Reproductive effort in Danish mudsnails (Hydrobiidae)

Summary The four species of Hydrobiid mudsnails found in Danish brackish water habitats are ranked with respect to reproductive effort. The biology of the species indicate that Hydrobia neglecta and Potamopyrgus jenkinsi, showing the highest reproductive effort are also more r-selected than the other species, H. ventrosa and H. ulvae.     

Character displacement and coexistence in mud snails (Hydrobiidae)

Summary Populations of coexisting and of allopatrically occurring species of hydrobiid snails (Hydrobia ulvae, H. neglecta, H. ventrosa and Potamopyrgus jenkinsi) have been studied in 90 localities within three different areas. When H. ventrosa coexists with H. ulvae they show character displacement, i.e., the average body size of the former is smaller and that of the latter is larger. When these species live alone they are of approximately the same size. It is shown here that the size ratio between the coexisting species usually found (1.3–1.5) allows stable coexistence based on food praticles size selectivity alone. Variation in the degree of character displacement from locality to locality is explained by different degrees of genetical isolation of the populations. Coexisting H. ventrosa and H. ulvae have shorter, more well-defined periods of reproduction than they do when they occur alone. H. neglecta is larger than coexisting H. ventrosa and smaller than coexisting H. ulvae. Due to the patchy distribution and the fluctuating populations of this species, and due to the fact that pure H. neglecta populations are rare, data on this species are difficult to interpret. Potamopyrgus jenkinsi shows a different food particle size selection than the Hydrobia spp. of identical sizes. Competitive interactions between P. jenkinsi and coexisting Hydrobia spp. are therefore probably weak. In accordance with this, P. jenkinsi does not show character displacement when coexisting with, e.g., H. ventrosa.The fact that the major study area, the Limfjord, is only 150 years old as a marine habitat as well as various more recent man-made changes of the coast line allow estimates of the time scale of the microevolutionary changes which lead to character displacement.     

THE EARLY LIFE-CYCLE STAGES OF HYDROBIA VENTROSA AND HYDROBIA NEGLECTA WITH OBSERVATIONS ON POTAMOPYRGUS JENKINSI

The egg mass of Hydrobia neglecta contains a single egg, whilethat of H. ventrosa contains up to three eggs. At hatching H.neglecta has a significantly greater shell length than H. ventrosaand the late embryos and young snails of the two species canbe separated according to the pigmentation of the head region.The young snails can also be separated on the basis of surfaceornamentation of the shell and comparisons with the closelyrelated H. ulvae and Potamopyrgus jenkinsi show how the basicpattern of shell sculpturing differs in the four species. Thesedifferences are discussed in relation to the mode of development. (Received 14 February 1980;     

The effect of different primary producers on Hydrobia ulvae population dynamics: a case study in a temperate intertidal estuary

The effect of macroalgal blooms and the consequent disappearance of Zostera noltii meadows on Hydrobia ulvae population dynamics and production was studied in the Mondego estuary based on data obtained from January 1993 to September 1995. Sampling was carried out at a non-eutrophicated area, covered with Z. noltii, and also at an eutrophicated area, where seasonal Enteromorpha spp. blooms occur.Stable populations represented by individuals of all age classes were found only at the Z. noltii meadows throughout the study period. On the contrary, at the eutrophicated area, during most of the time, solely juveniles were present, with adults appearing only during the macroalgal bloom (>1.5 mm width).During the algal bloom (e.g. 1993), H. ulvae population density was clearly higher in the eutrophicated area due to the combined effect of stronger benthic recruitments (99% of veliger larvae newly recruited) and dispersion of juveniles proceeding from the Z. noltii meadows to this area. On the other hand, in the absence of macroalgae (spring of 1994), 98.9% of veliger larvae was recruited in the Z. noltii meadows. Therefore, H. ulvae seems to respond rapidly to macroalgal dynamics and its presence at the eutrophicated area depends on the existence of green macroalgae.H. ulvae presented the same benthic recruitment pattern at the two sampling areas, with new cohorts being produced in March, June, July and September. Depending on the time of the year in which the recruitment took place, cohorts showed different growth rhythms. However, after 12 months they reached a similar size.A three-generation life cycle involving a short-lived (16 months), fast growing spring generation, a medium growing (17-19 months) summer generation and a longer-lived (20 months) slower growing generation that overwinters is identified.As a general trend, productivity and mean population standing biomass were higher at the Z. noltii meadows, during the entire study, except for a short period, during the macroalgal bloom, when production was higher at the eutrophicated area. On the contrary, P/B? ratios were higher at the eutrophicated area. According to our results, H. ulvae population structure and yearly productivity are clearly affected by eutrophication, namely by the dynamics of macroalgal blooms.In the long run, we may infer that, following the disappearance of the Z. noltii meadows, due to eutrophication, H. ulvae would also tend to disappear, since reproductive adults were almost exclusively found in this area.     

Ingestion rate of the deposit-feeder Hydrobia ulvae (Gastropoda) on epipelic diatoms: effect of cell size and algal biomass

The effects of cell size of epipelic diatoms and sediment Chl a content (as an index of algal biomass) on the ingestion rate of Hydrobia ulvae adults and juveniles were investigated in experimental microcosms. Results showed that both adults and juveniles ingested small and large diatoms without exhibiting cell size selection behaviour. The functional response of H. ulvae, juveniles (<4 mm) and adults (>4 mm), over a wide range of sediment Chl a content, was characterized by an increase of the ingestion rate according to a power law. Ingestion rate varied from 0.75 to 10 ng Chl a ind⁻¹ h⁻¹ for juveniles and from 1 to 52 ng Chl a ind⁻¹ h⁻¹ for adults, in the range ca. 10-100 μg Chl a (g dry weight sediment)⁻¹. The ingestion rate was about three times higher for adults than for juveniles. Based on these experimental results, we further proposed a mechanistic approach, using an individual based-model, to identify simple feeding mechanisms that might be involved in H. ulvae functional response.     

Environmental and behavioural factors affecting activity in the intertidal gastropod Hydrobia ulvae

Laboratory microcosms were used to investigate the mud snail Hydrobia ulvae (Pennant) bioturbation activities and behavioural changes in response to snail density, algal food, sediment moisture content, light regime and water cover conditions. Density-dependent kinetics of bioturbated muddy areas were described by von Bertalanffy equations, which provided reliable estimates of mud surface covering rates by snail tracks (m² h⁻¹ snail⁻¹). Snails need a wet habitat to be active either covered by seawater or by moving in fluid layers for low-tide conditions. Light and microphytobenthic biomass, which are less potent to affect snail activity, are positively interrelated to increase covering rates in the tested chl a concentrations within the range of 1-15 μg g⁻¹. Experimental results suggested us the relevance of microphytobenthos migration processes in affecting crawling activities of H. ulvae that appeared to adjust their foraging efforts in response to benthic algal biomass. Behavioural processes of H. ulvae, in terms of floating, crawling, burrowing and inactive snails, were described using a Markov model. Finally, an empirical model based on von Bertalanffy equations was proposed to describe kinetics of sediment covering by snail tracks under the influences of snail density, sediment moisture content, chl a concentrations and the four combinations of presence/absence of light and seawater. This model should provide a base for further development of a hydrosedimentary model to simulate the effects of H. ulvae bioturbation activities on the resuspension of the intertidal cohesive sediment-water interface for various in situ conditions.     

The feeding biology of Hydrobia ventrosa (Montagu). I. The assimilation of different components of the food

The assimilation efficiencies for 7 species of diatoms, two of blue-green algae, and one bacterium have been measured in the deposit-feeding prosobranch Hydrobia ventrosa (Montagu). Two methods have been used, a chemical and a radiotracer one.The results of the first method were difficult to evaluate due to the fact that the time taken to clear the gut is ill-defined. The second method gave more consistent results.All diatoms are assimilated efficiently (60–71 %) as is the bacterium (about 75 %). The blue-green algae are less efficiently utilized; an Oscillatoria sp. was assimilated with an efficiency of about 50 % whereas a Chroococcus sp. was only assimilated with an efficiency of 8 %. It is concluded that resource partitioning based on differential utilization of micro-organisms cannot play an important rôle in the co-existence of Hydrobia ventrosa with other deposit-feeders.     

Peculiarities of the Structure of and Trends in the Macrobenthos Community of the Ermolinskaya Bay Lagoon Ecosystem,Kandalaksha Bay,White Sea

The species composition and spatial and trophic structures of the macrobenthos communities of the Ermolinskaya Bay were studied, and the main changes that had occurred since previous explorations (the 1960s) were analyzed. The most significant changes in the spatial structure of the macrobenthos community (species diversity, population density and biomass, and similarities of the species’ distribution) were observed with regard to the sea level zone: the most pronounced at the upper littoral, followed by the middle littoral, lower littoral, and sublittoral. The longitudinal gradient of the community structure (from the innermost area seaward) was pronounced less strongly. Altogether, four macrobenthos communities can be identified in Ermolinskaya Bay; they differ in species composition, diversity, population density and biomass, and trophic structure: (1) community of the upper littoral with the dominance of collecting deposit feeders, Hydrobia ulvae, and scrapers, Littorina saxatilis; (2) community of the middle littoral with the prevalence of deposit feeders, Hydrobia ulvae and Macoma balthica, seston feeders, Mya arenaria, and, to a lesser extent, Mytilus edulis; (3) community of the lower littoral with the domination of the filtering organisms, Mytilus edulis, to a lesser extent collecting deposit feeders, Hydrobia ulvae and Macoma balthica, and scrapers, Littorina littorea; and (4) community of the sublittoral, where the leading positions belong to the group of collecting deposit feeders, Macoma balthica, to a lesser extent also Hydrobia ulvae (in the innermost area) and Capitella capitata, Arenicola marina (in the marine part of the bay). The gradual siltation of the bay and the reduction of its connection to the sea have led to the development of a littoral complex of species in the sublittoral, whereas the species typical at the sublittoral in the 1960s are now mainly found at the outlet of the bay.     

Uptake of glycine and release of primary amines by the polychaete Nereis virens (Sars) and the mud snail Hydrobia neglecta muus

Uptake of glycine and release of primary amines by the polychaete Nereis virens (Sars) and the mud snail Hydrobia neglecta Muus was investigated. In a flow system, Nereis virens had a net influx of glycine of 32 to 39 nmol·g^?1 fresh wt·h from a 5-μM solution of glycine. Simultaneously, an efflux of primary amines of 14 nmol·g^?1·h^?1 was measured. The experiments indicated that glycine uptake and release of primary amines are two independent processes. Glycine was absorved by Hydrobia neglecta from 50-μM solutions at an influx of 175 nmol·g^?1 organic wt·h^?1, and simultaneously, there was an independent efflux of 25 nmol primary amine·g^?1·h^?1. In the experiments, released primary amines from both species increased the ambient concentration of primary amines, but after various periods of time these concentrations apparently stabilized. Finally, the significance of micro-organisms in amino-acid uptake experiments is considered.     

ORGANIC CARBON, ENVIRONMENTAL FACTORS AND GROWTH IN THE SMALL EURYHALINE GASTROPOD, POTAMOPYRGUS JENKINSI (SMITH)

A method of analysing small invertebrates for organic carbon(in the range 6.0 to 1000 µg) is described. The relationshipsbetween organic carbon content, total dry weight and shell lengthwere detemined for Potamopyrgus jenkinsi living in differentcombinations of salinity and temperature. Differences in organiccarbon content were found in relation to weight at differentgrowth temperatures. This work is a further confirmation ofthe usefulness of organic carbon content as a measure of thegrowth or biomass of small invertebrates. ^*Present Address: School of Biology, University College, P.M.B.5323, Port Harcourt, Nigeria. (Received 27 April 1977;     

Temperature and salinity tolerances of larval Californian grunion, Leuresthes tenuis (Ayres): A comparison with gulf grunion, L. Sardina (Jenkins & Evermann)

Temperature and salinity tolerances were determined for larval California grunion, Leuresthes tenuis (Ayres), and compared with previous data for Gulf of California grunion, L. sardina (Jenkins & Evermann). Larvae of similar age and acclimation history showed little interspecific difference in thermal tolerance, as measured by half-hour LT₅₀ values for 20–30 day old late postlarvae acclimated at various temperatures, and by upper and lower incipient lethal temperatures for 18°C-acclimated prolarvae. The upper incipient lethal temperature differed by 1 deg.-C (32°C for L. tenuis, 31°C for L. sardina), while the lower incipient lethal temperature for the 18°C acclimated prolarvae of both species was 7.5°C. L. tenuis larvae were much less euryhaline than L. sardina, with incipient lethal salinities of 4.2–41 %. for prolarvae and 8.6–38 %. for 20-day-old postlarvae; comparable values for L. sardina are 4–67.5 %. and 5–57.5 %. Both species show a decrease in temperature and salinity tolerance with age. The larvae of these disjunct congeners show a significant physiological divergence in euryhalinity but not in overall temperature tolerance. These tolerances are discussed in relation to the respective geographic ranges and behavioral responses of the two species.     

Predation of the shelduck Tadorna tadorna on the mud snail Hydrobia ulvae

Many waders and ducks rely exclusively on invertebrates for their food and can deplete populations of these invertebrates. The mudsnail Hydrobia ulvae and the shelduck Tadorna tadorna are both widespread and abundant and in previous studies H. ulvae has been identified as the main food source for shelduck. The objective of the study was to quantify the predation of shelduck of H. ulvae on an intertidal mud flat in north Wales in order to estimate the contribution of shelduck predation to H. ulvae mortality and to estimate the importance of H. ulvae in the diet of shelduck. The contribution of shelduck towards the observed mortality was estimated from the number of H. ulvae per dropping, the number of droppings produced per hour and the number of shelduck found at the study site. Shelduck predation hardly made a contribution to the observed mortality of H. ulvae. Examination of the energetic requirements of shelduck shows that H. ulvae cannot be a significant part of the diet. In conclusion, the predation of shelduck on H. ulvae was irrelevant for both the population dynamics of the mud snail and for the energy requirements of shelduck in the study area.     

Post-feeding thermophily in lizards (Lialis burtonis Gray,Pygopodidae): Laboratory studies can provide misleading results

Many ectothermic vertebrates raise their preferred body temperature after feeding, likely expediting digestion. However, most studies documenting this phenomenon have relied upon laboratory thermal gradients, which grossly oversimplify an animal's environment. We explored the validity of thermal gradient methodology by investigating post-prandial thermophily in an Australian pygopodid lizard (Lialis burtonis Gray). Mean body temperatures did not differ between fed and unfed lizards in field enclosures. Feeding influenced body temperature in a thermal gradient, but in opposite directions depending on details of the methodology. When we introduced L. burtonis into the gradient at the warmer end, fed lizards had higher body temperatures than unfed conspecifics. However, the opposite was true when lizards were introduced at the cooler end. These contrasting results indicate that lizards with food in their stomachs did not seek out higher temperatures, but instead were more sedentary than unfed lizards. Our study highlights the need for caution in interpreting thermal gradient results unaccompanied by field data, and it demonstrates how minor changes in equipment design or procedures can significantly alter conclusions from laboratory studies.     

Effect of population density and presence of a potential competitor on the growth rate of the mud snail Hydrobia ulvae (Pennant)

The mud snail Hydrobia ulvae (Pennant) and the amphipod Corophium arenarium Crawford were used in a series of laboratory and field experiments to investigate the hypothesis that natural populations of such deposit-feeders live under conditions of resource limitation. Growth rates of Hydrobia confined at several different densities within cages in the field and in laboratory microcosms indicated that density-dependent reduction of growth rate and increased mortality occurred at densities found in natural populations. Similar experiments failed to provide any evidence that Corophium mortality or behaviour was modified by the presence of Hydrobia or members of its own species; nor did the presence of Corophium cause a reduction in growth rate of Hydrobia.     

Thermal preference and tolerance of Megastrea (Lithopoma) undosa (Wood, 1828; Gastropoda: Turbinidae)

The thermoregulatory behavior of the wavy turban snail Megastrea (Lithopoma) undosa was determined in a horizontal thermal gradient and was 16.31 in day cycle and 14.4 °C in night cycle. Displacement velocity of adults was 29.3±4.2 cm h⁻¹ during the light phase and 26.1±3.2 cm h⁻¹ during the dark phase. The critical thermal maxima of the wavy turban snail were determined. As a measure of thermal tolerance, snails were subjected to increasing water temperatures at a rate of 1 °C every 30 min until they were detached from the substrate. The critical thermal maximum at 50% was 29.7 °C.     

The Distribution of Hydrobiids and the Effects of Sediment Characteristics on the Population Dynamics of Hydrobia ventrosa in a Coastal Region of the Southern Baltic

The composition of the hydrobiid fauna in a coastal water of the southern Baltic was studied in 1990/91 at three stations with different sediment characteristics (grain size, organic content). Four species were found (Hydrobia ventrosa, H. ulvae, H. neglecta and Potamopyrgus jenkinsi). H. ventrosa was dominant at all stations. The mean hydrobiid abundances were 4,000 (stn. C), 18,000 (stn. B) and 40,000 (stn. A) ind./m² respectively. Abundance was found to correlate with the organic content of the sediment. The reproduction and growth of H. ventrosa was studied. Two reproduction phases were observed, one in summer and the other in late autumn. The autumn reproduction peaks were staggered owing to the different growth rates at the various stations. Possible correlations between the differences in shell size of H. ventrosa at the various stations and both sediment characteristics and abundance are discussed.     

Within-lake dispersal of the prosobranch snails,Viviparus ater and Potamopyrgus jenkinsi

Summary The gastropod species Viviparus ater and Potamopyrgus jenkinsi have both colonized Lake Zürich relatively recently. If it is assumed that they have dispersed in the Lake starting from one location, a model of random diffusion is insufficient to explain their present distribution in Lake Zürich. It is suggested that both species were dispersed by humans. In P. jenkinsi, passive dispersal by birds, fish or on floating algae may also have occurred. To estimate the rate of within-lake dispersal, life tables were constructed and the diffusivity estimated for the two species. Per capita population growth rates were calculated to be 0.00337/d for V. ater and 0.0159/d for P. jenkinsi. Diffusivity was close to 1 m²/d in both species.     

Evidence for competition between mudsnails (Hydrodiidae): a field experiment

The paper describes an experimental investigation of competition between Hydrobia ulvae and H. ventrosa using enclosed populations at a site at which the species coexist naturally. lntraspecific competition is more intense than interspecific competition and may have a regulatory influence on snail densities. Other experimental studies of competition between mudsnails are reviewed and found to infer strongly the occurrence of competition in natural field populations. However, features of the species' field distributions previously attributed to interspecific competition can be more convincingly explained by other processes.     

The circatidal rhythm of the estuarine gastropod Hydrobia ulvae (Gastropoda: Hydrobiidae)

Intertidal animals display a suite of cyclic behaviours that evolved as adaptations to the predictable cycle of inundation and exposure. In estuarine habitats, mud snails from the genus Hydrobia are among the most abundant grazers, and have received considerable attention with respect to the behavioural mechanisms mediating locomotion, dispersal, and feeding, although the nature of the control of these processes has remained elusive. In particular, it is not clear whether endogenous activity patterns are related to periodic changes of microphytobenthos biomass at the sediment surface, or whether they are timed to the tidal cycle at all. In the present study, we address the crawling activity of Hydrobia ulvae under constant conditions, as well as the effects of individual size and previous short‐term exposure to tides of different range, by recording immersed individual snails under constant dark conditions. We show that the species displays an overt circatidal pattern of crawling, with activity peaks around high water, and that the start of inundation may act as an entrainment agent of the rhythm. Moreover, the results obtained indicate that smaller snails display higher levels of activity, although neither the size nor previous in situ influence of tidal range has an effect on the period and on the amplitude of the rhythm. These findings suggest that fluctuations of microphytobenthos biomass are not a sufficiently strong selective pressure to have shaped locomotor activity in H. ulvae. Moreover, feeding of H. ulvae should take place mostly during high water and be independent of periodic fluctuations of microphytobenthos biomass at the surface of the sediment. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 439–450.     

Biogeographical patterns of marine larval trematode parasites in two intermediate snail hosts in Europe

Aim We used published inventories of trematodes in Littorina littorea (L.) and Hydrobia ulvae (Pennant) in European seas to search for two basic biogeographical patterns in the spatial occurrence of various trematode species: (1) do parasite distribution and richness patterns in the two host snails overlap with known ecoregions of free‐living organisms; and (2) does trematode species richness in the snails follow latitudinal or longitudinal gradients? Location North East Atlantic. Methods We used multidimensional scaling (MDS), analysis of similarity (ANOSIM) and analysis of variance (ANOVA) to test whether there were overlaps of parasite distribution and richness with known ecoregions of free‐living organisms. In addition, we used linear regression analyses to test whether trematode richness in snails (corrected for sampling effort) was correlated with the latitude or longitude of the sampling sites. Results When corrected for sampling effort, mean trematode species richness per site did not differ among the different ecoregions in L. littorea. In contrast, in H. ulvae, mean species richness was much lower for sites from the Celtic Sea compared with sites from the Baltic Sea and the North Sea. Based on the results of MDS analyses, trematode species composition was distinct among ecoregions; in particular, communities from the Baltic Sea differed markedly from communities in the Celtic Sea, for both snail species. Latitude and longitude were not significantly correlated with parasite species richness in either snail species. Most trematode species had restricted distributions, and only three species in L. littorea and five species in H. ulvae occurred at more than 50% of the sites. Main conclusions There is more structure in the large‐scale distribution of trematodes in gastropods than one would expect from the large‐scale dispersal capabilities of their bird and fish final hosts. We propose mechanisms based both on limited dispersal via fish and bird final hosts and on gradients in environmental factors to explain the observed patterns.