Reproductive synchrony in the intertidal amphipod Corophium volutator

Following logic of the mate-availability hypothesis, females are expected to show asynchronous reproduction in those species where operational sex ratios are female-biased and under circumstances where an individual female is sexually receptive only for short durations. We show that females of the intertidal amphipod Corophium volutator are receptive to mating only for a few days following their moult and are unable to hasten onset of moulting in the presence of a male. Despite meeting the conditions of the mate-availability hypothesis, reproduction was synchronous for female C. volutator across spatial and temporal scales relevant to mate-searching abilities of males. As such, some females are not expected to mate between moults, which coincide with their ability to mate. However, females do moult frequently (relative to males) which should increase their likelihood of mating over their lifetimes. It is unlikely that seasonal constraints, predation, or competition can account for the high degree of synchrony among breeding female amphipods. We suggest that dispersal of females or their offspring may constrain activity of females, as they moulted almost entirely during spring tides (although not always during the same set of spring tides). Female reproductive synchrony also has implications for reproductive behaviour of males, in particular, the possibility of harem-defence polygyny.     

Reproductive behavior of intersexes of an intertidal amphipod Corophium volutator

Abstract. Intersexes are common in crustaceans. Typically, these intersexes are sterile or function as females, but prior evidence from laboratory experiments suggests that intersexes of a key species of gammaridean amphipod, Corophium volutator , might function as males. We observed that intersexes of C. volutator behaved as males by crawling (mate-searching) on a mudflat during ebb tides and pairing in burrows with female amphipods. In the laboratory, intersexes and males did not differ in aspects of crawling such as movement rate and measures of burrow investigation. I`ntersexuality was costly in that intersexes crawled less often than males on a mudflat, formed fewer pairs with females than males, and remained in tandem less often with receptive females than males. The use of PCR-based identification methods failed to identify the presence of transovarial, feminizing, microsporidian parasites as a major cause of intersexuality in this species in that infected females did not produce broods that contained more intersexes than broods produced by uninfected females. Because intersexes may be mistaken as females, the percentage of functional males in amphipod populations may be underestimated: an important consideration given male limitation in populations of C. volutator. The occurrence of intersexes has significant implications for studies on the evolution and ecology of sex ratios, and the use of crustaceans as indicators of environmental quality.     

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.     

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.     

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.     

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.     

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.     

Laboratory Studies of Parasite Transmission Aspects between Hydrobia spp. (Gastropoda) and Corophium volutator (Amphipoda)

Aspects of parasite transmission between Hydrobia spp. and Corophium volutator, first and second intermediate host of digenetic trematodes, were investigated under laboratory conditions. H. ventrosa is used as an intermediate host by several trematode species. Under laboratory conditions the most frequently observed emergence from H. ventrosa was of cercariae of Maritrema subdolum. The number of cercariae shed per day varied considerably. It was observed that 30 cercariae on average and up to 450 cercariae at maximum can emerge from a single H. ventrosa per day. Cercarial production continued until the death of the snails. The life-span of cercariae of the species M. subdolum decreased as the water temperature increased. It can be concluded that under natural conditions the cercariae, after emerging, have a maximum period of 1 day in which to seek out their second intermediate host C. volutator. Almost all specimens of C. volutator exposed to cercariae of the species M. subdolum died within the test period of 6 days. High average cercarial densities caused short life-spans (<50 h), while at lower densities longer survival times were possible. For cercariae of other Trematoda species, we were unable to find any equally clear evidence of a reduction in the survival rate of C. volutator within the test period. Mortality of C volutator, and other effects of infestation, as observed in our experiment, can be assumed to be a result of the penetration process of the cercariae but they also can be attributed to the mesocercariae.     

A laboratory-based comparison of osmoregulatory ability at different water temperatures and salinities in the stenothermic isopod Saduria entomon (Linnaeus, 1758) and the eurythermic amphipod Corophium volutator (Pallas, 1766) from the Baltic Sea

The aim of the investigation was a comparison of osmoregulatory ability at different water temperatures and salinities by the stenothermic isopod Saduria entomon (Linnaeus, 1758) and the eurythermic amphipod Corophium volutator (Pallas, 1766) from the Baltic Sea. The experiments were performed under laboratory conditions at different water temperatures (5, 10 and 15?°C) and salinities (3, 6.6, 15 and 25 PSU for S. entomon and 3, 6.3, 15 and 25 PSU for C. volutator). The osmotic concentrations of the crustaceans’ haemolymph were determined using the melting point method. Temperature had a statistically significant effect on the osmoregulation in S. entomon, but had generally no statistical influence on the osmoregulation in C. volutator. This physiological information regarding adaptation provides a basis for predicting the distribution of these species in changing environments. The implications of the results for the comparative adaptability of the two species are considered.     

Anthropocene invasion of an ecosystem engineer: resolving the history of Corophium volutator (Amphipoda: Corophiidae) in the North Atlantic

Resolving the natural histories of species is important for the interpretation of ecological patterns, as it provides evolutionary context for the interactions between organisms and their environment. Despite playing an integral role on the intertidal mudflats of the North Atlantic as an abundant food source for predators and as an ecosystem engineer that alters the soft sediment environment, no previous studies have provided empirical evidence to determine the biogeographical origin of the amphipod Corophium volutator. To resolve its status as introduced or indigenous in Europe and North America, we analyzed sequence data for two mitochondrial loci and two nuclear markers, aiming to determine whether the present range of C. volutator is the result of unresolved taxonomy, persistence in glacial refugia, natural trans‐Atlantic dispersal, or human‐mediated introduction. Our results demonstrate a reduced genetic diversity in North American populations that is a subsample of diversity in European populations, with coalescent analyses of mitochondrial and nuclear DNA supporting different models of multiple introductions from Europe to the Bay of Fundy and Gulf of Maine in North America. These results suggest that C. volutator was introduced to North America prior to the first surveys of local biota in the 20th Century, which has broad implications for interpretations of community and ecosystem interactions in the North Atlantic intertidal. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 288–304.     

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.     

Rain increases the energy cost of bat flight

Similar to insects, birds and pterosaurs, bats have evolved powered flight. But in contrast to other flying taxa, only bats are furry. Here, we asked whether flight is impaired when bat pelage and wing membranes get wet. We studied the metabolism of short flights in Carollia sowelli, a bat that is exposed to heavy and frequent rainfall in neotropical rainforests. We expected bats to encounter higher thermoregulatory costs, or to suffer from lowered aerodynamic properties when pelage and wing membranes catch moisture. Therefore, we predicted that wet bats face higher flight costs than dry ones. We quantified the flight metabolism in three treatments: dry bats, wet bats and no rain, wet bats and rain. Dry bats showed metabolic rates predicted by allometry. However, flight metabolism increased twofold when bats were wet, or when they were additionally exposed to rain. We conclude that bats may not avoid rain only because of sensory constraints imposed by raindrops on echolocation, but also because of energetic constraints.     

The spring-mass model and the energy cost of treadmill running

During running, the behaviour of the support leg was studied by modelling the runner using an oscillating system composed of a spring (the leg) and of a mass (the body mass). This model was applied to eight middle-distance runners running on a level treadmill at a velocity corresponding to 90% of their maximal aerobic velocity [mean 5.10 (SD 0.33) m · s⁻¹]. Their energy cost of running (C _r ), was determined from the measurement of O₂ consumption. The work, the stiffness and the resonant frequency of both legs were computed from measurements performed with a kinematic arm. The C _r was significantly related to the stiffness (P < 0.05, r = −0.80) and the absolute difference between the resonant frequency and the step frequency (P < 0.05, r = 0.79) computed for the leg producing the highest positive work. Neither of these significant relationships were obtained when analysing data from the other leg probably because of the work asymmetry observed between legs. It was concluded that the spring-mass model is a good approach further to understand mechanisms underlying the interindividual differences in C _r . Accepted: 18 August 1997     

Energy cost and energy sources in karate

Energy costs and energy sources in karate (wado style) were studied in eight male practitioners (age 23.8 years, mass. 72.3 kg, maximal oxygen consumption (VO_2max) 36.8 ml · min^–1 · kg^–1) performing six katas (formal, organized movement sequences) of increasing duration (from approximately. 10 s to approximately 80 s). Oxygen consumption (VO₂) was determined during pre-exercise rest, the exercise period and the first 270 s of recovery in five consecutive expired gas collections. A blood sample for lactate (la^–) analysis was taken 5 min after the end of exercise. The overall amount of O₂ consumed during the exercise and in the following recovery increased linearly with the duration of exercise (t) from approximately 1.51 (for t equal to 10.5 s (SD 1.6)) to approximately 5.81, for t equal to 81.5 s (SD 1.0). The energy release from la^– production (VO_21a ^–) calculated assuming that an increase of 1 mmol · l^–1 la^– corresponded to a VO₂ of 3 mlO₂ · kg^–1 was negligible for t equal to or less than 20 s and increased to 17.3 ml · kg^–1 (la^– = 5.8 mmol · l^–1 above resting values) for t equal approximately to 80 s. The overall energy requirement (VO_2eq) as given by the sum of VO₂ and VO_2la ^– was described by VO_2eq = 0.87 + 0.071 · t (n = 64; r ² = 0.91), where VO_2eq is in litres and t in seconds. This equation shows that the metabolic power (VO_2eq · t ^–1) for this karate style is very high: from approximately 9.51 · min^–1 for t equal to 10 s to approximately 4.91 · min^–1 for t equal to 80 s, i.e. from 3.5 to 1.8 times the subjects' VO_2max. The fraction of VO_2eq derived from the amount of O₂ consumed during the exercise increased from 11% for t equal to 10 s to 41 % for t equal to 80 s whereas VO_21a ^– was negligible far t equal to or less than 20 s and increased to 13 % o for t equal to 80 s. The remaining fraction (from 90% for t equal to 10 s to 46% for t equal to 80 s), corresponding to the amount of O₂ consumed in the recovery after exercise, is derived from anaerobic alactic sources, i.e. from net splitting of high energy phosphates during the exercise.