Can environmental tolerances explain convergent patterns of distribution in endemic spring snails from opposite sides of the Australian arid zone?

Patterns of distribution are influenced by species environmental requirements and limits, but experimental tests are needed to discern whether correlates of abundance directly affect survival and success. Springs in Australia’s arid interior support a wide diversity of gastropods only found in springs, and these species show dichotomous patterns of distribution. “Amphibious” species are broadly distributed across many springs and microhabitats, and “aquatic” species confined to the deepest pool areas within large springs. This pattern appears to be driven by the interaction between different environmental conditions in different microhabitats and the environmental tolerances of each endemic snail species. Factorial experiments were used to test whether conditions in the environmentally extreme and variable tail area of springs (considering pH, conductivity, temperature and desiccation potential, alone and in synergistic scenarios) elicited lethal or sub-lethal responses in spring snails endemic to springs on opposite sides of the Australian arid zone. All species restricted to spring pools were able to endure 24 h exposed to the average tail conditions, alone and in combination, but most suffered mortalities when subjected to extremes, and mortalities occurred sooner in the most restricted species when elevated pH and conductivity were experienced in combination. Responses of species from different locations are similar, but pattern of distribution in the field were not correlated with tolerance of environmental extremes—with the “amphibious” species from the sub-tropics being far more sensitive than its arid counterpart. These findings suggest that environmental variance within springs can influence patterns of distribution and abundance, particularly when extremes are experienced simultaneously over sustained time periods. But despite similarities in responses across species from these two spring complexes, no simple generalisations linking distribution and tolerance were discernible.     

Parasites and salinity: costly tradeoffs in a threatened species

Parasites and environmental conditions can have direct and indirect effects on individuals. We explore the relationship between salinity and parasites in an endemic New Mexico State threatened fish, the White Sands pupfish (Cyprinodon tularosa). Spatial variation in salinity limits the distribution of the endemic springsnail (Juturnia tularosae) within Salt Creek, a small desert stream. The springsnail is the presumed intermediate host for trematodes that infect the White Sands pupfish, and trematode prevalence and intensity in pupfish are positively associated with the springsnail. Salinity and parasites both have negative impacts on pupfish, but in areas of high salinity, pupfish can effectively escape parasites. Pupfish trematodes were absent from sites lacking snails. At the upstream site, the absence of parasites and lower variance in salinity were correlated with larger pupfish that were in better condition than pupfish at either the middle or lower sites. Springsnails were present in the middle section, an area with moderate salinity, and all pupfish had trematodes (median abundance 847 trematodes/fish). Lipid levels and condition were lowest in fish from the middle site. Additionally, fewer older fish indicated an increased mortality rate. At the lower site, springsnails were absent due to high salinity; pupfish trematode abundance was much lower (six trematodes/fish), and fish condition was intermediate. An additional experiment revealed that snail activity and survival were significantly reduced at high salinities commonly present at the lower site. Although both high salinity and parasites significantly affect pupfish, parasites might be more detrimental.     

Imprints of glacial history and current environment on correlations between endemic plant and invertebrate species richness

Aim We evaluate how closely diversity patterns of endemic species of vascular plants, beetles, butterflies, molluscs and spiders are correlated with each other, and to what extent similar environmental requirements or survival in common glacial refugia and comparable dispersal limitations account for their existing congruence. Location Austria. Methods We calculated pairwise correlations among species numbers of the five taxonomic groups in 1405 cells of a 3′ × 5′ raster (c. 35 km²) using the raw data as well as the residuals of regression models that accounted for: (1) environmental variables, (2) environmental variables and the occurrence of potential refugia during the Last Glacial Maximum, or (3) environmental variables, refugia and spatial filters. Results Pairwise cross‐taxonomic group Spearman’s rank correlations in the raw data were significantly positive in most cases, but only moderate (0.3 < ρ < 0.5) to weak (ρ < 0.3) throughout. Correlations were closest between plants and beetles, plants and butterflies, and plants and snails, respectively, whereas the distribution of endemic spiders was largely uncorrelated with those of the other groups. Environmental variables explained only a moderate proportion of the variance in endemic richness patterns, and the response of individual groups to environmental gradients was only partly consistent. The inclusion of refugium locations and the spatial filters increased the goodness of model fit for all five taxonomic groups. Moreover, removing the effects of environmental conditions reduced congruence in endemic richness patterns to a lesser extent than did filtering the influence of refugium locations and spatial autocorrelation, except for spiders, which are probably the least dispersal‐limited of the five groups. Main conclusions The moderate to weak congruence of endemic richness patterns clearly limits the usefulness of a surrogacy approach for designating areas for the protection of regional endemics. On the other hand, our results suggest that dispersal limitations still shape the distributions of many endemic plant, snail, beetle and butterfly species, even at the regional scale; that is, survival in shared refugia and subsequent restricted spread retain a detectable signal in existing correlations. Concentrating conservation efforts on well‐known Pleistocene refugia hence appears to be a reasonable first step towards a strategy for protecting regional endemics of at least the less mobile invertebrate groups.     

Invertebrate community composition differs between restored and natural vernal pools

The loss of freshwater wetlands worldwide has underscored the importance of restoration to enhance biodiversity and functional objectives. While aquatic invertebrate communities within restored perennial freshwaters are well studied, few studies have occurred in the greatly reduced habitat of seasonal wetlands, such as vernal pools. California vernal pools have experienced high habitat loss and support many threatened or endangered invertebrate species. We compared 90 natural and 90 restored vernal pools of different ages across 10 sites throughout California and Southern Oregon using the Sars' method. Large branchiopod abundance, total invertebrate abundance, class richness, and community composition were assessed between pool types (natural vs. restored) and along environmental gradients (e.g. site, pool depth, surface area, age since restoration). Large branchiopod and total invertebrate abundance were 215 and 274% higher in natural pools than restored pools, but class richness was not different. Community composition was significantly different and driven by greater abundances of vernal pool fairy shrimp, San Diego fairy shrimp, Ostracoda, Cladocera, and Copepoda in natural pools. Few environmental or habitat variables explained patterns in richness or abundance. Our work demonstrates that restored pools, even those decades old, are different than natural pools. Future mitigation and monitoring guidelines for restored vernal pools should include quantitative evaluations for aquatic invertebrates. Restored pools are not adequate compensation for lost natural pools because they do not have the same ecological functions and values.     

Spatial and environmental effects on a rock‐pool metacommunity depend on landscape setting and dispersal mode

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Spatial and vertical patterns in the tidepool fish assemblage on the island of O`ahu

The microtides, wave regimes, and relative isolation of the Hawaiian archipelago may provide unique environmental and biogeographic effects that shape the structure of tidepool fishes. We sampled fishes across a narrow gradient at low tide from 6 sites on the island of O`ahu. We tested predictions of the hypotheses that environmental conditions (pool depth, volume, macroalgal cover, temperature, and salinity) would result in a vertically structured tidepool fish assemblage unique to basalt or limestone rocky shores. 343 fish were recorded from 40 pools, and 19 species from 10 families were identified. Tidepool fish diversity (H’: O`ahu = 2.4; Sites Average = 0.0–0.9) was typical for tropical islands, with members from Gobiidae (5 species), Blenniidae (4 species), Pomacentridae (3 species), Acanthuridae (2 species) and Kuhliidae (2 species) among the most common. Endemism (32%) was higher than other well studied assemblages yet similar to Hawaiian reef fishes (25%). Assemblage abundance varied among shores with basalt or limestone substrate, among sites, and vertically among high, mid, and low pools. In general, blenniids occurred at higher proportions on limestone shores and gobiids were more common on basalt shores. High pools were characterized by an abundance of a small sized (29.0 mm median standard length) blenniid Istiblennius zebra, while the blenniid Entomacrodus marmoratus and wrasses Thalassoma spp. were more common in low pools. Temperature was the best environmental predictor of assemblages and this relationship warrants further investigation. Our findings indicate that assemblages can vary across a narrow geographical range and intertidal shore.     

Microhabitat use by foraging white-clawed crayfish (<Emphasis Type="Italic">Austropotamobius pallipes</Emphasis>) in stream pools in the NE Iberian Peninsula

The white-clawed crayfish (Austropotamobius pallipes) is an endangered species across most of its distribution range, and information on its ecological requirements is needed to implement effective conservation measures. Its habitat use has been studied in different areas and at various spatial scales. However, being a nocturnal species, there is scarce information on its habitat selection during foraging periods. In this work we analyse nocturnal habitat use of white-clawed crayfish in pools of a small stream in the northeastern Iberian Peninsula at two different scales: (1) microhabitat selection and (2) pool characteristics. Large crayfish showed a clear positive selection for deeper microhabitats, a selection pattern that was weaker for medium-sized crayfish and absent for small ones. On the other hand, crayfish of all sizes avoided cobble and boulder microhabitats and positively selected fine substrate and more exposed microhabitats. Crayfish abundance in pools was positively influenced by pool area, pool depth and the availability of fine substrates, especially silt. While studies on white-clawed crayfish habitat use have often stressed the importance of rough substrates as crayfish refuge, our results show that fine substrates are positively selected by foraging crayfish of all size classes and promoted active crayfish abundance in pools. These apparently contradictory results may be due to the differences in microhabitat preferences exhibited by active and inactive crayfish. Thus, our results help to better complete the picture of white-clawed crayfish habitat requirements.     

SPATIAL PARADIGMS OF LOTIC DIATOM DISTRIBUTION: A LANDSCAPE ECOLOGY PERSPECTIVE

Spatial distributional patterns of benthic diatoms and their relation to current velocity were investigated in an unshaded cobble-bottom reach of White Creek (Washington County, NY). On 27 August 1999, diatoms were sampled and current velocity and depth were measured on a regular square sampling grid with a grain size of 0.01 m², interval of 0.5 m, and extent of 16 m². The relative abundance of the 18 common diatom species enumerated in the 81 samples was subjected to detrended correspondence analysis (DCA). The first axis (DCA1) explained 51% of the variance in diatom data and separated the samples according to current regimes. The spatial autocorrelation of DCA1 sample scores in deposition and erosion regions of White Creek was determined by Moran's I statistic to indicate patch size. In White Creek the patch length of all diatom communities was more than 3.1 m, whereas the patch width was 1 m in the deposition region and 0.5 m in the erosion region. There were 5 dominant diatom taxa, Achnanthes minutissima Kütz. et vars, Fragilaria capucina Dezmazières et vars, F. crotonensis Kitt., Diatoma vulgaris Bory, and Synedra ulna (Nitz.) Ehr. et vars. The patch length of the dominant species varied from 1 to more than 4.1 m, whereas the patch width, if defined, was 0.5 m. Achnanthes minutissima and F. capucina, the two diatom species with the highest relative abundance, displayed spatially structured patches of low abundance and comparatively random patches of high abundance, suggesting broad scale abiotic control of species performance in low abundance regions and finer scale biotic control of high abundance areas. Another objective of this study was to test the hypothesis that higher current velocities, which generally impede immigration, would increase randomness and complexity (i.e. homogeneity of diatom distributional patterns). The spatial complexity in low versus high velocity transects was determined by calculating the respective fractal dimension (D) of DCA1 scores. D of DCA1 was higher in the higher current velocity transects, suggesting that spatial complexity and homogeneity of diatom communities increased in faster currents. Partial canonical correspondence analysis was conducted on diatom, environmental, and spatial data to assess how much of the variance in species distribution could be attributed to environmental (current velocity and depth) versus spatial factors. The variance of species data, explained by the environment (exclusively current velocity), was 38%; whereas space alone contributed only 10%, indicating that 1) current velocity was the major factor that controlled diatom distribution in streams and 2) there were other spatially dependent variables, most likely biotic, but their role in shaping diatom communities was minor.     

Ground‐dwelling beetle assemblages in the northern Cape Floristic Region: Patterns,correlates and implications

Abstract Recent studies have both shown and predicted that global climate change will have a substantial influence on biodiversity. This is true especially of a global biodiversity hotspot, the Cape Floristic Region. Although the effects of predicted changes have been widely assessed for plants, little is known about how insect diversity in the region might be affected. In particular, patterns in and the correlates of diversity in the region are poorly understood, and therefore the likely affects of a changing abiotic environment on this significant group of organisms are not clear. Therefore, we investigate patterns in, and correlates of, epigaeic beetle (Tenebrionidae and Carabidae) diversity in one of the most climate change‐sensitive areas in the Cape Floristic Region, the Cederberg. In particular, we determine whether epigaeic beetle assemblage structure differs between the main vegetation types in the Cederberg (Strandveld, Mountain Fynbos and Succulent Karoo), how restricted these beetles are to specific vegetation types, and which environmental variables might be associated with site‐related differences in beetle richness and abundance. Sampling was undertaken during October 2002 and 2003 across an altitudinal gradient ranging from sea level (Lambert's Bay) to approximately 2000 m above sea level (Sneeukop, Cederberg) and down again to 500 m above sea level (Wupperthal) using pitfall traps. The environmental correlates of abundance and species density in the epigaeic beetles were similar to those identified previously for ants across the transect, with both taxa being positively related to several temperature variables. Several species showed habitat specificity and fidelity, and clear distinctions existed between the vegetation types across the transect. A larger proportion of the variance in tenebrionid species density was explained by environmental variables and spatial factors than for carabids. The most likely explanation for this difference is that the correlates might well reflect collinear historical processes, rather than a causal relationship between contemporary environmental variables and species density. If this is the case, it suggests that caution should be exercised when interpreting environmental correlates of species density, and making climate change predictions based on these correlates.     

Variability of Symbiodinium Communities in Waters,Sediments, and Corals of Thermally Distinct Reef Pools in American Samoa

Reef-building corals host assemblages of symbiotic algae (Symbiodinium spp.) whose diversity and abundance may fluctuate under different conditions, potentially facilitating acclimatization to environmental change. The composition of free-living Symbiodinium in reef waters and sediments may also be environmentally labile and may influence symbiotic assemblages by mediating supply and dispersal. The magnitude and spatial scales of environmental influence over Symbiodinium composition in different reef habitat compartments are, however, not well understood. We used pyrosequencing to compare Symbiodinium in sediments, water, and ten coral species between two backreef pools in American Samoa with contrasting thermal environments. We found distinct compartmental assemblages of clades A, C, D, F, and/or G Symbiodinium types, with strong differences between pools in water, sediments, and two coral species. In the pool with higher and more variable temperatures, abundance of various clade A and C types differed compared to the other pool, while abundance of D types was lower in sediments but higher in water and in Pavona venosa, revealing an altered habitat distribution and potential linkages among compartments. The lack of between-pool effects in other coral species was due to either low overall variability (in the case of Porites) or high within-pool variability. Symbiodinium communities in water and sediment also showed within-pool structure, indicating that environmental influences may operate over multiple, small spatial scales. This work suggests that Symbiodinium composition is highly labile in reef waters, sediments, and some corals, but the underlying drivers and functional consequences of this plasticity require further testing with high spatial resolution biological and environmental sampling.     

Characterization of microsatellite loci in the endemic mound spring snail Fonscochlea accepta and cross species amplification in four other hydrobiid snails

We identified and characterized nine microsatellite primer pairs from Fonscochlea accepta, a species of hydrobiid snail endemic to artesian mound springs associated with the Great Artesian Basin in arid South Australia. The loci were highly polymorphic, with five to 30 alleles per locus. Gene diversity, estimated as expected heterozygosity ranged from 0.364 to 0.851, and was generally matched by levels of observed heterozygosity (0.373–0.829). Cross‐species amplification trials with four other hydrobiid species associated with these southern mound springs showed that these primers will be useful for genetic analyses of these other endemic snails.     

Predator–prey relationships within natural,restored, and created vernal pools

We performed a multiyear monitoring study to compare amphibian habitat quality among four natural, four restored, and six created pools. We used successful reproduction and metamorphosis of two vernal pool indicator species, the wood frog and spotted salamander, to represent desired outcomes. Ordination techniques were used to identify the aspects of habitat quality that were most correlated with desired outcomes. Previously published results indicated that pool depth, volume, and hydroperiod were among the best predictors of success, regardless of pool type. Observations in the first few years of monitoring also suggested that pools with longer hydroperiods had a greater abundance of aquatic predators of eggs and larvae of indicator species. This follow‐up study further explores and compares predator–prey relationships among pool types. We quantified within‐pool predator and prey abundance and diversity and collected another year of data on the reproductive success of indicator species. Our results confirmed that mean predator abundance was eight times higher in pools with longer hydroperiods. We documented a 96% decrease in wood frog survival rates in a semi‐permanent, natural pool following a 41% decrease in overhead canopy cover and an increase in green frog abundance. At the same time, wood frog reproductive success increased in nearby restored pools with lower predator abundance. Pools with the highest mean survival rates for the two indicator species combined were short‐ or long‐cycle pools (i.e. hydroperiod of 12–35 weeks) with low predator abundance (i.e. <1 organism L^?1) and greater proportions of arthropod prey relative to other food items.     

Evaluating the potential causes of range limits of birds of the Colombian Andes

Aim To evaluate how factors acting at different spatial scales influence range limits in bird species of the Colombian Andes. Location Andes Mountains of Colombia. Methods We used Maxent , a climate envelope model (CEM), and environmental and geographic information to study range‐filling (i.e. the extent to which a species occurs in all the areas in which it is predicted to occur) in 70 range‐restricted bird species of the Colombian Andes. Environmental data were taken from the WorldClim database, and species occurrence data were taken from museum data collated by the BioMap project, an observational database, and the literature. We evaluated how climate and geographic barriers may shape range limits at two scales. Results At a broad extent (i.e. across the three main cordilleras within the Colombian Andes), we find that CEMs predict there to be suitable environmental conditions for particular species in regions where the species is absent, possibly as a result of dispersal limitation or biotic interactions. In contrast, at a finer scale (within a given cordillera), species generally occur across the entire area predicted to be suitable by a given CEM. Geographic discontinuities within cordilleras do not generally correspond to range limits; instead, range limits correspond to changes in environmental conditions. Main conclusions Our results suggest that different mechanisms influence the presence of species at different scales. Dispersal limitation, potentially combined with species interactions, may influence range limits at a broad extent (the entire Colombian Andes), while strong environmental gradients correspond to range limits at a finer scale (within a cordillera).     

Richness and structure of plant communities in temporary pools from western Morocco: influence of human activities

Temporary pools are numerous in coastal plains of Atlantic Morocco and have a rich and diverse flora. These habitats are increasingly under pressure by man impact through grazing by domestic livestock and the development of annual crops in their catchments. The objectives of this work were to evaluate the species richness and the species composition of the vegetation of a sample of 30 pools in this region, in order to assess the structure of the vegetation within pools and to evaluate the role of environmental and anthropogenic factors in their richness and species composition. The results highlighted the species richness of the pools with 300 species found among which annual species were heavily dominant. The structure of the vegetation varied within pools according to a topographical/flooding gradient and between pools in relation with the land use in the surrounding areas. Three vegetation belts were recognized from the centre to the periphery of the pool. The total species richness per pool was not found significantly related to environmental or anthropogenic factors. The species characteristics of the pools were found mostly in the centre and their number affected by hydrological factors and land use in the pool and in the surrounding areas. The species characteristics of woodland habitats and of agriculture crops were found mostly in the peripheral zone. Temporary pools maintain specific communities of both aquatic and amphibious species, probably because of the selection induced by water level variations. The peripheral zone, although often dominated by terrestrial species, is clearly interpreted as part of the pool. It contains amphibious species highly characteristic of the pool for which the irregular flooding is a key environmental factor which decreases competition.     

Regional and local drivers of macroinvertebrate assemblages in boreal springs

Aim To identify the most important environmental drivers of benthic macroinvertebrate assemblages in boreal springs at different spatial scales, and to assess how well benthic assemblages correspond to terrestrially derived ecoregions. Location Finland. Methods Benthic invertebrates were sampled from 153 springs across four boreal ecoregions of Finland, and these data were used to analyse patterns in assemblage variation in relation to environmental factors. Species data were classified using hierarchical divisive clustering (twinspan ) and ordinated using non‐metric multidimensional scaling. The prediction success of the species and environmental data into a priori (ecoregions) and a posteriori (twinspan ) groups was compared using discriminant function analysis. Indicator species analysis was used to identify indicator taxa for both a priori and a posteriori assemblage types. Results The main patterns in assemblage clusters were related to large‐scale geographical variation in temperature. A secondary gradient in species data reflected variation in local habitat structure, particularly abundance of minerogenic spring brooks. Water chemistry variables were only weakly related to assemblage variation. Several indicator species representing southern faunistic elements in boreal springs were identified. Discriminant function analysis showed poorer success in classifying sites into ecoregions based on environmental than on species data. Similarly, when classifying springs into the twinspan groups, classification based on species data vastly outperformed that based on environmental data. Main conclusions A latitudinal zonation pattern of spring assemblages driven by regional thermal conditions is documented, closely paralleling corresponding latitudinal patterns in both terrestrial and freshwater assemblages in Fennoscandia. The importance of local‐scale environmental variables increased with decreasing spatial extent. Ecoregions provide an initial stratification scheme for the bioassessment of benthic macroinvertebrates of North European springs. Our results imply that climate warming, landscape disturbance and degradation of spring habitat pose serious threats to spring biodiversity in northern Europe, especially to its already threatened southern faunistic elements.     

Seasonal drought limits tree species across the Neotropics

Within the tropics, the species richness of tree communities is strongly and positively associated with precipitation. Previous research has suggested that this macroecological pattern is driven by the negative effect of water‐stress on the physiological processes of most tree species. This implies that the range limits of taxa are defined by their ability to occur under dry conditions, and thus in terms of species distributions predicts a nested pattern of taxa distribution from wet to dry areas. However, this ‘dry‐tolerance’ hypothesis has yet to be adequately tested at large spatial and taxonomic scales. Here, using a dataset of 531 inventory plots of closed canopy forest distributed across the western Neotropics we investigated how precipitation, evaluated both as mean annual precipitation and as the maximum climatological water deficit, influences the distribution of tropical tree species, genera and families. We find that the distributions of tree taxa are indeed nested along precipitation gradients in the western Neotropics. Taxa tolerant to seasonal drought are disproportionally widespread across the precipitation gradient, with most reaching even the wettest climates sampled; however, most taxa analysed are restricted to wet areas. Our results suggest that the ‘dry tolerance' hypothesis has broad applicability in the world's most species‐rich forests. In addition, the large number of species restricted to wetter conditions strongly indicates that an increased frequency of drought could severely threaten biodiversity in this region. Overall, this study establishes a baseline for exploring how tropical forest tree composition may change in response to current and future environmental changes in this region.     

Environmental and spatial filters of zooplankton metacommunities in shallow pools in high‐elevation peatlands in the tropical Andes

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Extraordinary micro-endemism in Australian desert spring amphipods

Increasing pressure for water in the Australian arid zone is placing enormous stress on the diverse endemic communities inhabiting desert springs. Detailed information about the evolutionary processes occurring within and between individual endemic species will help to develop effective and biologically relevant management strategies this fragile ecosystem. To help determine conservation priorities, we documented the genetic structure of the endemic freshwater amphipod populations in springs fed by the Great Artesian Basin in central Australia. Phylogenetic and phylogeographic history and genetic diversity measures were examined using nuclear and mitochondrial DNA from approximately 500 chiltoniid amphipods across an entire group of springs. Pronounced genetic diversity was identified, demonstrating that levels of endemism have been grossly underestimated in these amphipods. Using the GMYC model, 13 genetically divergent lineages were recognized as Evolutionarily Significant Units (ESUs), all of which could be considered as separate species. The results show that due to the highly fragmented ecosystem, these taxa have highly restricted distributions. Many of the identified ESUs are endemic to a very small number of already degraded springs, with the rarest existing in single springs. Despite their extraordinarily small ranges, most ESUs showed relative demographic stability and high levels of genetic diversity, and genetic diversity was not directly linked to habitat extent. The relatively robust genetic health of ESUs does not preclude them from endangerment, as their limited distributions ensure they will be highly vulnerable to future water extraction.     

Periphyton responses to invertebrate grazing and riparian canopy in three northern California coastal streams

SUMMARY. 1. Field experiments were conducted to examine the impact of grazing invertebrates on periphyton biomass in twenty-one pools across three northern California coastal streams (U.S.A.): Big Sulphur Creek, the Rice Fork of the Eel River, and Big Canyon Creek. Periphyton accrual on artificial substrate tiles was compared in each stream between two treatments: those elevated slightly above the stream bottom to reduce access by grazers (= platforms) and those placed directly on the stream bottom to allow access by grazers (=controls).
2. Crawling invertebrate grazers (cased caddisflies and snails) were numerically dominant in each stream (86% of all grazers in Big Sulphur Creek, 61% in the Rice Fork, 84% in Big Canyon Creek). Platforms effectively excluded crawling grazers, but were less effective in excluding swimming mayfly grazers (Baetidae).
3. Periphyton biomass (as AFDM) on tiles was significantly lower on controls compared to platforms for the Rice Fork, an open-canopy stream, and Big Sulphur Creek, a stream with a heterogeneous canopy. In contrast, no grazer impact was found for Big Canyon Creek, a densely shaded stream. Here, extremely low periphyton biomass occurred for both treatments throughout the 60 day study.
4. The influence of riparian canopy on periphyton growth (i.e. accrual on platforms), grazer impact on periphyton, and grazer abundance was examined for Big Sulphur Creek. As canopy increased (15–98% cover), periphyton biomass on platforms decreased. In contrast, canopy had little influence on periphyton accrual on controls; apparently, grazers could maintain low periphyton standing crops across the full range of canopy levels. The abundance of one grazer species, the caddisfly Gumaga nigricula , was highest in open, sunlit stream pools; abundance of two other prominent grazers, Helicopsyche borealis (Trichoptera) and Centroptilum convexum (Ephemeroptera), however, was unrelated to canopy.