Disturbance-mediated consumer assemblages determine fish community structure and moderate top-down influences through bottom-up constraints.

Disturbance is a strong structuring force that can influence the strength of species interactions at all trophic levels, but controls on the contributions to community structure of top-down and bottom-up processes across such gradients remain poorly understood. Changes in the composition of predator and consumer assemblages, and their associated traits, across gradients of environmental harshness (e.g. flooding), are likely to be a particularly important influence on the strength of top-down control and may drive bottom-up constraints. We examined how consumers with particular traits, and the predators that consumed them, varied across a gradient of stream flooding disturbance and used experiments to assess the predation impact on those contrasting consumer communities (ultimately quantifying how flood disturbance altered the strength of top-down control). Consumer community composition and mobility were strongly related to flood disturbance; the biomass and drift of protected primary consumers (i.e. those with morphological defences) decreased with increasing flood disturbance. Predatory fish species had different disturbance niches, and path analysis identified that both direct flood disturbance effects and indirect bottom-up constraints of flood disturbance on consumers influenced predatory fish composition and biomass. Fish generally fed most effectively on consumer types associated with their particular niche, but all fish were strongly size-selective when feeding on protected consumers. Although protected consumers did not grow large enough to escape predation, an in situ experiment showed protected consumers were at a reduced risk of predation as disturbance increased compared to unprotected consumers. Overall, top-down control declined with flood disturbance, but the effect depended on consumer traits. Predatory fish were only capable of exerting top-down control on protected consumers in benign habitats but impacted unprotected consumers across a larger range of the disturbance gradient. Collectively our findings suggest that a shift towards a more disturbed state will probably result in reduced predator impacts and a weakening of top-down control. Moreover, predicted increases in the frequency and intensity of climatic events causing disturbance, such as flooding, are likely to result in a community shift that disproportionately impacts protected consumers and the predators that utilize them as prey through the subsequent bottom-up constraints.

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.

Increases in disturbance and reductions in habitat size interact to suppress predator body size.

Food webs are strongly size-structured so will be vulnerable to changes in environmental factors that affect large predators. However, mechanistic understanding of environmental controls of top predator size is poorly developed. We used streams to investigate how predator body size is altered by three fundamental climate change stressors: reductions in habitat size, increases in disturbance and warmer temperatures. Using new survey data from 74 streams, we showed that habitat size and disturbance were the most important stressors influencing predator body size. A synergistic interaction between that habitat size and disturbance due to flooding meant the sizes of predatory fishes peaked in large, benign habitats and their body size decreased as habitats became either smaller or harsher. These patterns were supported by experiments indicating that habitat-size reductions and increased flood disturbance decreased both the abundance and biomass of large predators. This research indicates that interacting climate change stressors can influence predator body size, resulting in smaller predators than would be predicted from examining an environmental factor in isolation. Thus, climate-induced changes to key interacting environmental factors are likely to have synergistic impacts on predator body size which, because of their influence on the strength of biological interactions, will have far-reaching effects on food-web responses to global environmental change.

Limitation and facilitation of one of the world's most invasive fish: an intercontinental comparison.

Purposeful species introductions offer opportunities to inform our understanding of both invasion success and conservation hurdles. We evaluated factors determining the energetic limitations of brown trout (Salmo trutta) in both their native and introduced ranges. Our focus was on brown trout because they are nearly globally distributed, considered one of the world's worst invaders, yet imperiled in much of their native habitat. We synthesized and compared data describing temperature regime, diet, growth, and maximum body size across multiple spatial and temporal scales, from country (both exotic and native habitats) and major geographic area (MGA) to rivers and years within MGA. Using these data as inputs, we next used bioenergetic efficiency (BioEff), a relative scalar representing a realized percentage of maximum possible consumption (0-100%) as our primary response variable and a multi-scale, nested, mixed statistical model (GLIMMIX) to evaluate variation among and within spatial scales and as a function of density and elevation. MGA and year (the residual) explained the greatest proportion of variance in BioEff. Temperature varied widely among MGA and was a strong driver of variation in BioEff. We observed surprisingly little variation in the diet of brown trout, except the overwhelming influence of the switch to piscivory observed only in exotic MGA. We observed only a weak signal of density-dependent effects on BioEff; however, BioEff remained < 50% at densities > 2.5 fish/m2. The trajectory of BioEff across the life span of the fish elucidated the substantial variation in performance among MGAs; the maximum body size attained by brown trout was consistently below 400 mm in native habitat but reached approximately 600 mm outside their native range, where brown trout grew rapidly, feeding in part on naive prey fishes. The integrative, physiological approach, in combination with the intercontinental and comparative nature of our study, allowed us to overcome challenges associated with context-dependent variation in determining invasion success. Overall our results indicate "growth plasticity across the life span" was important for facilitating invasion, and should be added to lists of factors characterizing successful invaders.

Pulse-dose application of chelated copper to a river for Didymosphenia geminata control: effects on macroinvertebrates and fish.

A 1-h pulse-dose of a chelated Cu formulation (Gemex™; New Zealand) was applied to a river to test efficacy against the invasive mat-forming diatom Didymosphenia geminata (didymo) and to provide information on nontarget species effects that could not be adequately predicted from laboratory and experimental mesocosm studies. Intensive sampling allowed characterization of doses achieved at multiple downstream locations, and concurrent application of rhodamine dye allowed quantification of dispersion, adsorption, and dilution processes. The target dose of 10 to 20 mg Cu/L for 60 min was achieved at least 0.9 km downstream at sites with contrasting levels of didymo mat development. Adsorptive losses of Gemex were 12%/km where didymo was mostly nonvisible and approximately 36%/km where substantial didymo mats were present. At 0.9 km downstream, Cu concentrations peaked at 12 mg/L, and didymo was <5% viable (down from 65-72%) for ≥21 d posttreatment. Viability data indicate that elimination of nonvisible infestations is possible and that suppression of early-stage infestations (≤40% cover, ≤4.5 mm thick) could be achieved after repeated applications. After a single Gemex application, no significant accumulation of Cu was noted in the sediments six weeks posttreatment, but Cu concentrations remained high in algal mats (109-367 mg/kg dry wt). Long-term effects on the nontarget algal, invertebrate, or fish communities were minimal, although significant localized trout mortalities, not predicted by prior laboratory exposures, occurred on the treatment day. Extended Gemex exposure in low-hardness waters might have caused the mortalities, although changes in chelated Cu speciation also possibly contributed. The present study integrates effects on resident biota with dosage data, including changes in pH, in a natural waterway.

Dual influences of ecosystem size and disturbance on food chain length in streams.

The number of trophic transfers occurring between basal resources and top predators, food chain length (FCL), varies widely in the world's ecosystems for reasons that are poorly understood, particularly for stream ecosystems. Available evidence indicates that FCL is set by energetic constraints, environmental stochasticity, or ecosystem size effects, although no single explanation has yet accounted for FCL patterns in a broad sense. Further, whether environmental disturbance can influence FCL has been debated on both theoretical and empirical grounds for quite some time. Using data from sixteen South Island, New Zealand streams, we determined whether the so-called ecosystem size, disturbance, or resource availability hypotheses could account for FCL variation in high country fluvial environments. Stable isotope-based estimates of maximum trophic position ranged from 2.6 to 4.2 and averaged 3.5, a value on par with the global FCL average for streams. Model-selection results indicated that stream size and disturbance regime best explained across-site patterns in FCL, although resource availability was negatively correlated with our measure of disturbance; FCL approached its maximum in large, stable springs and was <3.5 trophic levels in small, fishless and/or disturbed streams. Community data indicate that size influenced FCL, primarily through its influence on local fish species richness (i.e., via trophic level additions and/or insertions), whereas disturbance did so via an effect on the relative availability of intermediate predators (i.e., predatory invertebrates) as prey for fishes. Overall, our results demonstrate that disturbance can have an important food web-structuring role in stream ecosystems, and further imply that pluralistic explanations are needed to fully understand the range of structural variation observed for real food webs.

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).