Impacts of introduced brown and rainbow trout on benthic invertebrate communities in shallow New Zealand lakes

1. Brown and rainbow trout have been introduced to many inland waters in New Zealand, but research on the impacts on native communities has focused mainly on streams. The purpose of this study was to compare the benthic communities of trout and troutless lakes. Based on previous studies in North America and Europe, we predicted that the benthic biomass, and especially the abundance of large invertebrates, would be lower in lakes with trout as compared to those without. We surveyed the invertebrate fauna of 43 shallow, high‐elevation lakes (26 with and 17 without trout) in four geographic clusters on the central South Island and then conducted a detailed quantitative study of invertebrate biomass and community structure in 12 of these lakes. 2. Benthic community composition and diversity of lakes with and without trout were nearly identical and biomass was as high or higher in the lakes with as without trout. There was no evidence that trout have caused local extinctions of benthic invertebrates. Although the proportional abundance of large‐bodied aquatic was slightly lower in lakes with than without trout, the abundance of several groups of large‐bodied benthic taxa (dragonflies, caddisflies and water bugs) did not differ. 3. Our findings are in contrast to those in North American and Europe where trout introductions into previously troutless lakes have led to declines in the abundance of benthic invertebrates, especially large‐bodied taxa. We propose that the modest effects of trout in New Zealand could be explained by (i) the high areal extent of submergent vegetation that acts as a benthic refuge, (ii) low intensity of trout predation on benthic communities and/or (iii) characteristics of the benthic invertebrates that make them relatively invulnerable to fish predation. 4. Regardless of the relative importance of these hypotheses, our results emphasise that the same invertebrates occurred in all of the lakes, regardless of size, elevation and presence of trout, suggesting habitat generalists dominate the benthic fauna in shallow New Zealand lakes.     

Direct and indirect effects in microcosm communities of protists

Increased complexity in biological communities can increase the variety of interactions among species, but the relative strengths and long-term consequences of various direct and indirect interactions require further investigation. I studied interactions among four species of protists by monitoring their population dynamics when they were cultured either together or in seven different subset communities. Two protists were bacterivores (Chilomonas and Tetrahymena) and two were predators (Actinosphaerium and Euplotes). Actinosphaerium was omnivorous, and could eat both predatory Euplotes and bacterivores. Three indirect effects occurred among the four species of protists, including indirect facilitation of one predator by the other, apparent competition between bacterivores, and indirect facilitation of one bacterivore by the omnivorous predator. Community structure and invasibility depended on both direct and indirect effects; thus both can be mechanisms for assembly rules.     

The effects of stream acidity on benthic invertebrate communities in the south-eastern United States

1. Patterns in invertebrate community structure were determined from analysis of benthic samples taken quarterly for 1 year from four sites in the Great Smoky Mountains National Park (GSMNP). Sites ranged in baseflow pH from 4.5 to 6.8, with a corresponding range in inorganic monomeric aluminium concentrations from 3 to 197 μgl^?1. 2. Total invertebrate density (excluding Chironomidae) was correlated with pH (R²= 0.90, P<0.05). Density of Ephemeroptera was more highly correlated with pH than density of invertebrates in other taxonomic orders. Invertebrate density (total and density of different orders) was generally more strongly correlated with pH than to inorganic monomeric aluminium concentrations. 3. Species richness declined from sixty-nine species at the highest pH site to thirty-three species at the lowest pH site. Richness of Ephemeroptera and Trichoptera was positively correlated with pH (R²= 0.96 in both cases). Trichopteran richness was also negatively correlated with inorganic monomeric Al concentrations (R²= 0.96, P< 0.05). Plecopteran richness was not related to stream acidity. 4. Not all species of Ephemeroptera were negatively affected by low stream water pH. At low pH sites, Ameletus lineatus Traver (Family: Siphonoluridae) was relatively more abundant compared to other species of Ephemeroptera, and had greater biomass and greater weight per individual length than at higher pH sites. 5. Differences in invertebrate community parameters among sites were not based on changes in food abundance. Although grazer densities were positively correlated with pH (R²= 0.93, P < 0.05), differences in density were not related to differences in food. To the contrary, lower grazer densities occurred at the lowest pH site where measured food abundance (algae) was highest. Likewise, shredder density increased with increasing pH but was correlated with standing stocks of benthic organic matter (BOM), which had generally higher standing stocks at lower pH sites. 6. Transplants of two species of Ephemeroptera between sites differing in pH (6.4 v 5.0) resulted in increased mortality of organisms placed at lower pH sites. In another transplant experiment, differences in mortality were not observed, although surviving invertebrates at low pH sites had ×10 higher body burdens of aluminium following the transplant than invertebrates from the reference site. 7. These results indicate that direct effects of pH and Al (affecting survival) were more important than indirect effects of food availability in determining changes in invertebrate community structure between the study sites.     

水质对东江流域附生硅藻群落的影响

运用相关分析(CA),典型对应分析(CCA),加权平均分析(WA)研究了影响东江流域附生硅藻的主要水质因子及对应的最适生态范围。CA显示,13项水质指标中溶解氧(Dissolved Oxygen,DO)、氨氮(NH4-N)、亚硝氮(NO2-N)、氯化物(Cl)、五日生化需氧量(BOD5)、磷酸盐(PO4-P)与其它指标间的相关性较强(P0.05),其余指标间相关性较低(P0.05)。以相关性较弱的p H、电导率(Cond.)、硅酸盐(Si O2)、总磷(TP)、总氮(TN)、硝氮(NO3-N)、化学需氧量(CODKMn O4)进行CCA分析,排序显示影响附生硅藻的水质因子依次为Cond.、NO3-N、CODKMn O4、TP、TN,对应的生态最适范围分别为39.20—642.00μs/cm、0.46—2.77mg/L、1.0—5.5mg/L、0.02—0.64mg/L、0.48—3.43mg/L。研究表明,Cond、NO3-N、CODKMn O4、TP、TN是影响东江附生硅藻分布的主要水质因子,Cyclostephanos invisitatus、Gomphonema olivaceum、Nitzschia acicularis、N.capitellata、N.intermedia可以指示东江较高的电导率,Aulacoseira ambigua、A.lirata、Cyclotella meneghiniana、Cyclostephanos invisitatus、Eunotia minor、Nitzschia acicularis、N.Clausii、Pinnularia subcapitata是东江TN、TP污染的指示种。Nitzschia acicularis对TN、TP的耐受性最高,Navicula eidrigiana对TN、TP敏感性最强。     

Effects of urbanization on stream benthic invertebrate communities in Central Amazon

Urbanization and its physical and chemical effects on aquatic environments influence invertebrate communities negatively. Yet, it is not clear how urbanization affects inter-annual variation of invertebrate assemblages in streams. We 1) evaluated urbanization effects on the ecological conditions (biotic and abiotic) of streams in Manaus and 2) analyzed invertebrate community variation over time (between 2003 and 2010). Data on abiotic variables and invertebrates from 2003 were obtained from a previous study. In 2010 we sampled abiotic variables and invertebrate communities in the same low-order urban streams sampled in 2003 (n = 40). We recorded high values of total nitrogen, total phosphorous, deforestation, total impervious area (TIA), water temperature, pH, and electrical conductivity in the most urbanized streams, as compared to the least-impacted ones. In contrast, the least-impacted streams had high dissolved oxygen concentrations. Water quality was poorer in 2010 than in 2003: oxygen concentration was lower and total nitrogen, total phosphorous, deforestation, and TIA significantly higher in 2010. We recorded higher inter-annual variation of abiotic variables in the most-impacted streams as compared to the least-impacted streams. EPT (%, Ephemeroptera, Plecoptera, and Trichoptera) and richness metrics decreased with urbanization. On the other hand, % OP (percent of Oligochaeta and Psychodidae) increased with urbanization. Observed and EPT richness and% OP increased between 2003 and 2010. On the other hand, rarefied richness decreased between years. Increases of observed and EPT richness between 2003 and 2010 were related to low inter-annual variability in streams conditions; however, differences of% OP and rarefied richness were not related to inter-annual variability in environmental conditions. The degree of urbanization did not explain the magnitude of the within-stream difference of invertebrate communities between 2003 and 2010. The increased effects of urbanization represented by the abiotic variables sampled and the reduction of invertebrate richness and increased dominance of tolerant taxa indicate that public policy is not enough to protect or mitigate human impacts on the urban water systems under study.     

Predation,sediment stability and food availability as determinants of the benthic invertebrate fauna in two shallow lakes

The sedimentary benthos of a series of shallow, eutrophicated lakes, the Norfolk Broads is, in general, low not only in number of species but unexpectedly in number of individuals. In two of the lakes, Hudsons Bay and Hoveton Great Broad, chironomids and oligochaetes dominated the fauna. Hudsons Bay has an extensive stand of water lilies (Nuphar lutea); Hoveton Great Broad does not. There were significant relationships between number of chironomids and of Potamothrix hammoniensis with organic content of the sediments, but these were due not to food availability but to the structure imparted to the otherwise fluid sediment by the organic matter. Sediment stabilised in plastic bowls developed much larger populations of oligochaetes than found in the unrestricted sediment. Protection of the community from fish predation resulted in a further major increase in numbers. Sediment stability and predation rather than food supply were the major determinants of these benthic populations.     

Effects of macrophyte growth forms on invertebrate communities in saline lakes of the Wyoming High Plains

In saline lakes, areal cover and both species and structural diversity of macrophytes often decline as salinity increases. To assess effects of the loss of certain macrophyte growth forms, we characterized benthic and epiphytic invertebrates in three growth forms (thin-stemmed emergents, erect aquatics, and low macroalgae) in oligosaline lakes (0.8–4.2 mS cm⁻¹) of the Wyoming High Plains, USA. We also measured the biomass and taxonomic composition of epiphytic and benthic invertebrates in two erect aquatics with very similar structure that are found in both oligosaline (Potamogeton pectinatus) and mesosaline (9.3–23.5 mS cm⁻¹) (Ruppia maritima) lakes. Although total biomass of epiphytic invertebrates varied among oligosaline lakes, the relative distribution of biomass among growth forms was similar. For epiphytic invertebrates, biomass per unit area of lake was lowest in emergents and equivalent in erect aquatics and low macroalgae; biomass per unit volume of macrophyte habitat was greatest in low macroalgae. For benthic invertebrates, biomass was less beneath low macroalgae than other growth forms. Taxonomic composition did not differ appreciably between growth forms for either benthic or epiphytic invertebrates, except that epiphytic gastropods were more abundant in erect aquatics. Total biomass of epiphytic and benthic invertebrates for the same growth form (erect aquatic) did not differ between oligosaline (Potamogeton pectinatus) and mesosaline (Ruppia maritima) lakes, but taxonomic composition did change. In the oligosaline to mesosaline range, direct toxic effects of salinity appeared important for some major taxa such as gastropods and amphipods. However, indirect effects of salinity, such as loss of macrophyte cover and typically higher nutrient levels at greater salinities, probably have larger impacts on total invertebrate biomass lake-wide.     

Direct and indirect effects of solar ultraviolet-B radiation on long-term decomposition

As a result of stratospheric ozone depletion, more solar ultraviolet-B radiation (UV-B, 280–315 nm) is reaching the Earth's surface. Enhanced levels of UV-B may, in turn, alter ecosystem processes such as decomposition. Solar UV-B radiation could affect decomposition both indirectly, by changes in the chemical composition of leaves during growth, or directly by photochemical breakdown of litter and through changes in decomposer communities exposed to sunlight. In this experiment, we studied indirect and direct effects of solar UV-B radiation on decomposition of barley (Hordeum vulgare). We used barley straw and leaf litter grown under reduced UV-B (20% of ambient UV-B) or under near-ambient UV-B (90% of ambient UV-B) in Buenos Aires, Argentina, and decomposed the litter under reduced or near-ambient solar UV-B for 29 months in Tierra del Fuego, Argentina. We found that the UV-B treatment applied during growth decreased the decay rate. On the other hand, there was a marginally significant direct effect of elevated UV-B during the early stages of decomposition, suggesting increased mass loss. The effect of UV-B during growth on decomposition was likely the result of changes in plant litter chemical composition. Near-ambient UV-B received during plant growth decreased the concentrations of nitrogen, soluble carbohydrates, and N/P ratio, and increased the concentrations of phosphorus, cellulose, UV-B-absorbing compounds, and lignin/N ratio. Thus, solar UV-B radiation affects the decomposition of barley litter directly and indirectly, and indirect effects are persistent for the whole decomposition period.     

The effect of drought on benthic invertebrate communities in a lowland river

Benthic macroinvertebrates were sampled at regular intervals from rural and urban sections of the River Roding, Essex, England, in 1975 and 1976. During the latter year a severe drought led to a marked decline in flows and to desiccation of parts of the river bed. In general, drought conditions resulted in an increase in invertebrate populations and possible reasons for this are presented. A considerable number of individuals of certain groups such as cased caddisfly larvae and prosobranch molluscs were, however, eliminated from the river at this time, mainly as a result of stranding and chemical changes in the environment. The effect of reduced flows on river faunas is briefly discussed.     

Direct and indirect effects of termites on savanna tree-seedling growth

Termites are considered to be ecosystem engineers because they modify their biophysical environments. We tested the effects of soil-nutrient alteration in termite-enriched soils compared with termite-free open savannas. We also tested whether non-nutrient alterations (soil disturbance) induced by termites led to changes in tree-seedling growth. Soil samples taken from termite-enriched soils and adjacent open savanna sites in KwaZulu-Natal, South Africa were analyzed for nitrogen, pH, organic carbon and water-holding capacity. Seeds from three dominant tree species, Acacia sieberiana, Celtis africana and Ziziphus mucronata, were grown in soils taken from termite-enriched soils and adjacent savannas. Overall, organic carbon and nitrogen content were higher in termite-enriched soils than in adjacent savannas. We found that these differences in nutrients did not directly affect seedling growth rates or final height. However, C. africana had increased growth rates in compacted termite-enriched soils, while A. sieberiana seedlings were taller in non-compacted soils. We conclude that the indirect effects of disturbance by termites may be as important as the direct effects of increased nutrients for growth of savanna trees.     

Effects of livestock exclusion on in‐stream habitat and benthic invertebrate assemblages in montane streams

1. Stream and riparian ecosystems in arid montane areas, like the interior western United States, are often just narrow mesic strands, but support diverse and productive habitats. Meadows along many such streams have long been used for rangeland grazing, and, while impacts to riparian areas are relatively well known, the effect of livestock grazing on aquatic life in streams has received less attention. 2. Attempts to link grazing impacts to disturbance have been hindered by the lack of spatial and temporal replication. In this study, we compared channel features and benthic macroinvertebrate communities (i) between 16 stream reaches on two grazed allotments and between 22 reaches on two allotments where livestock had been completely removed for 4 years, (ii) before and after the 4‐year grazing respite at a subset of eight sites and (iii) inside and outside of small‐scale fenced grazing exclosures (eight pairings; 10+ year exclosures) in the meadows of the Golden Trout Wilderness, California (U.S.A.). 3. We evaluated grazing disturbance at the reach scale in terms of the effects of livestock trampling on per cent bank erosion and found that macroinvertebrate richness metrics were negatively correlated with bank erosion, while the percentage of tolerant taxa increased. 4. All macroinvertebrate richness metrics were significantly lower in grazed areas. Bank angle, temperature, fine sediment cover and erosion were higher in grazed areas, while riparian cover was lower. Regression models identified riparian cover, in‐stream substratum, bank conditions and bankfull width‐to‐depth ratios as the most important for explaining variability in macroinvertebrate richness metrics. 5. Small‐scale grazing exclosures showed no improvements for in‐stream communities and only moderate positive effects on riparian vegetation. In contrast, metrics of macroinvertebrate richness increased significantly after a 4‐year period of no grazing. 6. The success of grazing removal reported here suggests that short‐term removal of livestock at the larger, allotment meadow spatial scale is more effective than long‐term, but small‐scale, local riparian area fencing, and yields promising results in achieving stream channel, riparian and aquatic biological recovery.     

Hydromorphological restoration of running waters: effects on benthic invertebrate assemblages

1. River restoration has received considerable attention, with much recent focus on restoring river hydromorphology to improve impoverished aquatic communities. However, we still lack a clear understanding of the response of aquatic biota to river restoration. 2. We studied the effects of hydromorphological restoration on benthic invertebrate assemblages in 25 river sites in Germany using standardised methods. Restoration efforts were primarily aimed to restore habitat heterogeneity; correspondingly, habitat diversity increased at most sites. 3. Similarity of benthic invertebrate assemblages between restored and unrestored river sections was low (mean similarity was 0.32 for Jaccard and 0.46 for Sørensen). Community‐based metrics, such as the percentage of Ephemeroptera, Plecoptera and Trichoptera taxa, also differed between restored and unrestored sections. 4. Only three of the 25 restored sections were classified as having ‘good ecological quality’ class according to the European Water Framework Directive criteria; hence, poor water quality is probably one factor impeding recolonisation. 5. Our results show that isolated restoration measures do not necessarily result in positive effects on aquatic biota and that better understanding of the interconnectedness within a catchment is required before we can adequately predict biotic responses to structural river restoration.     

Characterization of bacterial communities in four freshwater lakes differing in nutrient load and food web structure

The phylogenetic composition of bacterioplankton communities in the water column of four shallow eutrophic lakes was analyzed by partially sequencing cloned 16S rRNA genes and by PCR-DGGE analysis. The four lakes differed in nutrient load and food web structure: two were in a clearwater state and had dense stands of submerged macrophytes, while two others were in a turbid state characterized by the occurrence of phytoplankton blooms. One turbid and one clearwater lake had very high nutrient levels (total phosphorus > 100 microg/l), while the other lakes were less nutrient rich (total phosphorus < 100 microg/l). Cluster analysis, multidimensional scaling and ANOSIM (analysis of similarity) were used to investigate differences among the bacterial community composition in the four lakes. Our results show that each lake has its own distinct bacterioplankton community. The samples of lake Blankaart differed substantially from those of the other lakes; this pattern was consistent throughout the year of study. The bacterioplankton community composition in lake Blankaart seems to be less diverse and less stable than in the other three lakes. Clone library results reveal that Actinobacteria strongly dominated the bacterial community in lake Blankaart. The relative abundance of Betaproteobacteria was low, whereas this group was dominant in the other three lakes. Turbid lakes had a higher representation of Cyanobacteria, while clearwater lakes were characterized by more representatives of the Bacteroidetes. Correlating our DGGE data with environmental parameters, using the BIOENV procedure, suggests that differences are partly related to the equilibrium state of the lake.     

Complex indirect effects of epiphytic bromeliads on the invertebrate food webs of their support tree

Ecosystem engineers are species that affect others through the provision of habitat rather than consumptive resources. The extent to which ecosystem engineers can indirectly affect entire food webs, however, is poorly understood. Epiphytic tank bromeliads (Bromeliaceae) are ecosystem engineers that are common throughout the Neotropics, and are associated with a variety of predatory arthropods. Here, we examine if bromeliads, by increasing predator densities, indirectly benefit their support tree through reduction in herbivorous insects and leaf damage. We observed and manipulated bromeliad densities in Costa Rican orange orchards, and measured impacts on leaf damage and arboreal and bromeliad invertebrate communities in two different seasons. Our results show that bromeliads are associated with predatory and herbivorous invertebrates but not leaf damage. Bromeliads were correlated with increased densities of their associated predators, especially ants and hunting spiders, but we could not confirm a causal link. Associations with bromeliads changed over time, with seasonal shifts interfering with responses to our manipulations. Bromeliads had a reduced association with predators in the dry season. Moreover, a null association between bromeliads and herbivorous invertebrates in the dry season unexpectedly became positive in the wet season. In summary, we have only limited evidence that bromeliads indirectly promote the top‐down control of arboreal herbivores; instead, our manipulations suggest that bromeliads increase herbivore densities in the wet season. This research suggests that although bromeliads may act as ecosystem engineers, indirectly influencing the invertebrate food web on support trees, their effects are trophically complex and seasonally dependent. Abstract in Spanish is available with online material.     

The response of two estuarine benthic communities to the quantity and quality of food

Experimental manipulations of food supply were performed on soft sediment cores from two European estuaries, the Westerscheldt and the Gironde, with a view to determining benthic macrofaunal community response. Over a period of twenty weeks in a laboratory mesocosm system, both communities showed losses in terms of numbers of individuals and small, but non-significant, losses in terms of numbers of species. Whereas no effect of the different types of foods or the dose levels at which they were supplied was detected for the Westerscheldt benthic community, that of the Gironde showed some significant response. This was largely attributed to the differential mortality of spionid polychaetes across the dose levels used, with the highest dose, equivalent to 200 g C m^–2 yr^–1, only just maintaining their initial population densities. The results are discussed in terms of the importance of lateral advection of food particles at the benthic boundary layer and the general insufficiency of many estimates of carbon input to shallow benthic systems.     

The independent and interactive effects of fine sediment and flow on benthic invertebrate communities characteristic of small upland streams

SUMMARY 1. Floods are an important mechanism of disturbance operating in streams that can markedly influence the abundance and diversity of benthic fauna. In upland streams many studies cite the scouring effects of fine transported sediments as a potential source of disturbance to the biota during spates, but few studies have sought to test this hypothesis critically.
2. Here we used a series of eight artificial streams to test whether high suspended-sediment concentrations influenced the short-term response of benthic invertebrate fauna to increases in flow. In an experiment designed to simulate a small spate, flow and sediment loads were each manipulated to examine their independent and interactive effects. Benthic invertebrates were sampled before and after the manipulation, and drift samples were taken at regular intervals during the experiment. The experiment was repeated twice, once at the end of winter (June, 1998), and once in summer (February, 1999).
3. Flow increases caused large increases in the number and diversity of drifting animals, and significant declines in the numbers and diversity of organisms found in benthic samples, but these declines were apparently not affected by the addition of fine sediment. The addition of sediment alone had little effect on the fauna. These results were consistent across both experiments.
4. The results suggest that flow increases alone can disturb benthic fauna, and that neither substrate movement nor suspended sediment increases are necessary for floods and spates to disturb the benthic assemblage. However, as argued elsewhere, the effects of flow increases are likely to be contingent upon the presence or absence of local flow refugia, which can allow animals to escape the shear forces that would otherwise remove them from the surface or interstitial areas of the streambed.     

Effects of salinity gradients on benthic invertebrate and diatom communities in a German lowland river

The Lippe is a lowland river located in the Western part of Germany and has been heavily impacted by coal mining activities ever since. Although mining activities significantly decreased during the last decades, the associated discharge of salt-enriched mine water into the river still poses a persistent threat to the local benthic invertebrate and diatom communities. To analyze the effect of salt pollution on invertebrate and diatom species, biological and chemical data were compiled for this study from a publicly available database. Changes in the community composition due to increased salt concentrations were explored by Non-Metrical-Multidimensional Scaling. Indicator species and salinity thresholds for single species and communities were identified using the method TITAN (Threshold Indicator Taxa Analysis). The method is an analytical approach to detect changes in frequency and abundance of species along an environmental gradient by combining the methods of change point analysis (nCPA) and indicator species analysis. The obtained salinity preferences and individual and community thresholds were compared to the literature and existing salinity classifications. For both diatoms and benthic invertebrates, Non-Metrical-Multidimensional Scaling showed a clear split between samples of high and low salinities. Significant salinity thresholds were determined for 50 invertebrate and 58 diatom species of which 23 respectively 18 species were described as ‘reliable’ indicators according to the specifications given by Baker and King (2010). A majority of salt-tolerant indicator organisms were invasive species. For both organism groups, major changes in community composition were detected at a conductivity value exceeding 900 μS/cm. A reduction of the average salinity to below this threshold may have positive effects on the overall species richness and the persistence of sensitive taxa in the river Lippe. Individual and community thresholds may however be data-dependent to a certain degree and subjected to fluctuations considering the potential interdependencies between salinity and additional physico-chemical and environmental parameters (e.g. water temperature, lime content).     

Damaging UV radiation and invertebrate predation: conflicting selective pressures for zooplankton vertical distribution in the water column of low DOC lakes

In nature most organisms have to manage conflicting demands of food gathering, predator avoidance, and finding a favorable abiotic environment (oxygen, temperature, etc.) in order to maximize their fitness. In the vertical water column of lakes with high solar ultraviolet radiation (UV) and invertebrate predators, zooplankton face two particularly strong and conflicting selective pressures. During daylight hours invertebrate predators often induce an upward vertical migration of zooplankton prey while potentially damaging UV forces a downward migration. We used 2.2 m long columns suspended vertically in a lake to conduct 2×2 factorial experiments to examine patterns of depth selection behavior by zooplankton in the presence and absence of both the invertebrate predator Chaoborus and UV. We hypothesized that Chaoborus and UV both affect the distribution of zooplankton and a combination of both factors would lead to a narrowing of depth distribution. We found that when Chaoborus were present zooplankton tended to be distributed at shallower depths in the columns, while in the presence of UV they exhibited a deeper distribution. Chaoborus themselves were always found near the bottom of the columns regardless of the UV treatment. Simultaneous exposure to predators and UV resulted in a peak of zooplankton (especially Daphnia catawba) distribution at intermediate depths. In a significant number of cases, depth range was narrowed in response to Chaoborus, UV, or both.     

Direct and indirect effects of area,energy and habitat heterogeneity on breeding bird communities

Aim To compare the ability of island biogeography theory, niche theory and species–energy theory to explain patterns of species richness and density for breeding bird communities across islands with contrasting characteristics. Location Thirty forested islands in two freshwater lakes in the boreal forest zone of northern Sweden (65°55′ N to 66°09′ N; 17°43′ E to 17°55′ E). Methods We performed bird censuses on 30 lake islands that have each previously been well characterized in terms of size, isolation, habitat heterogeneity (plant diversity and forest age), net primary productivity (NPP), and invertebrate prey abundance. To test the relative abilities of island biogeography theory, niche theory and species–energy theory to describe bird community patterns, we used both traditional statistical approaches (linear and multiple regressions) and structural equation modelling (SEM; in which both direct and indirect influences can be quantified). Results Using regression‐based approaches, area and bird abundance were the two most important predictors of bird species richness. However, when the data were analysed by SEM, area was not found to exert a direct effect on bird species richness. Instead, terrestrial prey abundance was the strongest predictor of bird abundance, and bird abundance in combination with NPP was the best predictor of bird species richness. Area was only of indirect importance through its positive effect on terrestrial prey abundance, but habitat heterogeneity and spatial subsidies (emerging aquatic insects) also showed important indirect influences. Thus, our results provided the strongest support for species–energy theory. Main conclusions Our results suggest that, by using statistical approaches that allow for analyses of both direct and indirect influences, a seemingly direct influence of area on species richness can be explained by greater energy availability on larger islands. As such, animal community patterns that seem to be in line with island biogeography theory may be primarily driven by energy availability. Our results also point to the need to consider several aspects of habitat quality (e.g. heterogeneity, NPP, prey availability and spatial subsidies) for successful management of breeding bird diversity at local spatial scales and in fragmented or insular habitats.