Gryphus Vitreus Research Articles

How long does a brachiopod shell last on a seafloor? Modern mid‐bathyal environments as taphonomic analogues of continental shelves prior to the Mesozoic Marine Revolution

AbstractCarbonate skeletal remains are altered and disintegrate at yearly to decadal scales in present‐day shallow‐marine environments with intense bioerosion and dissolution. Present‐day brachiopod death assemblages are invariably characterized by poor preservation on continental shelves, and abundant articulated shells of brachiopods with complete brachidia are thus not expected to be preserved if not rapidly buried. However, such preservation is paradoxically observed in shallow‐water Palaeozoic and Mesozoic brachiopod assemblages. Here, we show that a bathyal death assemblage time‐averaged to several millennia (Adriatic Sea) consists of sediment‐filled articulated shells of Gryphus vitreus with complete brachidia. Post‐mortem age distributions indicate that disintegration half‐lives exceed several centuries (c. 500–1700 years). The high frequency of articulated but centuries‐old shells (>50%) and the fitting of taphonomic models to post‐mortem ages indicate that disarticulation half‐life is unusually long (c. 200 years). Rapid sediment filling of shells: (1) inhibited disarticulation, loop fragmentation and colonization by coelobites; and (2) induced precipitation of ferromanganese oxides at redox fronts within shells. Sediment‐filled articulated shells, however, still resided at the sediment–water interface as indicated by encrusters and sponges that infested them after death. Sediment‐filled shells disintegrated through bioerosion and physical wear when residence in the taphonomically active zone exceeded c. 2000 years. We suggest that the articulation paradox is driven by the Mesozoic Marine Revolution (MMR) that escalated predation, bioturbation and organic matter recycling, all intensifying shell disintegration. A scenario with slow disarticulation in bathyal environments may have lead to preservation of articulated shells in shallow‐water assemblages prior to the MMR.

Brachiopod Fauna from the Deep Mediterranean Sea: Distribution Patterns and Ecological Preferences

Compared to their fossil counterparts, living brachiopods are investigated far less often, due to their occurrence in remote environments such as dark caves or deep environments. Due to the scarcity of studies targeting in situ brachiopods’ populations, large-scale information on their distribution and ecological preferences is still lacking, especially on hardgrounds. The extensive employment of remotely operated vehicles (ROVs), however, has opened up the chance to better explore this taxon’s diversity and ecology in the mesophotic and bathyal zones. The analysis of over 600 h of video footage collected from 624 sites, from 40 m to 1825 m, located along the Ligurian and Tyrrhenian coasts of Italy and the Sicily Channel, allowed for a large-scale investigation. The four identified species, Novocrania anomala, Gryphus vitreus, Megerlia truncata and Terebratulina retusa, emerged as common macrofaunal components of the explored habitats, especially between 150 m and 250 m, with high occurrences in the northern areas, especially on offshore seamounts. All species can form dense aggregations of individuals, with M. truncata showing the densest populations on steep rocky terraces (up to 773 individuals m−2). Except for G. vitreus, the only species also recorded on soft bottoms, the others were found exclusively on hardgrounds, with N. anomala showing a peculiar ability to exploit anthropogenic substrates such as terracotta amphorae. No stable species-specific associations were noted, even if numerous species were frequently observed together. Although brachiopods do not show the conspicuous tridimensionality of large filter-feeders, their substrate occupancy and their role in pelagic–benthic processes support their importance in deep-sea Mediterranean ecosystems.

A SAMPLING STRATEGY FOR RECENT AND FOSSIL BRACHIOPODS: SELECTING THE OPTIMAL SHELL SEGMENT FOR GEOCHEMICAL ANALYSES

Recent and fossil brachiopod shells have a long record as biomineral archives for (palaeo)climatic and (palaeo)environmental reconstructions, as they lack or exhibit limited vital effects in their calcite shell and generally are quite resistant to diagenetic alteration. Despite this, only few studies address the issue of identifying the best or optimal part of the shell for geochemical analyses. We investigated the link between ontogeny and geochemical signatures recorded in different parts of the shell. To reach this aim, we analysed the elemental (Ca, Mg, Sr, Na) and stable isotope (δ18O, δ13C) compositions of five recent brachiopod species (Magellania venosa, Liothyrella uva, Aerothyris kerguelensis, Liothyrella neozelanica and Gryphus vitreus), spanning broad geographical and environmental ranges (Chile, Antarctica, Indian Ocean, New Zealand and Italy) and having different shell layer successions (two-layer and three-layer shells). We observed similar patterns in the ventral and dorsal valves of these two groups, but different ontogenetic trends by the two- and three-layer shells in their trace element and stable isotope records. Our investigation led us to conclude that the optimal region to sample for geochemical and isotope analyses is the middle part of the mid-section of the shell, avoiding the primary layer, posterior and anterior parts as well as the outermost part of the secondary layer in recent brachiopods. Also, the outermost and innermost rims of shells should be avoided due to diagenetic impacts on fossil brachiopods.

Initiating DNA barcoding of Eastern Mediterranean deep-sea biota

Preliminary results of DNA barcoding survey of deep-sea mega-faunal biota are presented, collected by trawl and gillnet off the Israeli coast (SE Mediterranean, depths 700 to 1500 meters) during 2012-2013. 846 organisms were identified to 37 species, mainly fish and decapod crustaceans. The most abundant species were the blackmouth catshark Galeus melastomus, the cosmopolitan decapod Polycheles typhlops and the bivalve Abra longicallus. Two species were sampled for the first time from the southern Levant- the long armed chiroteuthid squid Chiroteuthis veranyi and the common mora, Mora moro. Four of the 18 fish species and two of the 10 crustacean species were abundant, representing 78% and 61%, respectively, of the organisms collected. Most other species are represented by fewer than 10 individuals. PCR products for the cytochrome c oxidase sub unit I (COI) gene for the 37 species were successfully sequenced. The identified and vouchered individuals are stored at the Steinhardt Museum of Natural History (Tel Aviv University, Israel) and their COI sequences were uploaded into the BoLD universal data center as part of the national marine barcoding project. The COI sequences of Acanthephyra eximia, Gryphus vitreus, Galeodea echinophora, Mesothuria intestinalis and Astropecten irregularis, constitute first records of these species in BoLD. When compared to the COI sequences in BoLD, the present results reveal some inconsistency in species identification, an outcome that should be taken into consideration primarily once the taxonomical verifications of collected taxa are elusive. This study is the first step in DNA barcoding of the Levant’s little-known benthic deep-sea fauna.

Megabenthic assemblages in the continental shelf edge and upper slope of the Menorca Channel, Western Mediterranean Sea

Highly diverse megabenthic assemblages dominated by passive and active suspension feeders have been recently reported in shelf edge environments of the Mediterranean Sea. Due to their frequent association with species of commercial interest, these assemblages have been heavily impacted by fishing. The vulnerability and low resilience of these assemblages, composed mainly by long-living and slow-growing species, have motivated the implementation of management measures such as the restriction of bottom trawling, and the establishment of large protected areas embracing these environments. The Menorca Channel is one of such areas recently included in the European Union Natura 2000 network. Quantitative analysis of video transects recorded at 95–360 m depth by manned submersible and remotely operated vehicles were used to characterize megabenthic assemblages and to assess their geographical and bathymetric distribution. Six different assemblages were identified, mainly segregated by substrate type and depth. Hard substrates hosted coral gardens and sponge grounds, whereas soft sediments were mainly characterized by large extensions of the crinoid Leptometra phalangium and the brachiopod Gryphus vitreus. The good preservation of most of the observed assemblages is probably related to a low bottom trawling pressure, which mainly concentrates deeper on the adjacent continental slope. Because of their biological and ecological value, management and conservation measures need to be established to preserve these benthic assemblages.

Overcoming the fragility - X-ray computed micro-tomography elucidates brachiopod endoskeletons.

IntroductionThe calcareous shells of brachiopods offer a wealth of informative characters for taxonomic and phylogenetic investigations. In particular scanning electron microscopy (SEM) has been used for decades to visualise internal structures of the shell. However, to produce informative SEM data, brachiopod shells need to be opened after chemical removal of the soft tissue. This preparation occasionally damages the shell. Additionally, skeletal elements of taxonomic/systematic interest such as calcareous spicules which are loosely embedded in the lophophore and mantle connective tissue become disintegrated during the preparation process.ResultsUsing a nondestructive micro-computed tomography (μCT) approach, the entire fragile endoskeleton of brachiopods is documented for the first time. New insights on the structure and position of tissue-bound skeletal elements (spicules) are given as add ons to existing descriptions of brachiopod shell anatomy, thereby enhancing the quality and quantity of informative characters needed for both taxonomic and phylogenetic studies. Here, we present five modern, articulated brachiopods (Rectocalathis schemmgregoryi n. gen., n. sp., Eucalathis sp., Gryphus vitreus, Liothyrella neozelanica and Terebratulina retusa) that were X-rayed using a Phoenix Nanotom XS 180 NF. We provide links to download 3D models of these species, and additional five species with spicules can be accessed in the Supplemental Material. In total, 17 brachiopod genera covering all modern articulated subgroups and 2 inarticulated genera were X-rayed for morphological analysis. Rectocalathis schemmgregoryi n. gen., n. sp. is fully described.ConclusionMicro-CT is an excellent non-destructive tool for investigating calcified structures in the exo- and endoskeletons of brachiopods. With high quality images and interactive 3D models, this study provides a comprehensive description of the profound differences in shell anatomy, facilitates the detection of new delicate morphological characters of the endoskeleton and gives new insights into the body plan of modern brachiopods.Electronic supplementary materialThe online version of this article (doi:10.1186/s12983-014-0065-x) contains supplementary material, which is available to authorized users.

Nanometer Scale Microstructure and Microtexture of Biological Materials Revealed by High Spatial Resolution (15 to 5 kV) EBSD

High resolution EBSD analysis was carried out under specific experimental conditions (15 to 5 kV) on the skeleton of the modern carbonate brachiopod Gryphus vitreus and resolved nano- to microscale preferred crystallographic orientation patterns undetcted so far. As biologic superstructures are formed by controlled nanoparticle assembly it is essential to resolve their internal structure and texture with the highest possible spatial resolution. Low kV EBSD (15 kV and at 5 kV) provides the required resolution. We observe in the investigated carbonate skeletons a strongly interlocking microstructure of concave/convex grains. The interface topology of the interdigitating structure reaches below the micrometer scale. Individual grains reach sizes up to 20 µm (or even more) in one dimension. They show a mosaic spread of several degrees such that they must be addressed as mesocrystals. Even though the shell consists of three different microstructures with completely different crystal morphologies and grain boundary topologies the crystallographic texture of the three layers is similar. This indicates that distinct control mechanisms prevail when the shell is formed.

Interdigitating biocalcite dendrites form a 3-D jigsaw structure in brachiopod shells

We report a newly discovered dense microstructure of dendrite-like biocalcite that is formed by marine organisms. High spatial resolution electron backscatter diffraction (EBSD) was carried out under specific analytical conditions (15 and 10 kV) on the primary layer of the modern brachiopod Gryphus vitreus. The primary layer of modern brachiopods, previously termed nanocrystalline, is formed by an array of concave/convex calcite grains with interdigitated recesses and protrusions of abutting crystals without any cavities in or between the dendrites. The interface topology of this structure ranges from a few tens of nanometres to tens of micrometres, giving a nanoscale structure to the material fabric. The dendritic grains show a spread of crystallographic orientation of several degrees and can thus be referred to as mesocrystals. Individual dendritic mesocrystals reach sizes in one dimension larger than 20 μm. The preferred crystallographic orientation is similar in the primary and adjacent fibrous shell layers, even though these two layers show completely different crystal morphologies and grain boundary topologies. This observation indicates that two separate control mechanisms are active when the primary and the fibrous shell layers are formed. We propose a growth model for the interdigitated dendritic calcite grain structure based on a precursor of vesicles filled with amorphous calcium carbonate (ACC).

Stable isotope profiles (Ca, O, C) through modern brachiopod shells of T. septentrionalis and G. vitreus: Implications for calcium isotope paleo-ocean chemistry

Brachiopod shells are widely used as an archive to reconstruct elemental and isotopic composition of seawater. Studies, focused on oxygen and carbon isotopes over the last decades, are increasingly extending to the emerging calcium isotope system. To date, only little attention has been paid to test the reliability of fossil brachiopods on their modern counterparts. In this context, the present study investigates two modern brachiopods, Terebratulina septentrionalis (eastern Canada, 5–30 m depth, 7.1 °C seasonal temperature variation, two-layer shell) and Gryphus vitreus (northern Mediterranean, 200 m depth, constant all-year round temperature, three-layer shell). Both species were sampled along the ontogenetic growth direction and calcium, oxygen, and carbon isotopes as well as elemental concentration were measured. Calcium isotopes were analyzed on TIMS. The elemental composition was analyzed by LA-ICP-MS and ICP-AES. The results indicate an intra-specimen δ 44/40Ca variation ranging from 0.16 to 0.33‰, pointing to a fairly homogenous distribution of calcium isotopes in brachiopod shells. However, in the light of the suggested 0.7‰ increase in calcium isotopes over the Phanerozoic such intra-specimen variations constrain ocean reconstruction. δ 44/40Ca values of T. septentrionalis do not seem to be affected by growth rate. Calcium isotopic values of G. vitreus are heavy in the central part of the shell and trend towards lighter values in peripheral areas approaching the maximum isotopic composition of T. septentrionalis. The maximum inter-species δ 44/40Ca difference of 0.62‰ between T. septentrionalis and G. vitreus indicates that care should be taken when using different taxa, species with different strontium content or brachiopods with specialized shell structure, such as G. vitreus, for ocean water reconstruction in terms of Ca isotopic composition. T. septentrionalis may record Ca isotopic fractionation related to seasonal seawater temperature variations in its shell but this is difficult to resolve at the current analytical precision. Average δ 18O-derived temperatures of the two investigated species are close to on-site measured temperatures.

Shell matrices of Recent rhynchonelliform brachiopods: microstructures and glycosylation studies

ABSTRACTLike most metazoan biomineralisations, the brachiopod shell is the end product of a biologically controlled calcification process. The main agent of the control is the extracellular matrix, which is secreted by the outer mantle epithelium. This matrix mediates the calcification process by allowing crystal nucleation and elongation in specific orientations and finally, by stopping crystal growth. The proteinaceous moiety of brachiopod shell matrices has been extensively studied. Less known are the post-translational modifications that occur in these matrices, in particular glycosylations. In this comparison of five species of Recent articulated brachiopods, the ratio of soluble to insoluble organic matrix varies between the species. Polydisperse macromolecular materials occur in each of these species with discrete proteins of 50 kDa in Notosaria nigricans, Calloria inconspicua and Neothyris lenticularis, 37 kDa in Terebratulina retusa and Gryphus vitreus and 20–25 kDa in N. nigricans. Protein mixtures from all five species respond differently to anionic stains (Stains-All and Alcian Blue). PAS staining results in a positive smear in C. inconspicua and T. retusa and highlights low molecular weight glycoproteins in C. inconspicua. The polysaccharide composition of the soluble matrix of T. retusa is different from the others due to high proportions of arabinose and low proportions of fucose. In all cases, polysaccharide composition of the insoluble matrix is dominated by glucose and glucosamine. Insoluble matrices have more glucose and xylose and less galactosamine and glucosamine than the corresponding soluble matrix. Relatively high amounts of glucosamine may suggest the presence of chitin in the shell matrix of rhynchonelliform brachiopods.

Drilling predation on Gryphus vitreus (Brachiopoda) off the French Mediterranean coasts

A total of 145 valves of Gryphus vitreus with naticid drillholes have been recorded from samples during 21 cruises in the Northwestern Mediterranean, out of about 30,000 empty valves. Drilling was lower than 1% in all stations, except in one station where it reached up to 25.5%. Selectivity of drillhole site by the naticid Euspirella pulchella seems to apply in G. vitreus as drillholes are mainly concentrated in the relatively meaty distal half of the shell, which is also the thickest part of the valve (about 0.6–2 mm). Attacks occur generally on the ventral valve (86.2%) according to the life position of Gryphus. The predation pressure on G. vitreus by drilling appears exceptional in that it has hardly any influence on the evolution of the species. This contradicts several previous statements on the fossil record.

Distributional patterns along the Mediterranean continental margin (upper bathyal) using Gryphus vitreus (Brachiopoda) densities

The Gryphus vitreus biocoenosis occurs in two types of profiles of the bottom topography on the continental slope. The density of G. vitreus can be divided into five zones directly related to the velocity of the bottom-current. The direction of the current over the Gryphus vitreus biocoenosis suggests three models according to the distributional limits of the density zones; the upper limit is always the continental shelf-edge (100–120 m) and the zonal limits are related to the slope gradient below 150 m. The bathymetric and spatial distribution of the density zones are analysed in relation to the current velocity and direction.

Offshore brachiopods investigated by submersible

The distributional limits and densities of the brachiopod Gryphus vitreus (Born) are investigated for the first time by submersible, along transects over the “Banc du Magaud” (east of Hyères islands, Provence, France). The offshore bottoms, on which the G. vitreus biocoenosis occurs (110 to 250 m), can be divided into three zones. The highest density (up to 600 individuals·m −2) is observed in large ripplemarks under strong bottom currents (1.5-2 knots) in Zone 2 (130–180 m), while the density is lower (< 10·m −2) in Zone 1 (110–130m) and Zone 3 (180–250 m, on a Quaternary thanatocoenosis) where the currents are weaker (0.5-1 knot). The ecological requirements of G. vitreus, a stenotopic species, are high hydrodynamic conditions (inducing a weak sedimentation, high nutrients supplies, sandy sediment with small hard substrata) and an annual constancy of temperature and salinity. The arrangement of the shells in relation to the current (Zone 2) is analysed. Paleoecological implications for branchiopods are suggested. A green alga is recorded to depths of 190 m.

Distribution et syn�cologie des fonds � Gryphus vitreus (Brachiopoda) en Corse

In the Mediterranean Sea, the brachiopod Gryphus vitreus (Born) was collected off Corsica by dredging and trawling during three cruises (June 1983; April and July 1984) and from along the French continental coast by dredging during many cruises (1975–1985). The upper limit of its bathymetric distribution lies between 115 and 125 m (rarely, from 90 m), the lower limit between 140 and 200 m depth, forming a belt along the edge of the continental shelf. The horizontal distribution of G. vitreus varies from ∼250 m to >4 km, and is directly related to geomorphological and hydrodynamic variations resulting from local topography and/or geographical location. The G. vitreus bottoms are bounded on the continental slope by the bathyal mud biocoenosis and on the shelf by the coastal detritic or shelf-edge detritic biocoenosis, both circalittoral. The bottoms correspond to a bathylittoral ecosystem characterized by (1) prevailing environmental conditions which result in a biotope characterized by coarse detritic sediment clogged by fine sand and mud with a high content of small hard substrates, by constant moderate bottom-currents which become weaker with increasing depth, and by small annual variations in temperature and salinity; the occurrence of an ostreobium sp. alga in the shell of G. vitreus indicates the involvement of the phytal system, as defined by Peres (1982); (2) the G. vitreus biocoenosis, comprised of characteristic macrobenthic stocks (particularly polychaetes, mollusks, and epifaunal species on small hard substrates). Whatever the geomorphology, neighbouring circalittoral and bathyal biocoenoses have little influence on the G. vitreus biocoenosis, which displays a comparatively greater species richness and density. However, immigration of circalittoral species is encouraged by detrital sediments and water currents. The occurrence of this ecosystem as a distinct belt intercalated between circalittoral and bathyal zones is discussed. The ecosystem is considered to be “bathylittoral”.

Notes on the Shell Musculature of Gryphus vitreus (Born).

In work on fossil brachiopods the present writer has found that descriptions and more particularly figures of the musculature of living froms are insufficiently explicit for parallels to be drawn with fossil specimens. These notes are the result of dissections of specimens of the living Terebratulinid Gryphus vitreus (Born) from the Marine Biological Station at Naples, and are intended primarily to elucidate the accompanying figures, which it is hoped might be of some value to palaeontologists.