The control of burrowing and the migratory behaviour of Donax denticulatus (Bivalvia: Tellinacea)

Donax denticulatus burrows in a similar manner to other infaunal bivalves, but also reacts to stimulation by activity and so proved to be suitable material for experiments on the control of locomotion. Tactile responses of the foot initiate burrowing and pedal stretch receptors control the duration of each digging cycle. Prevention of adduction of the valves, removal of ligament or cardinal hinge teeth, had little effect on burrowing and it is suggested that part of the digging cycle is programmed within the nervous system without peripheral feedback. Experiments on migratory behaviour are discussed and indicate that no intrinsic mechanism is required for its control beyond the adaptation of behavioural responses, common to most bivalves, to different physical conditions.     

Evolutionary morphology of oblique ribs of bivalves

Fossil bivalves bearing oblique ribs first appeared in the Mid Ordovician but their diversity remained low during the Palaeozoic. The diversity soon increased after the Early Triassic, peaking in the Early Cretaceous. The Palaeozoic–Mesozoic record is dominated by burrowing bivalves (mainly pholadomyoids and trigonioids), which developed oblique ribs with symmetric profiles, probably adapted for shell reinforcement, although there are indications that the ribs of trigonioids also enhanced burrowing efficiency. After the Paleocene, the main groups of burrowing bivalves were veneroids (primarily tellinoideans and lucinoideans) and nuculoids, which generated oblique ribs of the shingled type, adapted to increase burrowing efficiency. The inferred change in function at the Mesozoic/Cenozoic boundary can be correlated with an increase in mean mobility of the bivalve faunas bearing oblique ribs through time. This implies a major ecological cause for the observed temporal patterns, which forced bivalve faunas to burrow more rapidly and efficiently. In particular, either the Phanerozoic increase in the diversity of durophagous predators or the accelerating rate of sediment reworking (both being a consequence of the Mesozoic Marine Revolution), or both, could have provided the necessary evolutionary force.     

Comparative taphonomy of three bivalve species from a mass shell accumulation in the intertidal regime of North Sea tidal flats

This study comprises a comparative taphonomic analysis of three endobenthic bivalves (Mya arenaria, Cerastoderma edule, and Macoma balthica) derived from a mass accumulation of mainly vertically packed shells from the “Wurster Watt” in Lower Saxony’s Wadden Sea, German Bight. Bulk samples from two transects were analyzed with respect to taxonomic composition, left/right valve presence, counts and weight percentages of taxa, and size-frequency distributions. Taphonomic features including abrasion, fragmentation, encrustation and bioerosion were subjected to a semi-quantitative analysis. Taphonomic results show significant differences with respect to bivalve taxa as well as between transects. Mya arenaria, a large endobenthic bivalve, shows the greatest amount of fragmentation and is often encrusted by balanids and bryozoans. The smaller and more robust Cerastoderma edule has the greatest values for bioerosion especially by the polychaete Polydora ciliata. Macoma balthica, which has the thinnest valves, shows the highest values for abrasion, but low values for the rest of the measured taphonomic features. Taphonomic differences between the taxa, samples and transects are related to (1) the origin of the bivalves (from living populations or exposed colonization horizons), (2) the different size and morphology of the valves (themselves related to mode of life), (3) the taphonomic trajectories of the different bivalves, as well as (4) the varying depositional environment of the two transects.     

THE MANTLE AND SHELL OF SOLEMYA PARKINSONI (PROTOBRANCHIA: BIVALVIA)

The shell of Solemya exhibits considerable flexibility which is further enhanced by the marked extension of the periostracum beyond the calcareous portions of the valves. This fcature, more than any other, has made possible the habit, unique among bivalves, of burrowing deep within the substrate without direct contact with the water above. The inner calcareous layer of tho valves is restricted to a small area near the umbones while the outer calcareous layer is thin and contains a high proportion of organic material. The shell conchiolin consists mainly of protein, varying in composition, but much of it strengthcned by quinone-tanning, and in ccrtain regions probably by the presence of appreciable quantities of chitin. The ligament, although superficially resembling an amphidetic structure, is opisthodetic, the extcnsion anterior to the umbones consisting of anterior outer layer only.
The mantle is characterized by an extension of the outer fold of the mantle margin which has effected equally both the inner and outer surfaces of this fold. The secretory epithelium and the modified pallial musculature, contraction of which results in the intucking and plaiting of the periostracum, is dcscribed. Simple tubular oil glands open at the mantlo margin and are responsible for the water-repellent nature of the periostracum.
The form of the mantlelshell and that of the enclosed body are discussed and compared with those of other bivalves in which elongation of the mantle/shell is achieved in a different way. It is concluded that the mantlelshell of Solemya is of little value in determining its relationships, and that the greatly elongatod ligament, the edentulous hinge and the flexible shell are all adaptations to a specialized mode of life.     

A Comparative Study of Bivalves Which Bore Mainly by Mechanical Means

This account of the boring mechanisms of those bivalve groupswhich bore mainly by mechanical means attempts to show partlyby reference to published accounts of boring and partly fromour own recent observations of certain characteristics of theboring process in the Pholadidae and Petricolidae, that in contrastto the movements of burrowing forms from which originally allthe boring movements derive, the process of boring makes fewdemands on the hydrodynamic system of the bivalve. The characteristicsof the boring process are closely related to the movements inmodern forms having epifaunal or infaunal habits, supportingthe suggestions of Yonge (1963) concerning the origin of thishabit in the Bivalvia. In all groups in which boringis mechanical,the shell forms the boring tool. However, in those groups inwhich boring has its origin in the epifaunal habit, the majorforce applied to the shell in abrading the burrow isprovidedby contractions of the pedal or byssal retractor muscles. Inthe Adesmacea alone, where boring has been derived from a deepburrowing habit, the adductor muscles provide the major forcein abrasion, and the basic digging cycle has become specializedby the addition of the rocking action of the valves which succeedsretraction. In the former group the ligament is retained andprovides the strong outward force with which the shell is heldagainst the wall of the burrow. In the latter group, the ligamentis reduced, allowing the valves to rock, but here the reciprocalaction of the adductors allows the valves to diverge anteriorlyas the large posterior retractor muscle contracts. In the morespecialized species, water pressure plays a minor role, themaximum pressures recorded being associated with actions subordinateto those involved primarily in abrasion, such as rotation inthe burrow or expulsion of debris from the burrow aspseudofeces.The least specialized borers, such as Petricola, resemble burrowingforms in the importance of the hydrodynamic role of the bodyfluids. In all groups there is a tendency for hypertrophy totake place in the muscles which produce the main boring effect,and for their action to be applied with maximum mechanical advantageagainst a fulcrum provided in most cases by the foot.     

Burrowing criteria and burrowing mode adjustment in bivalves to varying geoenvironmental conditions in intertidal flats and beaches

The response of bivalves to their abiotic environment has been widely studied in relation to hydroenvironmental conditions, sediment types and sediment grain sizes. However, the possible role of varying geoenvironmental conditions in their habitats remains poorly understood. Here, we show that the hardness of the surficial intertidal sediments varies by a factor of 20-50 due to suction development and suction-induced void state changes in the essentially saturated states of intertidal flats and beaches. We investigated the response of two species of bivalves, Ruditapes philippinarum and Donax semigranosus, in the laboratory by simulating such prevailing geoenvironmental conditions in the field. The experimental results demonstrate that the bivalve responses depended strongly on the varying geoenvironmental conditions. Notably, both bivalves consistently shifted their burrowing modes, reducing the burrowing angle and burial depth, in response to increasing hardness, to compensate for the excessive energy required for burrowing, as explained by a proposed conceptual model. This burrowing mode adjustment was accompanied by two burrowing criteria below or above which the bivalves accomplished vertical burrowing or failed to burrow, respectively. The suitable and fatal conditions differed markedly with species and shell lengths. The acute sensitivities of the observed bivalve responses to geoenvironmental changes revealed two distinctive mechanisms accounting for the adult-juvenile spatial distributions of Ruditapes philippinarum and the behavioral adaptation to a rapidly changing geoenvironment of Donax semigranosus. The present results may provide a rational basis by which to understand the ensuing, and to predict future, bivalve responses to geoenvironmental changes in intertidal zones.     

The functional role of burrowing bivalves in freshwater ecosystems

1. Freshwater systems are losing biodiversity at a rapid rate, yet we know little about the functional role of most of this biodiversity. The ecosystem roles of freshwater burrowing bivalves have been particularly understudied. Here we summarize what is known about the functional role of burrowing bivalves in the orders Unionoida and Veneroida in lakes and streams globally. 2. Bivalves filter phytoplankton, bacteria and particulate organic matter from the water column. Corbicula and sphaeriids also remove organic matter from the sediment by deposit feeding, as may some unionids. Filtration rate varies with bivalve species and size, temperature, particle size and concentration, and flow regime. 3. Bivalves affect nutrient dynamics in freshwater systems, through excretion as well as biodeposition of faeces and pseudofaeces. Excretion rates are both size and species dependent, are influenced by reproductive stage, and vary greatly with temperature and food availability. 4. Bioturbation of sediments through bivalve movements increases sediment water and oxygen content and releases nutrients from the sediment to the water column. The physical presence of bivalve shells creates habitat for epiphytic and epizoic organisms, and stabilizes sediment and provides refugia for benthic fauna. Biodeposition of faeces and pseudofaeces can alter the composition of benthic communities. 5. There is conflicting evidence concerning the role of resource limitation in structuring bivalve communities. Control by bivalves of primary production is most likely when their biomass is large relative to the water volume and where hydrologic residence time is long. Future studies should consider exactly what bivalves feed upon, whether feeding varies seasonally and with habitat, and whether significant overlap in diet occurs. In particular, we need a clearer picture of the importance of suspension versus deposit feeding and the potential advantages and tradeoffs between these two feeding modes. 6. In North America, native burrowing bivalves (Unionidae) are declining at a catastrophic rate. This significant loss of benthic biomass, coupled with the invasion of an exotic burrowing bivalve (Corbicula), may result in large alterations of ecosystem processes and functions.     

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.     

The functional morphology of the abyssal Limopsis cristata (Arcoida: Limopsidae) with a discussion on the evolution of the more advanced bivalve foot

The bivalve Limopsis cristata pursues a semi‐endobenthic life in abyssal soft sediments. It attaches to particles by up to three byssal threads and filter feeds by inhaling water from posterior and anterior directions. Because of partial burial, however, only the latter is functionally significant. Complex ciliary currents in the mantle cavity concerned with the rejection of unwanted particles keep most material out of the simplified intestine. It is generally considered that the ligament is the constraining force in arcoid evolution. This may be true in part, but the lack of pallial fusions and the retention of anterior and posterior inhalant flows are more powerful limits to radiation in the Arcoida. In the deep sea, the Limopsidae has radiated into many micro‐niches through micro‐morphological adaptations. Loss of the arcoid ‘heel’ has resulted in the union of the separate rejectory currents of the visceral mass and foot, creating a single discharge point in more advanced bivalves. This greatly simplifies the rejectory roles of the visceral mass and foot and is thus of functional and selective advantage.     

Transport of juvenile clams: effects of species and sediment grain size

Erosion and transport of juvenile benthic invertebrates, including bivalves, have the potential to alter patterns of distribution and abundance during the early post-settlement period. However, the factors influencing rates of postlarval dispersal are not well understood. Both hydrodynamics and behaviour (e.g. burrowing) are likely to play a role in determining patterns of transport of juvenile bivalves. To determine the relationship between sediment transport and bivalve dispersal, experiments were conducted in a racetrack flume to examine the effect of grain size, flow, and clam size on rates of erosion of two species of juvenile clams (Mya arenaria and Mercenaria mercenaria). Results of the experiments were compared to predictions of erosion thresholds based on the physical characteristics of the sediment and clams. Erosion of Mercenaria was greater than Mya, the opposite of predictions based on Mercenaria's greater density, indicating the importance of burrowing behaviour. In most cases, erosion also was greater in the finer sand, in contrast to the predicted similarity of erosion thresholds of the two sediments. However, clam erosion did increase with increasing shear velocity and decrease with clam size, as expected. The results of this study indicate that both hydrodynamics and behaviour play roles in the transport of these two species of juvenile bivalves and that their vulnerability to passive erosion cannot be predicted solely from knowledge of sediment transport.     

A CAUTIONARY TALE: RAZOR SHELLS, ACORN BARNACLES AND PALAEOECOLOGY

Abstract:  The extant, introduced razor shell Ensis americanus (Binney) is a burrowing inhabitant of sandy, shallow-water substrates off the North Sea and Wadden Sea coasts of The Netherlands and adjacent areas. Three articulated shells with broken valves, collected from the strandline at Zandvoort, Noord-Holland, in April 2006 have dense skeletozoan infestations of the barnacle Balanus crenatus Brugière on the outer and inner surfaces of all valves. Such infestations must have occurred after death of the bivalve, decomposition of the soft tissues (but not the ligament) and disinterment of the shells. Larvae of B. crenatus settle in the spring, suggesting that these infestations are perhaps less than a year old, and testifying to the post-mortem persistence of the ligament and the density of skeletozoan infestation after a geologically brief duration. Such specimens would be a palaeoecological conundrum if fossilized; however, the ligament is likely to rot before final burial and the valves break further. Fossil specimens would probably be interpreted as valves that became encrusted on their inner and outer surfaces (balanuliths) during a long post-disarticulation residence on the sea floor.     

The burrowing process of Dentalium (Scaphopoda)

Investigations of the burrowing activity of Dentalium , using cine film and electronic recording techniques, have shown it to penetrate the sand in a series of steps, each termed a "digging cycle". Cycles involve first, pedal dilation, second, retraction followed by extension and probing of the foot. The epipodial lobes are elevated during pedal dilation and form a secure pedal anchor so that at retraction the shell is drawn down over the foot.
A comparison of the burrowing process in the Scaphopoda with that of the Bivalvia indicates that essentially the same mechanisms and sequence of activities are involved, for in both digging consists of the integration of pedal protraction and retraction with the application of shell and pedal anchors. The principal differences, such as the absence in Dentalium of water jets to loosen the sand and high pressures in the pedal haemocoele, are related to the form of their shell. The strength of the pedal anchor was determined and, relative to the weight of Dentalium , is comparable to that of bivalves. In contrast the probing force was relatively weak since the shell anchor of Dentalium , which holds the shell still during probing, is largely limited to its own weight, whereas that attained by the Bivalvia is principally due to the valves being pressed against the substrate by the opening moment of the ligament.     

Biology and functional morphology of the pallial organs of the Antarctic bivalve Mysella charcoti (Lamy, 1906) (Galeommatoidea: Lasaeidae)

Mysella charcoti is an Antarctic lasaeid bivalve and the most frequently encountered mollusc in Admiralty Bay, King George Island, South Shetlands. The behaviour of the species in aquaria, combined with analyses of the gross and microscopic morphologies and functioning of the organs in the mantle cavity of living and preserved specimens have allowed an understanding of important aspects of its biology. The role of the foot and its ciliature during the processes of dislodgement and burrowing within the sediment are described. The species is a free-living, shallow-burrower, with a predominantly deposit-feeding habit and derives part of its food from the labile settled organic deposits carried into the mantle cavity by the anterior–posterior current of water. Pedal sweep-feeding was not detected. M. charcoti is the first known lasaeid with ctenidia formed of the descending lamellae of the inner demibranchs only, a feature probably related to its highly specialised brooding habit.     

Infaunal hydraulics generate porewater pressure signals

Many activities by infauna, including burrowing and feeding, involve hydraulic mechanisms. We expected these activities to generate low-frequency pressure waves that would propagate through sediments and be detectable at some distance from the source. Pressure sensors in intertidal sediments recorded large-amplitude porewater pressure signals. Laboratory recordings of single individuals allowed us to identify characteristic signals of arenicolid and nereidid polychaetes and tellinid bivalves. In the bivalve Macoma nasuta, these high-amplitude signals were associated with burrowing, expulsion of pseudofeces, and siphon relocation. In the polychaetes Neanthes brandti and Abarenicola pacifica, the high-amplitude pressure signals were associated with burrowing, burrow construction, burrow ventilation, and defecation. These signals were detectable in the field at distances of at least 20 cm. Since the waveforms are species-specific as well as activity-specific, they may provide a mechanism for prey detection, for predator avoidance, for competitor detection, and perhaps even for mate detection.     

Multimarker response to salinity stress in two estuarine bivalves of different genetic diversity: Mya arenaria and Limecola balthica from the Gulf of Gdańsk (southern Baltic Sea)

The estuarine bivalves Limecola balthica and Mya arenaria are common inhabitants of marine soft bottom habitats in the Northern Hemisphere. Both species are able to live under a wide range of environmental conditions including variable salinity. However, in L. balthica there is high genetic variability, and populations are often genetically adapted to local conditions. By contrast, genetic diversity in M. arenaria is low across the species’ geographic range, which attests to acclimatization to different conditions. We hypothesized that individuals of M. arenaria should perform better under osmotic stress. We tested this hypothesis by performing a 5‐week experiment that exposed individuals of both clam species to hypo‐ and hyperosmotic conditions. A multiple biomarker approach that included physiological, biochemical, and histological markers was used to assess bivalve performance. Exposure to the different salinities induced biological responses that particularly affected respiratory activity in both species tested, but these responses were much more pronounced in individuals of L. balthica. The results confirmed the hypothesis that the phenotypic plasticity of M. arenaria was more pronounced and reflected a different strategy of adapting to heterogeneous habitats.     

Epibiont habitation patterns and their implications for life habits and orientation among trigoniid bivalves

The bivalve superfamily Trigoniacea has persisted from the Late Paleozoic to the Recent. Late Jurassic and terminal Cretaceous mass extinctions decimated this once-dominant group in shallow marine facies; only a single genus with seven species survives today in the Austral Province. Trigoniacea retain a vestigial byssus and primitive but efficient schizodont dentition. They have been widely considered as infaunal bivalves, burrowing with a very large foot to shallow depths, with inhalant and exhalant apertures at or slightly below the sediment-water interface (SWI). Yet the Trigoniacea are poorly adapted for this life habit. The mantle in living species is unfused and non-siphonate, and some fossil Trigoniacea have permanent shell gapes over these apertures, enhancing the probability of sediment fouling of feeding and respiratory structures. Some living Neotrigonia , e.g., N. margaritacea , solve this problem by having a semi-infaunal life habit, with the inhalant and exhalant apertures elevated above the SWI and the zone of active sediment transport. Semi-infaunal species commonly have epibionts cohabiting the exposed posterior-posteroventral portion of the shell. Numerous well-preserved species of South American Mesozoic Trigoniacea have phototropically and geotropically oriented epibionts on co-attached valves, strongly suggesting a semi-infaunal life mode for at least some members of these taxa. These shell symbionts allow orientation of extinct trigoniid shells relative to the SWI during life, as well as analysis of their depth of burial. Careful analyses of the kinds, size classes, orientation, and dispersion of various epibionts on fossil Trigoniacea thus yield important new information on their life habits, and demonstrate that semi-infaunal life modes were far more common than previously supposed.     

THE BIOLOGY AND FUNCTIONAL MORPHOLOGY OF MYOCHAMA ANOMIOIDES STUTCHBURY, 1830 (BIVALVIA: ANOMALODESMATA: PANDOROIDEA), WITH REFERENCE TO CEMENTATION

The small, exclusively Australasian, anomalodesmatan familyMychamidae comprises only two genera; the shallow-burrowingMyadora and the cementing Myochama. This paper describes theanatomy ad cementing behaviour of Myochama anomioides and drawscomparisons with Myadora. The anatomy of Myochama anomioides is little different fromthat previously described for Myadora, except that they aremirror images. Valve inequality is not reflected in the organsof the mantle cavity in either taxon. Such differences whichare present, for example the reduction of the foot in Myochama,mostly relate to the adoption of a sessile habit. There arefew idfferences in mantle folds of the cementing and non-cementinggenus, except that in M. anomioides the right mantle fold, whichsecretes the cemented valve, is thicker and less well-developedthan the left. During the cementation process, the periostracumsecreted by the right fold is thinner and has a quilted appearance. Individuals of Myochama anomioides cement by their right valveonce they have recahed a size of 1.2—3.9 mm. They appear tohave a preference for attaching to the posterior portions ofa diversity of living, shallow infaunal bivalves. The pronounced stereotypicorientation they adopt suggests that these hosts are most oftenalive at the time of colonization and that the mychamids benefit fromthe relationship. The relationship, however, is not obligate. Theyare capable of attaching to other shelly or rock debris, butdo so at a lager size, presumaby whe the preferred substrataare not available. The thin layer of extra-periostracal cementlacks the calcereous crystalline nature of oyster cement, insteadof being largely composed of organic material. This cement ispresumably secreted by glands within the mantle, but these havenot been identified. Indeed, the mantle lacks arenophilic glandswhich might have been thought a suitable candidate for supplyingcement. (Received 14 December 1999; accepted 4 February 2000)     

A kinetic approach to assess oxidative metabolism related features in the bivalve Mya arenaria

Electron paramagnetic resonance uses the resonant microwave radiation absorption of paramagnetic substances to detect highly reactive and, therefore, short-lived oxygen and nitrogen centered radicals. Previously, steady state concentrations of nitric oxide, ascorbyl radical (A·) and the labile iron pool (LIP) were determined in digestive gland of freshly collected animals from the North Sea bivalve Mya arenaria. The application of a simple kinetic analysis of these data based on elemental reactions allowed us to estimate the steady state concentrations of superoxide anion, the rate of A· disappearance and the content of unsaturated lipids. This analysis applied to a marine invertebrate opens the possibility of a mechanistic understanding of the complexity of free radical and LIP interactions in a metabolically slow, cold water organism under unstressed conditions. This data can be further used as a basis to assess the cellular response to stress in a simple system as the bivalve M. arenaria that can then be compared to cells of higher organisms.     

Pressures generated by the pumping mechanism of some ciliary filter-feeders

An apparatus is described that enables measurements to be made of the low pressures generated by ciliary filter-feeders. Such pressures range from 0.1 to 5.2 mm of water (10 to 520 dyn cm^?2) with the higher pressures in bivalves with long siphons. In these bivalves the frictional resistance of the siphons accounts for a major part of the total resistance to water flow. Pressures observed in bivalves with short siphons, in tunicates, and in sponges are all similar. Hydrodynamical equations have been used to see whether the pressures can be predicted from the dimensions and pumping rate of the bivalve Mya arenaria L.     

Dynamics of burrowing and pedal extension in Donax serra (Mollusca: Bivalvia)

Donax serra burrows powerfully and rapidly in the same manner as other infaunal bivalves except that the turgidity of the foot is increased by a standing pressure of up to 3 kPa in the pedal haemocoel. The standing pressure, upon which probing and adduction pulses are superimposed, is derived from muscular tension acting on the blood in the pedal haemocoel, which is held at near constant volume by the closure of Keber's valve. Blood flow through the foot is thus reduced to a minimum at the time of maximal pedal activity and it is possible that pedal respiration is anaerobic during burrowing.
Extension of the foot is brought about by blood flow and antagonistic musclc action using blood in the pedal haemocoel as the fluid of a classical fluid-muscle system. There is no evidence of the presence of a muscular antagonistic system causing pedal protraction as occurs in the columellar muscle of some cyrtosome molluscs.