Palatability of macrophytes to the invasive freshwater snail Pomacea canaliculata: differential effects of multiple plant traits

1. By selective grazing, invasive grazers can alter macrophyte‐herbivore relationships in shallow freshwater bodies. Evaluating the palatability of macrophytes and understanding the determinants of plant palatability can help predict grazing impact. In no‐choice feeding assays, we tested the palatability of 21 species of freshwater macrophytes to the invasive freshwater apple snail Pomacea canaliculata. 2. Daily feeding rate varied greatly with plant species, ranging from 1.1 to 22% of snail body mass. We assessed six plant properties and examined their correlation with feeding rate. Total nitrogen content was positively related, and C:N ratio and dry matter content (DMC) negatively related, to snail feeding rate. There was no significant correlation between snail feeding rate and plant phenolic content, but the feeding rate on Myriophyllum aquaticum (the plant with the highest phenolic content) was very low. 3. We repeated the feeding assays for 15 species that were not palatable as fresh leaves with reconstituted plant tissues formed by mixing ground up dried leaves with agar. The feeding rate still differed greatly among macrophyte species. Phragmites australis and Vallisneria natans (two species with the highest DMC) were eaten much more as reconstituted plant than as fresh leaves, indicating that structure (i.e. DMC) may be important in their defence against snail herbivory. For two plants (M. aquaticum and Alternanthera philoxeroides) that had moderate amounts of nitrogen/phosphorus but were consumed very little as fresh and reconstituted tissues, we incorporated their extracts into a palatable agar‐based food. The extracts from both species greatly reduced snail feeding rate, indicating the presence of chemical defences in these two species. 4. These results indicated that feeding was affected by several plant traits. The snail favoured plants with a high nitrogen content and avoided plants with a high DMC. Only a few plants possessed chemical feeding deterrents that were effective against this snail. Given the invasive spread of P. canaliculata in Asia, ecologists and managers should consider plant palatability when selecting plants for use in wetland restoration and when predicting the impact of further invasion by this species.     

Seasonal variability in the grazing potential of the invasive amphipod Gammarus tigrinus and the native amphipod Gammarus salinus (Amphipoda: Crustacea) in the northern Baltic Sea

Mesograzers are known to reduce the biomass of their host plant and modify the structure of the whole macrophyte community in many ecosystems. Thus, the introduction of an efficient mesograzer may destabilize macrophyte community and also affect the native grazers. We estimated how large proportion of macrophyte production are consumed by the alien gammarid G. tigrinus and the native gammarid G. salinus in the species poor ecosystem of the northern Baltic Sea. We analysed whether G. tigrinus consumes different diet as the native G. salinus and whether the effect of G. tigrinus on the survival of the native G. salinus is macrophyte species specific. Grazing experiments showed that there was a clear difference in the grazing rates of gammarids among the studied macrophyte species in summer and autumn but not in spring. The grazing rates were significantly higher in the prevailing macrophyte Pilayella littoralis as compared to other macrophytes. The grazing was inversely related to the diurnal net photosynthetic values of macrophytes. The gammarid amphipods potentially removed only a minor part of plant primary production except for summer and autumn when grazing of a few perennial species exceeded macrophyte production. Macrophyte species and presence of G. salinus had no effect on the survival of G. tigrinus. The presence of G. tigrinus, however, reduced the survival of the native gammarids within P. littoralis in summer. To conclude it is likely that both native and alien gammarid amphipods do not exert significant pressure on the macroalgal communities in the northern Baltic Sea. Competitive interactions between G. tigrinus and G. salinus within the prevailing macrophyte P. littoralis is the likely explanation of the decline of the native gammarid amphipods after the establishment of G. tigrinus in the northern Baltic Sea.     

Determinants of macrophyte palatability to the pond snail Lymnaea stagnalis

Summary 1. This study aimed to identify the chemical and structural determinants of macrophyte palatability to the pond snail Lymnaea stagnalis . Eleven macrophyte species were investigated, and one of them ( Potamogeton lucens ) was collected at four sites, on two different dates in the year, to study palatability determinants at an intra-specific level.
2. Plant palatability to L. stagnalis was determined through non-choice feeding assays. Dry matter content (DMC), total phenolic content and protein content were measured for each macrophyte species. These parameters, and soluble carbohydrate content, were also measured for each sample of P. lucens .
3. The palatability of macrophytes was positively related to their protein content (between species only) and negatively related to their DMC (both between species and within P. lucens ). No simple relationship was found between the palatability of macrophytes and their phenolic content, but highly palatable macrophytes consistently exhibited a low phenolic content.
4. These results emphasise that macrophyte palatability is a multifactorial attribute, potentially depending on both structural and chemical traits. Because some of these traits were correlated, further investigations are required to assess their respective influence on macrophyte palatability.     

Induced chemical defenses in a freshwater macrophyte suppress herbivore fitness and the growth of associated microbes

The freshwater macrophyte Cabomba caroliniana induces a chemical defense when attacked by either the crayfish Procambrus clarkii or the snail Pomacea canaliculata. Induction by either consumer lowers the palatability of the plant to both consumers. When offered food ad libitum, snails feeding on non-induced C. caroliniana grew 2.6–2.7 times more than those feeding on induced C. caroliniana. Because snails fed less on induced plants, this could be a behavioral effect (reduced feeding), a physiological effect of the induced metabolites on the consumer, or both. To assess these possibilities, we made artificial diets with lipid extracts of induced versus non-induced C. caroliniana and restricted control snails to consuming only as much as treatment snails consumed. Growth measured as shell diameter was significantly lower on the diet containing extract from induced, as opposed to non-induced, plants; change in snail mass was more variable and showed a similar, but non-significant, trend. Thus, snails may reduce feeding on induced plants to avoid suppression of fitness. The induced defenses also suppressed growth of co-occurring microbes that might attack the plant through herbivore-generated feeding scars. When two bacteria and three fungi isolated from C. caroliniana surfaces were cultured with the lipid extract from induced and non-induced C. caroliniana, both extracts inhibited the microbes, but the induced extract was more potent against three of the five potential pathogens. Thus, induced plant defenses can act against both direct consumers and microbes that might invade the plant indirectly through herbivore-generated wounds.     

Relative effects of nutrient enrichment and grazing on epiphyte-macrophyte (Zostera marina L.) dynamics

The independent and interactive effects of nutrient concentration and epiphyte grazers on epiphyte biomass and macrophyte growth and production were examined in Zostera marina L. (eelgrass) microcosms. Experiments were conducted during early summer, late summer, fall, and spring in a greenhouse on the York River estuary of Chesapeake Bay. Nutrient treatments consisted of ambient or enriched (3× ambient) concentrations of inorganic nitrogen (ammonium nitrate) and phosphate. Grazer treatments consisted of the presence or absence of field densities of isopods, amphipods, and gastropods. epiphyte biomass increased with both grazer removal and nutrient enrichment during summer and spring experiments. The effect of grazers was stronger than that of nutrients. There was little epiphyte response to treatment during the fall, a result possibly of high ambient nutrient concentrations and low grazing pressure. Under low grazer densities of early summer, macrophyte production (g m^–2 d^–1) was reduced by grazer removal and nutrient enrichment independently. Under high grazer densities of late summer, macrophyte production was reduced by enrichment only with grazers absent. During spring and fall there were no macrophyte responses to treatment. The relative influence of epiphytes on macrophyte production may have been related to seasonally changing water temperature and macrophyte requirements for light and inorganic carbon.     

Breeding biology of Tenagomysis tasmaniae Fenton,Anisomysis mixta australis (Zimmer) and Paramesopodopsis rufa Fenton from south-eastern Tasmania (Crustacea: Mysidacea)

The number, size and developmental stage of young in the brood pouch of female Tenagomysis tasmaniae, Anisomysis mixta australis and Paramesopodopsis rufa was recorded throughout the year. Breeding was intensive from spring till the end of autumn for the three species. Calculation of the egg ratio for each species showed that their major reproductive peaks occurred during spring and summer. A winter depression in the breeding cycle was observed for T. tasmaniae and P. rufa, but A. mixta australis ceased breeding during winter. Seasonal variation in the length of gravid females and number of young carried was evident for these three species. Females were longer in spring and summer and carried more young than in autumn and winter. A linear relationship between female length and brood size was demonstrated for each species; annual and seasonal equations were calculated for females carrying each developmental stage. The seasonal equations showed that for a female of given length fecundity was greater during spring than any other season. Natality was estimated to be highest during late spring, summer and early autumn for the three species. No seasonal variation in the size of eggs was evident for the three species. The reproduction pattern of T. tasmaniae, A. mixta australis and P. rufa appears to be very similar to that reported for the majority of iteroparous coastal temperate mysids throughout the world.     

Effects of macrophytes on feeding and life-history traits of the invasive apple snail Pomacea canaliculata

1. Biological invasions have become a serious threat to ecosystems worldwide. Various factors can contribute to the success of biological invasion. We examined how different macrophyte food affected feeding and life‐history traits of the invasive herbivorous snail Pomacea canaliculata, and whether differences in snail life‐history traits could explain its successful infestation of agricultural and non‐agricultural wetlands in Asia. 2. We tested five cultivated and five wild semi‐aquatic macrophytes. Snail daily feeding rate varied substantially with plant species, ranging from 1.3% to 22% of its body mass. Snails fed with four (Amaranthus gangeticus, Apium graveolens dulce, Ipomoea aquatica and Nasturtium officinale) of the five cultivated macrophyte species exhibited high survivorship, fast growth and high fecundity. Snails fed with Colocasia esculenta, however, grew poorly, did not reproduce and eventually died. 3. Of the five wild species (Eichhornia crassipes, Ludwigia adscendens, Murdannia nudiflora, Myriophyllum aquaticum and Polygonum hydropiper), M. nudiflora supported a high snail survival, but snails had slower growth and lower fecundity than those reared on the four palatable cultivated species. Snails fed with L. adscendens grew substantially slower than those fed with M. nudiflora, and produced only a small clutch of eggs. Snails fed with E. crassipes, M. aquaticum and P. hydropiper had very low survivorship, grew very little and did not reproduce. 4. We determined six plant properties and their correlation with the feeding, growth and reproduction of the apple snails. Cultivated macrophytes in general had a higher nutritional value and lower physical and chemical defences. Phenolic content was negatively correlated with snail feeding rate, while plant nitrogen and phosphorus contents were positively correlated with snail egg production and growth, respectively. 5. These results indicate that, due to their higher nutritional value and lower chemical and physical defences, cultivated macrophytes are in general desirable for the apple snail which may partly explain its successful invasion into wet agricultural areas in Asia. This snail may also selectively graze poorly defended wild macrophytes in non‐agricultural wetlands, leading to changes in floral diversity and wetland functioning. Management of this and other apple snails with similar life‐history traits should thus focus on the prevention of their further spread.     

Habitat choice in rotifera communities of three shallow lakes: impact of macrophyte substratum and season

The distribution of rotifer communities between emergent (Typha angustifolia) and submerged (Chara tomentosa) vegetation and a comparatively open water zone were compared during the spring, summer and autumn seasons at three macrophyte-dominated lakes. This survey identified 107 rotifera species of which 58% of the taxonomical structure was common for the three examined lakes. Stoneworts with a more complicated spatial and morphological structure (having a much longer stem length than the narrow leaf cattail), supported higher rotifer densities. The stem length appeared to be the best predictor of all the macrophyte parameters and pH and chlorophyll a for the chemical variables, for explaining the variation of rotifer densities using the stepwise multiple forward regressions. The distribution of pelagic species did not differ between particular sites, which may have reflected the behavioural requirements of those rotifers. Some of them remained in the open water zone while others seeking an anti-predator refuge, gathered within macrophyte stands during the daytime. Moreover, there were nine Chara-associated species recorded and only one Typha-associated species was noted. The similarity of rotifer communities was most strongly influenced by particular habitat and season. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

Indirect effects of fish on macrophytes in Bays Mountain Lake: evidence for a littoral trophic cascade

Summary We began this experiment to test specific hypotheses regarding direct and indirect effects of fish predation on the littoral macroinvertebrate community of Bays Mountain Lake, Tennessee. We used 24 m² enclosures in which we manipulated the presence and absence of large redear sunfish (Lepomis microlophus>150 mm SL), and small sunfish (L. macrochirus and L. microlophus <50 mm SL) over a 16-mo period. Here we report on effects of fish predation on gastropod grazers that appear to cascade to periphyton and macrophytes.Both large redear sunfish and small sunfish maintained low snail biomass, but snails in fish-free controls increased significantly during the first 2-mo of the experiment. By late summer of the first year of the experiment, the difference in biomass between enclosures with and without fish had increased dramatically (>10×). Midway through the second summer of the experiment, we noted apparent differences in the abundance of periphyton between enclosures containing fish and those that did not. We also noted differences in the macrophyte distribution among enclosures. To document these responses, we estimated periphyton cover, biovolume and cell size frequencies as well as macrophyte distributions among enclosures at the end of the experiment. When fish were absent, periphyton percent cover was significantly reduced compared to when fish were present. Periphyton cell-size distributions in enclosures without fish were skewed toward small cells (only 12% were greater than 200 m³), which is consistent with intense snail grazing. The macrophyte Najas flexilis had more than 60 x higher biomass in the fish-free enclosures than in enclosures containing fish; Potamogeton diversifolius was found only in fish-free enclosures. These results suggest a chain of strong interactions (i.e. from fish to snails to periphyton to macrophytes) that may be important in lake littoral systems. This contrasts sharply with earlier predictions based on cascading trophic interactions that propose that fish predation on snails would enhance macrophyte biomass.     

Effects of grazing by fish and waterfowl on the biomass and species composition of submerged macrophytes

Biomanipulation improved water transparency of Lake Zwemlust (The Netherlands) drastically. Before biomanipulation no submerged vegetation was present in the lake, but in summer 1987, directly after the measure, submerged macrophyte stands developed following a clear-water phase caused by high zooplankton grazing in spring. During the summers of 1988 and 1989 Elodea nuttallii was the most dominant species and reached a high biomass, but in the summers of 1990 and 1991 Ceratophyllum demersum became dominant. The total macrophyte biomass decreased in 1990 and 1991. In 1992 and 1993 C. demersum and E. nuttallii were nearly absent and Potamogeton berchtholdii became the dominant species, declining to very low abundance during late summer. Successively algal blooms appeared in autumn of those years reaching chlorophyll-a concentrations between 60–130 µg l^–1. However, in experimental cages placed on the lake bottom, serving as exclosures for larger fish and birds, E. nuttallii still reached a high abundance during 1992 and 1993. Herbivory by coots (Fulica atra) in autumn/winter, and by rudd (Scardinius erythrophthalmus) in summer, most probably caused the decrease in total abundance of macrophytes and the shift in species composition.     

Trophic Cascades Uncoupled in a Coastal Marsh Ecosystem

The Mediterranean population of the exotic eastern mosquitofish Gambusia holbrooki (Agassiz 1859) (Osteichthyes, Poeciliidae) has been held responsible for causing eutrophication due to zooplankton removal and phytoplankton enhancement, however no experimental evidence exists of this. To test this allegation, an enclosure experiment was conducted in spring in an oligohaline coastal marsh. The manipulation of fish density had profound effects on zooplankton, whose density greatly decreased when the occurrence of mosquitofish increased. Cladocerans and ostracods were more affected by mosquitofish than cyclopoid copepods, whilst rotifer density was not modified. Changes in zooplankton density did not cascade to lower trophic levels as no differences were observed between the chlorophyll concentration in fish and fish-less enclosures. This is because zooplankton was dominated by species with low filter-feeding rates, such as small cladocerans. In consequence, the total macrophyte standing crop was not affected. The only benthic macroinvertebrate species whose density increased in the absence of eastern mosquitofish was the mud snail P. acuta. Higher numbers of snails explain why the standing crop of the filamentous green algae Oedogonium sp. decreased in fish-less enclosures. The density of chironomid midge larvae did not increase in fish-less enclosures, because eastern mosquitofish forage on them mainly during summer, when zooplankton has already been depleted; nor were damselflies, probably because they are too large. Nitrogen concentration decreased after fish exclusion, but phosphorus concentration remain unchanged. In conclusion, it was found that the eastern mosquitofish affect zooplankton of the Mediterranean oligohaline lagoons considerably, but they do not enhance phytoplankton growth, because the system is bottom-controlled by submerged macrophytes.     

Upward cascading effects of nutrients: shifts in a benthic microalgal community and a negative herbivore response

We evaluated the effects of nutrient addition on interactions between the benthic microalgal community and a dominant herbivorous gastropod, Cerithidea californica (California horn snail), on tidal flats in Mugu Lagoon, southern California, USA. We crossed snail and nutrient (N and P) addition treatments in enclosures on two tidal flats varying from 71 to 92% sand content in a temporally replicated experiment (summer 2000, fall 2000, spring 2001). Diatom biomass increased slightly (~30%) in response to nutrient treatments but was not affected by snails. Blooms of cyanobacteria (up to 200%) and purple sulfur bacteria (up to 400%) occurred in response to nutrient enrichment, particularly in the sandier site, but only cyanobacterial biomass decreased in response to snail grazing. Snail mortality was 2–5 times higher in response to nutrient addition, especially in the sandier site, corresponding to a relative increase in cyanobacterial biomass. Nutrient-related snail mortality occurred only in the spring and summer, when the snails were most actively feeding on the microalgal community. Inactive snails in the fall showed no response to nutrient-induced cyanobacterial growths. This study demonstrated strongly negative upward cascading effects of nutrient enrichment through the food chain. The strength of this upward cascade was closely linked to sediment type and microalgal community composition.     

Food preference of freshwater invertebrates: comparing fresh and decomposed angiosperm and a filamentous alga

1. Fresh and decomposed Mougeotia sp. (a filamentous green alga) and Elodea nuttallii (a vascular plant) were offered as food to three species of aquatic macroinvertebrates (Lymnaea peregra, Asellus meridianus and Endochironomus albipennis) to test: (i) if filamentous algae are preferred to aquatic higher plants (hereafter, called ‘macrophytes’) and (ii), as is known for higher plants, if decomposition also results in greater palatability of filamentous algae. 2. Compared with the alga, the macrophyte in both states was of higher nutritional value. Conditioning improved the nutritional value of both food types, but especially of the macrophyte. 3. Both fresh alga and fresh macrophyte were eaten little by all animals except A. meridianus feeding upon the macrophyte. Consumption was higher for both plants in their decomposed state. However, L. peregra consumed more decomposed macrophyte than the decomposed alga. Both decomposed plants were eaten most by E. albipennis followed by A. meridianus and L. peregra. 4. Digestibility of both plants, but especially of the macrophyte, increased significantly after decomposition. The assimilation efficiencies of all animals on the fresh E. nuttallii were higher than on fresh Mougeotia sp. After decomposition, the efficiency increased significantly only on the alga. Consequently, both decomposed plants were assimilated with similar efficiency by all test animals. 5. Amongst aquatic macrophytes, the increase of their consumption and digestibility upon decomposition has hitherto been known only for vascular plants but not for filamentous algae.     

The distribution of submerged aquatic macrophyte biomass in a eutrophic stream,Badfish Creek: the effect of environment

The distribution of biomass of the macrophyte community in Badfish Creek was examined in three sections (A–C) totaling ten kilometers. Biomass samples were taken in a stratified-random manner, with sediment characteristics, depth, current velocity and incident light measured at each site to correlate individual biomass samples with environmental factors. Total community biomass decreased in the downstream section (C), with the biomass ofElodea canadensis decreasing abruptly below section A. The only environmental factors which were correlated with the decrease in macrophyte community biomass, especially that ofPotamogeton pectinatus, in section C was the increase in substrate heterogeneity and sand substrates which lacked surface gravel. The change in substrate was related to channelization. Considering the stream as a whole, the biomass of the dominant species,Potamogeton pectinatus, was correlated with incident light. Other species present wereCeratophyllum demersum andCladophora glomerata. Analysis of riparian vegetation type indicates that tree cover significantly reduced macrophyte biomass by incident light reduction.     

Nitrogen,phenolic acids,and other feeding cues for salt marsh detritivores

Summary A number of feeding cues determine the palatability of detritus to detritivorous invertebrates. In salt marsh detritus the feeding cues include flavor provided by phenolics such as ferulic acid, pH, salinity, and nitrogen content. We examined the feeding responses to each cue by using experimental manipulations where we changed concentrations of these chemical cues. The palatability of detritus of the salt marsh grass Spartina alterniflora to the detritivorous snail Melampus bidentatus is reduced by increases in ferulic acid in the detritus. The acidity of the acid is partly responsible for inhibition of feeding, but other flavors of the ferulic acid are the major factor. Changes in salinity makes detritus more or less palatable to different species of detritivores. Available nitrogen confers greater palatability to detritus.In the field the feeding cues are all present simultaneously, and detritivores feed based on a hierarchy of cues. For M. bidentatus the presence of sufficient available nitrogen overwhelms the response to ferulic acid, the second most important cue. Salinity and pH, although used as discriminatory cues by themselves, are located lower in the hierarchy than nitrogen and ferulic acid.As detritus ages in the marsh, nitrogen content changes. Similarly, detritus from eutrophied environments shows enhanced nitrogen content. The differences in palatability of new and old detritus, and from eutrophic and non-eutrophic environments, suggests that detritivores respond not to total nitrogen but rather to the availabel nitrogen fractions. The increase in lignin found in old and eutrophic detritus may increase the binding of available nitrogen, and hence reduce the palatability of detritus.     

The Effect of Seasonal Shifts in Temperature on the Lipid Composition of Marine Macrophytes

The comparative study of lipid composition was carried out in four species of marine algae, Ahnfeltia tobuchiensis, Laminaria japonica, Sargassum pallidum, and Ulva fenestrata, as well as a higher plant grass wrack (Zostera marina). Plants were collected in the Japan Sea in spring at 2.9 and 5.5°C and in summer at 23°C. The main lipid components of membranes were determined, and the general patterns of the ratio of phospholipids (PL), glycolipids (GL), betaine (BL), and neutral (NL) lipids were discerned. The relative content of NL in all species (except A. tobuchiensis) was higher in summer. The level of triacylglycerols was as high as 18–37%. The content of individual classes of PL and GL varied between the spring and summer samples, the relative content of PL being higher in spring. In most species, the ratio of PL to GL decreased in summer. The content of free sterols did not depend on the season. The molar ratios of phosphatidylcholine and diacylglycerol-o-(hydroxymethyl)-(N,N,N-trimethyl)homoserine to free sterols varied from 0.9 to 1.7. The seasonal changes of lipid composition were apparently related to macrophyte adaptation to water temperature and to biology of their development.     

Optimal diet theory: behavior of a starved predatory snail

Summary The tenets of optimal foraging theory are used to contrast the behavior of the predatory snail Acantina spirata when feeding on the barnacles Balanus glandula and Chthamalus fissus under conditions of satiation and starvation. As predicted in optimal diet models, A. spirata is less selective (ratio of attack frequency on a prey species to number of individuals available) when the higher ranking prey has low abundance. When given a choice, starved snails attack both barnacle species equally, whereas satiated individuals preferentially attack B. glandula, the more profitable prey (ash-free dry weight of barnacles ingested per unit handling time). Under starvation conditions, equal attack frequency does not result in equal prey species consumption because Acanthina spirata is more successful at attacking C. fissus than B. glandula.The assumption of constant prey encounter rates in optimal diet models is not met when A. spirata goes from a state of satiation to starvation. The encounter rate on B. glandula is lowered due to a decrease in attack success. A loss of feeding skills in starved A. spirata is responsible for the greater difficulty snails have in gaining access through the opercular plates of B. glandula.Behavioral changes in A. spirata as snails pass from satiation to hunger translate into an energetic disadvantage during feeding for hungry snails for two reasons. First, higher prey handling times result in a decreased rate of biomass intake. Second, alteration in the relative attack frequency between barnacle species, combined with a decrease in attack success on the more profitable prey leads to more frequent ingestion of the less profitable prey.     

Clear,crashing, turbid and back – long‐term changes in macrophyte assemblages in a shallow lake

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Predation induced changes in behavior and growth rate in three populations of the intertidal snail, Littorina sitkana (Philippi)

We investigated the sublethal effects of a predatory crab, Cancer productus (Randall), on the behavior and growth of its snail prey, Littorina sitkana, by setting up controlled rearing and prey-size selection experiments. L. sitkana were collected from three sites on San Juan Island, WA, USA. These sites varied in snail size, abundance, and vertical distribution, and in the abundance of the crab predator C. productus. Snails from all three populations were raised for 34 days under the following treatments: no-crab control, a non-feeding C. productus encased in mesh box, and an encased C. productus feeding on L. sitkana. The non-feeding crab treatment did not affect snail foraging behavior or growth rate in comparison with the no-crab control. In contrast, the presence of a feeding crab elicited escape behavior in the snails, halted grazing, and consequently reduced growth rates. A population difference in escape behavior was observed: upward migration in snails from rocky shores and hiding in crevices in snails from a mud flat. It thus appears that chemicals leaching from crushed conspecific snails, rather than the presence of the crab predator, act as the “alarm substance” to which L. sitkana react. The magnitude of the growth depression in the presence of feeding crabs was 85%, with no difference among the three populations. Once the feeding crab stimulus was removed, snails in all populations resumed normal growth, suggesting that this response to feeding predators is reversible with changing environmental conditions. Laboratory experiments were set up to determine if all size classes of L. sitkana are equally susceptible to C. productus predation. C. productus consistently selected the largest of three size classes of L. sitkana. These results suggest that slow growth rate and small size in L. sitkana may actually be an adaptation for coexisting with high C. productus abundance, rather than simply a cost of escape behavior.     

Spatial and temporal distribution of the zoobenthos community in a woodland pond (Switzerland)

Spatial and temporal changes of zoobenthos composition and density were assessed in a woodland pond, near Geneva (Switzerland), by monthly sampling (during 15 months), on the three main substrates: a submerged macrophyte (Chara sp.), an emergent macrophyte (Typha latifolia stems) and allochthonous detritus (oak leaves).Many taxa showed preferences for one or two of the substrates, as expressed by differences in densities, but few taxa presented an exclusivity for one substrate. Zoobenthos densities (per m² of pond bottom) were largely dominated on all substrates and seasons by Chironomidae and Oligochaeta. Other dense macroinvertebrates included Ostracoda (on Chara and Typha stems), Gastropoda (on Typha stems), Ephemeroptera (all substrates), and Ceratopogonidae (on Typha stems).The magnitude of temporal differences in densities was expressed by the ratio of maximal to minimal monthly densities and reached values between 2.8 to 11.8. On all substrates the highest monthly densities were observed in summer or beginning of autumn. The temporal changes in densities were strongly influenced by the life cycles of the invertebrates: presence of numerous young individuals in summer (for example Cypridopsis vidua, Cloeon dipterum, Caenis horaria, Ferrissia wautieri), and emergence (for example Chironomidae in April). Fluctuations in densities, especially on Chara and leaves, were also attributed to modification of substrate condition (surface availability, stage in the decomposition or senescence process). Temporal fluctuations of densities were compared with other lentic water bodies: highest densities showed a common trend, occurring between end of spring and beginning of autumn.