Taphonomy of Bouchardia rosea (Rhynchonelliformea, Brachiopoda) shells from Ubatuba Bay, Brazil: implications for the use of taphonomic signatures in (paleo)environmental analysis

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Viés Analítico em Estudos Tafonômicos com Macroinvertebrados: Implicações (Paleo)Ambientais e (Paleo)Ecológicas

A review of recent literature shows that most taphonomic studies of Holocene and fossil macrovertebrates are not methodologically standardized. Hence, results from distinct studies are not comparable, even among researches sharing virtually identical goals, targeting the same biological group of similar age and depositional environment. The effects of the shell size in the taphonomic analysis are still poorly understood. In order to study this issue, the taphonomic signatures (articulation, valve type, fragmentation, abrasion, corrosion, edge modification, color alteration, bioerosion and encrustation) of brachiopod shells (Bouchardia rosea (Mawe)), from Ubatuba Bay in the northern coast of São Paulo State, were investigated according to the sieve sizes. In the study area, 14 collecting stations were sampled via Van Veen grab sampler, along a bathymetric gradient, ranging from 0 to 35 m of depth. Bulk samples were sieved through 8 mm, 6 mm, and 2 mm mesh sizes, yielding a total of 5.204 shells. The results indicate that, when taphonomic signatures were independently analyzed per size classes (8 mm, 6 mm, and 2 mm), the taphonomic signatures are recorded in a complex and random way. Additionally, cluster analysis showed that the similarity among the clusters vary according to the considered sieve size. Thus, the sieve size plays an important role in the distribution of taphonomic signatures in shells of distinct sizes. These results suggest that the concentration of the taphonomic analysis on one class (e.g., the largest sieve size, 8 mm) is not always the best method. Rather, the total data (all sieves included) seems more accurate in recording the whole spectrum of taphonomic processes recorded in shells of a given assemblage.

Braquiópodes (Rhynchonelliformea, Bouchardioidea) neógenos da bacia de Pelotas (RS) e seu significado paleoambiental

The occurrence of brachiopods in Cenozoic rocks of the Pelotas Basin is known since 1862. In spite of that, detailed systematic and taphonomic studies are still missing. Investigations made a half century ago, have suggested that these brachiopods could belong to Bouchardia cf. zitteli, a species found in the San Julian Formation, Late Oligocene, Argentina. Our data suggest that those brachiopods may resemble Bouchardia transplatina. In the Uruguayan portion of the Pelotas Basin B. transplatina is known in rocks of the Camacho Formation, Miocene. In addition, small recrystallized shells of brachiopods were also recovered from three Petrobras boreholes (2PJ-1-RS, 2PN-1-RS, and 2GA-1-RS) from the Pelotas Basin. Brachiopods come from the interval of 130 to 150 meters within the Miocene Henryhowella evax Zone. Despite the degree of taphonomic modiication of those brachiopod shells they indubitably belong to Bouchardia sp. This is noteworthy for various reasons: 1- Bouchardia is a brachiopod with warm water afinities. Presently, extant members of this genus are unknown in latitudes up to 34[degree]S, with the main records at 23[degree]S. 2- Although occurring in depths down to 200 meters, the living member (Bouchardia rosea) of this genus is most abundant in shallow platformal, nutrient-rich waters. 3- The occurrence of Bouchardia in the Miocene of the Pelotas Basin indicates that, at least to the interval of Henryhowella evax Zone, warm waters of the Brazilian currents prevail. This interpretation is in strong accordance with other paleoeoceanographic and paleoclimatic data offered by various groups of co-occurring microfossils, such as ostracodes and foraminifers.

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.

Drill holes in shells of Permian benthic invertebrates

Newly discovered benthic fossils and specimens illustrated in the paleontological literature indicate that drilling predators (or parasites) were present in the Permian. New field data from southern Brazil document the first drill holes ever reported for Permian bivalve mollusks. In addition, a literature review revealed drill holes in shells of articulate brachiopods from Russia, Greece, and West Texas. Holes range in size from 0.1 to 5.8 mm and are typically round, cylindrical, singular penetrations perpendicular to the valve surface. Incomplete, healed, and multiple holes are absent. Drilling frequency, a proxy for predation intensity, is very low: less than 1 percent (this estimate may be seriously affected by taphonomic and monographic biases). Literature data suggest that frequency of drilled specimens varied significantly among higher brachiopod taxa. The geography and stratigraphy of drilled specimens indicate that drilling organisms were worldwide in their occurrence and continuously present in marine ecosystems throughout the Permian. This report is consistent with other recent studies indicating that although drillers were continuously present throughout the Phanerozoic, drilling intensity was lower in the Late Paleozoic and early Mesozoic.

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.

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.

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.

The temporal resolution of epibiont assemblages: Are they ecological snapshots or overexposures?

The effects of time averaging on the fossil record of soft-substrate marine faunas have been investigated in great detail, but the temporal resolution of epibiont assemblages has been inferred only from limited-duration deployment experiments. Individually dated shells provide insight into the temporal resolution of epibiont assemblages and the taphonomic history of their hosts over decades to centuries. Epibiont abundance and richness were evaluated for 86 dated valves of the rhynchonelliform brachiopod Bouchardia rosea collected from the inner shelf. Maximum abundance occurred on shells less than 400 yr old, and maximum diversity was attained within a century. Taphonomic evidence does not support models of live-host colonization, net accumulation, or erasure of epibionts over time. Encrustation appears to have occurred during a brief interval between host death and burial, with no evidence of significant recolonization of exhumed shells. Epibiont assemblages of individually dated shells preserve ecological snapshots, despite host-shell time averaging, and may record long-term ecological changes or anthropogenic environmental changes. Unless the ages of individual shells are directly estimated, however, pooling shells of different ages artificially reduces the temporal resolution of their encrusting assemblages to that of their hosts, an artifact of analytical time averaging. © 2006 by The University of Chicago. All rights reserved.

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.

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.

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.

Seawater carbonate chemistry and shell weights of Laternula elliptica, Yoldia eightsi, Nacella concinna and Liothyrella uva during experiments, 2009

Antarctic calcified macroorganisms are particularly vulnerable to ocean acidification because many are weakly calcified, the dissolution rates of calcium carbonate are inversely related to temperature, and high latitude seas are predicted to become undersaturated in aragonite by the year 2100. We examined the post-mortem dissolution rates of aragonitic and calcitic shells from four species of Antarctic benthic marine invertebrates (two bivalves, one limpet, one brachiopod) and the thallus of a limpet shell-encrusting coralline alga exposed to acidified pH (7.4) or non-acidified pH (8.2) seawater at a constant temperature of 4 C. Within a period of only 14-35 days, shells of all four species held in pH 7.4 seawater had suffered significant dissolution. Despite calcite being 35% less soluble in seawater than aragonite, there was surprisingly, no consistent pattern of calcitic shells having slower dissolution rates than aragonitic shells. Outer surfaces of shells held in pH 7.4 seawater exhibited deterioration by day 35, and by day 56 there was exposure of aragonitic or calcitic prisms within the shell architecture of three of the macroinvertebrate species. Dissolution of coralline algae was confirmed by differences in weight loss in limpet shells with and without coralline algae. By day 56, thalli of the coralline alga held in pH 7.4 displayed a loss of definition of the conceptacle pores and cracking was evident at the zone of interface with limpet shells. Experimental studies are needed to evaluate whether there are adequate compensatory mechanisms in these and other calcified Antarctic benthic macroorganisms to cope with anticipated ocean acidification. In their absence, these organisms, and the communities they comprise, are likely to be among the first to experience the cascading impacts of ocean acidification.

Brachiopods in Early Carboniferous

Changes in Mississippian global paleogeography derived from the reconfiguration of the continents, a reversal in ocean currents and global cooling. Although the tectonic and climatic changes are well-documented, their effects on the distribution of brachiopod fauna are poorly documented. Here we present systematic quantitative analyses on global paleobiogeography based on a global brachiopod database from the Mississippian (i.e., Tournaisian, Visean, and Serpukhovian). The dataset consists of 2123 species of 344 brachiopod genera from 1156 localities. Our results reveal that global provincialism was not evident during the Tournaisian and Visean Stages. Two realms, i.e., the Gondwanan and Paleoequatorial Realms, are recognized during the Tournaisian. The Paleoequatorial Realm dominates during the Visean Stage, whereas the Gondwanan Realm is not documented due to the absence of data points. In contrast to the early and middle Mississippian stages, faunal provincialism is greatly enhanced in the Serpukhovian Stage with Paleotethyan and North American realms easily distinguished. This indicates that the Rheic Ocean was closed before the Serpukhovian due to the collision between Gondwana and Laurussia, that disrupted faunal interchange between the Paleotethys and North America. In addition, the paleolatitude-related thermal gradient was enhanced and the Boreal Realm was distinguished from the Paleotethyan Realm during the onset of the Late Palaeozoic Ice Age (LPIA) in the Serpukhovian. The paleolatitude diversity gradient pattern further shows a distinct shift of diversity center from the southern tropic zone in the Tournaisian and Visean to the northern tropic zone in the Serpukhovian.

Evolution of the last koninckinids (Athyridida, Koninckinidae), a precursor signal of the early Toarcian mass extinction event in the Western Tethys

Koninckinids are a suitable group to shed light on the biotic crisis suffered by brachiopod fauna in the Early Jurassic. Koninckinid fauna recorded in the late Pliensbachian–early Toarcian from the easternmost Subbetic basin is analyzed and identified as a precursor signal for one of the most conspicuous mass extinction events of the Phylum Brachiopoda, a multi-phased interval with episodes of changing environmental conditions, whose onset can be detected from the Elisa–Mirabile subzones up to the early Toarcian extinction boundary in the lowermost Serpentinum Zone (T-OAE). The koninckinid fauna had a previously well-established migration pattern from the intra-Tethyan to the NW-European basins but a first phase with a progressive warming episode in the Pliensbachian–Toarcian transition triggered a koninckinid fauna exodus from the eastern/central Tethys toward the westernmost Mediterranean margins. A second stage shows an adaptive response to more adverse conditions in the westernmost Tethyan margins and finally, an escape and extinction phase is detected in the Atlantic areas from the mid-Polymorphum Zone onwards up to their global extinction in the lowermost Serpentinum Zone. This migration pattern is independent of the paleogeographic bioprovinciality and is unrelated to a facies-controlled pattern. The anoxic/suboxic environmental conditions should only be considered as a minor factor of partial control since well-oxygenated habitats are noted in the intra-Tethyan basins and this factor is noticeable only in the second westward migratory stage (with dwarf taxa and oligotypical assemblages). The analysis of cold-seep proxies in the Subbetic deposits suggests a radiation that is independent of methane releases in the Subbetic basin.