Biotic interactions recorded in shells of recent rhynchonelliform brachiopods from San Juan Island, USA

Biotic interactions between brachiopods and spionid polychaete worms, collected around San Juan Islands (USA), were documented using observations from live-collected individuals and traces of bioerosion found in dead brachiopod shells. Specimens of Terebratalia tranversa (Sowerby), Terebratulina unguicula (Carpenter), Laqueus californianus (Koch), and Hemithiris psittacea (Gmelin) were collected from rocky and muddy substrates, from sites ranging from 14.7-93.3 m in depth. Out of 1,131 specimens, 91 shells showed traces of bioerosion represented by horizontal tubes. Tubes are U-shaped, straight or slightly curved, sometimes branched, with both tube openings communicating externally. on internal surfaces of infested shells, blisters are observed. All brachiopod species yielded tubes, except for H. psittacea. Tubes are significantly more frequent on live specimens, and occur preferentially on larger, ventral valves. This pattern suggests selectivity by the infester rather than a taphonomic bias. Given the mode of life of studied brachiopods (epifaunal, sessile, attached to the substrate, lying on dorsal valve), ventral valves of living specimens should offer the most advantageous location for suspension-feeding infesters. Frequent infestation of brachiopods by parasitic spionids is ecologically and commercially noteworthy because farmed molluscs are also commonly infested by parasitic polychaetes. In addition, brachiopod shells are among the most common marine macroscopic fossils found in the Phanerozoic fossil record. From a paleontological perspective, spionid-infested brachiopod shells may be a prime target for studying parasite-host interactions over evolutionary time scales.

Abundant brachiopods on a tropical, upwelling-influenced shelf (Southeast Brazilian Bight, South Atlantic)

Rhynchonelliform brachiopods were diverse and often dominant benthos of tropical seas in the Paleozoic. In contrast, they are believed to be rare in open habitats of modern oceans, especially at low latitudes. This study documents numerous occurrences of rhynchonelliform brachiopods on a modern tropical shelf, particularly in areas influenced by upwelling. Extensive sampling of the outer shelf and coastal bays of the Southeast Brazilian Bight revealed dense populations of terebratulid brachiopods (>10(3) individuals /m(2) of seafloor) between 24 and 26 S. on the outer shelf, brachiopods are more abundant than bivalves and gastropods combined. However, brachiopod diversity is low: only four species belonging to the genera Bouchardia, Terebratulina, Argyrotheca, and Platidia were identified among over 16000 examined specimens. Brachiopods occur preferentially on carbonate bottoms and include two substrate-related associations: Bouchardia (40-70% CaCO3, weight content) and Terebratulina-Argyrotheca (70-95% CaCO3). All four species display a broad bathymetric range that contrasts with a narrow depth tolerance postulated for many Paleozoic rhynchonelliforms. The most abundant populations occur in the depth range between 100 and 200 m, and coincide with zones of shelf-break upwelling, where relatively colder and nutrient-rich water masses of the South Atlantic Central Water are brought upward by cyclonic meanders of the South Brazil Current (a western boundary current that flows poleward along the coast of Brazil). This is consistent with previous biological and paleontological studies that suggest upwelling may play a role in sustaining brachiopod-dominated benthic associations. The presence of abundant brachiopods in the open habitats of the tropical shelf of the western South Atlantic contrasts with current understanding of their latitudinal distribution and points to major gaps in our knowledge of their present-day biogeography. The ecological importance of rhynchonelliform brachiopods in modern oceans and their role as producers of biogenic sedimentary particles may be underestimated.

Biotic interaction between spionid polychaetes and bouchardiid brachiopods: Paleoecological, taphonomic and evolutionary implications

Shells of Bouchardia rosea (Brachiopoda, Rhynchonelliformea) are abundant in Late Holocene death assemblages of the Ubatuba Bight, Brazil, SW Atlantic. This genus is also known from multiple localities in the Cenozoic fossil record of South America. A total of 1211 valves of B. rosea, 2086 shells of sympatric bivalve mollusks (14 nearshore localities ranging in depth from 0 to 30 m), 80 shells of Bouchardia zitteli, San Julián Formation, Paleogene, Argentina, and 135 shells of Bouchardia transplatina, Camacho Formation, Neogene, Uruguay were examined for bioerosion traces. All examined bouchardiid shells represent shallow-water, subtropical marine settings. Out of 1211 brachiopod shells of B. rosea, 1201 represent dead individuals. A total of 149 dead specimens displayed polychaete traces (Caulostrepsis). Live polychaetes were found inside Caulostrepsis borings in 10 life-collected brachiopods, indicating a syn-vivo interaction (Caulostrepsis traces in dead shells of B. rosea were always empty). The long and coiled peristomial palps, large chaetae on both sides of the 5th segment, and flanged pygidium found in the polychaetes are characteristic of the polychaete genus Polydora (Spionidae). The fact that 100% of the Caulostrepsis found in living brachiopods were still inhabited by the trace-making spionids, whereas none was found in dead hosts, implies active biotic interaction between the two living organisms rather than colonization of dead brachiopod shells. The absence of blisters, the lack of valve/site stereotypy, and the fact that tubes open only externally are all suggestive of a commensal relationship. These data document a new host group (bouchardiid rhynchonelliform brachiopods) with which spionids can interact (interestingly, spionid-infested sympatric bivalves have not been found in the study area despite extensive sampling). The syn-vivo interaction indicates that substantial bioerosion may occur when the host is alive. Thus, the presence of such bioerosion traces on fossil shells need not imply a prolonged post-mortem exposure of shells on the sea floor. Also, none of the Paleogene and Neogene Bouchardia species included any ichnological evidence for spionid infestation. This indicates that the Spionidae/ Bouchardia association may be geologically young, although the lack of older records may also reflect limited sampling and/or taphonomic biases.

Colonization of a 'Lost World': Encrustation patterns in modern subtropical brachiopod assemblages.

The encrustation of Paleozoic rhynchonelliform brachiopods has been studied for decades, but modern brachiopods have not received similar scrutiny. The discovery of abundant subtropical brachiopods from the Southeast Brazilian Bight provides an unprecedented opportunity to assess epibiont abundance, diversity, and encrustation patterns in modern brachiopod assemblages. Across the outer shelf, encrustation frequencies vary among taxa, from mean values of 0.45% for Platidia to 9.3% for Argyrotheca. Encrustation frequencies for Bouchardia increase from 1.6% on the outer shelf to 84% on the inner shelf Larger valves are encrusted more frequently, and epibionts preferentially colonize valve interiors. Increased encrustation on the inner shelf may reflect the greater surface area of larger hosts, longer exposure of dead shells, water-mass characteristics, sedimentation rates, productivity, or other factors that vary with depth. Inner-shelf brachiopods exhibit encrustation frequencies comparable to those reported for epifaunal bivalves. The epibiont fauna is dominated by bryozoans and serpulids, with minor roles played by spirorbids, bivalves, barnacles, foraminifera, algae, and other taxa. Epibiont abundance at each site is highly variable, but sites are similar in rank importance of epibiont taxa. A different suite of epibionts colonized Paleozoic brachiopods, but similar patterns of encrustation have been observed, including preferential settlement according to valve morphology. These results provide a baseline for evaluating the encrustation of modern bivalves and ancient brachiopods, and may elucidate the macroevolutionary history of epibionts and their relationship to their hosts.

Comparative analysis of drilling frequencies in recent brachiopod-mollusk associations from the Southern Brazilian Shelf

Over 14,000 specimens-5,204 brachiopods, 9,137 bivalves, and 178 gastropods-acquired from 30 collecting stations (0 to 45 m depth) in the Ubatuba and Picinguaba bays, southern Brazil, were compared for drilling frequencies. Beveled (countersunk) circular-to-subcircular borings (Oichnus-like drill holes) were found in diverse bivalves but also in the rhynchonelliform brachiopod Bouchardia rosea-a small, semi-infaunal to epifaunal, free-lying species that dominates the brachiopod fauna of the southern Brazilian shelf. Drill holes in bivalve mollusks and brachiopods are comparable in their morphology, average diameter, and diameter range, indicating attacks by a single type of drilling organism. Drill holes in brachiopods were rare (0.4%) and found only at five sampling sites. Drillings in bivalves were over 10 times as frequent as in brachiopods, but the average drilling frequency was still low (5.6%) compared to typical boring frequencies of Cenozoic mollusks. Some common bivalve species, however, were drilled at frequencies up to 50 times higher than those observed for shells of B. rosea from the same samples. Due to scarcity of drilled brachiopods, it is not possible to evaluate if the driller displayed a nonrandom (stereotyped) site, size, or valve preference. Drilled brachiopods may record (1) naticid or muricid predation, (2) predation by other drillers, (3) parasitic drillings, and (4) mistaken or opportunistic attacks. Low drilling frequency in brachiopods is consistent with recent reports on ancient and modern examples. The scarcity of drilling in brachiopods, coupled with much higher drilling frequencies observed in sympatric bivalves, suggests that drilling in brachiopods may have been due to facultative or erroneous attacks. The drilling frequencies observed here for the brachiopod-bivalve assemblages are remarkably similar to those reported for Permian brachiopod-bivalves associations. This report adds to the growing evidence for an intriguing macroecological stasis: multiple meta-analytical surveys of present-day and fossil rhynchonelliform brachiopods conducted in recent years also point to persistent scarcity and low intensity of biotic interactions between brachiopods and drilling organisms throughout their evolutionary history.

Life mode of in situ Conularia in a Middle Devonian epibole

Exceptionally abundant specimens of Conularia aff. desiderata Hall occur in multiple marine obrution deposits, in a single sixth-order parasequence composed of argillaceous and silty very fine sandstone, in the Otsego Member of the Mount Marion Formation (Middle Devonian, Givetian) in eastern New York State, USA. Associated fossils consist mostly of rhynchonelliform brachiopods but also include bivalve molluscs, orthoconic nautiloids, linguliform brachiopods and gastropods. Many of the brachiopods, bivalve molluscs and conulariids have been buried in situ. Conulariids buried in situ are oriented with their aperture facing obliquely upward and with their long axis inclined at up to 87degree to bedding. Most specimens are solitary, but some occur in V-like pairs or in radial clusters consisting of three specimens, with the component specimens being about equally long or (less frequently) substantially different in length. The compacted apical end of Conularia buried in situ generally rests upon argillaceous sandstone. With one possible exception, none of the examined specimens terminates in a schott (apical wall), and internal schotts appear to be absent. The apical ends of specimens in V-like pairs and radial clusters show no direct evidence of interconnection of their periderms. The apical, middle or apertural region of some inclined specimens abuts or is in close lateral proximity to a recumbent conulariid or to one or more spiriferid brachiopods, some of which have been buried in their original life orientation. The azimuthal bearings of Conularia and nautiloid long axes and the directions in which conulariids open are nonrandom, with conulariids being preferentially aligned between 350 and 50degree and with their apertural end facing north-east, and nautiloids being preferentially aligned between 30 and 70degree. Otsego Member Conularia were erect or semi-erect, epifaunal or partially infaunal animals, the apical end of which rested upon very fine bottom sediment. The origin of V-like pairs and radial clusters remains enigmatic, but it is probable that production of schotts was not a regular feature of this animal's life history. Finally, conulariids and associated fauna were occasionally smothered by distal storm deposits, under the influence of relatively weak bottom currents. © The Palaeontological Association.

Recent brachiopods from the southern Brazilian shelf: Palaeontological and biogeographical implications

Until recently, the rhynchonelliform (articulated) brachiopod fauna from the Brazilian continental shelf (western South Atlantic) was represented only by the endemic species Bouchardia rosea (Mawe), reported from coastal waters of the states of São Paulo and Rio de Janeiro. The present study, based on samples from coastal (<30 m), shelf, and continental slope waters (99-485 m), documents the South Atlantic brachiopod fauna and shows that this fauna is more widespread, diverse, and cosmopolitan than previously thought. Based on a total of 16,177 specimens, the following brachiopods have been identified: Bouchardia rosea (Family Bouchardiidae), Platidia anomioides (Family Platidiidae), Argyrotheca cf. cuneata (Family Megathyrididae), and Terebratulina sp. (Family Cancellothyrididae). In coastal settings, the fauna is overwhelmingly dominated by Bouchardia rosea. Rare juvenile (<2 mm) specimens of Argyrotheca cf. cuneata were also found at two shallow-water sites. In shelf settings (100-200 m), the fauna is more diverse and includes Bouchardia rosea, Terebratulina sp., Argyrotheca cf. cuneata, and Platidia anomioides. Notably, Bouchardia rosea was found in waters as deep as 485 m, extending the known bathymetric range of this genus. Also, the record of this brachiopod in waters of the state of Parana is the southernmost known occurrence of this species. The genera Platidia and Terebratulina are documented here for the first time for the western South Atlantic. The Brazilian brachiopod fauna shares similarities with those from the Atlantic and Indian shelves of southern Africa, and from the Antarctic, Caribbean and Mediterranean waters. The present-day brachiopods of the western South Atlantic are much more cosmopolitan than previously thought and their Cenozoic palaeobiogeographic history has to be reconsidered from that perspective.

Biogeochemistry of Paleozoic brachiopods from New York State and Ontario

A comprehensive elemental, isotopic and microstructural analyses was undertaken of brachiopod calcites from the Hamilton Group (Middle Devonian), Clinton Group (Middle Silurian) and Middle to Upper Ordovician strata of Ontario and New York State. The majority of specimens were microstructurally and chemically preserved in a pristine state, although a number of specimens show some degree of post-depositional alteration. Brachiopod calcites from the Hamilton and Clinton Groups were altered by marine derived waters whereas Trenton Group (Middle Ordovician) brachiopods altered in meteorically derived fluids. Analysis of the elemental and isotopic compositions of pristine Hamilton Group brachiopods indicates there are several chemical relationships inherent to brachiopod calcite. Taxonomic differentiation of Mg, Sr and Na contents was evident in three co-occuring species from the Hamilton Group. Mean Mg contents of pristine brachiopods were respectively Athyris spiriferoides (1309ppm), Mucrospirifer mucronatus (1035ppm) and Mediospirifer audacula (789ppm). Similarly, taxonomic differentiation of shell calcite compositions was observed in co-occuring brachiopods from the Clinton Group (Middle Silurian) and the Trenton Group (Middle Ordovician). The taxonomic control of elemental regulation into shell calcite is probably related to the slightly different physiological systems and secretory mechanisms. A relationship was observed in Hamilton Group species between the depth of respective brachiopod communities and their Mg, Sr and Na contents. These elements were depleted in the shell calcites of deeper brachiopods compared to their counterparts in shallower reaches. Apparently shell calcite elemental composition is related to environmental conditions of the depositional setting, which may have controlled the secretory regime, mineral morphology of shell calcite and precipitation rates of each species. Despite the change in Mg, Sr and Na contents between beds and formations in response to environmental conditions, the taxonomic differentiation of shell calcite composition is maintained. Thus, it may be possible to predict relative depth changes in paleoenvironmental reconstructions using brachiopod calcite. This relationship of brachiopod chemistry to depth was also tested within a transgressiveregressive (T-R) cycle in the Rochester Shale Formation (Middle Silurian). Decreasing Mg, Sr and Na contents were observed in the transition from the shallow carbonates of the Irondequoit Formation to the deeper shales of the lowest 2 m of Rochester Shale. However, no isotopic and elemental trends were observed within the entire T-R cycle which suggests that either the water conditions did not change significantly or that the cycle is illusory. A similar relationship was observed between the Fe and Mn chemistries of shell calcite and redox/paleo-oxygen conditions. Hamilton Group brachiopods analysed from deeper areas of the shelf are enriched in Mn and Fe relative to those from shallow zones. The presence of black shales and dysaerobic faunas, during deposition of the Hamilton Group, suggests that the waters of the northern Appalachian Basin were stratified. The deeper brachiopods were marginally positioned above an oxycline and their shell calcites reflect periodic incursions of oxygen depleted water. Furthermore, analysis of Dalmanella from the black shales of the Collingwood Shale (Upper Ordovician) in comparison to those from the carbonates of the Verulam Formation (Middle Ordovician) confirm the relationship of Fe and Mn contents to periodic but not permanent incursions of low oxygen waters. The isotopic compositions of brachiopod calcite found in Hamilton Group (813C; +2.5% 0 to +5.5% 0; 8180 -2.50/00 to -4.00/00) and Clinton Group (813C; +4.00/00 to +6.0; 8180; -1.8% 0 to -3.60/ 00) are heavier than previously reported. Uncorrected paleotemperatures (assuming normal salinity, 0% 0 SMOW and no fractionation effects) derived from these isotopic values suggest that the Clinton sea temperature (Middle Silurian) ranged from 18°C to 28°C and Hamilton seas (Middle Devonian) ranged between 24°C and 29°C. In addition, the isotopic variation of brachiopod shell calcite is significant and is related to environmental conditions. Within a single time-correlative shell bed (the Demissa Bed; Hamilton Group) a positive isotopic shift of 2-2.5% 0 in 013C compositions and a positive shift of 1.0-1.50/00 in 0180 composition of shell calcite is observed, corresponding with a deepening of brachiopod habitats toward the axis of the Appalachian Basin. Moroever, a faunal succession from deeper Ambocoelia dominated brachiopod association to a shallow Tropidoleptus dominated assocation is reflected by isotopic shifts of 1.0-1.50/00. Although, other studies have emphasized the significance of ±20/oo shifts in brachiopod isotopic compositions, the recognition of isotopic variability in brachiopod calcite within single beds and within depositional settings such as the Appalachian Basin has important implications for the interpretation of secular isotopic trends. A significant proportion of the variation observed isotopic distribution during the Paleozoic is related to environmental conditions within the depositional setting.

Diagenesis and geochemical stratigraphy of Upper Pennsylvanian madera brachiopods, New Mexico

Owing to the fact that low-Mg calcite fossil shells are so important in paleoceanographic research, 249 brachiopod, cement and matrix specimens from two neighboring localities (Jemez Springs and Battleship Rock), of the Upper Pennsylvanian Madera Formation were analyzed. Of which, about 86% of the Madera brachiopods are preserved in their pristine mineralogy, microstructure and geochemistry. Cement and matrix samples, in contrast, have been subjected to complete but variable post-deposition~1 alteration. It is confirmed that the stable isotope data of brachiopods are much better than that of matrix material in defining depositional parameters. Because there is no uniform or constant relationship between the two data bases (e.g., from 0.1 to 3.0%0 for 0180 and from 0.2 to 6.7%0 for 013C in this study), it is not possible to make corrections for the matrix data. Regarding the two stratigraphic sections, elemental and petrographic analyses suggest that Jemez Springs is closer to Penasco Uplift than Battleship Rock. Seawater at Jemez Springs is more aerobic, and the water chemistry is more influenced by continental sources than that at Battleship Rock. In addition, there is a relatively stronger dolomitization in the mid-section of the Battleship Rock. Results further suggest that no significant biogenic fractionation or vital effects occurred during their shell secretion, suggesting that the Madera brachiopods incorporated oxygen and carbon isotopes in equilibrium with the ambient seawater. This conclusion is not only drawn from the temporal and spatial analyses, but also supported by brachiopod inter-generic comparison (Composita and Neospirifer) and statistical analysis ( t-test).

Aspartic acid racemization dating of Holocene brachiopods and bivalves from the southern Brazilian shelf, South Atlantic

The extent of racemization of aspartic acid (Asp) has been used to estimate the ages of 9 shells of the epifaunal calcitic brachiopod Bouchardia rosea and 9 shells of the infaunal aragonitic bivalve Semele casali. Both taxa were collected concurrently from the same sites at depths of 10 m and 30 m off the coast of Brazil. Asp D/L values show an excellent correlation with radiocarbon age at both sites and for both taxa (r(Site)(2) (9) (B. rosea) = 0.97 r(Site)(2) (1) (B.) (rosea) = 0.997, r(Site)(2) (9) (S.) (casali) = 0.9998, r(2) (Site) (1) (S.casali) = 0.93). The Asp ratios plotted against reservoir-corrected AMS radiocarbon ages over the time span of multiple millennia can thus be used to develop reliable and precise geochronologies not only for aragonitic mollusks (widely used for dating previously), but also for calcitic brachiopods. At each collection site, Bouchardia specimens display consistently higher D/L values than specimens of Semele. Thermal differences between sites are also notable and in agreement with theoretical expectations, as extents of racemization for both taxa are greater at the warmer, shallower site than at the cooler, deeper one. In late Holocene marine settings, concurrent time series of aragonitic and calcitic shells can be assembled using Asp racemization dating, and parallel multi-centennial to multi-millennial records can be developed simultaneously for multiple biomineral systems. (c) 2006 University of Washington. All rights reserved.

Neogene and Pleistocene fossil brachiopods of the Eastern Islands. Current appointments.

[EN] Numerous specimens of fossil brachiopods have been found in the different fossiliferous outcrops of the Canary Islands. These fossils have been found in the deposits of Mio-Pliocene age of the eastern Canary Islands, described and illustrated in the work of Meco et ali. 2005 and in the outcrops interpreted as a tsunami deposits  in Piedra Alta, Lanzarote, belonging to the Marine Isotope Stage 11 dated to circa 330 ka. 4 species of fossil brachiopods have been identificated: Terebratula sinuous Brocchi 1814, Lacazella mediterranea Risso 1826 Terebratulina caputserpentis (Zbyszewski, 1957) and Thecidium cf . digitatum (Sowerby 1823). These fossils provides stratigraphic and paleoclimatic taxonomic information. Furthermore, in order to compare the fossil brachiopods with present in the Canary Island, a reference collection is defined with specimens obtained from marine sediment surveys at Gran Canaria, La Palma and El Hierro, identifying 3 species: Argyrotheca barrettiatia (Davidson, 1866), Megerlia truncata (Linaeus 1767 ) and Pajaudina atlantica (Logan 1988).

Mediterranean brachiopods: morphological, ecological and phylogenetic aspects

The Brachiopoda of the Marine Protected Area “Secche di Tor Paterno”, Central Tyrrhenian Sea, have been investigated in order to give a first glance of the diversity of the brachiopods of this area and provide a new report on the Mediterranean Brachiopod fauna. Four species were reported: Novocrania anomala (Müller, 1776), Megathiris detruncata (Gmelin, 1790), Joania cordata (Risso,1826) and Argyrotheca cuneata (Risso,1826). For all the four species a morphological analysis was carried out. For the two most abundant species, J.cordata and A. cuneata, a morphometric study, based on thickness/width and length/width scattergrams, was carried out, in order to investigate their variability. Size-frequency distributions relative to the three dimensions of the shell were also computed, aimed at a evaluation of population dynamics of these two species. The results showed that, for both species, the parameters which most determine the rise of the shell during the growth of animal are width and length and that frequency distributions are mainly bi- or plurymodal and that they are difficult to interpret, as reported by other studies. Analysis of drill holes found on the shell of some specimens of the two same species revealed a predatory origin and that three different predators are responsible for them. Partial sequences of two different genetic markers, the Internal Transcribed Spacer 1 (ITS1) and the cytochrome oxidase subunit 1 (COI), were used to investigate the phylogenetic relationship between two populations of the eurybathic brachiopod species Gryphus vitreus (Born,1778) across the strait of Gibraltar. This represents the first genetic population study on brachiopods. Results from AMOVA and Bayesian analysis performed on 31 specimens highlighted no genetic differentiation indicating a likely panmixia, dispite the lecitotrophic development of the species.