtually identical in three different brachiopod genera, indicating homology. 5. Two carotenoids are present in Terebratella sanguinea: canthaxanthin and the tentatively identified monoacetylinic analogue of astaxanthin.     

Immunological responses from brachiopod skeletal macromolecules; a new technique for assessing taxonomic relationships using shells

The phylogenetic relationships of living, calcareous-shelled brachiopods have been assessed on the basis of immunological responses from intracrystalline macromolecules. Calculations of immunological distances between representatives of the order Terebratulida have revealed a primary threefold division which correlates precisely with a proposed subdivision of the order into three superfamilies but refutes attempts to establish a fourth superfamily. This conclusion was confirmed by carrying out immunological investigations of small shell fragments from other brachiopod genera which were so rare that no antisera could be prepared. The immunological results also indicate a fundamental subdivision of the long-looped brachiopods, with one group perhaps being derived from short-looped terebratuloids rather than long-looped terebratelloids. Sero-taxonomy of skeletal macromolecules provides an ideal method of acquiring molecular phylogenetic data in many groups because a large number of taxa can be surveyed in a short period of time, and microscopic pieces of shell contain sufficient antigenic determinants for many reactions. The technique can also be applied to specimens which have been stored without special treatment in museum collections, making the technique particularly applicable to rare taxa for which no other form of molecular data is available. □ Immunology, Brachiopoda, taxonomy, intracrystalline molecules.     

Mantle canals on brachiopod interareas and their significance in brachiopod classification

Wright, A.D. 1994 10 15: Mantle canals on brachiopod interareas and their significance in brachiopod classification.
Mantle canals have been located on the internal surface of the interareas in several clitambonitacean brachiopod genera. This indicates that, in contrast to the cardinal areas of Recent terebratulides and rhynchonellides, these surfaces were lined with mantle, with no fusion of the mantle lobes at the lateral ends of the hinge line, and the coelomic cavity confined to a median zone at the posterior of the shell. The discovery provides additional support for the view of Jaanusson (1971; Smiths. Contr. Paleobiol. 3 ) that the Beecher (1891) classification has considerable merit, and indicates that the calcareous brachiopods may be subdivided into the three subclasses Craniformea, Protremata and Telotremata. Brachiopoda, classification, Clitambonitacea, interareas, mantle canals, Protremata .     

Supraordinal brachiopod classification

Three controversial problems of brachiopod supraordinal classification are discussed: the position of brachiopods in the classification of Metazoa, their classification at phylum and class level, and the classification of the articulate brachiopods. The position of brachiopods in the system of Metazoa remains uncertain. There are no strong reasons for changing the traditional division of the phylum Brachiopoda into the classes Inarticulata and Articulata. The class Articulata is divided into the subclasses Orthata, Strophomenata, Spiriferata, and Terebratulata.     

Structure of the brachiopod lophophore

Data on the development, structure, and functional morphology of the brachiopod lophophore are analyzed. The common origin of the tentacle apparatus in Lophophorata from the postoral ciliary band of the larva is shown. The brachiopod lophophore is based on the brachial axis consisting of the brachial fold running along the row of tentacles. The brachial axis may be attached to the brachial (dorsal) mantle lobe (trocholophe, schizolophe, and ptycholophe lophophores) or extend freely into the mantle cavity to form coiling brachia (spirolophe, zygolophe, and plectolophe lophophores). The circulation of water flows through the mantle cavity in the brachiopods with attached and free lophophores is described. A new hypothesis on the sorting of particles suspended in water during filtration is proposed.     

Diagenetic fate of bioapatite in linguliform brachiopods: multiple apatite phases in shells of Cambrian lingulate brachiopod Ungula ingrica (Eichwald)

Fossil skeletal apatites vary in their composition and can yield mixed biochemical, environmental and diagenetic information. Thus, it is important to evaluate the diagenesis spatially inside the skeleton. We study the cross sections of shells of the Furongian lingulate brachiopod Ungula ingrica from Estonia using the Attenuated Total Reflectance – Fourier Transform Infrared (ATR‐FTIR) microspectroscopic and energy dispersive spectroscopic (EDS) mapping and show for the first time that different structural laminae of the shell have different chemical compositions. Compact laminae are rich in PO₄^3?, Na, Mg and poor in F and Ca. Porous (baculate) laminae are rich in carbonate anions, Ca and F, but contain less Na and Mg. The ATR‐FTIR spectra show further differences in the ν₂ carbonate region, where the IR band at 872 cm^?1 in compact laminae is replaced by a strong band at 864 cm^?1 in baculate laminae. The changes in shell apatite suggest different origins of the apatite phases. Compact laminae are likely chemically less altered and could potentially carry more reliable palaeoenvironmental or geochemical information than the apatite in baculate laminae, which is mostly authigenic in its origin.     

Ordovician trimerellacean brachiopod shell beds

The large, thick-shelled, inarticulate brachiopod Eodinobolus forms many conspicuous deposits of shells in the Upper Ordovician limestones of central western New South Wales. Both in situ and reworked shell beds arc preserved at recurrent intervals through the successions, in similar facies of both transgressivc and regressive phases of deposition. In situ shell beds arc best developed in transgressivc sequences, with up to four generations of shells exhibited in the individual in situ beds. These monotypic and very low diversity shell beds are interpreted as having formed in marginal marine, quiet water conditions: (1) on the fringes of an offshore island (in part the Molong High of the Tasman Orogen), with the island still providing a fairly continuous supply of terrigenous material: and, (2) after submergence of the island, on the resulting terrigenous-free, major offshore Bahamas-like platform. This may imply that the shell beds developed in different salinity regimes. Possibly Eodinobolus was capable of tolerating a wider than normal range of salinity, from slightly brackish through normal marine, even to marginally hypersaline. However, in both settings, Eodinobolus, in its role as the dominant member of the respective pioneer community, colonized similar substrates in the low energy mud zone. This appears to suggest depositional environments most directly analogous to those of Palaeozoic virgianid pentamerides, and perhaps also comparable with some modern marginal marine oyster and mussel-bed occurrences. ?Ordovician, Brachiopoda, Eodinobolus, palaeoecology, facies, shell beds. New South Wales.     

Functional adaptation of spiriferide brachiopod morphology

It has been suggested that spiriferide morphologies have evolved to adapt to a variety of environmental conditions. Through a computational fluid dynamics approach, we examined how the spiriferide original form was optimized for a lotic condition, specifically addressing the functionalization of the Devonian spiriferide brachiopod Paraspirifer bownockeri to generate passive feeding flows. The results using four models, each of which differed in the development of the spiriferide shell depression, i.e. sulcus, showed that a deeper sulcus functions to create strong spiral flows so as to align on the feeding organ inside the shell. Among the sulcus‐developed models, only the mimic of the natural form could generate comparative slow flows with a stable inflow area. The fossil record of spiriferides shows a morphological trade‐off between the development of the sulcus and wing form. We concluded that spiriferide shells with such a morphological combination evolved to produce various feeding strategies, resulting in diversification.     

Functional morphological analysis of evolution of ribbing in pliocene viviparid shells from Croatia

Posilović, H., & Bajraktarević, Z. 2010: Functional morphological analysis of evolution of ribbing in pliocene viviparid shells from Croatia. Lethaia, 10.1111/j.1502‐3931.2009.00209.x. This paper provides a functional analysis of Pliocene freshwater gastropods from the phylogenetic lineage of Viviparus neumayri to Viviparus ornatus. On this section of the phylogenetic line, it is possible to follow the continuous evolution of the shell ornamentation (shouldering and rib formation) from ancestral V. neumayri with smooth shell to V. ornatus with spiral ribs. The shell morphology and possible structural function of rib development is discussed from the analytical point of view, but also by Finite Element Modelling. Spiral rib development in the viviparids is not correlated with shell thickening, but rather with strengthening of the shell is achieved through development of spiral ribs, with direct biomechanical and evoloutionary significance. □Croatia, functional morphology, gastropod shell evolution, Pliocene, viviparid evolution.     

Paterimitra pyramidalis from South Australia: scleritome,shell structure and evolution of a lower Cambrian stem group brachiopod

The tommotiid Paterimitra pyramidalis Laurie, 1986, is redescribed based on well‐preserved material from the lower Cambrian Wilkawillina, Wirrapowie and Ajax limestones of the Flinders Ranges, South Australia. The material shows that the scleritome of Paterimitra pyramidalis includes three sclerite morphotypes (S1, S2 and L). Detailed shell microstructure studies show striking similarities with both the paterinid brachiopod Askepasma toddense and the tommotiid Eccentrotheca helenia, which strengthens the suggested evolutionary link between tommotiids and brachiopods. Based on the partly articulated specimens and similarities in shell microstructure and sclerite morphology with Eccentrotheca, Paterimitra pyramidalis is reconstructed as a tube‐dwelling, epifaunal, sessile, filter‐feeder with an organic pedicle‐like attachment structure. The proposed reconstruction of the scleritome comprises a basal unit composed of one S1 and one S2 sclerite, as well as an unresolved number of L sclerites lining a coniform tubular structure.     

Ribbing patterns in the brachiopod Diceromyonia

Ribbing patterns in two collections of the Ordovician brachiopod Diceromyonia from central North America are compared using the system invented by Bancroft (1928; Manchester Memoirs 72 ) for naming ribs. Simple and nearly symmetric arrangements of frequently occurring branches characterize both collections, with much of the complexity evident in larger valves due to irregular occurrence of low-frequency branches on either side of major ribs. Examination of median sectors reveals little support for Bancroft's derivation of the rhipidomelloid midrib from isorthoid forms, and an alternative non-evolutionary interpretation is proposed in which median patterns appear as predictable outcomes of a simple space-filling process. The approximately 2500 ribs identified in this study are treated as a population in their own right: orderly patterns which emerge are not inferred but are simple summaries of complete counts, obtained by hand. This approach is quick and painless compared to methods using order relations to assess ribbing differences in dalmanellid brachiopods. For this reason, it may encourage systematic study of ribbing patterns among these taxa and lead ultimately to better understanding of the utility of ribbing in their taxonomy.