The importance of acid episodes in determining faunal distributions in Welsh streams

SUMMARY. 1. In regional studies of surface-water acidification, annual means of chemical variables are often used to describe differences and change. Outputs from hydrochemical models are often in the form of mean values, which are used in biological models, and these, in turn, are usually derived from responses to mean conditions. Thus, biological forecasts are constrained to ignore the possible effects of the short-term variations in water chemistry which characterize acid streams. This approach requires appraisal.
2. Here, regional Welsh data and daily records from a smaller number of streams were used to investigate the estimation of pH parameters. Variations in aluminium concentration in relation to pH were also assessed. Empirical relationships between invertebrate assemblages, fish populations, mean stream chemistry and measures of fluctuation in pH and aluminium concentration were explored.
3. In general, pH or Al variability and mean pH or Al in Welsh streams were closely related, so that the biological influences of episodes could not easily be separated from those of chronic conditions. Mean pH and mean aluminium concentration were the most effective pH and aluminium statistics used in multivariate models of trout density, which were not improved by including other pH or aluminium variables. For models of invertebrate assemblages based on mean pH or mean aluminium, the inclusion of variables related to episodicity (e.g. pH minimum, aluminium maximum) gave moderate increases in precision.
4. This analysis indicates that it is reasonable to use means of stream chemical variables in biological models of acidification. Consideration of chemical variability could give improvements in some cases, but at the expense of increased model complexity and effort in parameter selection. Nevertheless, we emphasize the need for accurate calibration of both biological and hydrochemical models.     

The relative importance of refugia in determining the drift and habitat selection of predaceous stoneflies in a sandy-bottomed stream

Patch structure in sandy, compared to rocky streams, is characterized by isolated snags that can only be colonized by drifting. By measuring drift from patches (snags) of various quality we determined the factors that influence habitat selection and drift of the predaceous stoneflies Acroneuria abnormis and Paragnetina fumosa. The presence of refugia (loose bark and leaf packs) was more important than hunger level and modified the effects of increased predator densities and aggressive interactions. Stoneflies concentrated to 8x natural densities with access to refugia remained longer on snags than a single stonefly without access to refugia. During periods of activity, refugia were defended with larger stoneflies always displacing smaller nymphs. During long periods of inactivity, two and sometimes three nymphs would rest side-by-side sharing the same refuge. Hunger level (starved versus satiated stoneflies), an indirect measure of a predator's response to prey availability, had no significant effect on drift or habitat selection regardless of the presence of refugia. Stonefly predators had a uniform distribution while their prey were clumped. Drift was deliberate and almost always delayed until night, usually at dusk or dawn. An examination of previous research plus the results of this study suggest that non-predatory intra- and interspecific interactions can be an important mechanism causing drift in streams.     

The importance of meiofauna to lotic ecosystem functioning

• 1 Although meiofauna occur in large numbers in many streams, almost nothing is known about their functional role.
• 2 In other systems, meiofauna influence microbial and organic matter dynamics through consumption and bioturbation. Given that these are important processes in streams, meiofauna have the potential to influence lotic function by changing the quality and availability of organic matter as well as the number and biotic activity of benthic microbes. Selective feeding by meiofauna has the potential to alter the availability of nutrients and organic carbon.
• 3 Meiofauna generally contribute only a small amount to metazoan production and biomass in streams, although exceptions occur. Within a stream, the relative importance of meiofauna may reflect whether the temporary or permanent meiofauna dominate the meiobenthos as well as the season when samples are collected.
• 4 We suggest stream conditions (small sediment grain size, restricted interstitial flow) under which meiofauna have the greatest likelihood of influencing stream ecosystem function.
• 5 Important areas for future research include addressing whether meiofauna feed selectively, whether meiofauna are links or sinks for carbon in streams, and whether bioturbation by meiofauna influences stream ecosystem processes in a predictable manner.

     

Spatially heterogeneous refugia and predation risk in intertidal salt marshes

The effect of habitat structure on interactions between predators and prey may vary spatially. In estuarine salt marshes, heterogeneity in refuge quality derives from spatial variation in vegetation structure and in tidal inundation. We investigated whether predation by blue crabs on periwinkle snails was influenced by distance from the seaward edge of the salt marsh and by characteristics of the primary habitat structure, smooth cordgrass ( Spartina alterniflora ). Spartina may provide refuge for snails and interfere with foraging by crabs. Furthermore, predation risk should decline with distance from the seaward edge because landward regions require more travel time for crabs during tidal inundation. We investigated these processes using a comparative survey of snails and habitat traits, an experiment that assessed the crab population and measured predation risk, and a size-structured model that estimated encounter rates. Taken together, these approaches indicated that predation risk for snails was lower where Spartina was present and was lower in a landward direction. Furthermore, Spartina architecture and distance from the seaward edge interacted. The strength of the predation gradient between seaward and landward regions of the marsh was greater where Spartina was tall or dense. These predation gradients emerge because vegetation and distance inland decrease encounter rates between crabs and snails. This study suggests that habitat modification, a process not uncommon in salt marshes, may have consequences for interactions among intertidal fauna.     

Gut passage and insect grazer selectivity of lotic diatoms

SUMMARY 1. Grazing experiments were conducted in a small Kentucky stream to determine if epilithic diatom taxa differed in susceptibility to grazing by the larval caddisfly, Neophylax autumnus , and whether ingested taxa varied in digestibility. Live/dead ratios of diatom cells from N. autumnus faecal material were compared with live/dead ratios of cells taken from adjacent epilithic habitats to determine diatom digestibility. Selectivity was studied by comparing relative abundances of diatom taxa on substrata with the relative abundances of these taxa in grazer faecal material.
2. 73% of the diatoms collected from epilithic habitats contained intact chloroplasts and were characterized as live cells. Only 42% of diatoms eliminated in caddisfly faeces were living. The five dominant diatom taxa did not differ in digestibility. Elimination of viable diatoms in grazer faecal material may contribute to diatom drift in streams, and thus supply individuals for downstream recolonization. Failure to document live/dead ratios of diatoms from grazed substrata can lead to overestimation of the digestibility of taxa and misinterpretation of results.
3. Grazers in this investigation were more successful at ingesting large, high-profile diatom taxa (i.e. Cymbella, Meridion and Gomphonema ) and less able to remove small, adnate forms ( Achnanthes minutissima ). Such 'selectivity'may be one mechanism by which A. minutissima maintains dominance in this system.     

The relative importance of deforestation,precipitation change,and temperature sensitivity in determining the future distributions and diversity of Amazonian plant species

Tropical forests are threatened by many human disturbances – two of the most important of which are deforestation and climate change. To mitigate the impacts of these disturbances, it is important to understand their potential effects on the distributions of species. In the tropics, such understanding has been hindered by poor knowledge of the current distributions and range limits of most species. Here, we use herbarium collection records to model the current and future distributions of ca. 3000 Amazonian plant species. We project these distributions into the future under a range of different scenarios related to the magnitude of climate change and extent of deforestation as well as the response of species to changes in temperature, precipitation, and atmospheric concentrations of CO₂ . We find that the future of Amazonian diversity will be dependant primarily on the ability of species to tolerate or adapt to rising temperatures. If the thermal niches of tropical plant species are fixed and incapable of expanding under rapid warming, then the negative effects of climate change will overshadow the effects of deforestation, greatly reducing the area of suitable habitat available to most species and potentially leading to massive losses of biodiversity throughout the Amazon. If tropical species are generally capable of tolerating warmer temperatures, rates of habitat loss will be greatly reduced but many parts of Amazonia may still experience rapid losses of diversity, with the effects of enhanced seasonal water stress being similar in magnitude to the effects of deforestation.     

The relative importance of climate and habitat in determining the distributions of species at different spatial scales: a case study with ground beetles in Great Britain

Experimental studies have shown that many species show preferences for different climatic conditions, or may die in unsuitable conditions. Climate envelope models have been used frequently in recent years to predict the presence and absence of species at large spatial scales. However, many authors have postulated that the distributions of species at smaller spatial scales are determined by factors such as habitat availability and biotic interactions. Climatic effects are often assumed by modellers to be unimportant at fine resolutions, but few studies have actually tested this. We sampled the distributions of 20 beetle species of the family Carabidae across three study sites by pitfall trapping, and at the national scale from monitoring data. Statistical models were constructed to determine which of two sets of environmental variables (temperature or broad habitat type) best accounted for the observed data at the three sites and at the national (Great Britain) scale. High‐resolution temperature variables frequently produced better models (as determined by AIC) than habitat features when modelling the distributions of species at a local scale, within the three study sites. Conversely, habitat was always a better predictor than temperature when describing species’ distributions at a coarse scale within Great Britain. Northerly species were most likely to occur in cool micro‐sites within the study sites, whereas southerly species were most likely to occur in warm micro‐sites. Effects of microclimate were not limited to species at the edges of their distribution, and fine‐resolution temperature surfaces should therefore ideally be utilised when undertaking climate‐envelope modelling.     

Competition, predation, and the distributions of four desert anurans

Several studies have shown that larval competition and susceptibility to predation affect distributions of amphibian assemblages across ephemeral and perennial habitats. However, few studies have examined mechanisms affecting distribution patterns and site use of anurans adapted to highly ephemeral habitats. This study examines hypotheses about competition and predation as mechanisms creating non-overlapping patterns of site use in four anurans that breed in highly ephemeral habitats: Scaphiopus couchii (Couch's spade-foot toad), Gastrophryne olivacea (narrow-mouthed toad), Bufo speciosus (Texas toad), and Bufo punctatus (red-spotted toad). These four anurans showed a significantly nonrandom pattern of co-occurrence. Only 12% of 95 ephemeral breeding sites surveyed were occupied by more than one species. We tested the hypotheses that non-overlapping use of breeding sites was due to activity rates of their tadpoles that in turn reflect their competitive ability and susceptibility to predation. Tadpoles of S. couchii were significantly more active and more susceptible to predation than were tadpoles of G. olivacea, B. speciosus, and B. punctatus. The masses of G. olivacea, B. speciosus, and B. punctatus were less when they were reared with S. couchii, demonstrating the possible competitive dominance of S. couchii. These results suggest that the competitive ability of S. couchii may play a role in excluding G. olivacea, B. speciosus, and B. punctatus from very ephemeral breeding sites, and that susceptibility to predation could play a role in excluding S. couchii from breeding sites of longer duration that are more likely to be colonized by aquatic predators.     

Patterns of spatial resource use in lotic invertebrate assemblages

Spatial resource overlap by benthic invertebrates was investigated between stones in 11 freshwater habitats (10 streams and a wind-swept lake shore) of similar physicochemical nature but different thermal and hydrologic stability in the Cass-Craigieburn region, New Zealand. Both overlap and breadth of stone use by the invertebrate taxa decreased as stability declined, in part because of the lower number of species available with which to share resources at unstable sites. Although there was no evidence for resource partitioning of stones, communities at stable sites and on more stable substrates differed from those that would have been expected to accumulate by chance colonisation alone. Stone assemblages at unstable sites are dominated by the same general suite of species well suited to surviving disturbances, while conditions at stable sites allow more specialised taxa to colonise individual stones differentially.     

The relative importance of predation and competition in two reef fishes

Competition and predation may both strongly influence populations of reef fishes, but the importance of these processes relative to one another is poorly understood. I quantified the effects of predation and competition on the growth and survival of two temperate reef fishes, Lythrypnus dalli and Coryphopterus nicholsii, in field experiments in which I manipulated the densities of the two species and the abundance of predators (using exclosure cages) on small replicate patch reefs. I also evaluated the influence of predators on the behavior of the two species to help interpret the mechanisms of any predatory influences on growth or survival. Predation was much more important than competition (inter- or intraspecific) in Lythrypnus. For Coryphopterus, neither competition nor predation were particularly important. Behaviorally, both species responded to predators by reducing foraging rate and hiding. This altered behavior, however, had no repercussions for growth or survival of Coryphopterus. In contrast, Lythrypnus grew more slowly and suffered greater mortality when exposed to predators. Interspecific competition did not significantly influence either species. Intraspecific competition did not affect the growth of Coryphopterus, but survival tended to be lower at high densities. Growth of Lythrypnus was depressed by intraspecific competition, but survival was not, except that, in the presence of predators, survival was density dependent. In contrast to the historical emphasis placed on the role of competition, this study indicates that predation can be more important than competition in determining patterns of abundance of some reef fishes. For example, predators not only influenced foraging of both Lythrypnus and Coryphopterus, but they also reduced growth and survival of Lythrypnus, and therefore appear to help maintain the marked habitat segregation between the two species. Received: 16 June 1997 / Accepted: 3 December 1997     

The relative importance of factors determining genetic drift: mating system, spatial genetic structure, habitat and census size in Arabidopsis lyrata

? The mating system, dispersal and census size are predicted to determine the magnitude of genetic drift, but little is known about their relative importance in nature. ? We estimated the contributions of several population-level features to genetic drift in 18 populations of Arabidopsis lyrata. The factors were outcrossing rate, within-population spatial genetic structure, census size and substrate type. The expected heterozygosity (H(E)) at 10 microsatellite loci was taken to reflect the effective population size (N(e)) and the strength of genetic drift. ? The mating system explained most of the variation in H(E) (60%), followed by substrate (10%), genetic structure (9%) and census size (6%). The most outcrossing population had a +0.32 higher predicted H(E) than the most selfing population; the estimated N(e) of selfing populations was less than half that of outcrossing populations. Rocky outcrops supported populations with a +0.14 higher H(E) than did sandy substrates. The most structured population had a +0.24 higher H(E) than the least structured population, and the largest population had a +0.18 higher H(E) than the smallest population. ? This study illustrates the importance of outcrossing, genetic structure and the physical environment--together with census size--in maintaining H(E), and suggests that multiple population-level characteristics influence N(e) and the action of genetic drift.     

Salamander predation and vertical distributions of zooplankton

SUMMARY. 1. The impact of different predation regimes of zooplanktivorous larval tiger salamanders, Ambystoma tigrinum nebulosum , on the vertical temporal distributions of Daphnia pulex, Diaptomus nudus and larval Chaoborus flavicans was studied in limnetic enclosures in a pond in east-central Arizona. Zooplankton densities were monitored 1-2 h before sunset, at sunset, and 1-2 h after sunset at three depths (0-0.4, 1.0 and 2.0 m) in four salamander treatments (0, 20, 40 salamanders per enclosure, and unenclosed limnetic areas).
2. Densities and body size of Daphnia pulex were consistently greater in salamander-free enclosures than in other treatments. More Daphnia were generally nearer the surface (0-1.0 m) than along the bottom before and at sunset than after sunset in enclosures without salamanders. After sunset, Daphnia was by far most dense along the bottom. Densities were uniform among depths and sample times in treatments with salamanders. Salamanders moved throughout the water column. Frequencies of D. pulex spined morphs (with prominent dorsal crest) did not differ among salamander treatments.
3. Chaoborus densities were positively correlated with D. pulex distributions among depths and sample times in treatments. Spatio-temporal distributions of Chaoborus were influenced more by their daphnid prey than by risk of predation from larval salamanders.
4. Densities of Diaptomus nudus were highest in open water controls and lowest in enclosures without salamanders. Diaptomus was generally more dense between 0 and 1 m than at 2.0 m depths in all treatments, and vertical distributions were not dependent on salamander density.     

The microdistribution of some lotic insect predators in relation to their prey and to abiotic factors

SUMMARY. 1. The microdistribution of three species of insect predators in a Swedish stream was assessed using a multivariate statistical approach. Both abiotic factors and factors pertaining to the prey community were included in the analysis.
2. The factors most strongly correlated to the distribution of large Dinocras cephalotes were the densities of case less caddis larvae ( Wormaldia sp. and Hydropsyche siltalai ) and the median weight of stonefly prey. For small D. cephalotes , the density of H. siltalai and the percentage of moss cover were most important. Isoperla grammatica showed a significant correlation to blackfly density. Rhyacophila nubila occurred predominantly in patches with high prey biomass and with high densities and median weight of case less caddis larvae.
3. It was concluded that the quantity and quality of the prey, such as size, availability and species, influenced the predators'microdistribution. The distributional pattern differed between species of predators and was probably related to their different hunting strategies.     

Prey refugia and the distributions of two Sonoran Desert cacti

Summary Two small Sonoran Desert cacti, Mammillaria microcarpa and Echinocereus englemannii, are commonly found beneath canopies of the larger, tree-like cactus Opuntia fulgida. The mechanism leading to this distribution pattern is incidental to the mode of reproduction in O. fulgida. Opuntia fulgida propagates by means of easily-detached, spine-covered stem joints that accumulate beneath the parent plant. These accumulations of spines apparently deter mammalian herbivores that otherwise consume succulent tissues of the smaller cacti. Such incidental effects are little studied, but they may contribute substantially to structure within plant communities.     

Glacial refugia,haplotype distributions,and clonal richness of the Daphnia pulex complex in arctic Canada

As part of a large international Arctic biodiversity expedition (Tundra Northwest '99), we examined the distribution of members of the arctic Daphnia pulex complex (Cladocera, Anomopoda) from 121 tundra ponds, spread across 16 sites spanning a large portion of arctic Canada (i.e. from 62 degrees 22' N to 79 degrees 01' N; 66 degrees 45' W to 139 degrees 37' W). Using allozyme electrophoresis and mitochondrial (mt)DNA analyses, we examined the population genetic (clonal) structure of these populations. The following taxa were detected in this complex: Daphnia pulicaria, D. middendorffiana and D. tenebrosa. Clear geographical differences in mean clonal richness and diversity were observed, with most western sites exhibiting higher clonal richness and diversity, than sites in the eastern Canadian Arctic. For both the pulicaria group (i.e. D. pulicaria and D. middendorffiana) and D. tenebrosa, the highest mean regional clonal richness was detected from the southern section of Banks Island, an unglaciated site situated on the edge or directly in the eastern fringe of the Beringian glacial refuge. A significant negative correlation was found between geographical distance from the Beringian edge, and overall regional clonal richness (i.e. sites closer to the edge harboured greater clonal richness). These results clearly indicate that more recently deglaciated regions (i.e. eastern Canadian Arctic) harbour lower levels of clonal richness than western regions nearer Beringia. We discuss the role that glacial refugia have played in influencing both biotic and genetic diversity in arctic taxa.     

Relative importance of environment, human activity and spatial situation in determining the distribution of terrestrial mammal diversity in Argentina

Aim The partition of the geographical variation in Argentinian terrestrial mammal species richness (SR) into environmentally, human and spatially induced variation. Location Argentina, using the twenty‐three administrative provinces as the geographical units. Methods We recorded the number of terrestrial mammal species in each Argentinian province, and the number of species belonging to particular groups (Marsupialia, Placentaria, and among the latter, Xenarthra, Carnivora, Ungulates and Rodentia). We performed multiple regressions of each group's SR on environmental, human and spatial variables, to determine the amounts of variation explained by these factors. We then used a variance partitioning procedure to specify which proportion of the variation in SR is explained by each of the three factors exclusively and which proportions are attributable to interactions between factors. Results For marsupials, human activity explains the greatest part of the variation in SR. The purely environmental and purely human influences on all mammal SR explain a similarly high proportion of the variation in SR, whereas the purely spatial influence accounts for a smaller proportion of it. The exclusive interaction between human activity and space is negative in carnivores and rodents. For rodents, the interaction between environment and spatial situation is also negative. In the remaining placental groups, pure spatial autocorrelation explains a small proportion of the variation in SR. Main conclusions Environmental factors explain most of the variation in placental SR, while Marsupials seem to be mainly affected by human activity. However, for edentates, carnivores, and ungulates the pure human influence is more important than the pure spatial and environmental influences. Besides, human activity disrupts the spatial structure caused by the history and population dynamics of rodents and, to a lesser extent, of carnivores. The historical events and population dynamics on the one hand, and the environment on the other, cause rodent SR to vary in divergent directions. In the remaining placental groups the autocorrelation in SR is mainly the result of autocorrelation in the environmental and human variables.     

The interactive effects of competition and predation risk on dispersal in an insect

Dispersal dynamics have significant consequences for ecological and evolutionary processes. Previous work has demonstrated that dispersal can be context-dependent. However, factors affecting dispersal are typically considered in isolation, despite the probability that individuals make dispersal decisions in response to multiple, possibly interacting factors. We examined whether two ecological factors, predation risk and intraspecific competition, have interactive effects on dispersal dynamics. We performed a factorial experiment in mesocosms using backswimmers (Notonecta undulata), flight-capable, semi-aquatic insects. Emigration rates increased with density, and increased with predation risk at intermediate densities; however, predation had minimal effects on emigration at high and low densities. Our results indicate that factorial experiments may be required to understand dispersal dynamics under realistic ecological conditions.     

Fragmentation within and between wetland reserves: the importance of spatial scales for nest predation in reed buntings

Wetlands in many parts of the world are affected by fragmentation at multiple spatial scales. In Switzerland, most wetlands have been destroyed over the past two centuries and management of the remaining wetland reserves has intensified in the recent years leading to increased fragmentation of reed areas within reserves. Using four years of data on the reproductive performance of color-banded reed bunting Emberiza schoeniclus populations, we explored fragmentation effects on nest predation rates at four spatial scales ranging from the nest to the landscape scale. In the egg stage, predation rate was negatively related to vegetation cover, vegetation height and nest height, but positively linked to water cover and depth next to the nest (nest scale). Probability of predation declined with increasing size of reed patches containing the nests and distances of nests to the water and land sided reed edge, as well as with decreasing edge to area ratio (edge scale). There was a weak positive association between degree of fragmentation of reed patches within sites and nest predation rates (site scale). Finally, nest predation probability increased with distance to the nearest wetland (landscape scale). Jointly analyzing variables from different spatial scales revealed that a model combining variables from the nest, edge and landscape scale best explained predation probability in the egg stage. In the nestling stage, the single most important factor influencing nest predation probability was the distance to the nearest wetland (landscape scale), with nest predation decreasing with distance between sites. Our results show that the probability of nest predation in reed buntings is affected by fragmentation within and between wetland reserves and that the effects differ between breeding stages. Future management of wetland reserves should aim at sparing reed patches large and dense enough to provide safe nest sites for birds.