Threshold for particle entrainment into suspension

ABSTRACT Laboratory observations regarding the limit conditions for particle entrainment into suspension are presented. A high‐speed video system was used to investigate conditions for the entrainment of sediment particles and glass beads lying over a smooth boundary as well as over a rough bed. The results extend experimental conditions of previous studies towards finer particle sizes. A criterion for the limit of entrainment into suspension is proposed which is a function of the ratio between the flow shear velocity and particle settling velocity. Observations indicate that particles totally immersed within the viscous sublayer can be entrained into suspension by the flow, which contradicts the conclusions of previous researchers. A theoretical analysis of the entrainment process within the viscous sublayer, based on force–balance considerations, is used to show that this phenomenon is related to turbulent flow events of high instantaneous values of the Reynolds stress, in agreement with previous observations. In the case of experiments with a rough bed, a hiding effect was observed, which tends to preclude the entrainment of particles finer than the roughness elements. This implies that, as the ratio between particle and roughness element sizes becomes smaller, progressively higher bed shear stresses are required to entrain particles into suspension. On the other hand, an overexposure effect was also observed, which indicates that a particle moving on a smooth bed is more prone to be entrained than the same particle moving on a bed formed by identical particles.     

Packing and resistance to compaction of shells

The separated valves of bivalve molluscs, certain gastropods, and many brachiopods closely resemble the regular geometrical form known as the shell. The packing concentration of natural accumulations of such organic shells may be placed within bounds with the aid of packing models making use of ordered arrangements of equal geometrical shells, either conical, cylindrical or spherical. The low packing concentrations, comparable with 0·1–0·2, indicated by these models are confirmed by experiments using the shells of four common British species of bivalve or gastropod mollusc. Packings of these shells have a substantial intrinsic strength, and experimentally appear able to support without failing sedimentary overburdens equivalent to loads comparable with 1000 kg mass/m². The observed and predicted low packing concentrations suggests that natural shell beds can hold relatively very large amounts of pore fluids or mineral cements. The resistance to compaction of the packings means that the high original porosities of natural shell beds have a good chance of being permanently preserved.     

Bedload abrasion and the in situ fragmentation of bivalve shells

The aim of this study was to determine how Unio bivalve shells fragment within the channel of the Sakmara River (southern Urals, Russia). The Sakmara River has an abundant bivalve population and a highly variable flow regime which, at low flow, allowed much of the channel bed to be examined. A large data set of 1013 shells (Unio sp.) was examined and these were shown to have consistent patterns of orientation, aspect, shell abrasion, perforation and fracture. The close spatial relationship between areas of shell abrasion, shell perforation and shell fracture showed that they form part of a continuum whereby areas of abrasion evolve into perforations and perforations coalesce and enlarge into fractures. The mechanism of shell damage proposed is one of abrasion in place, whereby the shell remains stationary on the surface of the point bar and is impacted by bedload. Underpinning this process are the hydrodynamic properties of the bivalve shell, with consistency in the orientation and aspect of the valve in a flowing current producing consistency in the distribution of damage on the shell surface. Valves preferentially lie in a convex‐up position and orientate in the flow such that the umbo faces upstream. The elevated, upstream‐facing umbo region is exposed to particle impact and is the first to be abraded and perforated. The vulnerability of the umbo to perforation is greatly increased by the thinness of the shell at the umbo cavity. The in situ abrasion process is enhanced by the development of an armoured gravel bed which restricts valve mobility and maintains shells within the abrasion zone at the sediment–water interface. The in situ abrasion process shows that broken shells are not a reliable indicator of long distance transport. The study also raises the issue that tumbling barrel experiments, which are generally used to simulate shell abrasion, will not replicate the type of directionally focused sand‐blasting which appears to be the principal cause of shell fragmentation in the Sakmara River.     

Settling velocities and entrainment thresholds of biogenic sands (shell fragments) under unidirectional flow

Settling velocities and entrainment thresholds of biogenic sedimentary particles, under unidirectional flow conditions, are derived on the basis of settling tower and laboratory flume experiments. Material consisting predominantly of equant blocks (shell fragments of Cerastoderma edule , density, ρ _s=2800 kg m⁻³) or of mica-like flakes and elongate rods ( Mytilus edulis fragments, ρ _s=2720 kg m⁻³) are used in separate series of experiments. Differences in the measured settling and threshold properties are related primarily to particle shape. The selection of a characteristic length scale for non-spherical grains is investigated by comparing two approaches used to define the grain size ( D ) of the sediment samples: grain settling and sieve analysis that are used to derive data for the threshold criteria, in terms of the Shields and Movability diagrams. The empirical curves effectively predict the threshold conditions for any grain shape, provided that grain size is defined in terms of grain settling velocity. However, a functional distinction is made between the characteristic `hydraulic' grain size, defined by grain settling for grain transport applications, and the actual (physical) grain size defined by sieve analysis.     

Threshold of motion and settling velocities of mollusc shell debris: Influence of faunal composition

Bioclastic particles derived from mollusc shell debris can represent a significant fraction of sandy to gravelly sediments in temperate and cool‐water regions with high carbonate productivity. Their reworking and subsequent transport and deposition by waves and currents is highly dependent on the shape and density of the particles. In this study, the hydrodynamic behaviour of shell debris produced by eight mollusc species is investigated for several grain sizes in terms of settling velocity (measurements in a settling tube) and threshold of motion under unidirectional current (flume experiments using an acoustic profiler). Consistent interspecific differences in settling velocity and critical bed shear stress are found, related to differences in shell density, shell structure imaged by scanning electron microscopy and grain shape. Drag coefficients are proposed for each mollusc species, based on an interpolation of settling velocity data. Depending on the shell species, the critical bed shear stress values obtained for bioclastic particles fall within or slightly below empirical envelopes established for siliciclastic particles, despite very low settling velocity values. The results suggest that settling velocity, often used to describe the entrainment of sediment particles through the equivalent diameter, is not a suitable parameter to predict the initiation of motion of shell debris. The influence of the flat shape of bioclastic particles on the initiation of motion under oscillatory flows and during bedload and saltation transport is yet to be elucidated.     

Metabolic effects on stable carbon isotopic composition of freshwater bivalve shell Corbicula fluminea

The stable isotopic composition of the bivalve shell has been widely used to reconstruct the pa-laeo-climate and palaeo-environment. The climatic and environmental significance of carbon isotopic composition of the bivalve shell is still in dispute, and incorporation of metabolic carbon can obscure carbon isotope records of dis-solved inorganic carbon. This study deals with freshwater bivalve, Corbicula fluminea aragonite shell. The results indicated that the δ13C values of bivalve shells deposited out of equilibrium with the host water and showed an onto-genic decrease, indicating that there are metabolic effects and more metabolic carbon is incorporated into larger shells. The proportion of metabolic carbon of shells varies between 19.8% and 26.8%. However, δ13CS can still be used as qualitative indicators of δ13CDIC and environmental processes that occurred during shell growth.     

The entrainment factor in froth flotation: Model for particle size and other operating parameter effects

Accurately estimating entrainment is crucial when predicting flotation performance as it is essential for determining the concentrate grade achieved. It has been found previously that the amount of gangue entrained is proportional to the water recovery; this proportionality is referred to as the entrainment factor. Experimentally it has been found that entrainment is a strong function of particle size, as well as being dependent on other cell operating parameters such as froth depth and air rate.A simplified theoretical model for entrainment is developed which includes the effects of liquid motion and content, particle settling and particle dispersion. First, a detailed one-dimensional differential model for the entrainment factor is developed and solved numerically. Thereafter, a simplified analytical expression for the entrainment factor is produced which is a good approximation to the more detailed one-dimensional model. Both these models are shown to predict closely experimental trends for entrainment as a function of particle size and froth depth.     

Influences of organic matter and calcification rate on trace elements in aragonitic estuarine bivalve shells

A suite of elements (B, Na, Mg, S, K, Ca, V, Mn, Cr, Sr, and Ba) was measured in aragonitic shells of the estuarine bivalve Corbula amurensis, the Asian clam, using the Sensitive High-Resolution Ion MicroProbe with Reverse Geometry (SHRIMP RG). Our initial intent was to explore potential geochemical proxy relationships between shell chemistry and salinity (freshwater inflow) in northern San Francisco Bay (SFB). In the course of this study we observed variations in shell trace element to calcium ([M]/Ca) ratios that could only be attributed to internal biological processes. This paper discusses the nature and sources of internal trace element variability in C. amurensis shells related to the shell organic fraction and shell calcification rates. The average organic content of whole C. amurensis shells is 19%. After treating whole powdered shells with an oxidative cleaning procedure to remove organic matter, shells contained on average 33% less total Mg and 78% less total Mn. Within our analytical uncertainty, Sr and Ba contents were unchanged by the removal of organic matter. These results show that aragonitic C. amurensis shells have a large component of non-lattice-bound Mg and Mn that probably contribute to the dissimilarity of [M]/Ca profiles among five same-sized shells. Non-lattice-bound trace elements could complicate the development and application of geochemical proxy relationships in bivalve shells. Because B, Ba and Sr occur exclusively in shell aragonite, they are good candidates for external proxy relationships. [M]/Ca ratios were significantly different in prismatic and nacreous aragonite and in two valves of the same shell that had different crystal growth rates. Some part of these differences can be attributed to non-lattice-bound trace elements associated with the organic fraction. The differences in [M]/Ca ratios were also consistent with the calcification rate-dependent ion transport model developed by Carré et al. [Carré M., Bentaleb I., Bruguier O., Ordinola E., Barrett N. T. and Fontugne M. (2006) Calcification rate influence on trace element concentrations in aragonitic bivalve shells: evidences and mechanisms. Geochim. Cosmochim. Acta70, 4906-4920] which predicts that [M]/Ca ratios increase as calcification rates increase and Ca²⁺ channel specificity decreases. This result, in combination with the possibility that there were ontogenetic variations in growth rates among individuals younger than 2 years, underscores the need to develop an independent age model for C. amurensis shells. If growth-rate effects on lattice-bound [M]/Ca ratios can be constrained, it may yet be possible to develop high-resolution geochemical proxies for external solution chemistry in low-salinity regions of SFB.     

Environmental and biological controls on elemental (Mg/Ca, Sr/Ca and Mn/Ca) ratios in shells of the king scallop Pecten maximus

The relationship between potential elemental proxies (Mg/Ca, Sr/Ca and Mn/Ca ratios) and environmental factors was investigated for the bivalve Pecten maximus in a detailed field study undertaken in the Menai Strait, Wales, U.K. An age model constructed for each shell by comparison of measured and predicted oxygen-isotope ratios allowed comparison on a calendar time scale of shell elemental data with environmental variables, as well as estimation of shell growth rates. The seasonal variation of shell Mn/Ca ratios followed a similar pattern to one previously described for dissolved Mn²⁺ in the Menai Strait, although further calibration work is needed to validate such a relationship. Shell Sr/Ca ratios unexpectedly were found to co-vary most significantly with calcification temperature, whilst shell Mg/Ca ratios were the next most significant control. The temporal variation in the factors that control shell Sr/Ca ratios strongly suggest the former observation most likely to be the result of a secondary influence on shell Sr/Ca ratios by kinetic effects, the latter driven by seasonal variation in shell growth rate that is in turn influenced in part by seawater temperature. P. maximus shell Mg/Ca ratio to calcification temperature relationships exhibit an inverse correlation during autumn to early spring (October to March-April) and a positive correlation from late spring through summer (May-June to September). No clear explanation is evident for the former trend, but the similarity of the records from the three shells analysed indicate that it is a real signal and not a spurious observation. These observations confirm that application of the Mg/Ca proxy in P. maximus shells remains problematic, even for seasonal or absolute temperature reconstructions. For the range of calcification temperatures of 5-19 °C, our shell Mg/Ca ratios in P. maximus are approximately one-fourth those in inorganic calcite, half those in the bivalve Pinna nobilis, twice those in the bivalve Mytilus trossulus, and four to five times higher than Mg/Ca ratios in planktonic and benthonic foraminifera. Our findings further support observations that Mg/Ca ratios in bivalve shell calcite are an unreliable temperature proxy, as well as substantial taxon- and species-specific variation in Mg incorporation into bivalves and other calcifying organisms, with profound implications for the application of this geochemical proxy to the bivalve fossil record.     

The settling of consecutive spheres in viscoplastic fluids

One of the factors contributing to the uncertainties involved in the estimation of particle settling velocity in viscoplastic fluids is the time-dependent effect where the viscous parameters of the fluid change as a particle flows through and shears the medium. These changes, particularly at low shear Reynolds numbers, are reflected in the settling velocity of a following sphere that is released some time after an initial one, with the following sphere having a significantly greater velocity. This study found that changes in both fall velocity and equivalent viscosity can be correlated satisfactorily by a power law equation to the dimensionless form of the time interval between releases, and the rheogram shape factor for the fluid. A collision of particles occurs in cases where the time interval between releases is small, after which the particles combine and travel at a terminal velocity. A new variable, β, which takes into account the different surficial stress of the combined spheres, was introduced to the correlation of Wilson et al. [Wilson, K.C., Horsley, R.R., Kealy, T., Reizes, J.A., Horsley, M.R., 2003. Direct prediction of fall velocities in non-Newtonian materials. Int. J. Miner. Process. 71, 17–30] β was found to depend on the rheogram shape factor for the fluid and the shear Reynolds number for the particle. The validity of this approach was supported by experimental data.     

九种现代双壳类壳体物相组成的对比研究

利用X射线衍射分析技术,对黄东海常见的9种现代双壳类壳体进行了物相分析,结果表明这些双壳类壳体分别属于三种类型,即文石质壳、方解石质壳、文石+方解石混合质壳。其中,菲律宾蛤仔(Ruditapesphilippinarum)、中国蛤蜊(Mactrachinensis)、四角蛤蜊(Mactraveneriformis)、薄片镜蛤(Dosinialaminata)、毛蚶(Scapharcasubcrenata)属于文石质壳,长牡蛎(Crassostreagigas)、中国不等蛤(Anomiachinensis)、海湾扇贝(Argopectenirradieus)属于方解石质壳,紫贻贝(Mytilusedulis)属于文石+方解石质的混合质壳;双壳类壳体物相组成与其生活方式有着一定的联系,营底栖埋入式生活者趋于形成文石质壳,底栖固着生活方式者趋于形成方解石质壳或混合质壳;不同大小的菲律宾蛤仔(Ruditapesphilippinarum)的物相组成上几乎没有差别,表明其壳体物相组成不受生命效应的影响。     

考虑钙质砂细观颗粒形状影响的液体拖曳力系数试验

钙质砂作为南海岛礁填筑常用的岩土材料,其渗透性很大程度上决定着填筑后土体的固结和沉降。拖曳力系数是表达流体对土体颗粒表面力的参数,也是表征颗粒状土体渗透能力的一个重要参数,目前国内外对钙质砂拖曳力系数的研究十分有限。首先引入一个修正的三维参数 对钙质砂这种天然非规则颗粒材料的形状进行定量描述,然后开展一系列单个钙质砂颗粒在液体中沉降试验,利用高速相机记录颗粒沉降过程,结合图像处理技术获得颗粒沉降平衡速度Ut,进而计算出拖曳力系数CD和雷诺数Re,最后拟合出包含CD、Re及 三个参数的钙质砂拖曳力系数半经验模型。结果发现,在相同雷诺数条件下钙质砂的形状系数 越大,拖曳力系数越小。通过与其他研究结果对比发现,其表面微孔隙越发育,拖曳力系数越小的规律。该模型能够考虑不规则颗粒形状对拖曳力系数的影响,从而提高对土体渗透性预测的精度,对南海岛礁填筑工程中钙质砂固结和沉降的计算也具有重要意义。     

Stable isotopes and heavy metal distribution in Dreissena polymorpha (Zebra Mussels) from western basin of Lake Erie, Canada

 Dreissena polymorpha is an exotic freshwater bivalve species which was introduced into the Great Lakes system in the fall of 1985 through the release of ballast water from European freighters. Utilizing individual growth rings of the shells, the stable isotope distribution (δ¹⁸O and δ¹³C) was determined for the life history of selected samples which were collected from the western basin of Lake Erie. These bivalves deposit their shell in near equilibrium with the ambient water and thus reflect any annual variation of the system in the isotopic records held within their shells. Observed values for δ¹⁸O range from -6.64 to –9.46‰ with an average value of –7.69‰ PDB, while carbon values ranged from –0.80 to –4.67‰ with an average value of –1.76‰ PDB. Dreissena polymorpha shells incorporate metals into their shells during growth. Individual shell growth increments were analyzed for Pb, Fe, Mg, Mn, Cd, Cu, and V concentrations. The shells show increased uptake of certain metals during periods of isotopic enrichment which correspond with warmer water temperatures. Since metals are incorporated into the shells, the organism may be useful as a biomonitor of metal pollution within aquatic environments. Received: 31 October 1996 · Accepted: 21 May 1997     

The impact of mobile disarticulated shells ofCerastoderma edulis on the abrasion of a cohesive substrate

An annular laboratory flume was used to investigate the effect of mobile cockle shells on the erosion of a cohesive sediment bed. A standard clay bed was created and shells of differing sizes placed upon it. Flow in the flume was increased in increments and the onset of motion and the transport patterns of the cockles were monitored. The release of bed material to the water column was monitored and compared to controls made in the absence of shells (due only to the flow). The shells moved as bedload; first as surface creep (sliding) and then by rolling. The onset velocity of motion (U_c) of the shells was found to be directly related to the settling rate (W_s) in still water. The fluid-induced stresses did not cause any detectable erosion of the bed. The addition of even a single shell induced significant erosion rates (E). The erosion was found to be the result of abrasion rather than corrasion, as the shells never entered into saltation. There was a linear increase in erosion rate with increasing shell size, and an exponential increase in the suspended sediment concentration with time. The drag coefficients (C_d) for settling in traction were calculated. The ratio of the drag forces acting on the shells when settling and moving as traction was found to equal to 1/tan(ф) where ф is the friction angle.     

Stable carbon isotopes in freshwater mussel shells: Environmental record or marker for metabolic activity?

Mussel shells have been used in a number of paleoecological and environmental studies. The interpretation of stable carbon isotopic composition of shell material is still controversial. The carbon for shell carbonate precipitation can either be derived from ambient dissolved inorganic carbon (DIC), with shells recording environmental signals, or from metabolic CO₂, with the potential to disguise environmental signals. To gain insight into this question, we investigated four nearly 100-yr long-term records of aragonite shells from an extant freshwater bivalve species, the endangered freshwater pearl mussel (Margaritifera margaritifera L.). Single growth increments of the outer prismatic and the inner nacreous zones were successfully and easily separated with a simple heat treatment for chronological analyses of δ¹³C in single layers of each zone. Autocorrelation and semivariance statistical methods reveal that mussels show distinct individual signal patterns, which extend up to 25 yr. Signal patterns are reliably reproduced with replicate samples from defined layers within one shell and show similar patterns with a slight offset for inner nacreous and outer prismatic layers for individual animals. Mussels exposed to the same environmental conditions exhibit distinct and contradictory signature patterns, which do not match between individuals. This observation can only be explained by strong metabolic influences on shell precipitation. Environmental changes in pH, temperature, electric conductivity and atmospheric carbon signature had no or little (<5%) influence, whereas body tissue protein and body tissue δ¹³C signatures negatively correlated with the youngest produced shell δ¹³C signatures, indicating that respiration causes a preferential loss of light isotopes from body mass and an inverse enrichment in shell aragonite. Hence, the shells of the freshwater pearl mussel yield a long-term record of metabolic activity, whereas the use of δ¹³C in these shells as recorder for environmental signals is questionable. This may also be true for shells from other species, for which metabolic carbon incorporation has been acknowledged.     

Exploring aberrant bivalve shell ultrastructure and geochemistry as proxies for past sea water acidification

Throughout much of Earth's history, marine carbonates have represented one of the most important geological archives of environmental change. Several pivotal events during the Phanerozoic, such as mass extinctions or hyperthermal events have recently been associated with ocean acidification. Nevertheless, well‐defined geological proxies for past ocean acidification events are, at best, scarce. Here, experimental work explores the response of bivalve shell ultrastructure and isotope geochemistry (δ¹³C, δ¹⁸O and δ²⁶Mg) to stressful environments, in particular to sea water acidification. In this study, the common blue mussel, Mytilus edulis, was cultured (from early juvenile stages to one year of age) at four pH regimes (pH_NBS 7·2 to pH 8·0). Shell growth rate and ultrastructure of mainly the calcitic portion of the shells were compared between experimental treatments. Specimens exposed to low‐pH environments show patches of disordered calcitic fibre orientation in otherwise well‐structured shells. Furthermore, the electron backscattered diffraction analyses reveal that, under acidified conditions, the c‐axis of the calcite prisms exhibits a bimodal or multi‐modal distribution pattern. Similar shell disorder patterns have been reported from mytilids kept under naturally acidified sea water conditions. In contrast, this study found no evidence that different pH regimes affect shell carbon, oxygen or magnesium isotope ratios. Based on these observations, it is proposed that: (i) stressful environments, in this case low sea water pH, predictably affect bivalve biomineralization patterns; and (ii) these findings bear potential as a novel (petrographic) proxy for ancient sea water acidification. An assessment of the applicability of these data to well‐preserved fossil shell material from selected time intervals requires additional work.     

Trace metal bioaccumulation in the shells of mussels and clams at deep-sea hydrothermal vent fields

The bioaccumulation of trace metals in the carbonate shells of mussel and clams was investigated at seven hydrothermal vent fields of the Mid-Atlantic Ridge (Menez Gwen, Snake Pit, Rainbow, and Broken Spur) and the Eastern Pacific (9°N and 21°N at the East Pacific Rise and the southern trough of Guaymas Basin). Mineralogical analysis showed that the carbonate skeletons of the mytilid mussel Bathymodiolus sp. and the vesicomyid clam Calyptogena m. are composed mainly of calcite and aragonite, respectively. The first data were obtained for the content of a variety of elements in the bivalve carbonate shells from various hydrothermal vent sites. The analysis of the chemical compositions (including Fe, Mn, Zn, Cu, Cd, Pb, Ag, Ni, Cr, Co, As, Se, Sb, and Hg) of 35 shell samples and 14 water samples from the mollusk biotopes revealed the influences of environmental conditions and some biological parameters on the bioaccumulation of metals. Bivalve shells from hydrothermal fields with black smokers are enriched in Fe and Mn by a factor of 20–30 relative to the same species from the Menez Gwen low-temperature vent site. It was shown that the essential elements Fe, Mn, Ni, and Cu were more actively accumulated during the early ontogeny of the shells. The high concentration factors of most metals (n × 10²−n × 10⁴) indicate an efficient accumulation function of bivalve carbonate shells. Passive metal accumulation owing to adsorption on the shell surface was estimated to be no higher than 50% of the total amount, varying from 14% for Fe to 46% for Mn.     

A review on bivalve shell,a tool for reconstruction of paleo-climate and paleo-environment

Environmental information can be extracted from bivalve shell geochemistry. In this review, the latest research progress and the existing problems as well as the focus of future research on stable oxygen and carbon isotopic composition and trace elemental ratios of bivalve shell were investigated. Oxygen isotopic signatures of bivalve shell have been proved precipitate in equilibrium with their host water and used as a robust tool to reconstruct ancient water temperature. In order to reconstruct quantitative paleo-temperature information, the future research on bivalve shell oxygen isotope should combine with the study of ancient water oxygen isotope. Metabolic effect pre- vents the direct application of stable carbon isotopic composition of bivalve shell from extracting dissolved inorganic carbon information. The contribution of metabolic carbon to bivalve shell is species specific. Stable carbon isotopic composition of dissolved inorganic carbon could be reconstructed if the metabolic contribution could be accounted for. Explore ways to remove contribution of metabolic effects to bivalve shell is the future focus of the research. There is no consensus on the effect of temperature on bivalve trace element ratios, which may be also species spe- cific, each proxy should ideally be validated and calibrated for each species before being used to interpret past climate and environmental conditions. Future research needs to pay attention to the bivalve shell biomineralization process, which controlled bivalve shell trace element ratios. Cultivation experiment is an important way to obtain the relevant conclusions.     

Adaptive morphologies and guild structure in a high-diversity bivalve fauna from an early Campanian rocky shore, Ivö Klack (Sweden)

The bivalve fauna from a late early Campanian rocky shore at Ivö Klack (southern Sweden), comprises just over sixty species, a very high diversity in comparison to other Late Cretaceous and modern rocky shore bivalve assemblages. This high diversity is here considered to represent a reliable census of the fauna; only in part can it be explained by the cumulative effect of generations of bivalves inhabiting this coastal environment. The high density and diversity and the wide range of shell morphologies allow interpretation of different modes of life in this variable environment with many contrasting habitats. Study of the functional morphology of bivalve shells and comparison with extant relatives has resulted in a subdivision of the fauna into seven guilds and five habitats. The bivalve fauna represents a within-habitat, time-averaged assemblage to which none of the species was introduced from adjacent environments. It includes some of the most northerly known, very small rudistid bivalves, in addition to the oldest known occurrences of Mytilus and Barbatia in association with rocky shores. Bivalves constituted the most important invertebrate group inhabiting the late early Campanian rocky shore at Ivö Klack, in terms of diversity, density and biomass.     

A large metabolic carbon contribution to the δC record in marine aragonitic bivalve shells

It is well known that the incorporation of isotopically light metabolic carbon (C_M) significantly affects the stable carbon isotope (δ¹³C) signal recorded in biogenic carbonates. This can obscure the record of δ¹³C of seawater dissolved inorganic carbon (δ¹³C_DIC) potentially archived in the shell carbonate. To assess the C_M contribution to Mercenaria mercenaria shells collected in North Carolina, USA, we sampled seawater δ¹³C_DIC, tissue, hemolymph and shell δ¹³C. All shells showed an ontogenic decrease in shell δ¹³C, with as much as a 4‰ decrease over the lifespan of the clam. There was no apparent ontogenic change in food source indicated by soft tissue δ¹³C values, therefore a change in the respired δ¹³C value cannot be the cause of this decrease. Hemolymph δ¹³C, on the other hand, did exhibit a negative relationship with shell height indicating that respired CO₂ does influence the δ¹³C value of internal fluids and that the amount of respired CO₂ is related to the size or age of the bivalve. The percent metabolic C incorporated into the shell (%C_M) was significantly higher (up to 37%, with a range from 5% to 37%) than has been found in other bivalve shells, which usually contain less than 10%C_M. Interestingly, the hemolymph did contain less than 10%C_M, suggesting that complex fractionation might occur between hemolymph and calcifying fluids. Simple shell biometrics explained nearly 60% of the observed variability in %C_M, however, this is not robust enough to predict %C_M for fossil shells. Thus, the metabolic effect on shell δ¹³C cannot easily be accounted for to allow reliable δ¹³C_DIC reconstructions. However, there does seem to be a common effect of size, as all sites had indistinguishable slopes between the %C_M and shell height (+0.19% per mm of shell height).