antipredator behaviour chemical communication cohabitation gastropod great pond snail learning Ly... more antipredator behaviour chemical communication cohabitation gastropod great pond snail learning Lymnaea stagnalis phylogeny In aquatic habitats prey often show maximum responsiveness to predators when they are exposed to kairomones and alarm cues in combination. As alarm cues can derive from heterospecific as well as conspecific animals, prey may need to fine tune their responses depending on the taxa with which they co-occur. We used juveniles of the freshwater gastropod, Lymnaea stagnalis L. (great pond snail) to test whether the response to heterospecific alarm cues could be enhanced by cohabitation in the absence of predators, and therefore without the potential for cue association learning. Snails were raised from oviposition alongside one of four other pulmonate species, differing in their phylogenetic relatedness to L. stagnalis, that is, from intragenus to intrasuborder. The antipredator response to heterospecific alarm cues paired with fish kairomones was then compared with the response of L. stagnalis reared in isolation. Cohabitation increased the response to heterospecific alarm cues paired with fish kairomones only when L. stagnalis had developed in the presence of closely related (Lymnaea fusca and Radix balthica) rather than distantly related heterospecifics (Physella acuta and Planorbarius corneus). Hence, in L. stagnalis, learning to recognize heterospecific alarm cues can occur without the need for cue association but the learnt response is constrained by species' relatedness. This result suggests that differential responses observed in studies based on wild-caught animals in response to heterospecific alarm cues may be explained by learning through cohabitation alone. Ó
1. Hydrobiological changes were assessed along an altirudinal transect of eighteen to twenty-thre... more 1. Hydrobiological changes were assessed along an altirudinal transect of eighteen to twenty-three tributaries from 600 to 3750m in two adjacent river systems in east-central Nepal. The transect incorporated catchments under terraced agriculture at the lowest altitudes in the Likhu Khola, through streams in forest, alpine scrub and tundra at higher altitudes in Langtang.2. Diatoms, bryophytes, macroinvertebrates and fish all showed pronounced altitudinal changes in assemblage composition as shown by TWINSPAN and DECORANA. A few taxa were restricted to streams at high altitude, but many more occurred only at lower altitudes where taxon richness increased substantially despite catchment disturbance by terraced agriculture.3. Diatoms characteristic of lower altitude streams were mostly motile, epipelic or episammic Navicula and Nitzschia spp., which occur typically at greater electrolyte and nutrient concentrations. Those characteristic of higher and steeper sites included attached Fragilaria spp. and prostrate Achnanthes spp., tolerant of turbulent flow.4. Cover by bryophytes varied within catchment type; high altitude springs supported dense mats, unlike streams fed by ice and glaciers. Taxa confined to low altitudes included those characteristic of humid subtropical conditions.5. Invertebrate families occurring only at lower altitudes included a range of burrowers and pool dwellers. Numerically, filter feeding Hydropsychidae and Simuliidae dominated streams in terraced and forested catchments, whereas grazing baetid mayflies dominated higher altitude streams in scrub and tundra.6. The combined density and biomass of at least six fish species in the Likhu Khola were 23–250 (per 100m−2), and 86–1282 g wet mass (per 100 m−2), respectively. No fish were found in Langtang streams, probably because torrential headwaters prevented colonization.7. Our data confirm that altitudinal transitions in stream biota are pronounced in the Himalaya of Nepal, but are likely to reflect a wide array of potential influences.
1 Macroinvertebrates were sampled in the riffles of fifty-eight streams from three regions of the... more 1 Macroinvertebrates were sampled in the riffles of fifty-eight streams from three regions of the Himalaya (Anapurna, Langtang and Everest) in Nepal. A semi-quantitative method with identification to family level was used to describe communities on-site.2 Stream physicochemistry was assessed and the community structure of macroinvertebrates was related to chemistry, physiography (substratum composition, altitude and size), geographical location and the dominant land use in each catchment (terraced agriculture, forest or scrub). Community data were analysed by ordination (DECORANA) and classification (TWINSPAN).3 The concentration of cations in stream water decreased significantly with altitude. Chemistry also differed between regions; sites from Anapurna had a higher pH and conductivity than those in the other two areas.4 Communities were dominated by aquatic insect larvae, with Ephemeroptera, in particular the Baetidae, most numerous across sites.5 There were, nevertheless, differences in community structure between sites, which were related closely to stream physicochemistry. Ordination scores were strongly correlated with altitude, magnesium concentration and substratum composition. Classification was also linked to altitude and chemistry, differentiating high-altitude sites with low silica concentrations from others. Sites from the Anapurna and Everest regions, with their contrasting chemistry, were also separated.6 Community structure was also related to land use: streams draining catchments dominated by terraced agriculture had different communities from those in scrub or forest. This result was confounded, however, by the strong relationship between land use, altitude and chemistry; sites in terracing were at lower altitude, had higher concentrations of silica and a higher proportion of fine sediments than those in the other land-use types.7 Overall, our data indicate that natural features of the relief and geology in the Himalaya create strong gradients in their invertebrate faunas, but that activities of man may have an effect on stream structure and ecology through catchment management. Macroinvertebrates were sampled in the riffles of fifty-eight streams from three regions of the Himalaya (Anapurna, Langtang and Everest) in Nepal. A semi-quantitative method with identification to family level was used to describe communities on-site. Stream physicochemistry was assessed and the community structure of macroinvertebrates was related to chemistry, physiography (substratum composition, altitude and size), geographical location and the dominant land use in each catchment (terraced agriculture, forest or scrub). Community data were analysed by ordination (DECORANA) and classification (TWINSPAN). The concentration of cations in stream water decreased significantly with altitude. Chemistry also differed between regions; sites from Anapurna had a higher pH and conductivity than those in the other two areas. Communities were dominated by aquatic insect larvae, with Ephemeroptera, in particular the Baetidae, most numerous across sites. There were, nevertheless, differences in community structure between sites, which were related closely to stream physicochemistry. Ordination scores were strongly correlated with altitude, magnesium concentration and substratum composition. Classification was also linked to altitude and chemistry, differentiating high-altitude sites with low silica concentrations from others. Sites from the Anapurna and Everest regions, with their contrasting chemistry, were also separated. Community structure was also related to land use: streams draining catchments dominated by terraced agriculture had different communities from those in scrub or forest. This result was confounded, however, by the strong relationship between land use, altitude and chemistry; sites in terracing were at lower altitude, had higher concentrations of silica and a higher proportion of fine sediments than those in the other land-use types. Overall, our data indicate that natural features of the relief and geology in the Himalaya create strong gradients in their invertebrate faunas, but that activities of man may have an effect on stream structure and ecology through catchment management.
Interspecific recognition of alarm cues among guild members through ''eavesdropping'' may allow p... more Interspecific recognition of alarm cues among guild members through ''eavesdropping'' may allow prey to fine-tune antipredator responses. This process may be linked to taxonomic relatedness but might also be influenced by local adaptation to recognize alarm cues from sympatric species. We tested this hypothesis using antipredator responses of a freshwater gastropod Lymnaea stagnalis (L.) to alarm cues from damaged conspecific and 10 heterospecific gastropod species. As predicted, the magnitude of antipredator response decreased significantly with increasing phylogenetic distance, but increased when species were naturally sympatric (defined as species cohabiting in the same water body) with the source population of L. stagnalis. The responses to sympatric species were higher overall, and the relationship between genetic distance and alarm cue response was stronger when tested with sympatric species. This is the first study to demonstrate that population sympatry influences innate antipredator responses to alarm cues from intraguild members and suggests that responses based on phylogenetic relationships can be modified through local adaptation. Such adaptation to heterospecific alarm cues suggests that species could be at a disadvantage when they encounter novel intraguild members resulting from species invasion or range expansion due to a reduction in the presence of reliable information about predation risk.
SUMMARY. 1. Micro-arthropods were surveyed during October 1986 at thirty stony, stream riffle sit... more SUMMARY. 1. Micro-arthropods were surveyed during October 1986 at thirty stony, stream riffle sites in the Ashdown Forest, southern England.2. The importance of a number of physicochemical variables in determining both the distribution of micro-arthropod taxa and community structure was assessed.3. Acidic sites had an impoverished fauna. Total micro-arthropod species richness and densities were highest under circumneutral conditions. and the same patterns were shown by the Hydrachnellae, Harpucticoida and Cladocera. A number of species seemed indifferent to acidic conditions and were widespread and, as a group, the cyclopoid copepods showed no relationship with pH.4. Multiple regression showed that other environmental variables, in particular annual mean temperature and maximum discharge, were also important in explaining the between-site distribution of separate micro-arthropod groups and individual species.5. Other multivariate techniques (ordination, classification and multiple discriminant analysis) showed that impoundment linkage, source distance and conductivity, along with pH, were the most important variables in explaining patterns of species composition between sites.
1. Cross-species macroecological comparisons in freshwater invertebrates have been restricted by ... more 1. Cross-species macroecological comparisons in freshwater invertebrates have been restricted by a lack of large-scale distributional data, and robust phylogenies. Here, we use data from the OdonataCentral database to explore body length–range size and wing length–range size relationships in damselflies from the genus Enallagma; the recent publication of a phylogeny for this group meant that, as well as a cross-species analysis, we were able to assess relationships in a phylogenetically controlled manner.2. For cross-species comparisons, only wing length showed significant (positive) regression relationships with range size and occupancy, although the inclusion of body length in multiple regressions increased the fit of the models. Damselflies with larger wings relative to their body length had larger distributions, a result confirmed by a significant positive relationship between range size and residuals from the regression of wing size on body size.3. For the phylogenetically controlled analyses, only wing length contrast scores were significantly related to distribution patterns and entered into regression models; the significant positive relationships between wing length contrasts and both range size and occupancy contrasts suggested that evolutionary increases in wing length had occurred alongside range expansions.4. Together these results suggest that species of Enallagma with larger wings (both absolute and relative to body length) tend to be more widely distributed in North America and that the evolution of wing size may have played a role in range expansion. No such relationships were evident for body size. We discuss the potential importance of wing morphometrics for studying the evolutionary ecology of freshwater insects.
Freshwater snails and anuran tadpoles have been suggested to have their highest population densit... more Freshwater snails and anuran tadpoles have been suggested to have their highest population densities in ponds of intermediate size where abiotic disturbance (e.g. desiccation) is low and large predators absent. Both snails and tadpoles feed on periphytic algae and, thus, there should be a large potential for competitive interactions to occur between these two distantly related taxa. In a field experiment we examined the relative strength of competition between two closely related snail species, Lymnaea stagnalis and L. peregra, and between L. stagnalis and tadpoles of the common frog, Rana temporaria. Snail growth and egg production and tadpole size at and time to metamorphosis were determined. Effects on the common food source, periphyton, were monitored with the aid of artificial substrates. Periphyton dry weight was dramatically reduced in the presence of snails and/or tadpoles. There were no competitive effects on growth or egg production of the two snail species when they were coexisting. Mortality of L. peregra was high (95%) after reproduction, but independent of treatment. Growth of L. stagnalis was reduced only at the highest tadpole densities, whereas egg production was reduced both by intraspecific competition and by competition with tadpoles. Differences in egg production were retained after tadpole metamorphosis. Tadpole larval period increased, weight of metamorphosing frogs decreased and growth rate was reduced as a function of increasing tadpole density. However, contrary to expectation, snails had a positive effect on tadpole larval period, weight and growth rate. Further, in experimental containers without snails there was a dense growth of the filamentous green alga Cladophora sp. We suggest that the facilitative effects of snails on tadpoles are due to an “indirect mutualistic” mechanism, involving competition between food sources of different quality (microalgae and Cladophora sp.) and tadpoles being competitively dominant over snails for the preferred food source (microalgae). In the presence of tadpoles snails will be forced to feed on low-quality Cladophora, increasing nutrient turnover rates, which results in enhanced productivity of microalgae, increasing tadpole food resources. Thus, tadpoles have a negative effect on snails through resource depression, while snails facilitate tadpole growth through an indirect enhancement of food availability.
ABSTRACT First introduced by Ernst Haeckel in the nineteenth century, the use of developmental se... more ABSTRACT First introduced by Ernst Haeckel in the nineteenth century, the use of developmental sequences has recently seen a renaissance as part of the study of the evolutionary biology of embryos; here we review briefly the literature describing gastropod developmental sequences, appraising the extent to which it has contributed to this renaissance. Gastropods have figured extensively in studies of early development with cell lineage analysis available for numerous taxa. Phylogenetic comparisons of these data reveal strong evolutionary signals, particularly in relation to early cell divisions. In contrast, although the description of post cell division developmental stages, including functional elements of development, in gastropods is extensive, interspecific comparisons are rare and tend to focus instead on developmental mode. However, a recent comparison of the sequence of functional and morphological events in a clade of basommatophoran snails demonstrated several alterations in the timing of developmental events (i.e., heterochronies) across the phylogeny. Many gastropod groups may offer the potential to carry out similar investigations of the evolutionary importance of sequence heterochrony and to try to unravel the mechanistic basis for such patterns in developmental sequences.
SUMMARY. 1. Micro-arthropods were sampled seasonally (January, May, August and October) during 19... more SUMMARY. 1. Micro-arthropods were sampled seasonally (January, May, August and October) during 1986 from ten, stony riffle sites on streams in the Ashdown Forest of southern England, using both standard benthic and interstitial samplers.2. Total densities peaked at most sites in summer. Species richness reached a maximum at acid sites in summer but at cireumneutral sites in autumn, when Hydrachnellae and Cladocera were particularly species rich.3. Individual species showed no obvious differences in seasonally between sites; the majority peaking in summer or autumn, regardless of pH. However, cyclopoid copepods were particularly numerous at acid sites in summer, a pattern not observed at circumneutral sites.4. Multivariate ordination and classification of data sets from the separate seasons, and all four seasons combined, showed that mean site pH, conductivity, and aluminium and calcium concentrations were the most important variables explaining between-site variation in species composition. This clear distinction between the community structure at acidic and circumneutral sites was evident in all seasons except winter. Species composition was also more predictable throughout the year at low-pH sites.5. A number of species were taken consistently in interstitial samples and the cyclopoids Diacydops languidus and D. languidoides were restricted to the hyporheos at circumneutral sites. The similar faunal composition of the hyporheos and the epibenthos indicated that the separation of these communities was not well defined in Ashdown Forest streams.
1. Total species richness for an assemblage or site is a valuable measure in conservation monitor... more 1. Total species richness for an assemblage or site is a valuable measure in conservation monitoring and assessment, but protocols for sampling and species richness determination in wetland habitats such as ponds, bogs or mires remain largely unrefined. 2. Techniques for estimation of total richness of an assemblage based upon replicated sampling offer the opportunity to derive useful estimates of total richness based upon small numbers of samples, and limit sampling-derived disturbance which can be particularly problematic in small aquatic habitats. 3. We quantified the performance of eight of the most commonly encountered estimators of species richness for a variety of littoral zone macrofauna from ponds, comparing estimated richness to maximum richness derived from sampling. 4. Estimates using non-parametric techniques based on species incidence provided the most accurate and precise estimates. The estimators Chao 2 and incidence-based coverage estimator (ICE) from this category were reliable and consistent slight over-estimators; the abundance-based estimator Chao1 also performed well. 5. Species inventory based on relatively small numbers of samples might be significantly improved by use of non-parametric estimators for quantification of species richness. 6. Use of non-parametric estimators of species richness can assist biodiversity inventory by preventing erroneous rankings of habitat richness based upon observed species numbers from limited sampling.
SUMMARY.1The invertebrate colonization of a man-made stream, Flugströmmen, in southern Sweden was... more SUMMARY.1The invertebrate colonization of a man-made stream, Flugströmmen, in southern Sweden was monitored for 18 months in 1988 and 1989. Benthic samples were taken on twelve occasions from three sites (upstream, middle and downstream) and community structure was compared with that at ten natural, permanent reference sites nearby.2The number of species colonizing increased rapidly during the first 3 months. The increase was most rapid upstream but levelled off during the second year at this site, while numbers continued to increase downstream.3Simuliid species were the earliest colonizers and reached high densities at upstream and middle sites during the first year. Ephemeropteran and plecopteran species also occurred early on, whereas Coleoptera, Odonata and Trichoptera were, on average, slower to colonize. Blackfly densities decreased upstream after the first year and hydropsychids became numerically dominant.4The colonization order of functional feeding groups was as predicted: filter feeders first, grazers/collectors intermediate, predators and shredders last.5After a year, the community structure in Flugströmmen closely resembled that in lake-outlet streams situated in the area, although communities at the three sites within the stream were most similar to one another.6The possible role of competitive and predatory processes in determining the observed successional patterns are discussed.The invertebrate colonization of a man-made stream, Flugströmmen, in southern Sweden was monitored for 18 months in 1988 and 1989. Benthic samples were taken on twelve occasions from three sites (upstream, middle and downstream) and community structure was compared with that at ten natural, permanent reference sites nearby.The number of species colonizing increased rapidly during the first 3 months. The increase was most rapid upstream but levelled off during the second year at this site, while numbers continued to increase downstream.Simuliid species were the earliest colonizers and reached high densities at upstream and middle sites during the first year. Ephemeropteran and plecopteran species also occurred early on, whereas Coleoptera, Odonata and Trichoptera were, on average, slower to colonize. Blackfly densities decreased upstream after the first year and hydropsychids became numerically dominant.The colonization order of functional feeding groups was as predicted: filter feeders first, grazers/collectors intermediate, predators and shredders last.After a year, the community structure in Flugströmmen closely resembled that in lake-outlet streams situated in the area, although communities at the three sites within the stream were most similar to one another.The possible role of competitive and predatory processes in determining the observed successional patterns are discussed.
Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus ... more Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus for human settlement and are heavily exploited for water supplies, irrigation, electricity generation, and waste disposal. Lotic systems also have an intimate contact with their catchments and so land-use alterations affect them directly. Here long-term trends in the factors that currently impact running waters are reviewed with the aim of predicting what the main threats to rivers will be in the year 2025. The main ultimate factors forcing change in running waters (ecosystem destruction, physical habitat and water chemistry alteration, and the direct addition or removal of species) stem from proximate influences from urbanization, industry, land-use change and water-course alterations. Any one river is likely to be subjected to several types of impact, and the management of impacts on lotic systems is complicated by numerous links between different forms of anthropogenic effect. Long-term trends for different impacts vary. Concentrations of chemical pollutants such as toxins and nutrients have increased in rivers in developed countries over the past century, with recent reductions for some pollutants (e.g. metals, organic toxicants, acidification), and continued increases in others (e.g. nutrients); there are no long-term chemical data for developing countries. Dam construction increased rapidly during the twentieth century, peaking in the 1970s, and the number of reservoirs has stabilized since this time, whereas the transfer of exotic species between lotic systems continues to increase. Hence, there have been some success stories in the attempts to reduce the impacts from anthropogenic impacts in developed nations. Improvements in the pH status of running waters should continue with lower sulphurous emissions, although emissions of nitrous oxides are set to continue under current legislation and will continue to contribute to acidification and nutrient loadings. Climate change also will impact running waters through alterations in hydrology and thermal regimes, although precise predictions are problematic; effects are likely to vary between regions and to operate alongside rather than override those from other impacts. Effects from climate change may be more extreme over longer time scales (>50 years). The overriding pressure on running water ecosystems up to 2025 will stem from the predicted increase in the human population, with concomitant increases in urban development, industry, agricultural activities and water abstraction, diversion and damming. Future degradation could be substantial and rapid (c. 10 years) and will be concentrated in those areas of the world where resources for conservation are most limited and knowledge of lotic ecosystems most incomplete; damage will centre on lowland rivers, which are also relatively poorly studied. Changes in management practices and public awareness do appear to be benefiting running water ecosystems in developed countries, and could underpin conservation strategies in developing countries if they were implemented in a relevant way.
Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus ... more Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus for human settlement and are heavily exploited for water supplies, irrigation, electricity generation, and waste disposal. Lotic systems also have an intimate contact with their catchments and so land-use alterations affect them directly. Here long-term trends in the factors that currently impact running waters are reviewed with the aim of predicting what the main threats to rivers will be in the year 2025. The main ultimate factors forcing change in running waters (ecosystem destruction, physical habitat and water chemistry alteration, and the direct addition or removal of species) stem from proximate influences from urbanization, industry, land-use change and water-course alterations. Any one river is likely to be subjected to several types of impact, and the management of impacts on lotic systems is complicated by numerous links between different forms of anthropogenic effect. Long-term trends for different impacts vary. Concentrations of chemical pollutants such as toxins and nutrients have increased in rivers in developed countries over the past century, with recent reductions for some pollutants (e.g. metals, organic toxicants, acidification), and continued increases in others (e.g. nutrients); there are no long-term chemical data for developing countries. Dam construction increased rapidly during the twentieth century, peaking in the 1970s, and the number of reservoirs has stabilized since this time, whereas the transfer of exotic species between lotic systems continues to increase. Hence, there have been some success stories in the attempts to reduce the impacts from anthropogenic impacts in developed nations. Improvements in the pH status of running waters should continue with lower sulphurous emissions, although emissions of nitrous oxides are set to continue under current legislation and will continue to contribute to acidification and nutrient loadings. Climate change also will impact running waters through alterations in hydrology and thermal regimes, although precise predictions are problematic; effects are likely to vary between regions and to operate alongside rather than override those from other impacts. Effects from climate change may be more extreme over longer time scales (>50 years). The overriding pressure on running water ecosystems up to 2025 will stem from the predicted increase in the human population, with concomitant increases in urban development, industry, agricultural activities and water abstraction, diversion and damming. Future degradation could be substantial and rapid (c. 10 years) and will be concentrated in those areas of the world where resources for conservation are most limited and knowledge of lotic ecosystems most incomplete; damage will centre on lowland rivers, which are also relatively poorly studied. Changes in management practices and public awareness do appear to be benefiting running water ecosystems in developed countries, and could underpin conservation strategies in developing countries if they were implemented in a relevant way.
antipredator behaviour chemical communication cohabitation gastropod great pond snail learning Ly... more antipredator behaviour chemical communication cohabitation gastropod great pond snail learning Lymnaea stagnalis phylogeny In aquatic habitats prey often show maximum responsiveness to predators when they are exposed to kairomones and alarm cues in combination. As alarm cues can derive from heterospecific as well as conspecific animals, prey may need to fine tune their responses depending on the taxa with which they co-occur. We used juveniles of the freshwater gastropod, Lymnaea stagnalis L. (great pond snail) to test whether the response to heterospecific alarm cues could be enhanced by cohabitation in the absence of predators, and therefore without the potential for cue association learning. Snails were raised from oviposition alongside one of four other pulmonate species, differing in their phylogenetic relatedness to L. stagnalis, that is, from intragenus to intrasuborder. The antipredator response to heterospecific alarm cues paired with fish kairomones was then compared with the response of L. stagnalis reared in isolation. Cohabitation increased the response to heterospecific alarm cues paired with fish kairomones only when L. stagnalis had developed in the presence of closely related (Lymnaea fusca and Radix balthica) rather than distantly related heterospecifics (Physella acuta and Planorbarius corneus). Hence, in L. stagnalis, learning to recognize heterospecific alarm cues can occur without the need for cue association but the learnt response is constrained by species' relatedness. This result suggests that differential responses observed in studies based on wild-caught animals in response to heterospecific alarm cues may be explained by learning through cohabitation alone. Ó
1. Hydrobiological changes were assessed along an altirudinal transect of eighteen to twenty-thre... more 1. Hydrobiological changes were assessed along an altirudinal transect of eighteen to twenty-three tributaries from 600 to 3750m in two adjacent river systems in east-central Nepal. The transect incorporated catchments under terraced agriculture at the lowest altitudes in the Likhu Khola, through streams in forest, alpine scrub and tundra at higher altitudes in Langtang.2. Diatoms, bryophytes, macroinvertebrates and fish all showed pronounced altitudinal changes in assemblage composition as shown by TWINSPAN and DECORANA. A few taxa were restricted to streams at high altitude, but many more occurred only at lower altitudes where taxon richness increased substantially despite catchment disturbance by terraced agriculture.3. Diatoms characteristic of lower altitude streams were mostly motile, epipelic or episammic Navicula and Nitzschia spp., which occur typically at greater electrolyte and nutrient concentrations. Those characteristic of higher and steeper sites included attached Fragilaria spp. and prostrate Achnanthes spp., tolerant of turbulent flow.4. Cover by bryophytes varied within catchment type; high altitude springs supported dense mats, unlike streams fed by ice and glaciers. Taxa confined to low altitudes included those characteristic of humid subtropical conditions.5. Invertebrate families occurring only at lower altitudes included a range of burrowers and pool dwellers. Numerically, filter feeding Hydropsychidae and Simuliidae dominated streams in terraced and forested catchments, whereas grazing baetid mayflies dominated higher altitude streams in scrub and tundra.6. The combined density and biomass of at least six fish species in the Likhu Khola were 23–250 (per 100m−2), and 86–1282 g wet mass (per 100 m−2), respectively. No fish were found in Langtang streams, probably because torrential headwaters prevented colonization.7. Our data confirm that altitudinal transitions in stream biota are pronounced in the Himalaya of Nepal, but are likely to reflect a wide array of potential influences.
1 Macroinvertebrates were sampled in the riffles of fifty-eight streams from three regions of the... more 1 Macroinvertebrates were sampled in the riffles of fifty-eight streams from three regions of the Himalaya (Anapurna, Langtang and Everest) in Nepal. A semi-quantitative method with identification to family level was used to describe communities on-site.2 Stream physicochemistry was assessed and the community structure of macroinvertebrates was related to chemistry, physiography (substratum composition, altitude and size), geographical location and the dominant land use in each catchment (terraced agriculture, forest or scrub). Community data were analysed by ordination (DECORANA) and classification (TWINSPAN).3 The concentration of cations in stream water decreased significantly with altitude. Chemistry also differed between regions; sites from Anapurna had a higher pH and conductivity than those in the other two areas.4 Communities were dominated by aquatic insect larvae, with Ephemeroptera, in particular the Baetidae, most numerous across sites.5 There were, nevertheless, differences in community structure between sites, which were related closely to stream physicochemistry. Ordination scores were strongly correlated with altitude, magnesium concentration and substratum composition. Classification was also linked to altitude and chemistry, differentiating high-altitude sites with low silica concentrations from others. Sites from the Anapurna and Everest regions, with their contrasting chemistry, were also separated.6 Community structure was also related to land use: streams draining catchments dominated by terraced agriculture had different communities from those in scrub or forest. This result was confounded, however, by the strong relationship between land use, altitude and chemistry; sites in terracing were at lower altitude, had higher concentrations of silica and a higher proportion of fine sediments than those in the other land-use types.7 Overall, our data indicate that natural features of the relief and geology in the Himalaya create strong gradients in their invertebrate faunas, but that activities of man may have an effect on stream structure and ecology through catchment management. Macroinvertebrates were sampled in the riffles of fifty-eight streams from three regions of the Himalaya (Anapurna, Langtang and Everest) in Nepal. A semi-quantitative method with identification to family level was used to describe communities on-site. Stream physicochemistry was assessed and the community structure of macroinvertebrates was related to chemistry, physiography (substratum composition, altitude and size), geographical location and the dominant land use in each catchment (terraced agriculture, forest or scrub). Community data were analysed by ordination (DECORANA) and classification (TWINSPAN). The concentration of cations in stream water decreased significantly with altitude. Chemistry also differed between regions; sites from Anapurna had a higher pH and conductivity than those in the other two areas. Communities were dominated by aquatic insect larvae, with Ephemeroptera, in particular the Baetidae, most numerous across sites. There were, nevertheless, differences in community structure between sites, which were related closely to stream physicochemistry. Ordination scores were strongly correlated with altitude, magnesium concentration and substratum composition. Classification was also linked to altitude and chemistry, differentiating high-altitude sites with low silica concentrations from others. Sites from the Anapurna and Everest regions, with their contrasting chemistry, were also separated. Community structure was also related to land use: streams draining catchments dominated by terraced agriculture had different communities from those in scrub or forest. This result was confounded, however, by the strong relationship between land use, altitude and chemistry; sites in terracing were at lower altitude, had higher concentrations of silica and a higher proportion of fine sediments than those in the other land-use types. Overall, our data indicate that natural features of the relief and geology in the Himalaya create strong gradients in their invertebrate faunas, but that activities of man may have an effect on stream structure and ecology through catchment management.
Interspecific recognition of alarm cues among guild members through ''eavesdropping'' may allow p... more Interspecific recognition of alarm cues among guild members through ''eavesdropping'' may allow prey to fine-tune antipredator responses. This process may be linked to taxonomic relatedness but might also be influenced by local adaptation to recognize alarm cues from sympatric species. We tested this hypothesis using antipredator responses of a freshwater gastropod Lymnaea stagnalis (L.) to alarm cues from damaged conspecific and 10 heterospecific gastropod species. As predicted, the magnitude of antipredator response decreased significantly with increasing phylogenetic distance, but increased when species were naturally sympatric (defined as species cohabiting in the same water body) with the source population of L. stagnalis. The responses to sympatric species were higher overall, and the relationship between genetic distance and alarm cue response was stronger when tested with sympatric species. This is the first study to demonstrate that population sympatry influences innate antipredator responses to alarm cues from intraguild members and suggests that responses based on phylogenetic relationships can be modified through local adaptation. Such adaptation to heterospecific alarm cues suggests that species could be at a disadvantage when they encounter novel intraguild members resulting from species invasion or range expansion due to a reduction in the presence of reliable information about predation risk.
SUMMARY. 1. Micro-arthropods were surveyed during October 1986 at thirty stony, stream riffle sit... more SUMMARY. 1. Micro-arthropods were surveyed during October 1986 at thirty stony, stream riffle sites in the Ashdown Forest, southern England.2. The importance of a number of physicochemical variables in determining both the distribution of micro-arthropod taxa and community structure was assessed.3. Acidic sites had an impoverished fauna. Total micro-arthropod species richness and densities were highest under circumneutral conditions. and the same patterns were shown by the Hydrachnellae, Harpucticoida and Cladocera. A number of species seemed indifferent to acidic conditions and were widespread and, as a group, the cyclopoid copepods showed no relationship with pH.4. Multiple regression showed that other environmental variables, in particular annual mean temperature and maximum discharge, were also important in explaining the between-site distribution of separate micro-arthropod groups and individual species.5. Other multivariate techniques (ordination, classification and multiple discriminant analysis) showed that impoundment linkage, source distance and conductivity, along with pH, were the most important variables in explaining patterns of species composition between sites.
1. Cross-species macroecological comparisons in freshwater invertebrates have been restricted by ... more 1. Cross-species macroecological comparisons in freshwater invertebrates have been restricted by a lack of large-scale distributional data, and robust phylogenies. Here, we use data from the OdonataCentral database to explore body length–range size and wing length–range size relationships in damselflies from the genus Enallagma; the recent publication of a phylogeny for this group meant that, as well as a cross-species analysis, we were able to assess relationships in a phylogenetically controlled manner.2. For cross-species comparisons, only wing length showed significant (positive) regression relationships with range size and occupancy, although the inclusion of body length in multiple regressions increased the fit of the models. Damselflies with larger wings relative to their body length had larger distributions, a result confirmed by a significant positive relationship between range size and residuals from the regression of wing size on body size.3. For the phylogenetically controlled analyses, only wing length contrast scores were significantly related to distribution patterns and entered into regression models; the significant positive relationships between wing length contrasts and both range size and occupancy contrasts suggested that evolutionary increases in wing length had occurred alongside range expansions.4. Together these results suggest that species of Enallagma with larger wings (both absolute and relative to body length) tend to be more widely distributed in North America and that the evolution of wing size may have played a role in range expansion. No such relationships were evident for body size. We discuss the potential importance of wing morphometrics for studying the evolutionary ecology of freshwater insects.
Freshwater snails and anuran tadpoles have been suggested to have their highest population densit... more Freshwater snails and anuran tadpoles have been suggested to have their highest population densities in ponds of intermediate size where abiotic disturbance (e.g. desiccation) is low and large predators absent. Both snails and tadpoles feed on periphytic algae and, thus, there should be a large potential for competitive interactions to occur between these two distantly related taxa. In a field experiment we examined the relative strength of competition between two closely related snail species, Lymnaea stagnalis and L. peregra, and between L. stagnalis and tadpoles of the common frog, Rana temporaria. Snail growth and egg production and tadpole size at and time to metamorphosis were determined. Effects on the common food source, periphyton, were monitored with the aid of artificial substrates. Periphyton dry weight was dramatically reduced in the presence of snails and/or tadpoles. There were no competitive effects on growth or egg production of the two snail species when they were coexisting. Mortality of L. peregra was high (95%) after reproduction, but independent of treatment. Growth of L. stagnalis was reduced only at the highest tadpole densities, whereas egg production was reduced both by intraspecific competition and by competition with tadpoles. Differences in egg production were retained after tadpole metamorphosis. Tadpole larval period increased, weight of metamorphosing frogs decreased and growth rate was reduced as a function of increasing tadpole density. However, contrary to expectation, snails had a positive effect on tadpole larval period, weight and growth rate. Further, in experimental containers without snails there was a dense growth of the filamentous green alga Cladophora sp. We suggest that the facilitative effects of snails on tadpoles are due to an “indirect mutualistic” mechanism, involving competition between food sources of different quality (microalgae and Cladophora sp.) and tadpoles being competitively dominant over snails for the preferred food source (microalgae). In the presence of tadpoles snails will be forced to feed on low-quality Cladophora, increasing nutrient turnover rates, which results in enhanced productivity of microalgae, increasing tadpole food resources. Thus, tadpoles have a negative effect on snails through resource depression, while snails facilitate tadpole growth through an indirect enhancement of food availability.
ABSTRACT First introduced by Ernst Haeckel in the nineteenth century, the use of developmental se... more ABSTRACT First introduced by Ernst Haeckel in the nineteenth century, the use of developmental sequences has recently seen a renaissance as part of the study of the evolutionary biology of embryos; here we review briefly the literature describing gastropod developmental sequences, appraising the extent to which it has contributed to this renaissance. Gastropods have figured extensively in studies of early development with cell lineage analysis available for numerous taxa. Phylogenetic comparisons of these data reveal strong evolutionary signals, particularly in relation to early cell divisions. In contrast, although the description of post cell division developmental stages, including functional elements of development, in gastropods is extensive, interspecific comparisons are rare and tend to focus instead on developmental mode. However, a recent comparison of the sequence of functional and morphological events in a clade of basommatophoran snails demonstrated several alterations in the timing of developmental events (i.e., heterochronies) across the phylogeny. Many gastropod groups may offer the potential to carry out similar investigations of the evolutionary importance of sequence heterochrony and to try to unravel the mechanistic basis for such patterns in developmental sequences.
SUMMARY. 1. Micro-arthropods were sampled seasonally (January, May, August and October) during 19... more SUMMARY. 1. Micro-arthropods were sampled seasonally (January, May, August and October) during 1986 from ten, stony riffle sites on streams in the Ashdown Forest of southern England, using both standard benthic and interstitial samplers.2. Total densities peaked at most sites in summer. Species richness reached a maximum at acid sites in summer but at cireumneutral sites in autumn, when Hydrachnellae and Cladocera were particularly species rich.3. Individual species showed no obvious differences in seasonally between sites; the majority peaking in summer or autumn, regardless of pH. However, cyclopoid copepods were particularly numerous at acid sites in summer, a pattern not observed at circumneutral sites.4. Multivariate ordination and classification of data sets from the separate seasons, and all four seasons combined, showed that mean site pH, conductivity, and aluminium and calcium concentrations were the most important variables explaining between-site variation in species composition. This clear distinction between the community structure at acidic and circumneutral sites was evident in all seasons except winter. Species composition was also more predictable throughout the year at low-pH sites.5. A number of species were taken consistently in interstitial samples and the cyclopoids Diacydops languidus and D. languidoides were restricted to the hyporheos at circumneutral sites. The similar faunal composition of the hyporheos and the epibenthos indicated that the separation of these communities was not well defined in Ashdown Forest streams.
1. Total species richness for an assemblage or site is a valuable measure in conservation monitor... more 1. Total species richness for an assemblage or site is a valuable measure in conservation monitoring and assessment, but protocols for sampling and species richness determination in wetland habitats such as ponds, bogs or mires remain largely unrefined. 2. Techniques for estimation of total richness of an assemblage based upon replicated sampling offer the opportunity to derive useful estimates of total richness based upon small numbers of samples, and limit sampling-derived disturbance which can be particularly problematic in small aquatic habitats. 3. We quantified the performance of eight of the most commonly encountered estimators of species richness for a variety of littoral zone macrofauna from ponds, comparing estimated richness to maximum richness derived from sampling. 4. Estimates using non-parametric techniques based on species incidence provided the most accurate and precise estimates. The estimators Chao 2 and incidence-based coverage estimator (ICE) from this category were reliable and consistent slight over-estimators; the abundance-based estimator Chao1 also performed well. 5. Species inventory based on relatively small numbers of samples might be significantly improved by use of non-parametric estimators for quantification of species richness. 6. Use of non-parametric estimators of species richness can assist biodiversity inventory by preventing erroneous rankings of habitat richness based upon observed species numbers from limited sampling.
SUMMARY.1The invertebrate colonization of a man-made stream, Flugströmmen, in southern Sweden was... more SUMMARY.1The invertebrate colonization of a man-made stream, Flugströmmen, in southern Sweden was monitored for 18 months in 1988 and 1989. Benthic samples were taken on twelve occasions from three sites (upstream, middle and downstream) and community structure was compared with that at ten natural, permanent reference sites nearby.2The number of species colonizing increased rapidly during the first 3 months. The increase was most rapid upstream but levelled off during the second year at this site, while numbers continued to increase downstream.3Simuliid species were the earliest colonizers and reached high densities at upstream and middle sites during the first year. Ephemeropteran and plecopteran species also occurred early on, whereas Coleoptera, Odonata and Trichoptera were, on average, slower to colonize. Blackfly densities decreased upstream after the first year and hydropsychids became numerically dominant.4The colonization order of functional feeding groups was as predicted: filter feeders first, grazers/collectors intermediate, predators and shredders last.5After a year, the community structure in Flugströmmen closely resembled that in lake-outlet streams situated in the area, although communities at the three sites within the stream were most similar to one another.6The possible role of competitive and predatory processes in determining the observed successional patterns are discussed.The invertebrate colonization of a man-made stream, Flugströmmen, in southern Sweden was monitored for 18 months in 1988 and 1989. Benthic samples were taken on twelve occasions from three sites (upstream, middle and downstream) and community structure was compared with that at ten natural, permanent reference sites nearby.The number of species colonizing increased rapidly during the first 3 months. The increase was most rapid upstream but levelled off during the second year at this site, while numbers continued to increase downstream.Simuliid species were the earliest colonizers and reached high densities at upstream and middle sites during the first year. Ephemeropteran and plecopteran species also occurred early on, whereas Coleoptera, Odonata and Trichoptera were, on average, slower to colonize. Blackfly densities decreased upstream after the first year and hydropsychids became numerically dominant.The colonization order of functional feeding groups was as predicted: filter feeders first, grazers/collectors intermediate, predators and shredders last.After a year, the community structure in Flugströmmen closely resembled that in lake-outlet streams situated in the area, although communities at the three sites within the stream were most similar to one another.The possible role of competitive and predatory processes in determining the observed successional patterns are discussed.
Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus ... more Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus for human settlement and are heavily exploited for water supplies, irrigation, electricity generation, and waste disposal. Lotic systems also have an intimate contact with their catchments and so land-use alterations affect them directly. Here long-term trends in the factors that currently impact running waters are reviewed with the aim of predicting what the main threats to rivers will be in the year 2025. The main ultimate factors forcing change in running waters (ecosystem destruction, physical habitat and water chemistry alteration, and the direct addition or removal of species) stem from proximate influences from urbanization, industry, land-use change and water-course alterations. Any one river is likely to be subjected to several types of impact, and the management of impacts on lotic systems is complicated by numerous links between different forms of anthropogenic effect. Long-term trends for different impacts vary. Concentrations of chemical pollutants such as toxins and nutrients have increased in rivers in developed countries over the past century, with recent reductions for some pollutants (e.g. metals, organic toxicants, acidification), and continued increases in others (e.g. nutrients); there are no long-term chemical data for developing countries. Dam construction increased rapidly during the twentieth century, peaking in the 1970s, and the number of reservoirs has stabilized since this time, whereas the transfer of exotic species between lotic systems continues to increase. Hence, there have been some success stories in the attempts to reduce the impacts from anthropogenic impacts in developed nations. Improvements in the pH status of running waters should continue with lower sulphurous emissions, although emissions of nitrous oxides are set to continue under current legislation and will continue to contribute to acidification and nutrient loadings. Climate change also will impact running waters through alterations in hydrology and thermal regimes, although precise predictions are problematic; effects are likely to vary between regions and to operate alongside rather than override those from other impacts. Effects from climate change may be more extreme over longer time scales (>50 years). The overriding pressure on running water ecosystems up to 2025 will stem from the predicted increase in the human population, with concomitant increases in urban development, industry, agricultural activities and water abstraction, diversion and damming. Future degradation could be substantial and rapid (c. 10 years) and will be concentrated in those areas of the world where resources for conservation are most limited and knowledge of lotic ecosystems most incomplete; damage will centre on lowland rivers, which are also relatively poorly studied. Changes in management practices and public awareness do appear to be benefiting running water ecosystems in developed countries, and could underpin conservation strategies in developing countries if they were implemented in a relevant way.
Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus ... more Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus for human settlement and are heavily exploited for water supplies, irrigation, electricity generation, and waste disposal. Lotic systems also have an intimate contact with their catchments and so land-use alterations affect them directly. Here long-term trends in the factors that currently impact running waters are reviewed with the aim of predicting what the main threats to rivers will be in the year 2025. The main ultimate factors forcing change in running waters (ecosystem destruction, physical habitat and water chemistry alteration, and the direct addition or removal of species) stem from proximate influences from urbanization, industry, land-use change and water-course alterations. Any one river is likely to be subjected to several types of impact, and the management of impacts on lotic systems is complicated by numerous links between different forms of anthropogenic effect. Long-term trends for different impacts vary. Concentrations of chemical pollutants such as toxins and nutrients have increased in rivers in developed countries over the past century, with recent reductions for some pollutants (e.g. metals, organic toxicants, acidification), and continued increases in others (e.g. nutrients); there are no long-term chemical data for developing countries. Dam construction increased rapidly during the twentieth century, peaking in the 1970s, and the number of reservoirs has stabilized since this time, whereas the transfer of exotic species between lotic systems continues to increase. Hence, there have been some success stories in the attempts to reduce the impacts from anthropogenic impacts in developed nations. Improvements in the pH status of running waters should continue with lower sulphurous emissions, although emissions of nitrous oxides are set to continue under current legislation and will continue to contribute to acidification and nutrient loadings. Climate change also will impact running waters through alterations in hydrology and thermal regimes, although precise predictions are problematic; effects are likely to vary between regions and to operate alongside rather than override those from other impacts. Effects from climate change may be more extreme over longer time scales (>50 years). The overriding pressure on running water ecosystems up to 2025 will stem from the predicted increase in the human population, with concomitant increases in urban development, industry, agricultural activities and water abstraction, diversion and damming. Future degradation could be substantial and rapid (c. 10 years) and will be concentrated in those areas of the world where resources for conservation are most limited and knowledge of lotic ecosystems most incomplete; damage will centre on lowland rivers, which are also relatively poorly studied. Changes in management practices and public awareness do appear to be benefiting running water ecosystems in developed countries, and could underpin conservation strategies in developing countries if they were implemented in a relevant way.
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