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 in Hydrobia

Summary Fenchel's study of size variation in Hydrobiid snails in the Limfjord, Denmark, provides one of the most convincing cases of ecological character displacement available. In order to assess the generality of the phenomena within the Hydrobia genus, allopatric and sympatric Hydrobia ventrosa, H. neglecta and H. ulvae were collected from 24 coastal sites around Eastern England in July and October, 1982. Shell heights of 5,850 snails from 55 samples were recorded. These data were analysed for intraspecific allopatric-sympatric, and interspecific height differences. Such differences were not significant for H. ventrosa/H. neglecta, but the species exhibited parallel variation at sympatric sites. H. ventrosa/H. ulvae showed only significant interspecific differences, but in both allopatry and sympatry. The pattern of mean size variation for these species resembles that in the Limfjord. However the statistical analyses fail to provide support for character displacement. Differences in character states attributed to the process of character displacement may result from a number of other causes. Environmental conditions at sympatric and allopatric sites may act differentially on the heights of H. ulvae and H. ventrosa. The conditions that prevail at sympatric H. ulvae sites appear to lead to increased size in this species irrespective of the presence of H. ventrosa.     

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 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.     

The effect of larval trematodes on the survival rates of two species of mud snails (hydrobiidae) experimentally exposed to desiccation,freezing and anoxia

Digenetic trematodes are widespread among mud snails (Hydrobiidae) living in coastal lagoons and estuaries, but knowledge is generally lacking on their impact on these host organisms. We examined the survival rates of infected and non-infected experimental populations of two mud snail species,Hydrobia ventrosa (Montagu) andHydrobia neglecta Muus, exposed to desiccation, freezing and anoxia in the laboratory. Our experiments indicated that non-infected groups of both species had similar survival rates after being subjected to desiccation and anoxia, whereasH. ventrosa survived freezing better thanH. neglecta. However, infected groups ofH. neglecta specimens subjected to desiccation showed significantly lower survival rates than non-infected groups. Infected and non-infected snails of both species exposed to freezing and anoxia exhibited similar survival rates. The possible mechanisms by which parasites influence their hosts are discussed. It is unlikely that the parasites in the present case mediate the coexistence of the twoHydrobia-species, because the snail with the highest reproductive effort-H. neglecta-showed lower infection rates in situ than its congenerH. ventrosa.     

Tolerance and Resistance in Gastropod Mollusks Hydrobia ulvaeand H. ventrosafrom the White Sea to Abiotic Environmental Factors

Tolerance to salinity changes and resistance to desiccation, fresh water, and freezing were studied in two hydrobiid species from the White Sea, Hydrobia ulvae(Pennant) and H. ventrosaMontagu. It was shown that H. ventrosahas a greater tolerance to low salinities in the range of 6–10 but is less tolerant to high salinities (35–45). The interspecies differences in low salinity tolerance persisted after the acclimation of snails to 20, 16, and 12. A comparison of survival in fresh water and under desiccation conditions suggests that H. ulvaeis more resistant to these factors. In fresh water, LT₅₀was 30 and 60 days for H. ventrosaand H. ulvae, respectively. Under desiccation conditions, LT₅₀was 6 and 25 days for H. ventrosaand H. ulvae, respectively. At subzero temperatures, H. ventrosawas shown to be a superior survivor. Within the same species of mud snail, the detrimental effect of freezing depended on the salinity: survival decreased with decreasing salinity. These data suggest significant differences in the mechanisms of resistance and tolerance to abiotic environmental factors between the two species. Despite the partial overlapping of their ecological niches, the interspecies differences may play an important role in the distribution and spatial structure dynamics of coexisting populations of these species in the White Sea region.     

Salinity detection by Hydrobia ulvae (Pennant) and Potamopyrgus jenkinsi Smith (Gastropoda:Prosobranchia)

Both Hydrobia ulvae (Pennant) and Potamopyrgus jenkinsi Smith climb vertically in the laboratory but stop or crawl horizontally if presented with a sharp salinity drop. 50% of Hydrobia ulvae respond to a difference of 18–26% from the original solution. 50% of Potamopyrgus jenkinsi respond to a difference of 53–55%. In both species, the response is impaired at high temperatures, and in Hydrobia ulvae it is also impaired at low temperatures. From 9–22 °C no temperature influence is evident.From acclimation experiments it is suggested that the snails respond not to the gradient across the interface, but to the gradient between body fluids and the upper solution presented. Snails respond to non-ionic gradients when these are presented either as density gradients or as solutions separated by a semi-permeable membrane, and it is argued that they detect osmotic pressure directly. The sense organs are probably located in the tentacles.     

DISTRIBUTION AND PRODUCTION OF THREE HYDROBIA SPECIES (GASTROPODA: HYDROBIIDAE) IN A SHALLOW COASTAL LAGOON IN THE BAY OF CADIZ, SPAIN

The abundance, life span, growth and production of the mud snailsHydrobia minoricensis, H. ulvae and H. ventrosa in a semi-naturallagoon system were studied by taking monthly samples at threesites during 1991 and 1992. The most abundant species, H. minoricensisoccurred at mean densities of 12834 to 26264 snails m^–2(10.7 to 25.8g dry weigh m^–2), depending on the site.The least abundant species, H. ulvae, occurred at mean densitiesof 185 to 353 snails m^–2 (3.2 to 2.2g dry weight m^–2).The numerical abundance and biomass of the three Hydrobia specieswere positively related to the biomass of benthic macroalgae(P<0.01). Although H. ulvae egg capsules were recorded throughoutthe year, newly hatched snailsof this species were not observed,in contrast to the other two species. The early spring and summercohorts of H. minoricensis and H. ventrosa seemed to be themost numerous. The average life spans of these two species wereestimated to be about 18 and 13 months respectively. Annualproduction estimates for the whole lagoon system were 29.0 (6.3),5.5 (0.8) and 5.2 (1.0)g dry weight (ash-free dry weight) m^–2yr^–1 for H. minoricensis, H. ulvae and H. ventrosa respectively.The annual P/B ratio was about 2 for H. minoricensis and H.ventrosa. (Received 5 July 1994; accepted 5 October 1994)     

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.     

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.     

The occurrence and ecology of a marine hydrobiid mudsnail in the southern hemisphere: the Knysna Estuary, South Africa

Earlier published records of the gastropod Hydrobia from the warm temperate Knysna Lagoon on the Indian Ocean coast of South Africa are in error. Nevertheless, an animal hitherto identified as an Assiminea (Assimineidae) or as a Tomichia (Pomatiopsidae) is in fact a Hydrobia s.l., and it does occur intertidally within the Knysna system. The same species also occurs in salt pans in the cool temperate Great Berg Estuary on the Atlantic coast of South Africa. This extends the distribution of confirmed species of the dominant coastal hydrobiid mudsnails of the northern hemisphere into the southern hemisphere. H. knysnaensis (Krauss) occurs patchily at low density near the head of the Knysna Estuary in waters of low salinity. This appears to be a suboptimal habitat, and it is suggested that Hydrobia is restricted to this zone as a result of interference competition from potamidid mudwhelks and ocypodid crabs. The prevalence of potamidids and ocypodids in the tropics and in the temperate southern hemisphere may account for the rareness or absence of hydrobiids in these areas. Unlike its equally intertidal relative, the widespread North Atlantic/Arctic H. ulvae, H. knysnaensis develops directly like other Hydrobia.     

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.     

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.     

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.     

The availability of microorganisms attached to sediment particles as food for Hydrobia ventrosa Montagu (Gastropoda: Prosobranchia)

Summary The deposit-feeding prosobranch Hydrobia ventrosa Montagu feeds most rapidly upon sediment particles that pass through a 10 m sieve. Ingestion rate decreases with particles 80–125 m, then increases with larger particles, which are fed upon by scraping fine material from their surfaces. Hydrobia is capable of digesting diatoms and bacteria from sediment particles, but with generally lower efficiencies than reported when fed pure cultures. Digestion of microorganisms appears to be constrained by ability of the snail to detach cells from sediment particles; only those cells detached from sediment seem to be available for digestion. In contrast, the amphipod Corophium volutator is capable of utilizing most of the diatoms not digested by Hydrobia. For a given sediment, a constant number of microorganisms appear to be safe from digestion by H. ventrosa, and bacteria and microalgae over this amount constitutes the available food.     

Comparability of Cartesian Diver,Gilson, Warburg and Winkler methods of measuring the respiratory rates of aquatic invertebrates in ecological studies

Summary Comparison of Cartesian Diver-Winkler measurements of respiratory rates in larvae of the damselfly Pyrrhosoma nymphula, and Cartesian Diver-Gilson comparisons in the snail Potamopyrgus jenkinsi are descibed. Measurements of oxygen uptake using these different methods do not differ significantly for the same species of animal at the same temperature. Published data likewise suggest that Warburg and Winkler measurements can give similar results.All these methods, therefore, appear to be equally suitable for ecological work, providing they are used carefully and with a sound knowledge of the biology of the animal under investigation.     

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.     

Operculum closing as a defence against predatory leeches in four British freshwater prosobranch snails

The defence reaction of operculum closing in response to the presence of the molluscivorous leech Glossiphonia complanata (L.) and the non-molluscivorous Erpobdella octoculata (L.) was studied in four species of freshwater prosobranch gastropod. Bithynia tentaculata (L.) and Valvata piscinalis (Müller) can distinguish between the leeches, reacting only to G. complanata. V. piscinalis is capable of a greater degree of distance chemoreception of the leech ‘scent’. Valvata cristata Müller and Potamopyrgus jenkinsi (Smith) did not react to either leech. V. cristata may not be a potential prey item for G. complanata, while P. jenkinsi is fed on by the leech, but is a relative newcomer to the freshwater fauna. Animal Ecology Research Group, Department of Zoology, University of Oxford Commonwealth Forestry Institute, Department of Plant Sciences, University of Oxford     

Resource partitioning on basis of hydrolytic enzymes in deposit-feeding mud snails (Hydrobiidae)

Summary Crude extracts of hydrolytic enzymes from the related mud snailsHydrobia ulvae, H. ventrosa andH. neglecta are compared by use of six different methods and 18 natural carbohydrates. Synthetic substrates for linkage specific carbohydrases and trypsin-like activity were studied in addition to lysozyme-like activity.A significant hydrolysis was only observed with the reserve carbohydrates amylose, glycogen and laminaran. Many algal carbohydrates were, however, digested to some extent.The qualitative spectra are almost identical for the three species but significant quantitative differences were found. The findings are discussed in relation to information on the chemical composition of potential food items and it is concluded that the commonly observed coexistence of the snails can in part be explained by selection for microalgae and detritus particularly meeting their enzymatic potentials.