Analysis of the shell torsion phenomenon in the Bivalvia

The curious phenomenon of shell torsion arose independently among members of two separate lineages within the Pteriomorpha (Bivalvia), the Arcidae and the Bakevelliidae. In torted bivalves the shell is twisted about the hinge axis normal to the antero-posterior direction, and the usually two-dimensional plane containing the commissure is deformed into a complex three-dimensional surface. The fabrication of shell torsion is particularly amenable to mathematical analysis, and by utilizing computer simulation it is possible to tort a normal bivalve by introducing 'errors' of a particular nature into the morphogenetic program. Analysis of the pattern of torsion within the Arcidae and Bakevelliidae, and the examination of modem untorted arcids, indicate that the interference of the byssus with the normal bivalve morphogenetic program played a definitive role in the origination of the torsion. Though quite different adaptive pathways were utilized by the two groups, the fabricational principle producing the torsion remained the same. The resultant torted morphologies observed in the Arcidae and Bakevelliidae can be understood in terms of the interaction between the adaptive significance of the torsion and the historical, or preexisting morphology of the animals involved.     

Periostracal mineralization in the gastrochaenid bivalve Spengleria

We investigated the spikes on the outer shell surface of the endolithic gastrochaenid bivalve genus Spengleria with a view to understand the mechanism by which they form and evaluate their homology with spikes in other heterodont and palaeoheterodont bivalves. We discovered that spike formation varied in mechanism between different parts of the valve. In the posterior region, spikes form within the translucent layer of the periostracum but separated from the calcareous part of the shell. By contrast those spikes in the anterior and ventral region, despite also forming within the translucent periostracal layer, become incorporated into the outer shell layer. Spikes in the posterior area of Spengleria mytiloides form only on the outer surface of the periostracum and are therefore, not encased in periostracal material. Despite differences in construction between these gastrochaenid spikes and those of other heterodont and palaeoheterodont bivalves, all involve calcification of the inner translucent periostracal layer which may indicate a deeper homology.     

FIRST RECORD OF A HETERODONT BIVALVE (MOLLUSCA) FROM THE EARLY TRIASSIC: PALAEOECOLOGICAL SIGNIFICANCE AND IMPLICATIONS FOR THE 'LAZARUS PROBLEM'

Abstract:  The heterodont bivalve Sinbadiella pygmaea gen. et sp. nov. is described from the Sinbad Limestone Member (Olenekian, Smithian) of the Lower Triassic Moenkopi Formation of Utah (USA). The new taxon is tentatively assigned to the Lucinidae on the grounds of its external morphology, larval shell morphology and hinge characters. It is the only definite representative of the Superorder Heterodonta in the Olenekian, removing this major clade of bivalves from the long list of Early Triassic Lazarus taxa. Although at least locally very abundant, the new taxon has been overlooked by earlier workers, probably due to its minute size. Dwarfism is a widespread phenomenon in Early Triassic faunas, which has been attributed to ongoing reduction of primary production. If S .  pygmaea was a lucinid, a chemosymbiotic feeding mode can be inferred, which might explain why it was able to thrive in conditions that suppressed filter feeding heterodonts.     

A molecular phylogeny of bivalve mollusks: ancient radiations and divergences as revealed by mitochondrial genes

Background

Bivalves are very ancient and successful conchiferan mollusks (both in terms of species number and geographical distribution). Despite their importance in marine biota, their deep phylogenetic relationships were scarcely investigated from a molecular perspective, whereas much valuable work has been done on taxonomy, as well as phylogeny, of lower taxa.

Methodology/Principal Findings

Here we present a class-level bivalve phylogeny with a broad sample of 122 ingroup taxa, using four mitochondrial markers (MT-RNR1, MT-RNR2, MT-CO1, MT-CYB). Rigorous techniques have been exploited to set up the dataset, analyze phylogenetic signal, and infer a single final tree. In this study, we show the basal position of Opponobranchia to all Autobranchia, as well as of Palaeoheterodonta to the remaining Autobranchia, which we here propose to call Amarsipobranchia. Anomalodesmata were retrieved as monophyletic and basal to (Heterodonta + Pteriomorphia).

Conclusions/Significance

Bivalve morphological characters were traced onto the phylogenetic trees obtained from the molecular analysis; our analysis suggests that eulamellibranch gills and heterodont hinge are ancestral characters for all Autobranchia. This conclusion would entail a re-evaluation of bivalve symplesiomorphies.     

Interaction strength between a predator and dangerous prey: Sinistrofulgur predation on Mercenaria

The lack of direct empirical evidence of predator evolution in response to prey adaptation is a fundamental weakness of the arms race analogy of predator-prey coevolution. I examined the interaction between the predatory busyconine whelk Sinistrofulgur sinistrum and its bivalve prey Mercenaria mercenaria to evaluate whether reciprocal adaptation was likely in this predator-prey system. Thick-lipped whelks use their shell lip to chip open the shell of their prey, often resulting in breakage to their own shell. Thus, hard-shelled prey, such as Mercenaria, may be considered dangerous because they are able to inflict damage to the predator as a consequence of the interaction. The strength of interaction between whelks and their bivalve prey was viewed by regressing predator performance (the incidence of shell breakage in encounters with prey) on prey phenotype (a function of size). Interaction with Mercenaria of varying sizes has strong and predictable consequences (r²=0.946; p=0.028) for Sinistrofulgur. Predators that select large, thick bivalve prey increase the likelihood that their shell lip will be broken in the process of attempting to open their prey. Ecological consequences of feeding-induced breakage may include reduced growth rate, reproductive success, and survivorship. These results suggest that natural selection should favor predator phenotypes that reduce feeding-induced breakage when interactions with damage-inducing prey occur.     

Sphaeriid and corbiculid clams represent separate heterodont bivalve radiations into freshwater environments

Nine families of bivalve molluscs have undergone successful radiations in freshwater habitats, including three heterodont taxa: the Sphaeriidae, Corbiculidae, and Dreissenidae. Although the phylogenetic relationships of these freshwater heterodont families are controversial, most workers place the first two in the superfamily Corbiculoidea and assume that they represent a monophyletic grouping. We have tested competing phylogenetic hypotheses for the Corbiculoidea by constructing a representative molecular phylogeny, based on domains D1-D3 of the nuclear large subunit 28S rDNA, for 18 heterodont bivalves and for two oyster outgroup taxa. Our results do not support the monophyly of the Corbiculoidea and are consistent with the hypothesis that all three families of freshwater heterodonts represent independent colonization events by marine ancestors. Similarities in developmental mode specializations exhibited by some sphaeriids and corbiculids, such as sequential direct-developing broods, represent convergent adaptations to the freshwater environment. The corbiculid taxa form a clade with venerid and mactrid outgroups but we were not able to identify a putative marine outgroup for the sphaeriids.     

On bivalve phylogeny: a high-level analysis of the Bivalvia (Mollusca) based on combined morphology and DNA sequence data

Abstract. Bivalve classification has suffered in the past from the crossed-purpose discussions among paleontologists and neontologists, and many have based their proposals on single character systems. More recently, molecular biologists have investigated bivalve relationships by using only gene sequence data, ignoring paleontological and neontological data. In the present study we have compiled morphological and anatomical data with mostly new molecular evidence to provide a more stable and robust phylogenetic estimate for bivalve molluscs. The data here compiled consist of a morphological data set of 183 characters, and a molecular data set from 3 loci: 2 nuclear ribosomal genes (18S rRNA and 28S rRNA), and 1 mitochondrial coding gene (cytochrome c oxidase subunit I), totaling ∼3 Kb of sequence data for 76 molluscs (62 bivalves and 14 outgroup taxa). The data have been analyzed separately and in combination by using the direct optimization method of Wheeler (1996), and they have been evaluated under 12 analytical schemes. The combined analysis supports the monophyly of bivalves, paraphyly of protobranchiate bivalves, and monophyly of Autolamellibranchiata, Pteriomorphia, Heteroconchia, Palaeoheterodonta, and Heterodonta s.l., which includes the monophyletic taxon Anomalodesmata. These analyses strongly support the conclusion that Anomalodesmata should not receive a class status, and that the heterodont orders Myoida and Veneroida are not monophyletic. Among the most stable results of the analysis are the monophyly of Palaeoheterodonta, grouping the extant trigoniids with the freshwater unionids, and the sister-group relationship of the heterodont families Astartidae and Carditidae, which together constitute the sister taxon to the remaining heterodont bivalves. Internal relationships of the main bivalve groups are discussed on the basis of node support and clade stability.     

Phylogenetic implications and diagenetic stability of macromolecules from pleistocene and recent shells of Mercenaria mercenaria (mollusca,bivalvia)

The phylogeny and diagenesis of Pleistocene and Recent bivalves were studied immunologically by use of a conventional antiserum elicited against an EDTA‐soluble macromolecular extract from shells of the modern bivalve mollusc Mercenaria mercenaria. ELISA tests of the antiserum with shell fragments of a wide range of modern bivalves gave taxonomically significant results. The antiserum reacted with palaeoheterodonts and heterodonts but not with representatives of other bivalve subclasses. This phylogenetic reactivity was also apparent in tests with fossil shells, although the specificity and overall strength of the reaction were both reduced. Absorption of the antiserum with etched shell powders of various (palaeo)heterodonts yielded more specific antibody preparations.

Investigations of shell matrix diagenesis, using the anti‐Mercenaria serum, demonstrated that small amounts of original determinants could be detected even in fossils over one million years old. The reactivity of the serum with extracts of fossil Mercenaria decreased with sample age. The relationship between serum reactivity and the degree of amino acid racemization was almost linear. Clearly, the various determinants to which antibodies were elicited were being destroyed at different rates.     

Modelling various sculptures in the Cretaceous bivalve Inoceramus hobetsensis

The geometry of the external shell sculpture in the Late Cretaceous inoceramid bivalve Inoceramus hobetsensis Nagao & Matsumoto, 1939 was studied both empirically and theoretically. A large sample, collected from the Upper Cretaceous of Hokkaido, Japan, shows remarkably high intraspecific variation in the shell sculptural pattern. Quasi-commarginal ribs, slightly oblique to the external growth increments, occur in some specimens. These sculptures are commonly irregular in strength and spacing, and their features are successfully modelled by computer simulations when the commarginal ribs are superposed with nearly concentric divaricate rib. Computer models indicate that the divergent sculpture element, often found in other inoceramids, was present throughout the evolution of I. hobetsensis and was developing in the evolutionary lineage from I. hobetsensis nonsulcatus to I. hobetsensis hobetsensis, although it was only weakly expressed. The results also suggest that some apparently distinct sculptural patterns of I. hobetsensis are the result of minor changes in the morphogenetic program.     

淡水贝类贝壳多层构造形成研究

对几种淡水贝（包括蚌、螺）进行形态及组织学观察，并通过实验方法重现贝壳三种物质，即：角质、棱柱质、珍珠质的生成过程，结果表明：外套膜外表皮细胞是由相同类型细胞组成，这些相同细胞在不同的作用条件下形成贝壳多层构造。     

Effects of plasma from bivalve mollusk species on the in vitro proliferation of the protistan parasite Perkinsus marinus

The in vitro culture of the Eastern oyster parasite Perkinsus marinus has provided a unique opportunity to examine its susceptibility to putative recognition and effector defense mechanisms operative in refractory bivalve species. In this study, we report the effect of supplementing the culture medium with plasma from: (1) uninfected to heavily infected Eastern oysters; (2) oyster species considered to be disease-resistant; and (3) bivalve mollusk species that are naturally exposed to the parasite but show no signs of disease. We also examined in vitro the interaction between hemocytes from Crassostrea virginica and C. gigas and P. marinus trophozoites. Our results revealed a significant decrease (32%) in proliferation of P. marinus in the presence of plasma from heavily infected C. virginica oysters. The inhibitory effects were less pronounced with plasma from moderately infected and uninfected oysters. In contrast, plasma from C. rivularis and C. gigas enhanced P. marinus proliferation. Proliferation was significantly reduced in media supplemented with plasma from Mytilus edulis, Mercenaria mercenaria, and Anadara ovalis. The highest inhibitory activity was apparent in M. edulis, for which 5% plasma-supplemented medium reduced growth by 35% relative to the controls. M. edulis active component(s) was heat-stable, yet pronase-sensitive. The significantly higher uptake of live P. marinus trophozoites by hemocytes from C. virginica, relative to those from C. gigas, suggests a certain level of specificity in the recognition/endocytosis of the parasite by its natural bivalve host species.     

Barbatia mytiloides and the evolution of shell torsion in arcid pelecypods

Barbatia mytiloidrs (Brocchi) (Arcidae, Arcinae) from the Neogene of northwestern Italy shows a variable but significant amount of dextral shell torsion about the hinge axis. This phenomenon represents a morphological and functional convergence with the sinistrally torted ark Trisidos . Polyphyletic evolution of shell torsion in mud-dwelling, semi-infaunal descendants of epibyssate Barbatia is probably due to ventilation problems that forced these forms to lie with the posterior commissure horizontal. Depending on which valve happened to lie lowermost in the non-torted reclining or semi-infaunal ancestors, shell torsion became established in dextral or sinistral direction.     

Effects of the gastropod,Ilyanassa obsoleta(Say) and the bivalve,Mercenaria mercenaria (L.), on larval settlement and juvenile recruitment of infauna

Densities of a deposit-feeding gastropod, Ilyanassa obsoleta(Say) and a suspension-feeding bivalve, Mercenaria mercenaria (L.), were manipulated to test the importance of interactions between adults and settling larvae in affecting abundance patterns of infaunal invertebrates. Abundance of postlarvae and juveniles was significantly lower in the presence of Ilyanassa obsoleta than in snail-free controls. I. obsoleta caused a reduction in abundance of as much as 45%. Surface-dwelling tubiculous polychaetes, gastropods, harpacticoid copepods, and mobile subsurface-dwelling infauna all declined in abundance with increasing I. obsoleta density. Mercenaria mercenaria had little or no effect upon larval settlement. The effect of Ilyanassa obsoleta and Mercenaria mercenaria combined was equivalent to the sum of the effects of each species separately. Disturbance and ingestion of newly settled larvae by Ilyanassa obsoleta, larval settlement selectivity, and disturbance-induced emigration are probable mechanisms causing differences in abundance between I. obsoleta treatments.     

Articulated external fixation of the ankle: minimizing motion resistance by accurate axis alignment

This study describes how an optimal single hinge axis position can be established for the application of articulated external fixation to the ankle joint. By deliberately introducing various amounts of relative mal-alignment between the optimal talocrural joint axis and the actual fixator hinge axis, it was possible to measure the corresponding amounts of additional resistance to joint motion. In a cadaveric study of six ankle specimens, we determined the instant axis of rotation of the talocrural joint from 3-D kinematic data. acquired by an electromagnetic motion tracking system. For each specimen, an optimal fixator hinge position was calculated from these motion data. Compared to the intact natural joint, aligning the fixator along the optimized axis position caused a moderate increase in energy (0.14 J) needed to rotate the ankle through a prescribed plantar/dorsiflexion range. However, malpositioning the hinge by 10 mm caused more than five times that amount of increase in motion resistance. While articulated external fixation with limited internal fixation can establish a favorable environment for the repair of severe injuries such as tibial pilon fractures, the large additional resistance to motion accompanying a malpositioned fixator axis suggests the development of untoward intra-articular forces that could act to disturb fragment alignment.     

Intermediary metabolism and shell growth in the brachiopod Terebratalia transversa

Juvenile Terebratalia transversa (Brachiopoda) metabolize carbohydrates in the anterior-most marginal mantle at a rate of 0.46 μM glucose/g/hr (in vitro incubation of mantle in C¹⁴-glucose in a carrying medium of 10^-3 M non-radioactive glucose). The rate declines to 0.18μM glucose/g/hr in full-grown specimens. Carbohydrate metabolism in the marginal (anterior-most) mantle averages approximately 3.7 times greater than metabolism in (a portion of the ‘posterior’) mantle situated between the coelomic canals and the marginal mantle. This ratio remains constant in specimens of all sizes (i.e. an ontogenetic trend in the ratio is absent at p≤ 0.05). Organic acids are not detectable within the mantle (HPLC techniques) even after simulated anoxia (N₂ bubbling during mantle incubation). Glucose metabolism in vitro declines in both the marginal and ‘posterior’ mantles during anoxia and the metabolic ratio between marginal/‘posterior’ mantles becomes 1/1. We found no difference (at p≤ 0.05) in mean metabolic activity or in sue-related metabolic trends among populations from depths ranging between mean sea level and 70 m. However, the activity within the ‘posterior’ mantle was more variable in specimens from 70 m than in those from shallower habitats (10 m - mean sea level). The size of the specimens analyzed was most variable in the groups obtained from the shallowest habitats and least variable at 70 m depth. Our results may help define the energetics of fossil as well as living brachiopod shell growth. Brachiopod shell growth is known to be very slow relative to that of bivalves and our results indicate that this is a result of the animals' slow metabolism. The inflation of the valves in T. transversa is, in part, a function of the high ratio of intermediary metabolism in the marginal vs‘posterior’ mantle (i.e. parallels the relative growth rates at the shell margin vs‘posterior’ areas). We found that the bivalve, Chlamys hastata, which is commonly associated with T. transversa, has a lower ratio of metabolic activities in the ventral/dorsal mantle areas than the brachiopod has in the anterior/posterior. The difference produces a flatter shell in the bivalve in accord with allometric principles. The higher metabolic rate in the marginal vs‘posterior’ brachiopod mantle and its more pronounced decline with anaerobiosis is reflected in the greater definition of growth increments in the outer shell layer. Our results do not support recent generalizations that correlate shell thickness of a wide variety of invertebrates inversely with metabolic rate. Growth rate as determined from width of shell growth increments is a better index of metabolic rate. Although the genetic basis of glucose metabolism is unknown, the observed metabolic variability is consistent with suggestions that populations of marine organisms living in stable offshore environments are genetically more variable but morphologically more uniform than populations from shallow water. Furthermore, our results support suggestions that bivalved molluscs and brachiopods are very different metabolically, but the data are neutral with respect to theories of competitive exclusion of the two taxa throughout geologic history.     

Sublittoral attached epifaunal development in Buzzards Bay,Massachusetts

Summary Sixteen epifaunal species are identified encrusting dead valves of Mercenaria mercenaria, Mya arenaria, and Aequipecten irradians exposed for one year on the bottom of Buzzards Bay, Massachusetts.The three most common cheilostomatous bryozoans are Schizoporella unicormis, S. biaperta, and Parasmittina trispinosa. Balanus (Balanus) amphitrite neveus demonstrates a preferred orientation of the rostro-carinal axis parallel to the growth lines of Mercenaria mercenaria, Mya arenaria and Aequipecten irradians.No preference for difference bivalve shells is apparent in any of the epifauna examined. The lesser abundance of most species on Mercenaria mercenaria valves may be due to the more rapid burial of these heavier valves in shifting sediments of the study area.     

A clockwork mollusc: Ultradian rhythms in bivalve activity revealed by digital photography

Time-lapse digital images can be acquired and archived using web-cameras, allowing non-invasive analysis of behavior patterns of bivalve molluscs at ultradian (sub-daily) time-scales over long intervals. These records can be analyzed directly by a human operator or through properly calibrated image analysis software. Preliminary results using species of marine and freshwater bivalves identify several ultradian biological rhythms of similar duration. Wavelet analysis indicates strong periodicity in mantle and siphon activity in the 3 to 7 min range, with longer duration shell contraction periods at 60-90 min. The recurrence of these rhythms among marine and freshwater bivalve species maintained under constant (but differing) conditions suggests the influence of common intrinsic drivers (chemico-physical mechanisms or biological clocks). Sub-daily growth increments preserved in the shells of rapidly growing bivalve species are potentially related to these biological rhythms, with implications for shell growth, biomineralization, and the temporal resolution of paleoclimate proxy data.