Biomechanics and energy cost of the amphipod Corophium volutator filter-pump

The integrated function of the setal filter-basket and the pleopodal pump in the burrowing amphipod Corophium volutator was studied by video-microscopy in order to evaluate the energy costs of filter feeding. Video-microscope observations indicated that, in general, nine short, water-pumping beat cycles of the pleopods are succeeded by one slow cycle that coincides with cleaning of the setal filter and transient slowdown of inhalant water velocity. The position of the plumose setal filter on the second pair of gnathopods ensures that all water runs through the filter-basket. The fine V-shaped bristles on the setae enlarge the total filter area so that the velocity of water flowing through the filter is relatively slow, about 2.5 mm s(-1), giving rise to a resistance of about 2.9 mm H(2)O, which is the most important contribution to the total pressure drop in the system. In "standard" individuals of C. volutator with lengths of 3 and 6 mm, the normal operating pump pressure and pumping rate were, respectively, 2.6 and 3.1 mm H(2)O, and 18.3 and 85.5 ml h(-1); the overall pump efficiencies were 5.1% and 11.6%, respectively. These results show that the Corophium filter-pump is comparable to other low-pressure biological pumps in filter-feeding marine invertebrates, such as mussels, polychaetes, ascidians, and bryozoans.     

Surface deposit feeding versus filter feeding in the amphipod Corophium volutator

The previously indicated ability of the amphipod Corophium volutator to switch between deposit feeding and filter feeding was confirmed and studied in more detail in controlled laboratory experiments in which filtration rate measurements were combined with simultaneous video recordings of surface-feeding activity of the amphipod exposed to different known concentrations of algal (Tetraselmis sp.) cells. When algal cells were added to the ambient water, this stimulated C. volutator, buried in natural sediment or transferred to glass tubes, to commence filter feeding, which was maintained as long as the algal concentration was kept above a certain threshold level. However, shortly after the algal concentration was grazed below the threshold level, filter feeding was abandoned and replaced by surface deposit feeding, as evident from a video observed increase in surface scraping frequency. The average frequency of surface scraping was 0.64±0.27 min^-1, with a residence time of 3.7±1.4 s on the sediment surface where the amphipod grabbed material within a semicircle. Such detailed knowledge of filter feeding versus deposit feeding in C. volutator is of importance for a better understanding of the ecological role of this key organism in many shallow-water ecosystems where the feeding conditions are frequently changing.     

Parasites on an intertidal Corophium-bed: factors determining the phenology of microphallid trematodes in the intermediate host populations of the mud-snail Hydrobia ulvae and the amphipod Corophium volutator

The phenology of microphallid trematodes within their intermediate hostpopulations has been studied on an intertidal mud flat. The parasites usethe mud snail Hydrobia ulvae and the infaunal amphipod Corophium volutatoras first and secondary intermediate host, respectively. Migratory shorebirdsact as final hosts. Our results show a general trend of decline in thedensity of infected intermediate hosts during both spring and autumn, whichcould mainly be ascribed to shorebird predation. During summer the densityof both infected snails and infected amphipods increased considerably, witha culmination in June within the snail population (1000 infectedm-² and in August within the amphipod population (40 000infected m-². This time lag in parasite occurrence could berelated to (1) the development time of larval trematodes within the snails,(2) higher ambient temperatures in late summer increasing parasitetransmission between snails and amphipods during this period, and (3) ageneral increase in the Corophium population during late summer. Fromsamples collected between 1990 and 1995 it is shown that microphallidtrematodes occasionally may give rise to mass mortality in the amphipodpopulation. The prerequisites for such an event are a high parasiteprevalence within the first intermediate host population and unusually highambient temperatures, facilitating parasite transmission to the secondaryintermediate host, C. volutator.     

Genetic isolation of populations of the gammaridean amphipod, Corophium volutator, in the Bay of Fundy, Canada

Selection experiments suggest that evolutionary modifications in amphipod demography can respond to local environmental changes and that local races of amphipods may be common. We tested this hypothesis in mudflat populations of Corophium volutator in the Bay of Fundy, Canada. Due to the unique topography of the Bay of Fundy, distinctive environmental conditions are prevalent in different branches of the Bay, while the impact of shorebird predation has also been shown to vary between populations. Several hypotheses have been suggested to explain ecological evidence which indicates that Corophium volutator (Pallas), a common amphipod crustacean, exhibits extensive life history variation in Bay of Fundy populations. We used RAPD-PCR techniques to examine populations of C. volutator in an investigation of genetic isolation in marine environments. Our data suggest that variation in selection pressures have played a significant role in the genetic divergence of populations of C. volutator in the Bay of Fundy.     

The influence of Corophium volutator abundance on resuspension

Two experiments were performed to test the hypothesis that Corophium volutator affects the turbidity of water in estuaries through active resuspension of sediment. One experiment was done in a flume system under different flow velocities, and one in aquaria. A diatom film developed at the sediment surface in both experiments before Corophium was added. This diatom film was supposed to have a stabilising effect on the sediment. In both experiments, the concentration of suspended solids in the water column increased with the density of Corophium individuals. No effect of flow velocity on suspended solids concentration was found. This indicates that, in our flume experiment, active resuspension by Corophium was more important than physical resuspension, at least at low flow velocity (<20 cm s^-1) and in the presence of a diatom film. The critical erosion threshold decreased with increasing Corophium density in the aquarium experiment, indicating that indirect effects of Corophium grazing may become more important at high levels of bottom shear stress. The implications of our findings for suspended solids concentration in estuarine systems are discussed.     

Functional morphology of the stomach of Corophium volutator Pallas

Corophium volutator Pallas is a small amphipod crustacean which burrows in inter-tidal mud on the British coast, and feeds on organic detritus, mainly vegetable, by selecting particles from the mud.
The alimentary canal consists of foregut, midgut and hindgut. The midgut is produced into a pair of anterior dorsal caeca, a pair of ventral caeca and a pair of posterior dorsal caeca.
The cardiac stomach has a large number of chitinous plates or ridges beset with hooks and spines for the trituration of food.
The pyloric stomach has long fine bristles which form an eifective filter apparatus and allow only fine particles of food to pass into the midgut, where they are digested and absorbed.
The hepato-pancreas secrete digestive enzymes and store reserve food material as oil globules.     

The behaviour of Corophium volutator (Crustacea: Amphipoda)

The behaviour of Corophium volutator (Pallas) is outlined. Swimming, crawling, burrowing and feeding activities are described in detail. Animals usually swim on their backs. Every few seconds, swimming alternates with passive sinking. Animals can crawl over surfaces in and out of water. Out of water they do so by a looping motion using their second antennae and telson. When out of water animals crawl away from light and down slopes. In water they swim towards light. Burrowing is initiated by rapid beating of the pleopods; the animal then sinks below the surface by a concerted action of pereiopods, pleopods, telson, uropods and second antennae; within a few minutes, a shallow burrow is formed. The formation of permanent burrows is dependant on particle size, on adhesive properties of detritus and primary films on sand particles, and on a secretion produced by the animal itself. Individuals can turn about in permanent burrows. The species is essentially a detritus feeder. Animals normally feed only when in their burrows, by using their second antennae to scrape material from the substrate surface into the entrance of the burrow. This material is then transported to the mouth by the feeding appendages and respiratory current. The behaviour of small and large animals differs; small animals burrow rapidly and permanently, and do not emerge spontaneously; furthermore, they only swim occasionally. Large animals swim more frequently, spend more time on the substrate surface, and periodically move burrows. It is suggested that new habitats are colonized by large animals which have already bred once.     

Behavioral Response of Corophium volutator to Shorebird Predation in the Upper Bay of Fundy,Canada

Predator avoidance is an important component of predator-prey relationships and can affect prey availability for foraging animals. Each summer, the burrow-dwelling amphipod Corophium volutator is heavily preyed upon by Semipalmated Sandpipers (Calidris pusilla) on mudflats in the upper Bay of Fundy, Canada. We conducted three complementary studies to determine if adult C. volutator exhibit predator avoidance behavior in the presence of sandpipers. In a field experiment, we monitored vertical distribution of C. volutator adults in bird exclosures and adjacent control plots before sandpipers arrived and during their stopover. We also made polymer resin casts of C. volutator burrows in the field throughout the summer. Finally, we simulated shorebird pecking in a lab experiment and observed C. volutator behavior in their burrows. C. volutator adults were generally distributed deeper in the sediment later in the summer (after sandpipers arrived). In August, this response was detectably stronger in areas exposed to bird predation than in bird exclosures. During peak predator abundance, many C. volutator adults were beyond the reach of feeding sandpipers (>1.5 cm deep). However, burrow depth did not change significantly throughout the summer. Detailed behavioral observations indicated that C. volutator spent more time at the bottom of their burrow when exposed to a simulated predator compared to controls. This observed redistribution suggests that C. volutator adults move deeper into their burrows as an anti-predator response to the presence of sandpipers. This work has implications for predators that feed on burrow-dwelling invertebrates in soft-sediment ecosystems, as density may not accurately estimate prey availability.     

Tube-building in two epifaunal amphipod species,Corophium insidiosum andJassa falcata

Jassa falcata andCorophium insidiosum are epifaunal tube-building marine amphipods, whose niches overlap in habitat and food requirements. Laboratory experiments were conducted to study the influence of the quality of different available particulate substrates on settlement and tube-building behaviour of these two amphipod species. Our experiments suggest thatC. insidiosum is less specialized in this respect thanJ. falcata. C. insidiosum is able to use organic materials for tube-building such as artificial (Ulva spec. powder) or mixed natural detritus as well as inorganic material (coarse sand); whereasJ. falcata utilizes organic materials, but sand only to a very limited extent.     

Reproductive synchrony in captive macaques

Captive bonnet macaques housed at the California Primate Research Center reproduce seasonally. The chances of producing surviving infants were substantially reduced among females who conceived at the peak of the mating season compared with females whose conceptions were more isolated in time. Primiparous females and low ranking females were most consistently affected by the extent of reproductive synchrony. Behavioral data suggest that harassment of conceiving and pregnant females may have contributed to the correlation between the extent of reproductive synchrony and infant mortality as the levels of aggression toward females were highest during the months in which conceptions were most common.     

Behaviour and time allocation of the mud shrimp, Corophium volutator, during the tidal cycle: a laboratory study

Despite the numerous ecological and behavioural studies, relatively little is known about the behavioural dynamics of Corophium volutator during the tidal cycle. In the present study, the behaviour of C. volutator was observed in the laboratory in a tide-simulated aquarium, and time allocation of the different activities was quantified. Overall, the surface activity was low and showed a steep decline after submersion. A clear tide-based cyclic pattern was found, with a boost of activity immediately after first submersion, continuing during submersion, and shifting over a period with decreasing activity towards almost total inactivity at the end of emersion.     

Reproductive synchrony in a recovering bottlenecked sea turtle population

1. The assessment of species extinction risk has been well established for some time now. Assessing the potential for recovery in endangered species is however much more challenging, because complementary approaches are required to detect reliable signals of positive trends. 2. This study combines genetics, demography and behavioural data at three different time-scales to assess historical and recent population changes and evidence of reproductive synchrony in a small population of olive ridley sea turtle Lepidochelys olivacea. Lepidochelys is considered as the most extraordinary example of reproductive synchrony in reptiles, yet to date, it has only been reported in large populations. 3. Using Bayesian coalescent-based models on microsatellite nuclear DNA variability, we demonstrate that effective population size in olive ridleys nesting in French Guiana has dramatically declined by 99% over the last 20 centuries. This low current population size is further illustrated by the absence of genetic mitochondrial DNA diversity in the present nesting population. Yet, monitoring of nesting sites in French Guiana suggests a possible recovery of the population over the last decade. 4. Satellite telemetry shows that over the first 14 days of their 28-days inter-nesting interval, i.e. when eggs maturation is likely to occur, gravid females disperse over the continental shelf. They then gather together with a striking spatiotemporal consistency close to the nesting site, where they later emerge for their second nesting event. 5. Our results therefore suggest that reproductive synchrony also occurs in small populations. Olive ridleys may ensure this synchrony by adjusting the duration of the second half of their inter-nesting interval prior to landing, possibly through social mediation. 6. Such reproductive synchrony may be related to the maintenance of some species-specific strategy despite former collapse and may contribute to the present population recovery. The gregarious behaviour of reproductive individuals close to shore where human-induced perturbations occur is however a cause for conservation concern for this still poorly known species.     

Reproductive competition and the evolution of extreme birth synchrony in a cooperative mammal

Reproductive events in animal societies often show a high degree of temporal clustering, but the evolutionary causes of this synchronization are poorly understood. Here, we suggest that selection to avoid the negative effects of competition with other females has given rise to a remarkable degree of birth synchrony in the communal-breeding banded mongoose (Mungos mungo). Within banded mongoose groups, births are highly synchronous, with 64 per cent of females giving birth on exactly the same night. Our results indicate that this extreme synchrony arises because offspring suffer an increased risk of infanticide if their mother gives birth before other females, but suffer in competition with older littermates if their mother gives birth after them. These findings highlight the important influence that reproductive competition can have for the evolution of reproductive synchrony.     

Intense episodic predation by shorebirds may influence life history strategy of an intertidal amphipod

Life history theory suggests that selective predation on older or larger individuals in prey populations or a disturbance such as intense and episodic predation should lead to smaller size at maturity, and a tendency toward semelparity. Many populations of the intertidal amphipod, Corophium volutator , in the western Atlantic are subjected to an intense period of size-selective predation for about one month in summer, during the southward migration of shorebirds. We compared size at maturity and fecundity of populations of C. volutator from mudflats that are intensively used by shorebirds with populations that are visited by very few birds. We found that mature females were of similar size in May, but those from bird mudflats produced more offspring during the first reproductive episode. In July, females of the summer generation began to reproduce at a smaller size on bird mudflats, and as a consequence, produced fewer offspring that grew more slowly. The results of this correlative study suggest that shorebird predation has shaped C. volutator life history in two ways. First, females on bird mudflats concentrate their reproductive effort into a larger early brood, probably because later broods would come to maturity during the period of intense predation. Second, in summer, amphipods begin to reproduce at a smaller size so as to produce a brood before the arrival of the birds in mid-July. Predators are not present at the time that these responses occur, and thus they are probably not a plastic response to perceived predation risk. Thus the patterns we observed agree with the predictions of recent theories: individuals faced with a predictable disturbance alter their life cycles so as to minimize the effect of that disturbance.     

Reproductive synchrony,parental investment,and the evolutionary dynamics of sexual selection

Computer simulations showed that reproductive synchrony in the greater investing sex can evolve as a strategy for increasing the investment of mates. The potential for synchrony to increase in frequency for this reason decreased with (a) increasing population size, (b) decreasing importance of synchrony-enforced investment, (c) increasing variance in quality among members of the lesser investing sex, and (d) decreasing male: female sex ratio. By reducing the payoffs to desertion, synchrony may result in the evolution of uniquely effective forms of parental care by the lesser investing sex, making desertion unprofitable regardless of the availability of potential mates. Thus relative levels of optimal parental investment are subject to change through the action of sexual selection itself.     

Acute toxicity testing with the European estuarine amphipod <Emphasis Type="Italic">Corophium multisetosum</Emphasis>

This study reports on the use of the estuarine amphipod Corophium multisetosum in acute toxicity testing. The species was successfully acclimated to the laboratory and was used in a water-only whole effluent 96 h acute bioassay and in a 10 days whole estuarine sediment test. C. multisetosum response was compared to other species in 96 h bioassays, testing boiling cork effluent and iron filings lixiviates. The amphipod showed high sensitivity and the results were similar to those obtained with others species namely, the freshwater cladoceran Daphnia magna, the estuarine amphipod Gammarus chevreuxi and the seawater annelid Sabellaria alveolata. In a 10 days static exposure to natural impacted estuarine sediments, the response of the species was compared to a fertilization bioassay with the sea-urchin Paracentrotus lividus. Both species indicated the same sediment samples as the most potentially toxic. These samples were collected in sites where the resident macrofauna benthic community is also the most affected, with strong reduction of the species richness, abundance and biomass. The results revealed that C. multisetosum presents high potential to be used in routine acute toxicity testing in the estuarine environment.     

Stressed but stable: canopy loss decreased species synchrony and metabolic variability in an intertidal hard-bottom community

The temporal stability of aggregate community properties depends on the dynamics of the component species. Since species growth can compensate for the decline of other species, synchronous species dynamics can maintain stability (i.e. invariability) in aggregate properties such as community abundance and metabolism. In field experiments we tested the separate and interactive effects of two stressors associated with storminess--loss of a canopy-forming species and mechanical disturbances--on species synchrony and community respiration of intertidal hard-bottom communities on Helgoland Island, NE Atlantic. Treatments consisted of regular removal of the canopy-forming seaweed Fucus serratus and a mechanical disturbance applied once at the onset of the experiment in March 2006. The level of synchrony in species abundances was assessed from estimates of species percentage cover every three months until September 2007. Experiments at two sites consistently showed that canopy loss significantly reduced species synchrony. Mechanical disturbance had neither separate nor interactive effects on species synchrony. Accordingly, in situ measurements of CO(2)-fluxes showed that canopy loss, but not mechanical disturbances, significantly reduced net primary productivity and temporal variation in community respiration during emersion periods. Our results support the idea that compensatory dynamics may stabilise aggregate properties. They further suggest that the ecological consequences of the loss of a single structurally important species may be stronger than those derived from smaller-scale mechanical disturbances in natural ecosystems.     

Reproductive synchrony and annual variation in foetal growth rate in the long-fingered bat (Miniopterus schreibersii)

Mating in the long-fingered bat (Miniopterus schreibersii) from the East Cape region of South Africa occurred during a four-week period in April and May, and implantation was delayed until August. Analyses of foetal age and estimated dates of conception indicate that after implantation, the progression of pregnancy within the population was synchronous both within and between years. It is suggested that implantation, probably initiated by increasing day length, occurred at about the same time throughout the population and thus synchronized reproduction. In the final month of pregnancy foetal growth rate and, consequently, the timing of parturition in the three years of the study varied. These variations were correlated with variations in rainfall and it is suggested that the failure of the November rains in 1990 would have caused a reduction in insect abundance and that this was responsible for the decrease in foetal growth rate and the delay in parturition.