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.
Summary Amphidromy is a widespread migratory behavioural syndrome exhibited by fish (and some aquatic invertebrates) that spawn in fresh water and whose larvae migrate to pelagic marine (or lentic) habitats for a period of early growth, followed by a return migration to adult freshwater habitats. The fitness advantage of amphidromy has been the subject of prolonged debate, and we examined the hypothesis that amphidromy mainly increases fecundity through the production of small pelagic larvae. We compared egg size (a proxy for larval size) of closely related non‐migratory and amphidromous fish species in the families Cottidae, Galaxiidae, Eleotridae and Gobiidae. To examine how egg size changes in relation to body size within a taxonomic group, we also compared egg size and maximum body size across most species of New Zealand non‐migratory and amphidromous galaxiids. Non‐migratory species generally have relatively larger eggs than their amphidromous confamilial species. This particular trait has evolved independently several times in each of the four families of amphidromous fish that have given rise to significant freshwater radiations. Amongst the New Zealand galaxiids, mean egg diameter increased with maximum body length for both non‐migratory and amphidromous species; however, despite the considerably smaller relative body size of the non‐diadromous species, the rate of increase in egg diameter relative to the increase in body size is considerably higher in the non‐migratory fish. We propose that amphidromous fish maintain a high level of fecundity by producing small pelagic larvae. In contrast, the relatively large eggs and well‐developed larvae of non‐migratory species increase larval survival in what are often relatively harsh and unproductive freshwater habitats. Consequently, amphidromous species are likely to have a competitive advantage over their non‐migratory relatives when close to a pelagic habitat in which their larvae can grow and develop and then migrate upstream, releasing them from recruitment limitation and giving them a local reproductive advantage over their less fecund non‐migratory relatives. We argue that the persistence and distribution of both life‐history strategies across the landscape depends on the relative difference in the net reproductive return for each strategy in relation to distance from a pelagic larval habitat, as mediated by the relative costs of migration and egg size/fecundity relationships.
Tree-holes provide an important microhabitat that is used for feeding, roosting and breeding by numerous species around the world.Yet despite their ecological importance for many of New Zealand's endangered species, few studies have investigated the abundance or distribution of tree-holes in native forests. We used complementary ground and climbed tree surveys to determine the abundance, distribution and characteristics of tree-holes in undisturbed Nothofagus forest in the Lewis Pass, New Zealand. We found that hole-bearing trees were surprisingly abundant compared with many other studies, including Australian Eucalyptus species and American beech. In fact, we estimated as many as 3906 tree-holes per hectare, of which 963 holes per hectare were potentially large enough to provide roost sites for hole-nesting bats in New Zealand, while only eight holes per hectare were potentially suitable for specialist hole-nesting birds. This was of great interest as primary cavity-excavating animals are absent from New Zealand forests, compared with North America and Australia. Moreover, tree-hole formation in New Zealand is likely to be dominated by abiotic processes, such as branch breakage from windstorms and snow damage. As has been found in many other studies, tree-holes were not uniformly distributed throughout the forest. Tree-holes were significantly more abundant on the least abundant tree species, Nothofagus fusca, than on either N. menziesii or N. solandri. In addition to tree species, tree size was also an important factor influencing the structural characteristics of tree-holes and their abundance in this forest. Moreover, these trends were not fully evident without climbed tree surveys. Our results revealed that ground-based surveys consistently underestimated the number of tree-holes present on Nothofagus trees, and illustrate the importance of using climbed inspections where possible in tree-hole surveys. We compare our results with other studies overseas and discuss how these are linked to the biotic and abiotic processes involved in tree-hole formation. We consider the potential implications of our findings for New Zealand's hole-dwelling fauna and how stand dynamics and past and future forest management practices will influence the structural characteristics of tree-holes and their abundance in remnant forest throughout New Zealand.
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 ∼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.
1. Flood-related disturbances are predicted to be seriously altered by climate change effects, and this will have strong implications for stream communities. Predicting how and why community structure responds to changes in disturbance regimes will require measures of disturbance that are closely linked to community variability. A range of disturbance measures have been tested for their ability to explain patterns in stream periphyton and invertebrate assemblages, but assessments of fish have largely focussed on flow measures as predictors. Consequently, the mechanisms driving fish assemblage responses to disturbance are poorly known. 2. Stream disturbance was quantified using seven measures, three based on disturbance of the streambed (assessed using painted tracer particles), three associated with variation in stream discharge and one from a subjective measure of stream channel stability. Twenty streams were sampled on five occasions to evaluate which disturbance measure explained the most variation in fish assemblages and also to quantify the influence of disturbance on spatial and temporal assemblage variability. To determine whether disturbance affected fish directly or via its effect on benthic food supply (i.e. stream invertebrates), a one-off survey of 52 streams was conducted. 3. Path analysis on data from the 20 streams indicated that fish biomass responded most strongly to flow-mediated bed movement, not flow variability. Consequently, bed-movement measures were better predictors of spatial and temporal variability in fish biomass and assemblage structure than purely hydrological measures. Furthermore, variations in fish biomass and assemblage structure were higher in more disturbed streams. A second path analysis conducted on data from the 52 stream survey compared the effects of physical habitat and food-related components of bed disturbance on fish biomass. It indicated that both had a similar influence on disturbance-related reductions in fish biomass. 4. These results indicate that variation in fish assemblages can best be understood by measuring or predicting bed movement, because bed disturbance is likely to encompass more effectively major factors that influence stream fish, such as habitat availability and food supply. A shift in disturbance regime is likely to change the biomass of fish a habitat can support and fish assemblage structure.
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