Ecosystem-size relationships of river populations and communities.

Knowledge of ecosystem-size influences on river populations and communities is integral to the balancing of human and environmental needs for water. The multiple dimensions of dendritic river networks complicate understanding of ecosystem-size influences, but could be resolved by the development of scaling relationships. We highlight the importance of physical constraints limiting predator body sizes, movements, and population sizes in small rivers, and where river contraction limits space or creates stressful conditions affecting community stability and food webs. Investigations of the scaling and contingency of these processes will be insightful because of the underlying generality and scale independence of such relationships. Doing so will also pinpoint damaging water-management practices and identify which aspects of river size can be most usefully manipulated in river restoration.

Consequences of climate-induced range expansions on multiple ecosystem functions.

Climate-driven species range shifts and expansions are changing community composition, yet the functional consequences in natural systems are mostly unknown. By combining a 30-year survey of subalpine pond larval caddisfly assemblages with species-specific functional traits (nitrogen and phosphorus excretion, and detritus processing rates), we tested how three upslope range expansions affected species' relative contributions to caddisfly-driven nutrient supply and detritus processing. A subdominant resident species (Ag. deflata) consistently made large relative contributions to caddisfly-driven nitrogen supply throughout all range expansions, thus "regulating" the caddisfly-driven nitrogen supply. Whereas, phosphorus supply and detritus processing were regulated by the dominant resident species (L. externus) until the third range expansion (by N. hostilis). Since the third range expansion, N. hostilis's relative contribution to caddisfly-driven phosphorus supply increased, displacing L. externus's role in regulating caddisfly-driven phosphorus supply. Meanwhile, detritus processing contributions became similar among the dominant resident, subdominant residents, and range expanding species. Total ecosystem process rates did not change throughout any of the range expansions. Thus, shifts in species' relative functional roles may occur before shifts in total ecosystem process rates, and changes in species' functional roles may stabilize processes in ecosystems undergoing change.

Species-specific traits predict whole-assemblage detritus processing by pond invertebrates.

Functional trait diversity determines if ecosystem processes are sensitive to shifts in species abundances or composition. For example, trait variation suggests detritivores process detritus at different rates and make different contributions to whole-assemblage processing, which could be sensitive to compositional shifts. Here, we used a series of microcosm experiments to quantify species-specific coarse and fine particulate organic matter (CPOM and FPOM) processing for ten larval caddisfly species and three non-caddisfly species in high-elevation wetlands. We then compared trait-based models including life history, dietary, and extrinsic traits to determine which traits explained interspecific variation in detritus processing. Finally, we compared processing by mixed caddisfly assemblages in microcosms and natural ponds to additive predictions based on species-specific processing to determine if single-species effects are additive in multi-species assemblages. We found considerable interspecific variation in biomass-specific CPOM (13-fold differences) and FPOM (8-fold differences) processing. Furthermore, on a mass-specific basis, amphipods, chironomids, and caddisflies processed similar amounts of detritus, suggesting non-shredder taxa could process more than previously recognized. Trait models including dietary percent detritus, development rate, body size, and wetland hydroperiod explained 81 and 57% of interspecific variation in CPOM and FPOM processing, respectively. Finally, species-specific additive predictions were strikingly similar to mixed-assemblage processing in microcosms and natural ponds, with the largest difference being a 15% overestimate. Thus, additivity of species-specific processing suggests single-species rates may be useful for understanding functional consequences of shifting assemblages, and a trait-based approach to predicting species-specific processing could support generating additive predictions of whole-assemblage processing.

Regulation of open populations of a stream insect through larval density dependence.

In organisms with complex life cycles, the various stages occupy different habitats creating demographically open populations. The dynamics of these populations will depend on the occurrence and timing of stochastic influences relative to demographic density dependence, but understanding of these fundamentals, especially in the face of climate warming, has been hampered by the difficulty of empirical studies. Using a logically feasible organism, we conducted a replicated density-perturbation experiment to manipulate late-instar larvae of nine populations of a stream caddisfly, Zelandopsyche ingens, and measured the resulting abundance over 2 years covering the complete life cycle of one cohort to evaluate influences on dynamics. Negative density feedback occurred in the larval stage, and was sufficiently strong to counteract variation in abundance due to manipulation of larval density, adult caddis dispersal in the terrestrial environment as well as downstream drift of newly hatched and older larvae in the current. This supports theory indicating regulation of open populations must involve density dependence in local populations sufficient to offset variability associated with dispersal, especially during recruitment, and pinpoints the occurrence to late in the larval life cycle and driven by food resource abundance. There were large variations in adult, egg mass and early instar abundance that were not related to abundance in the previous stage, or the manipulation, pointing to large stochastic influences. Thus, the results also highlight the complementary nature of stochastic and deterministic influences on open populations. Such density dependence will enhance population persistence in situations where variable dispersal and transitioning between life stages frequently creates mismatches between abundance and the local availability of resources, such as might become more common with climate warming.

Perils of life on the edge: Climatic threats to global diversity patterns of wetland macroinvertebrates.

Climate change is rapidly driving global biodiversity declines. How wetland macroinvertebrate assemblages are responding is unclear, a concern given their vital function in these ecosystems. Using a data set from 769 minimally impacted depressional wetlands across the globe (467 temporary and 302 permanent), we evaluated how temperature and precipitation (average, range, variability) affects the richness and beta diversity of 144 macroinvertebrate families. To test the effects of climatic predictors on macroinvertebrate diversity, we fitted generalized additive mixed-effects models (GAMM) for family richness and generalized dissimilarity models (GDMs) for total beta diversity. We found non-linear relationships between family richness, beta diversity, and climate. Maximum temperature was the main climatic driver of wetland macroinvertebrate richness and beta diversity, but precipitation seasonality was also important. Assemblage responses to climatic variables also depended on wetland water permanency. Permanent wetlands from warmer regions had higher family richness than temporary wetlands. Interestingly, wetlands in cooler and dry-warm regions had the lowest taxonomic richness, but both kinds of wetlands supported unique assemblages. Our study suggests that climate change will have multiple effects on wetlands and their macroinvertebrate diversity, mostly via increases in maximum temperature, but also through changes in patterns of precipitation. The most vulnerable wetlands to climate change are likely those located in warm-dry regions, where entire macroinvertebrate assemblages would be extirpated. Montane and high-latitude wetlands (i.e., cooler regions) are also vulnerable to climate change, but we do not expect entire extirpations at the family level.

Habitat size influences community stability.

Mechanisms linked to demographic, biogeographic, and food-web processes thought to underpin community stability could be affected by habitat size, but the effects of habitat size on community stability remain relatively unknown. We investigated whether those habitat-size-dependent properties influenced community instability and vulnerability to perturbations caused by disturbance. This is particularly important given that human exploitation is contracting ecosystems, and abiotic perturbations are becoming more severe and frequent. We used a perturbation experiment in which 10 streams, spanning three orders of magnitude in habitat size, were subjected to simulated bed movement akin to a major flood disturbance event. We measured the resistance, resilience, and variability of basal resources, and population and community-level responses across the stream habitat-size gradient immediately before, and at 0.5, 5, 10, 20, and 40 d post-disturbance. Resistance to disturbance consistently increased with stream size in all response variables. In contrast, resilience was significantly higher in smaller streams for some response variables. However, this higher resilience of small ecosystems was insufficient to compensate for their lower resistance, and communities of smaller streams were thus more variable over time than those of larger streams. Compensatory dynamics of populations, especially for predators, stabilized some aspects of communities, but these mechanisms were unrelated to habitat size. Together, our results provide compelling evidence for the links between habitat size and community stability, and should motivate ecologists and managers to consider how changes in the size of habitats will alter the vulnerability of ecosystems to perturbations caused by environmental disturbance.

Predators balance consequences of climate-change-induced habitat shifts for range-shifting and resident species.

While many species distributions are shifting poleward or up in elevation in response to a changing climate, others are shifting their habitats along localized gradients in environmental conditions as abiotic conditions become more stressful. Whether species are moving across regional or local environmental gradients in response to climate change, range-shifting species become embedded in established communities of competitors and predators. The consequences of these shifts for both resident and shifting species are often unknown, as it can be difficult to isolate the effects of multiple species interactions. Using a model system of insects in high-elevation ponds in the Rocky Mountains of Colorado, we sought to disentangle the effects of predation and intraguild interactions on the survival and development of a semi-permanent pond resident caddisfly Limnephilus externus and the habitat-shifting caddis Asynarchus nigriculus that is being forced into semi-permanent ponds as temporary ponds dry too quickly to complete development. We conducted a manipulative in-situ pond cage experiment in which L. externus and A. nigriculus caddisfly larvae in single-species treatments and together were exposed to the presence/absence of predatory Dytiscus diving beetle larvae. This approach allowed us to isolate the effects of intraguild interactions and predation on the survival and development of both the resident and habitat-shifting species. We found that intraguild interactions had strong negative effects on the resident and habitat-shifting species. Intraguild interactions reduced the survival of the resident L. externus and increased the variation in survival of the shifting A. nigriculus. However, Dytiscus predators reduced these negative effects, stabilizing the community by increasing L. externus survival and reducing variation in A. nigriculus survival. We also found that intraguild interactions reduced L. externus biomass but resulted in increased A. nigriculus development. A. nigriculus development was also increased by predation. Our results show that strong intraguild interactions between resident and shifting species are likely to have negative consequences for both species. However, the presence of predators reduces these negative consequences of the habitat shift on both the resident and the shifting.

Does taxonomic and numerical resolution affect the assessment of invertebrate community structure in New World freshwater wetlands?

The efficiency of biodiversity assessments and biomonitoring studies is commonly challenged by limitations in taxonomic identification and quantification approaches. In this study, we assessed the effects of different taxonomic and numerical resolutions on a range of community structure metrics in invertebrate compositional data sets from six regions distributed across North and South America. We specifically assessed the degree of similarity in the metrics (richness, equitability, beta diversity, heterogeneity in community composition and congruence) for data sets identified to a coarse resolution (usually family level) and the finest taxonomic resolution practical (usually genus level, sometimes species or morphospecies) and by presence-absence and relative abundance numerical resolutions. Spearman correlations showed highly significant and positive associations between univariate metrics (richness and equitability) calculated for coarse- and finest-resolution datasets. Procrustes analysis detected significant congruence between composition datasets. Higher correlation coefficients were found for datasets with the same numerical resolutions regardless of the taxonomic level (about 90%), while the correlations for comparisons across numerical resolutions were consistently lower. Our findings indicate that family-level resolution can be used as a surrogate of finer taxonomic resolutions to calculate a range of biodiversity metrics commonly used to describe invertebrate community structure patterns in New World freshwater wetlands without significant loss of information. However, conclusions on biodiversity patterns derived from datasets with different numerical resolutions should be critically considered in studies on wetland invertebrates.

Elevation alters outcome of competition between resident and range-shifting species.

Species' geographic range shifts toward higher latitudes and elevations are among the most frequently reported consequences of climate change. However, the role of species interactions in setting range margins remains poorly understood. We used cage experiments in ponds to test competing hypotheses about the role of abiotic and biotic mechanisms for structuring range boundaries of an upslope range-shifting caddisfly Limnephilus picturatus. We found that competition with a ubiquitous species Limnephilus externus significantly decreased L. picturatus survival and emergence at subalpine elevations supporting the notion that species interactions play a critical role in determining upslope range limits. However, without competitors, L. picturatus survival was greater at high-elevation than low-elevation sites. This was contrary to decreases in body mass (a proxy for fecundity) with elevation regardless of the presence of competitors. We ultimately show that species interactions can be important for setting upslope range margins. Yet, our results also highlight the complications in defining what may be abiotically stressful for this species and the importance of considering multiple demographic variables. Understanding how species ranges will respond in a changing climate will require quantifying species interactions and how they are influenced by the abiotic context in which they play out.

Repeat disturbances have cumulative impacts on stream communities.

Climate change has altered disturbance regimes in many ecosystems, and predictions show that these trends are likely to continue. The frequency of disturbance events plays a particularly important role in communities by selecting for disturbance-tolerant taxa.However, ecologists have yet to disentangle the influence of disturbance frequency per se and time since last disturbance, because more frequently disturbed systems have also usually been disturbed more recently. Our understanding of the effects of repeated disturbances is therefore confounded by differences in successional processes.We used in-situ stream mesocosms to isolate and examine the effect of disturbance frequency on community composition. We applied substrate moving disturbances at five frequencies, with the last disturbance occurring on the same day across all treatments. Communities were then sampled after a recovery period of 9 days.Macroinvertebrate community composition reflected the gradient of disturbance frequency driven by differential vulnerability of taxa to disturbance. Diversity metrics, including family-level richness, decreased, reflecting a likely loss of functional diversity with increasing disturbance frequency. In contrast, overall abundance was unaffected by disturbance frequency as rapid recovery of the dominant taxon compensated for strong negative responses of disturbance-vulnerable taxa.We show that cumulative effects of repeated disturbances-not just the time communities have had to recover before sampling-alter communities, especially by disproportionately affecting rare taxa. Thus, the timing of past disturbances can have knock-on effects that determine how a system will respond to further change.

Landscape heterogeneity strengthens the relationship between ß-diversity and ecosystem function.

Consensus has emerged in the literature that increased biodiversity enhances the capacity of ecosystems to perform multiple functions. However, most biodiversity/ecosystem function studies focus on a single ecosystem, or on landscapes of homogenous ecosystems. Here, we investigate how increased landscape-level environmental dissimilarity may affect the relationship between different metrics of diversity (α, ß, or γ) and ecosystem function. We produced a suite of simulated landscapes, each of which contained four experimental outdoor aquatic mesocosms. Differences in temperature and nutrient conditions of the mesocosms allowed us to simulate landscapes containing a range of within-landscape environmental heterogeneities. We found that the variation in ecosystem functions was primarily controlled by environmental conditions, with diversity metrics accounting for a smaller (but significant) amount of variation in function. When landscapes were more homogeneous, α, ß, and γ diversity was not associated with differences in primary production, and only γ was associated with changes in decomposition. In these homogeneous landscapes, differences in these two ecosystem functions were most strongly related to nutrient and temperature conditions in the ecosystems. However, as landscape-level environmental dissimilarity increased, the relationship between α, ß, or γ and ecosystem functions strengthened, with ß being a greater predictor of variation in decomposition at the highest levels of environmental dissimilarity than α or γ. We propose that when all ecosystems in a landscape have similar environmental conditions, species sorting is likely to generate a single community composition that is well suited to those environmental conditions, ß is low, and the efficiency of diversity-ecosystem function couplings is similar across communities. Under this low ß, the effect of abiotic conditions on ecosystem function will be most apparent. However, when environmental conditions vary among ecosystems, species sorting pressures are different among ecosystems, producing different communities among locations in a landscape. These conditions lead to stronger relationships between ß and the magnitude of ecosystem functions. Our results illustrate that abiotic conditions and the homogeneity of communities influence ecosystem function expressed at the landscape scale.

Capacity to support predators scales with habitat size.

Habitat reduction could drive biodiversity loss if the capacity of food webs to support predators is undermined by habitat-size constraints on predator body size. Assuming that (i) available space restricts predator body size, (ii) mass-specific energy needs of predators scale with their body size, and (iii) energy availability scales with prey biomass, we predicted that predator biomass per unit area would scale with habitat size (quarter-power exponent) and prey biomass (three-quarter-power exponent). We found that total predator biomass scaled with habitat size and prey resources as expected across 29 New Zealand rivers, such that a unit of habitat in a small ecosystem supported less predator biomass than an equivalent unit in a large ecosystem. The lower energetic costs of large body size likely mean that a unit of prey resource supports more biomass of large-bodied predators compared to small-bodied predators. Thus, contracting habitat size reduces the predator mass that can be supported because of constraints on predator body size, and this may be a powerful mechanism exacerbating reductions in biodiversity due to habitat loss.

The body size dependence of trophic cascades.

Trophic cascades are indirect positive effects of predators on resources via control of intermediate consumers. Larger-bodied predators appear to induce stronger trophic cascades (a greater rebound of resource density toward carrying capacity), but how this happens is unknown because we lack a clear depiction of how the strength of trophic cascades is determined. Using consumer resource models, we first show that the strength of a trophic cascade has an upper limit set by the interaction strength between the basal trophic group and its consumer and that this limit is approached as the interaction strength between the consumer and its predator increases. We then express the strength of a trophic cascade explicitly in terms of predator body size and use two independent parameter sets to calculate how the strength of a trophic cascade depends on predator size. Both parameter sets predict a positive effect of predator size on the strength of a trophic cascade, driven mostly by the body size dependence of the interaction strength between the first two trophic levels. Our results support previous empirical findings and suggest that the loss of larger predators will have greater consequences on trophic control and biomass structure in food webs than the loss of smaller predators.

A bioenergetic framework for the temperature dependence of trophic interactions.

Changing temperature can substantially shift ecological communities by altering the strength and stability of trophic interactions. Because many ecological rates are constrained by temperature, new approaches are required to understand how simultaneous changes in multiple rates alter the relative performance of species and their trophic interactions. We develop an energetic approach to identify the relationship between biomass fluxes and standing biomass across trophic levels. Our approach links ecological rates and trophic dynamics to measure temperature-dependent changes to the strength of trophic interactions and determine how these changes alter food web stability. It accomplishes this by using biomass as a common energetic currency and isolating three temperature-dependent processes that are common to all consumer-resource interactions: biomass accumulation of the resource, resource consumption and consumer mortality. Using this framework, we clarify when and how temperature alters consumer to resource biomass ratios, equilibrium resilience, consumer variability, extinction risk and transient vs. equilibrium dynamics. Finally, we characterise key asymmetries in species responses to temperature that produce these distinct dynamic behaviours and identify when they are likely to emerge. Overall, our framework provides a mechanistic and more unified understanding of the temperature dependence of trophic dynamics in terms of ecological rates, biomass ratios and stability.

Increased temperature variation poses a greater risk to species than climate warming.

Increases in the frequency, severity and duration of temperature extremes are anticipated in the near future. Although recent work suggests that changes in temperature variation will have disproportionately greater effects on species than changes to the mean, much of climate change research in ecology has focused on the impacts of mean temperature change. Here, we couple fine-grained climate projections (2050-2059) to thermal performance data from 38 ectothermic invertebrate species and contrast projections with those of a simple model. We show that projections based on mean temperature change alone differ substantially from those incorporating changes to the variation, and to the mean and variation in concert. Although most species show increases in performance at greater mean temperatures, the effect of mean and variance change together yields a range of responses, with temperate species at greatest risk of performance declines. Our work highlights the importance of using fine-grained temporal data to incorporate the full extent of temperature variation when assessing and projecting performance.

Top-down control of prey increases with drying disturbance in ponds: a consequence of non-consumptive interactions?

1. Biotic interactions are often expected to decrease in intensity as abiotic conditions become more stressful to organisms. However, in many cases, food-web and habitat complexity also change with abiotic stress or disturbance, potentially altering patterns of species interactions across environmental gradients. 2. We used a combination of field assays and mesocosm experiments to investigate how disturbance from desiccation moderates top-down control of prey by predators across a gradient of pond duration in New Zealand. 3. Field manipulations of predator abundance in ponds led to an unexpected decrease in the top-down control of prey biomass by predatory invertebrates as pond duration increased (decreasing abiotic stress). Predatory fish, which are restricted to permanent ponds, had negligible effects on prey biomass. Mesocosm experiments further indicated the consumptive effects of fish are weak; a result that cannot be explained by an increase in physical habitat refugia in relatively more permanent ponds. 4. Manipulations of invertebrate predator diversity in mesocosms (both substitutive and additive treatments), and the addition of olfactory fish cues, revealed that strong non-consumptive effects of fish reduced predation by predatory invertebrates, and these effects overwhelmed the positive influence of invertebrate predator diversity on prey consumption. 5. These results suggest that decreases in top-down control with increasing pond permanence are likely a result of non-consumptive effects of fish weakening predation by invertebrate predators in the more complex food webs of permanent ponds. Therefore, considering non-consumptive effects of predators in complex food webs will likely improve the understanding of biotic interactions across environmental gradients.

Warming shifts top-down and bottom-up control of pond food web structure and function.

The effects of global and local environmental changes are transmitted through networks of interacting organisms to shape the structure of communities and the dynamics of ecosystems. We tested the impact of elevated temperature on the top-down and bottom-up forces structuring experimental freshwater pond food webs in western Canada over 16 months. Experimental warming was crossed with treatments manipulating the presence of planktivorous fish and eutrophication through enhanced nutrient supply. We found that higher temperatures produced top-heavy food webs with lower biomass of benthic and pelagic producers, equivalent biomass of zooplankton, zoobenthos and pelagic bacteria, and more pelagic viruses. Eutrophication increased the biomass of all organisms studied, while fish had cascading positive effects on periphyton, phytoplankton and bacteria, and reduced biomass of invertebrates. Surprisingly, virus biomass was reduced in the presence of fish, suggesting the possibility for complex mechanisms of top-down control of the lytic cycle. Warming reduced the effects of eutrophication on periphyton, and magnified the already strong effects of fish on phytoplankton and bacteria. Warming, fish and nutrients all increased whole-system rates of net production despite their distinct impacts on the distribution of biomass between producers and consumers, plankton and benthos, and microbes and macrobes. Our results indicate that warming exerts a host of indirect effects on aquatic food webs mediated through shifts in the magnitudes of top-down and bottom-up forcing.

Warming modifies trophic cascades and eutrophication in experimental freshwater communities.

Climate warming is occurring in concert with other anthropogenic changes to ecosystems. However, it is unknown whether and how warming alters the importance of top-down vs. bottom-up control over community productivity and variability. We performed a 16-month factorial experimental manipulation of warming, nutrient enrichment, and predator presence in replicated freshwater pond mesocosms to test their independent and interactive impacts. Warming strengthened trophic cascades from fish to primary producers, and it decreased the impact of eutrophication on the mean and temporal variation of phytoplankton biomass. These impacts varied seasonally, with higher temperatures leading to stronger trophic cascades in winter and weaker algae blooms under eutrophication in summer. Our results suggest that higher temperatures may shift the control of primary production in freshwater ponds toward stronger top-down and weaker bottom-up effects. The dampened temporal variability of algal biomass under eutrophication at higher temperatures suggests that warming may stabilize some ecosystem processes.

Nonlinear effects of consumer density on multiple ecosystem processes.

1. In the face of human-induced declines in the abundance of common species, ecologists have become interested in quantifying how changes in density affect rates of biophysical processes, hence ecosystem function. We manipulated the density of a dominant detritivore (the cased caddisfly, Limnephilus externus) in subalpine ponds to measure effects on the release of detritus-bound nutrients and energy. 2. Detritus decay rates (k, mass loss) increased threefold, and the loss of nitrogen (N) and phosphorus (P) from detrital substrates doubled across a range of historically observed caddisfly densities. Ammonium and total soluble phosphorus concentrations in the water column also increased with caddisfly density on some dates. Decay rates, nutrient release and the change in total detritivore biomass all exhibited threshold or declining responses at the highest densities. 3. We attributed these threshold responses in biophysical processes to intraspecific competition for limiting resources manifested at the population level, as density-dependent per-capita consumption, growth, development and case : body size in caddisflies was observed. Moreover, caddisflies increasingly grazed on algae at high densities, presumably in response to limiting detrital resources. 4. These results provide evidence that changes in population size of a common species will have nonlinear, threshold effects on the rates of biophysical processes at the ecosystem level. Given the ubiquity of negative density dependence in nature, nonlinear consumer density-ecosystem function relationships should be common across species and ecosystems.

Heavy metals: confounding factors in the response of New Zealand freshwater fish assemblages to natural and anthropogenic acidity.

Acidification of freshwaters is a global phenomenon, occurring both through natural leaching of organic acids and through human activities from industrial emissions and mining. The West Coast of the South Island, New Zealand, has both naturally acidic and acid mine drainage (AMD) streams enabling us to investigate the response of fish communities to a gradient of acidity in the presence and absence of additional stressors such as elevated concentrations of heavy metals. We surveyed a total of 42 streams ranging from highly acidic (pH 3.1) and high in heavy metals (10 mg L(-)(1) Fe; 38 mg L(-)(1) Al) to circum-neutral (pH 8.1) and low in metals (0.02 mg L(-)(1) Fe; 0.05 mg L(-)(1) Al). Marked differences in pH and metal tolerances were observed among the 15 species that we recorded. Five Galaxias species, Anguilla dieffenbachii and Anguillaaustralis were found in more acidic waters (pH<5), while bluegill bullies (Gobiomorphus hubbsi) and torrentfish (Cheimarrichthys fosteri) were least tolerant of low pH (minimum pH 6.2 and 5.5, respectively). Surprisingly, the strongest physicochemical predictor of fish diversity, density and biomass was dissolved metal concentrations (Fe, Al, Zn, Mn and Ni) rather than pH. No fish were detected in streams with dissolved metal concentrations >2.7 mg L(-)(1) and nine taxa were only found in streams with metal concentrations <1 mg L(-)(1). The importance of heavy metals as critical drivers of fish communities has not been previously reported in New Zealand, although the mechanism of the metal effects warrants further study. Our findings indicate that any remediation of AMD streams which seeks to enable fish recolonisation should aim to improve water quality by raising pH above approximately 4.5 and reducing concentrations of dissolved Al and Fe to <1.0 mg L(-)(1).