Protegulum and brephic shell of the earliest organophosphatic brachiopods

Early development stages imprinted on the shells of Cambrian brachiopods from the class Linguliformea (orders Paterinida, Lingulida, Acrotretida) were studied with scanning electron microscope based on a large collection from the Siberian Platform. Some specimens of all three orders preserved protegulum (embryonic shell); their brephic (juvenile) shells, also were studied. Many of them might lack larval development stage as it is known for the recent representatives of the family Lingulidae. But unlike recent lingulids, the surface of the juvenile dorsal valves of all studied linguliformeans bore two (rarely three) pairs of gentle elevations accommodating bundles of setae, which sometime preserve as groups of fine imprints along their margins. Recent Lingula and Glottidia lack setae in this stage. On the other hand, recent classes Craniformea and Rhynchonelliformea have setae, but they develop in the larval stage. Most of brachiopod groups have the setae in the adult stage but these setae have different origin, are short, located on the lateral and anterior ends of the shell, and grow from the marginal cells of the mantle.     

Growth of protein-doped rhombohedra in the calcitic shell of craniid brachiopods

The secondary shell of the living inarticulated brachiopod Neocrania consists of calcitic laminae interleaved with organic sheets, predominantly a 44 kDa protein with high levels of aspartic acid–asparagine and glutamic acid–glutamine. Laminae consist of tabular (01.4) rhombohedra that are composed of spherular or rhombohedral granules, ca. 30 nm in size. Rhombohedra increase by planar or spiral growth as granular, monolayered plates that commonly act as foundations for multilayered tablets up to 300 nm or so high. Rough (0k.l) faces, kinked by cleavage, usually develop at either end of the long diagonals of rhombohedra, the edges of which may support ramparts that can accrete centripetally to enclose any organic material situated at tablet centres. Induced degradation by proteinase K shows that sectors subtended by (0k.l) steps are doped by a fibrous polymer, identified as an exclusively intralaminar glycosylated 60 kDa protein. A 44 kDa protein has also been extracted from laminae and is presumably incorporated into tablets by centripetally growing ramparts.     

Common crystal nucleation mechanism in shell formation of two morphologically distinct calcite brachiopods

Closely related mineral-producing organisms share common biomineralisation processes. We demonstrate that, in cases of disparate mineral structures where crystal growth mechanisms are necessarily diverse, nucleation processes are the common underlying mechanism during shell formation. Detailed crystallography in the context of shell microstructure in two morphologically distinct calcite brachiopods indicates that, despite differences in shell growth and fabric, at the centre of growth, calcite crystals nucleate with the c-axis 0001 parallel to the shell surface. Such detailed contextual crystallography of biomineralisation using electron backscatter diffraction (EBSD) will have significant applications for future research in biological and medical sciences.     

Structure of the first-formed shell of the Middle Ordovician orthid-like brachiopods from the Leningrad Region

Shell structure of the first-formed shell of the Middle Ordovician orthid-like brachiopods from the Leningrad Region is described. The 190-μm-wide first-formed shell is composed of finely granular layer while 700-μm-wide first-formed shell is fibrous. Thus the order Orthida in the Early Paleozoic included brachiopods with both planktotrophic and lecithotrophic larvae in the ontogeny.     

Functional interpretation of inner shell layers in Triassic ceratid ammonites

The wrinkle layer the inner prismatic layer are described in three Triassic ceratid genera: Phyllocludiscites. Megaphyllites Proarcestes. Both layers have their counterparts in the shell wall of the recent Nautilus: the wrinkle layer corresponds to the mantle-adhesive layer the inner prismatic layer to the myostracal layer in Nautilus. A detailed structural functional comparison between these layers is given. The wrinkle layer is also compared with the oblique prismatic layer in recent gastropods.     

Devonian brachiopods of the orders Spiriferida and Spiriferinida of the European Russia and Transcaucasia: Systematics,shell microstructure,and microornament

The shell microstructure and microornament of 33 species of 28 spiriferid and spiriferinid genera from the Devonian of the East European Platform (Belarus, Central, Volga–Ural subregions), southern and middle Timan, and Transcaucasia are described.     

Microborings in Recent brachiopods and the functions of caeca

Microborings in the primary shell layer of Recent brachiopods are clearly seen to avoid endopunctamicroscopic canals pervading the shell fabric and housing papillose extcnlions of the mantle (the caeca). This avoidance confirms the suggestion that the caecal contents inhibit boring organisms (Owen & Williams 1969; Proc. R. Soc. Loud. B, 172 ), and as such the caecum can be considered as an important instrument in protecting the brachiopod shell. A comparison of the relative fecundity of co-habitating impunctate and cndopunctate New Zealand brachiopods provides indirect evidence that the caecum may indeed also function in a nutrient storage capacity. Brachiopods, microborings, primary shell layer, endopuncta, defence, storage.     

Biogeography of the recent brachiopods

The vertical, latitudinal, and circumcontinental zonality of the distribution of the species, genera, and families of recent brachiopods is considered. The distortions of the latitudinal and meridional symmetry of the biogeographic structure of the ocean are analyzed in view of the patterns of the global circulation of the surface and intermediate waters. Thus ancient faunas may be reconstructed based on data on the structural characteristics of the taxocene of recent brachiopods. The features of the paedomorphic evolution of brachiopods from the different families in extreme habitats (interstitial, underwater caverns, submarine rises, abyssal depths, hydrothermal areas, and margins of habitats) are discussed. The biogeographic structure of bottom dwellers is shown to simplify with depth as well as with simplification of the hydrological structure of the ocean. The important role of the bathyal oceanic zone (slopes of continents, islands, submarine mountains, ridges, and rises) in the preservation of faunal relicts is shown. The historical change from brachiopods to bivalves that occurred from the Paleozoic to the Mesozoic and Cenozoic is shown to have resulted not from competitive exclusion, but from complex and global changes in the plankton composition, which were unfavorable for articulate brachiopods, which had already developed specialized feeding habits, feeding on food that led to the production of almost no metabolic waste products; they had even partly lost their alimentary canal. The development of shelly plankton and, especially, of diatoms hampered the post-Paleozoic revival of large assemblages of articulate brachiopods in shallow-water habitats. The unfilled ecological niches were colonized by bivalves, which were widely adapted to feeding on live phyto-and zooplankton. Recent articulate brachiopods, which are adapted to feeding on the products of decay of dead plankton, form a belt of densely populated settlements of the organic biofilter outside the photic zone on the seaward edge of shelves and on the upper parts of the slopes of continents, islands, and submarine rises throughout the world.     

Origin and dispersal of the earliest brachiopods

Brachiopods first appeared at the very beginning of the Phanerozoic together with the first skeletal organisms. Most brachiopod taxa that arose in the first half of the Cambrian had a short temporal range and became completely extinct by the middle of the Middle Cambrian. Rigid articulation of the valves of brachiopods was provided by various structures, which also appeared in the Early Cambrian. This fact points to the importance of this feature for the formation of the whole group and at the same time testifies to the high variability of rigid articulation at the early stages of brachiopod evolution. This is a typical manifestation of archaic diversity in this animal phylum, which appeared very early in the Phanerozoic. Another important property of the archaic diversity of the early brachiopods was the large number of centers of diversification. As for the majority of groups, climatic zonality was the main factor determining the distribution of brachiopods at the beginning of the Phanerozoic. The main ecological types of brachiopods also appeared in the Early Cambrian.     

Environmental effects on adult shell size in Cepaea hortensis

In natural populations of the land snail Cepaea hortensis there is often a negative association between shell size and population density. The usual interpretation of such associations, both in C. hortensis and other species, is that they represent an ecophenotypic effect that is part of a system of density-dependent natality, rather than a genetically-based effect that is produced through density-related selection. However, this interpretation is usually based on supposition rather than evidence. Here we present the results of an experiment in which small juveniles collected from two sampling sites within a colony known to exhibit a negative size-density association were grown to maturity under constant laboratory conditions. The highly significant inter-site difference observed in the wild disappeared in the laboratory-reared snails, indicating the direct influence of environmental factors on size. Because of the nature of the data-sets from which density-dependent natality is inferred, this result has a clearer connection with such density dependence than conventional within-population heritability estimates.     

Predatory asteroids and the decline of the articulate brachiopods

Various causes, such as increased predation pressure, the lack of planktotrophic larvae, a 'resetting' of diversity, increased competition from benthic molluscs and the decline of the Palaeozoic fauna, have been suggested to explain the failure of the brachiopods to reradiate following the Permo-Triassic mass extinction. Increased predation pressure has hitherto appeared improbable, because typical predators of brachiopods, such as teleostean fish, brachyuran crabs and predatory gastropods, did not undergo major radiation until the late Mesozoic and early Cenozoic. However, new evidence strongly suggests that one important group of predators of shelly benthic organisms, the asteroids, underwent a major radiation at the beginning of the Mesozoic. Although asteroids appeared in the early Ordovician, they remained a minor element of the marine benthos during the Palaeozoic acme of the brachiopods. However, these early asteroids lacked four important requirements for active predation on a bivalved epifauna: muscular arms (evolved in the early Carboniferous); suckered tube feet, a flexible mouth frame and an eversible stomach (all evolved in the early Triassic). Thus radiation of the Subclass Neoasteroidea coincided with both their improved feeding capability and the decline of the articulates. The asteroids were the only group of predators of brachiopods that underwent a major adaptive radiation in the earliest Mesozoic. The asteroids may therefore have contributed to inhibiting a Mesozoic reradiation of the brachiopods. Epifaunal species lacking a muscular pedicle may have been particularly vulnerable. Unlike bivalve molluscs, modern brachiopods show only a limited range of adaptations to discourage asteroid predation. □ Asteroidea, Brachiopoda, evolution, predation, functional morphology.     

The phenomenon of the replication of ontogenetic immaturity in adult cats caused by the cerebrospinal fluid of kittens

Studies have been made on the electrical activity of muscles in the hindlimbs in kittens and adult cats before and after injection the liquor from 10-, 21-day and 1-month kittens into the lumbar part of the spinal cord. Analysis was made of the EMG at rest and that of the evoked reactions. It was shown that EMG of adult cats after injection of kitten liquor attained the pattern typical of the donor. The data obtained indicate the presence in kitten liquor of biologically active substances which account for the level of regulatory relationships. Transfer of these substances to adult animals results in the formation of a new level of coordinations of movements which corresponds to immature organism.     

Climatic control of brachiopods in the lower Namurian

The fauna and lithology of a marine band in the lower Edale Shales (Namurian E₂b) at Edale, Derbyshire, England, are described quantitatively. Two brachiopods, Rugosochonetes sp. and Productus hibernicus were the dominant elements of the fauna. The average age at death was about 3 years, although the maximum age of Rugosochonetes (ca. 7 years) was probably a little greater than that of Productus (ca. 5 years). Both sediment deposition and fauna were climatically controlled. The climate was seasonal from wet to dry. Quartz was deposited and Rugosochonetes sp. flourished during the wet periods. Carbonate deposition and Productus hibernicus were dominant during the dry periods.     

Independent similarity in the morphological evolution of brachiopods

The close similarity of the shell exterior of articulate brachiopods from different orders, which must be taken into account in taxonomic identifications and phylogenetic reconstructions, is analyzed. A possible mechanism of the appearance of such brachiopods in connection with the morphogenetic generality of the structurally similar organisms is evaluated.     

Pyramidal spiriferids (brachiopods) from the Middle and Upper Devonian of the Russian Plate: Morphology, systematics, and shell wall structure

The pyramidal spiriferids Thomasaria Stainbrook, 1945 and Pyramina Ljaschenko, 1969 from the collection of brachiopods of Ljaschenko are examined. New data on the shell microornamentation and inner structure support the validity of the genus Piramina and its type species P. oskolensis Ljaschenko. A new species of the genus Thomasaria, T. rotunda is described.     

The origin of the spiriferidine brachiopods

Wright, Anthony D. 197901 15: The origin of the spiriferidine brachiopods. Lethaia . Vol. 12. pp. 29–33. Oslo. ISSN 0024–1164.
In recent years doubts have been expressed as to whether the strophic spiriferidines should continue to be grouped with the non-strophic spiriferids in the order Spiriferida or not. The solution to this problem will only be found by establishing the origin of the spiriferidines. A characteristic feature of the spiriferidines is their well developed micro-ornament; this is shared also by the orthid Platystrophiinae, and together with the striking similarity in overall morphology (well recognized in the early nineteenth century) it suggests that here, in the Platystrophia plexus, is the ancestral stock of the strophic spirebearers.
The Platystrophünae share with the non-strophic Rhynchonellida the potential to develop a spire. It is here considered that the former gave rise to the spiriferidines and the latter to the non-strophic spire-bearers. These quite separate developments justify the separation of the spire-bearers into the two orders Spiriferida and Atrypida.     

Ultrastructure of the loop of terebratulide brachiopods

The folded and twisted calcareous ribbon, forming both the ascending and descending lamellae of the loop of Waltonia inconspicua (Sowerby), is a two-layered structure consisting of a wedge of regularly stacked secondary layer fibres that overlie a thin layer of non-fibrous calcite (herein termed brachiotest). On one surface, that facing into the mantle cavity, secondary fibrous mosaic predominates, but smooth, finely banded brachiotest occurs as a narrow marginal lip upon which secondary layer fibres proliferate and progressively overlap. This growing edge of the ribbon is secreted by long, folded epithelial cells with digitate extensions to their apical plasmalemmas, which are distinguishable from the cuboidal epithelium-secreting fibres and their membranous sheaths. The other surface, facing the body cavity and the brachial coelom, consists entirely of roughened brachiotest exhibiting prominent banding that is aligned parallel to the growing loop edge. This surface is overlain by microfilamentar epithelium acting as a holdfast for the connective tissue frame of the lophophore. The other edge of the ribbon consists of truncated sections of both secondary-layer fibres and brachiotest which bear signs of resorption consistent with the degenerated state of the associated epithelium. Growth of the Waltonia loop is controlled by these localized processes of secretion and resorption of the fibrous and brachiotest layers and is typical of all terebratulides so far studied. The brachiotest is not homologous with the non-fibrous primary shell secreted at the valve margin. □ Brachiopoda, Articulata, Terebratulida, ultrastructure, lophophore, loop.     

Size-frequency and population structure of brachiopods

The living terebratulids, Terabratulina unguicula, Terebratalia transversa, Laqueus vancouverensis, and the rhynchonelid Hemithiris psittacea were studied in the San Juan Islands, Washington, U.S.A. Those results and a review of a the literature lead to the conclusion that most brachiopod populations experience episodic recruitment at intervals which may be irregular. The occurrence of juveniles attached to adults, brooding, and bi- or multimodal size-frequency distributions demonstrate that, contrary to a previously suggested hypothesis, adult brachiopods do not generally exclude juveniles from the same area. The commony observed rarity of small individuals is regarded as a product of local recruitment failure due to patchy distribution of larvae; it does not justify the assumption that brachiopods are unaffected by high post-larval juvenile mortality. However, the frequent rarity of small individuals confirms that this cannot be used as a criterion of transport in assemblages of fossil brachiopods.     

Adaptive homoeomorphy in the brachiopods Tetractinella Bittner and Cheirothyris Rollier

The Middle Triassic athyraceanTetractinella and the Upper Jurassic terebratellaceanCheirothyris show a close homoeomorphic resemblance in their external shell form. The homoeomorphy is here interpreted as the result of the parallel evolution of an identical adaptive device. Structurally, the homoeomorphy is due to the development of two pairs of long, narrow radial deflections of the commissure, and consequently of two pairs of marginal projections and radial costae on the shell. Functionally, these projections are interpreted as sensory “antennae”, which carried the sensitive mantle edges outwards beyond the rest of the shell, and hence provided the brachiopods with early warning of the approach of potentially harmful agents in the environment.