Non-Native Fish Introductions and the Decline of the Mountain Yellow-Legged Frog from within Protected Areas

Abstract: One of the most puzzling aspects of the worldwide decline of amphibians is their disappearance from within protected areas. Because these areas are ostensibly undisturbed, habitat alterations are generally perceived as unlikely causes. The introduction of non-native fishes into protected areas, however, is a common practice throughout the world and may exert an important influence on amphibian distributions. We quantified the role of introduced fishes (several species of trout) in the decline of the mountain yellow-legged frog (   Rana muscosa ) in California's Sierra Nevada through surveys openface> 1700 sites in two adjacent and historically fishless protected areas that differed primarily in the distribution of introduced fish. Negative effects of fishes on the distribution of frogs were evident at three spatial scales. At the landscape scale, comparisons between the two protected areas indicated that fish distribution was strongly negatively correlated with the distribution of frogs. At the watershed scale, the percentage of total water-body surface area occupied by fishes was a highly significant predictor of the percentage of total water-body surface area occupied by frogs. At the scale of individual water bodies, frogs were three times more likely to be found and six times more abundant in fishless than in fish-containing waterbodies, after habitat effects were accounted for. The strong effect of introduced fishes on mountain yellow-legged frogs appears to result from the unique life history of this amphibian which frequently restricts larvae to deeper water bodies, the same habitats into which fishes have most frequently been introduced. Because fish populations in at least some Sierra Nevada lakes can be removed with minimal effort, our results suggest that the decline of the mountain yellow-legged frog might be relatively easy to reverse.     

Emerging infectious disease as a proximate cause of amphibian mass mortality

A newly discovered infectious disease of amphibians, chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis, is implicated in population declines and possible extinctions throughout the world. The purpose of our study was to examine the effects of B. dendrobatidis on the mountain yellow-legged frog (Rana muscosa) in the Sierra Nevada of California (USA). We (1) quantified the prevalence and incidence of B. dendrobatidis through repeat surveys of several hundred R. muscosa populations in the southern Sierra Nevada; (2) described the population-level effects of B. dendrobatidis on R. muscosa population abundance; and (3) compared the mortality rates of infected and uninfected R. muscosa individuals from pre- through post-metamorphosis using both laboratory and field experiments. Mouthpart inspections conducted in 144 and 132 R. muscosa populations in 2003 and 2004, respectively, indicated that 19% of R. muscosa populations in both years showed indications of chytridiomycosis. Sixteen percent of populations that were uninfected in 2003 became infected by 2004. Rana muscosa population sizes were reduced by an average of 88% following B. dendrobatidis outbreaks at six sites, but at seven B. dendrobatidis-negative sites, R. muscosa population sizes increased by an average of 45% over the same time period. In the laboratory, all infected R. muscosa developed fatal chytridiomycosis after metamorphosis, while all uninfected individuals remained healthy. In the field experiment in which R. muscosa tadpoles were caged at infected and uninfected sites, 96% of the individuals that metamorphosed at infected sites died vs. 5% at the uninfected sites. These studies indicate that chytridiomycosis causes high mortality in post-metamorphic R. muscosa, that this emerging disease is the proximate cause of numerous observed R. muscosa population declines, and that the disease threatens this species with extirpation at numerous sites in California's Sierra Nevada.     

Aquatic and terrestrial organic matter in the diet of stream consumers: implications for mercury bioaccumulation

The relative contribution of aquatic vs. terrestrial organic matter to the diet of consumers in fluvial environments and its effects on bioaccumulation of contaminants such as mercury (Hg) remain poorly understood. We used stable isotopes of carbon and nitrogen in a gradient approach (consumer isotope ratio vs. periphyton isotope ratio) across temperate streams that range in their pH to assess consumer reliance on aquatic (periphyton) vs. terrestrial (riparian vegetation) organic matter, and whether Hg concentrations in fish and their prey were related to these energy sources. Taxa varied in their use of the two sources, with grazing mayflies (Heptageniidae), predatory stoneflies (Perlidae), one species of water strider (Metrobates hesperius), and the fish blacknose dace (Rhinichthys atratulus) showing strong connections to aquatic sources, while Aquarius remigis water striders and brook trout (Salvelinus fontinalis) showed a weak link to in-stream production. The aquatic food source for consumers, periphyton, had higher Hg concentrations in low-pH waters, and pH was a much better predictor of Hg in predatory invertebrates that relied mainly on this food source vs. those that used terrestrial C. These findings suggest that stream biota relying mainly on dietary inputs from the riparian zone will be partially insulated from the effects of water chemistry on Hg availability. This has implications for the development of a whole-system understanding of nutrient and material cycling in streams, the choice of taxa in contaminant monitoring studies, and in understanding the fate of Hg in stream food webs.     

Restoring piscivorous fish populations in the Laurentian Great Lakes causes seabird dietary change

Ecosystem change often affects the structure of aquatic communities thereby regulating how much and by what pathways energy and critical nutrients flow through food webs. The availability of energy and essential nutrients to top predators such as seabirds that rely on resources near the water's surface will be affected by changes in pelagic prey abundance. Here, we present results from analysis of a 25-year data set documenting dietary change in a predatory seabird from the Laurentian Great Lakes. We reveal significant declines in trophic position and alterations in energy and nutrient flow over time. Temporal changes in seabird diet tracked decreases in pelagic prey fish abundance. As pelagic prey abundance declined, birds consumed less aquatic prey and more terrestrial food. This pattern was consistent across all five large lake ecosystems. Declines in prey fish abundance may have primarily been the result of predation by stocked piscivorous fishes, but other lake-specific factors were likely also important. Natural resource management activities can have unintended consequences for nontarget ecosystem components. Reductions in pelagic prey abundance have reduced the capacity of the Great Lakes to support the energetic requirements of surface-feeding seabirds. In an environment characterized by increasingly limited pelagic fish resources, they are being offered a Hobsonian choice: switch to less nutritious terrestrial prey or go hungry.     

Terrestrial, benthic, and pelagic resource use in lakes: results from a three-isotope Bayesian mixing model

Fluxes of organic matter across habitat boundaries are common in food webs. These fluxes may strongly influence community dynamics, depending on the extent to which they are used by consumers. Yet understanding of basal resource use by consumers is limited, because describing trophic pathways in complex food webs is difficult. We quantified resource use for zooplankton, zoobenthos, and fishes in four low-productivity lakes, using a Bayesian mixing model and measurements of hydrogen, carbon, and nitrogen stable isotope ratios. Multiple sources of uncertainty were explicitly incorporated into the model. As a result, posterior estimates of resource use were often broad distributions; nevertheless, clear patterns were evident. Zooplankton relied on terrestrial and pelagic primary production, while zoobenthos and fishes relied on terrestrial and benthic primary production. Across all consumer groups terrestrial reliance tended to be higher, and benthic reliance lower, in lakes where light penetration was low due to inputs of terrestrial dissolved organic carbon. These results support and refine an emerging consensus that terrestrial and benthic support of lake food webs can be substantial, and they imply that changes in the relative availability of basal resources drive the strength of cross-habitat trophic connections.     

Simulation of aquatic food web and species interactions by adaptive agents embodied with evolutionary computation: a conceptual framework

Individual-based and state variable-based adaptive agents (AA) are discussed regarding their relevance to different types of ecosystems. Individual-based AA proved applicable to a spatially explicit simulation of highly simplified terrestrial food webs. State variable-based AA with evolutionary computation (EC) embodied are suggested for the simulation of aquatic food webs and plankton species interactions. Embodiment of EC in AA can be achieved by evolving predictive rules (ER), differential equations (EDE) or artificial neural networks (ANN) derived from a diverse lake database. In order to provide ecosystem simulation with resilience to environmental change, agent banks can be created containing alternative agents for same species or functional groups from different lakes. State variable-based AA are currently tested for aquatic ecosytem simulation by means of a diverse lake database. It promises to overcome constraints by the rigidity of traditional lake ecosystem models.     

Benthic algal production across lake size gradients: interactions among morphometry, nutrients, and light

Attached algae play a minor role in conceptual and empirical models of lake ecosystem function but paradoxically form the energetic base of food webs that support a wide variety of fishes. To explore the apparent mismatch between perceived limits on contributions of periphyton to whole-lake primary production and its importance to consumers, we modeled the contribution of periphyton to whole-ecosystem primary production across lake size, shape, and nutrient gradients. The distribution of available benthic habitat for periphyton is influenced by the ratio of mean depth to maximum depth (DR = z/ z(max)). We modeled total phytoplankton production from water-column nutrient availability, z, and light. Periphyton production was a function of light-saturated photosynthesis (BPmax) and light availability at depth. The model demonstrated that depth ratio (DR) and light attenuation strongly determined the maximum possible contribution of benthic algae to lake production, and the benthic proportion of whole-lake primary production (BPf) declined with increasing nutrients. Shallow lakes (z < or =5 m) were insensitive to DR and were dominated by either benthic or pelagic primary productivity depending on trophic status. Moderately deep oligotrophic lakes had substantial contributions by benthic primary productivity at low depth ratios and when maximum benthic photosynthesis was moderate or high. Extremely large, deep lakes always had low fractional contributions of benthic primary production. An analysis of the world's largest lakes showed that the shapes of natural lakes shift increasingly toward lower depth ratios with increasing depth, maximizing the potential importance of littoral primary production in large-lake food webs. The repeatedly demonstrated importance of periphyton to lake food webs may reflect the combination of low depth ratios and high light penetration characteristic of large, oligotrophic lakes that in turn lead to substantial contributions of periphyton to autochthonous production.     

Multiple stressors and amphibian declines: dual impacts of pesticides and fish on yellow-legged frogs.

More than 40% of Earth's 5700+ amphibian species have undergone recent declines. Despite the likely involvement of multiple factors in driving these declines, most studies continue to focus on single stressors. In California (USA), separate studies have implicated either introduced fish or pesticides as causal agents. To date, however, no study has simultaneously evaluated the respective roles of these two potential stressors nor attempted to assess their relative importance, information critical for the development of effective conservation efforts and environmental policies. We examined the role and relative effect of fish and pesticides on the mountain yellow-legged frog (Rana muscosa) using unusually detailed data sets for a large portion of R. muscosa's historic range in California's Sierra Nevada. Habitat characteristics and presence/absence of R. muscosa and fish were quantified at each of 6831 sites during field surveys. Pesticide use upwind of each site was calculated from pesticide application records and predominant wind directions. Using generalized additive models, we found that, after accounting for habitat effects, the probability of R. muscosa presence was significantly reduced by both fish and pesticides, with the landscape-scale effect of pesticides much stronger than that of fish. The degree to which a site was sheltered from the predominant wind (and associated pesticides) was also a significant predictor of R. muscosa presence. Taken together, these results represent the strongest evidence to date that windborne pesticides are contributing to amphibian declines in pristine locations. Our results suggest that amphibian declines may have complex multi-factorial causes, and caution that single-factor studies that demonstrate the importance of one factor should not be used as evidence against the importance of other factors.     

Carryover aquatic effects on survival of metamorphic frogs during pond emigration.

In organisms with complex life cycles, physiological stressors during early life stages may have fitness-level impacts that are delayed into later stages or habitats. We tested the hypothesis that body size and date of metamorphosis, which are highly responsive to aquatic stressors, influence post-metamorphic survival and movement patterns in the terrestrial phase of an ephemeral pond-breeding frog by examining these traits in two populations of northern red-legged frogs (Rana aurora aurora). To increase variation of body size at metamorphosis, we manipulated food availability for 314 of 1045 uniquely marked tadpoles and estimated the probability that frogs survived and emigrated using concentric rings of drift fencing surrounding ponds and Bayesian capture-recapture modeling. The odds of surviving and emigrating from the ponds to the innermost drift fences, approximately 12 m, increased by factors of 2.20 (95% credibility intervals 1.39-4.23) and 2.54 (0.94-4.91) with each millimeter increase in snout-vent length and decreased by factors of 0.91 (0.85-0.96) and 0.89 (0.80-1.00) with each day's delay in metamorphosis for the two ponds. The odds of surviving and moving to the next ring of fencing, 12 m to approximately 40 m from the ponds, increased by a factor of 1.20 (0.45-4.06) with each millimeter increase in size. Our results demonstrated that body size and timing of metamorphosis relate strongly to the performance of newly metamorphosed frogs during their initial transition into terrestrial habitat. Carryover effects of aquatic stressors that reduce size and delay metamorphosis may have population-level impacts that are not expressed until terrestrial stages. Since changes in both aquatic and terrestrial systems are implicated in many amphibian declines, quantifying both immediate and delayed effects of stressors on demographic rates is critical to sound management.     

Measuring terrestrial subsidies to aquatic food webs using stable isotopes of hydrogen

Understanding river food webs requires distinguishing energy derived from primary production in the river itself (autochthonous) from that produced externally (allochthonous), yet there are no universally applicable and reliable techniques for doing so. We compared the natural abundance stable isotope ratios of hydrogen (deltaD) of allochthonous and autochthonous energy sources in four different aquatic ecosystems. We found that autochthonous organic matter is uniformly far more depleted in deuterium (lower deltaD values) than allochthonous: an average difference of approximately 100% per hundred. We also found that organisms at higher trophic levels, including both aquatic invertebrates and fish, have deltaD values intermediate between aquatic algae and terrestrial plants. The consistent differences between leaves and algae in deltaD among these four watersheds, along with the intermediate values in higher trophic levels, indicate that natural abundance hydrogen isotope signatures are a powerful tool for partitioning energy flow in aquatic ecosystems.     

Assessing Changes in Amphibian Population Dynamics Following Experimental Manipulations of Introduced Fish

Abstract: Sport‐fish introductions are now recognized as an important cause of amphibian decline, but few researchers have quantified the demographic responses of amphibians to current options in fisheries management designed to minimize effects on sensitive amphibians. Demographic analyses with mark–recapture data allow researchers to assess the relative importance of survival, local recruitment, and migration to changes in population densities. I conducted a 4‐year, replicated whole‐lake experiment in the Klamath Mountains of northern California (U.S.A.) to quantify changes in population density, survival, population growth rate, and recruitment of the Cascades frog (Rana cascadae) in response to manipulations of non‐native fish populations. I compared responses of the frogs in lakes where fish were removed, in lakes in their naturally fish‐free state, and in lakes where fish remained that were either stocked annually or no longer being stocked. Within 3 years of fish removals from 3 lakes, frog densities increased by a factor of 13.6. The survival of young adult frogs increased from 59% to 94%, and realized population growth and recruitment rates at the fish‐removal lakes were more than twice as high as the rates for fish‐free reference lakes and lakes that contained fish. Population growth in the fish‐removal lakes was likely due to better on‐site recruitment of frogs to later life stages rather than increased immigration. The effects on R. cascadae of suspending stocking were ambiguous and suggested no direct benefit to amphibians. With amphibians declining worldwide, these results show that active restoration can slow or reverse the decline of species affected by fish stocking within a short time frame.     

Effects of Introduced Salmonids on a Montane Population of Iberian Frogs

Abstract:  Amphibians are declining worldwide because of multiple factors, including human-mediated introduction of fishes into naturally fishless areas. Although several studies have focused on the effect of exotic fishes on native amphibians breeding in ponds or lakes, little is known about their effects on stream-breeding species. We studied the effects of introductions of native brown trout ( Salmo trutta ) and exotic brook trout ( Salvelinus fontinalis ) on the stream-breeding, endemic Iberian frog (  Rana iberica ) in a protected area in central Spain. We assessed occurrence patterns of tadpoles and salmonids and compared habitat use of the three species. We also determined experimentally whether chemical cues from salmonids elicited antipredator behavior in tadpoles. Finally, we assessed the relative influence of tadpole habitat preferences, differences in salmonid species, and invasion geography on tadpole occurrence. Despite widely overlapping habitat preferences, tadpoles and trout did not coexist, with the former restricted to fishless habitats. Tadpoles detected chemical cues from both trout species and reacted by decreasing their activity, although the response toward the native brown trout was stronger. The residual distribution of Iberian frogs in Peñalara is better explained by the geography of fish invasions than by the fish species involved. Measures such as fish extirpation from certain areas, aimed at recovering lost habitat and improving connectivity among remaining populations of Iberian frogs, seem critical for the species' long-term survival in central Spain.     

Evidence supporting the importance of terrestrial carbon in a large-river food web

Algal carbon has been increasingly recognized as the primary carbon source supporting large-river food webs; however, many of the studies that support this contention have focused on lotic main channels during low-flow periods. The flow variability and habitat-heterogeneity characteristic of these systems has the potential to significantly influence food web structure and must be integrated into models of large-river webs. We used stable-isotope analysis and IsoSource software to model terrestrial and algal sources of organic carbon supporting consumer taxa in the main channel and oxbow lakes of the Brazos River, Texas, USA, during a period of frequent hydrologic connectivity between these habitat types. Standardized sampling was conducted monthly to collect production sources and consumer species used in isotopic analysis. Predictability of hydrologic connections between habitat types was based on the previous 30 years of flow data. IsoSource mixing models identified terrestrial C3 macrophytes (riparian origin) as the primary carbon source supporting virtually all consumers in the main channel and most consumers in oxbow lakes. Small-bodied consumers (<100 mm) in oxbow lakes assimilated large fractions of algal carbon whereas this pattern was not apparent in the main channel. Estimates of detritivore trophic positions based on delta15N values indicated that terrestrial material was likely assimilated via invertebrates rather than directly from detritus. High flows in the river channel influenced algal standing stock, and differences in the importance of terrestrial and algal production sources among consumers in channel vs. oxbow habitats were associated with patterns of flooding. The importance of terrestrial material contradicts the findings of recent studies of large-river food webs that have emphasized the importance of algal carbon and indicates that there can be significant spatial, temporal, and taxonomic variation in carbon sources supporting consumers in large rivers.     

An unusual trophic subsidy and species dominance in a tropical stream

In classical theory, species are assumed to achieve dominance through competitive exclusion, but if food resources are limiting, cross-habitat trophic subsidies could also underpin dominance. The impact of dominant species on community dynamics may depend on the energy base of population size. We report on an unusual, spatially subsidized population of a tropical, stream-dwelling crab that dominates the benthic fauna of a Kenyan stream. Diet and stable isotope analyses indicated that this crab is a true omnivore, with terrestrial subsidies dominating both plant and animal resources. Unusually, the animal prey included almost no aquatic invertebrates. Instead, a single species of ant constituted approximately 35% of the annual diet (stomach contents analysis) and up to 90% of assimilated nitrogen (estimates from stable isotope analysis). Ants may be pivotal to enabling crab dominance, and this crab may be largely disconnected from the local trophic network for its dietary needs. The paucity of other invertebrates in the stream community suggests that this super-dominant crab is a strong interactor that suppresses aquatic invertebrate populations. Common stabilizing attributes of spatially subsidized food webs (e.g., asynchronous prey availability, wide feeding niche, consumer migration) were absent from this system, and although apparently stable, it may be vulnerable to disturbance in the donor habitat.     

太湖鼋头渚地区鹭类觅食生境研究

连续 2a实地调查了太湖鼋头渚地区鹭鸟的觅食生境 ,并搜集其雏鸟的反吐物 ,初步掌握了该地区不同鹭鸟对觅食生境的选择与食物组成情况。结果表明 ,无锡鼋头渚地区白鹭与夜鹭主要在鱼塘、湖泊觅食 ,白鹭在湖滨觅食 ,而夜鹭还会到湖中央水面觅食 ;池鹭觅食区域较广 ,主要在鱼塘。鹭群主要的食物类型是鱼类 ,白鹭食物几乎全为鱼类 ,夜鹭 95 %的食物为鱼类 ,兼有少量的蛙类、甲壳类和小型哺乳类动物。     

Lake to land subsidies: experimental addition of aquatic insects increases terrestrial arthropod densities

Aquatic insects are a common and important subsidy to terrestrial systems, yet little is known about how these inputs affect terrestrial food webs, especially around lakes. Myvatn, a lake in northern Iceland, has extraordinary midge (Chironomidae) emergences that result in large inputs of biomass and nutrients to terrestrial arthropod communities. We simulated this lake-to-land resource pulse by collecting midges from Myvatn and spreading their dried carcasses on 1-m2 plots at a nearby site that receives very little midge deposition. We hypothesized a positive bottom-up response of detritivores that would be transmitted to their predators and would persist into the following year. We sampled the arthropod community once per month for two consecutive summers. Midge addition resulted in significantly different arthropod communities and increased densities of some taxa in both years. Detritivores, specifically Diptera larvae, Collembola, and Acari increased in midge-addition plots, and so did some predators and parasitoids. Arthropod densities were still elevated a year after midge addition, and two years of midge addition further increased the density of higher-order consumers (e.g., Coleoptera and Hymenoptera). Midge addition increased arthropod biomass by 68% after one year and 108% after two years. By manipulating the nutrient pulse delivered by midges we were able to elucidate food web consequences of midge deposition and spatial and temporal dynamics that are difficult to determine based on comparative approaches alone. Resources cross ecosystem boundaries and are assimilated over time because of life-history strategies that connect aquatic and terrestrial food webs and these systems cannot be fully understood in isolation from each other.     

Factors Mediating Co‐Occurrence of an Economically Valuable Introduced Fish and Its Native Frog Prey

Habitat characteristics mediate predator–prey coexistence in many ecological systems but are seldom considered in species introductions. When economically important introduced predators are stocked despite known negative impacts on native species, understanding the role of refuges, landscape configurations, and community interactions can inform habitat management plans. We measured these factors in basins with introduced trout (Salmonidae) and the Cascades frog (Rana cascadae) to determine, which are responsible for observed patterns of co‐occurrence of this economically important predator and its native prey. Large, vegetated shallows were strongly correlated to co‐occurrence, and R. cascadae larvae occur in shallower water when fish are present, presumably to escape predation. The number of nearby breeding sites of R. cascadae was also correlated to co‐occurrence, but only when the western toad (Anaxyrus boreas) was present. Because A. boreas larvae are unpalatable to fish and resemble R. cascadae, they may provide protection from trout via Batesian mimicry. Although rescue‐effect dispersal from nearby populations may maintain co‐occurrence, within‐lake factors proved more important for predicting co‐occurrence. Learning which factors allow co‐occurrence between economically important introduced species and their native prey enables managers to make better‐informed stocking decisions. Factores que Median la Co‐Ocurrencia de un Pez Introducido con Valor Económico y su Presa, una Rana Nativa     

Comparing resource pulses in aquatic and terrestrial ecosystems

Resource pulses affect productivity and dynamics in a diversity of ecosystems, including islands, forests, streams, and lakes. Terrestrial and aquatic systems differ in food web structure and biogeochemistry; thus they may also differ in their responses to resource pulses. However, there has been a limited attempt to compare responses across ecosystem types. Here, we identify similarities and differences in the causes and consequences of resource pulses in terrestrial and aquatic systems. We propose that different patterns of food web and ecosystem structure in terrestrial and aquatic systems lead to different responses to resource pulses. Two predictions emerge from a comparison of resource pulses in the literature: (1) the bottom-up effects of resource pulses should transmit through aquatic food webs faster because of differences in the growth rates, life history, and stoichiometry of organisms in aquatic vs. terrestrial systems, and (2) the impacts of resource pulses should also persist longer in terrestrial systems because of longer generation times, the long-lived nature of many terrestrial resource pulses, and reduced top-down effects of consumers in terrestrial systems compared to aquatic systems. To examine these predictions, we use a case study of a resource pulse that affects both terrestrial and aquatic systems: the synchronous emergence of periodical cicadas (Magicicada spp.) in eastern North American forests. In general, studies that have examined the effects of periodical cicadas on terrestrial and aquatic systems support the prediction that resource pulses transmit more rapidly in aquatic systems; however, support for the prediction that resource pulse effects persist longer in terrestrial systems is equivocal. We conclude that there is a need to elucidate the indirect effects and long-term implications of resource pulses in both terrestrial and aquatic ecosystems.     

Allee effect limits colonization success of sexually reproducing zooplankton

Understanding the dynamics of populations at low density and the role of Allee effects is a priority due to concern about the decline of rare species and interest in colonization/invasion dynamics. Despite well-developed theory and observational support, experimental examinations of the Allee effect in natural systems are rare, partly because of logistical difficulties associated with experiments at low population density. We took advantage of fish introduction and removal in alpine lakes to experimentally test for the Allee effect at the whole-ecosystem scale. The large copepod Hesperodiaptomus shoshone is often extirpated from the water column by fish and sometimes fails to recover following fish disappearance, despite the presence of a long-lived egg bank. Population growth rate of this dioecious species may be limited by mate encounter rate, such that below some critical density a colonizing population will fail to establish. We conducted a multi-lake experiment in which H. shoshone was stocked at densities that bracketed our hypothesized critical density of 0.5-5 copoepods/m3. Successful recovery by the copepod was observed only in the lake with the highest initial density (3 copepods/m3). Copepods stocked into small cages at 3000 copepods/m3 survived and reproduced at rates comparable to natural populations, confirming that the lakes were suitable habitat for this species. In support of mate limitation as the mechanism underlying recovery failure, we found a significant positive relationship between mating success and density across experimental and natural H. shoshone populations. Furthermore, a mesocosm experiment provided evidence of increased per capita population growth rate with increasing population density in another diaptomid species, Skistodiaptomus pallidus. Together, these lines of evidence support the importance of the Allee effect to population recovery of H. shoshone in the Sierra Nevada, and to diaptomid copepods in general.     

Aufnahmepfade von POPs in Biota

POPs (persistent organic pollutants) associated with aquatic sediments can pose a risk to aquatic food chains, since they can be re-introduced to the food web. One major pathway is the bioaccumulation of POPs by endobenthic, sedimentingesting invertebrates (especially tubificid oligochaetes). These worms serve as food for benthivorous fish, which thereby ingest the sediment-borne chemicals and may accumulate contaminant concentrations far higher than from water exposure alone, and consequently transfer them to organisms of higher trophic levels. In order to evaluate such a potential biomagnification, a laboratory test was developed. It consisted of a two-step food chain including the sediment dwelling freshwater oligochaete Tubifex tubifex (Müller) and the three-spined stickleback (Gasterosteus aculeatus, Linné), a small teleost fish which often feeds primarily on benthic invertebrates. Artificial sediment and reconstituted water were used. To examine the influence of benthic prey on the bioaccumulation of a POP in the predator, fish were exposed to ¹⁴C-labelled hexachlorobenzene via spiked water, spiked sediment, pre-contaminated prey organisms, and to combinations of these exposure routes. Summarising the results of these experiments, it could be shown that the exposure to HCB via different routes resulted in a significantly higher accumulation in fish than an exposure to single pathways. It was concluded that the major uptake routes for fish were the overlying water and the food, whereas the contribution of spiked sediment itself was relatively small. HCB was biomagnified in the rested laboratory food chain. Therefore, concerning secondary poisoning, the environmental risk assessment of POPs like HCB should not be based on existing bioaccumulation tests alone, since they focus only on exposure via the water pathway. Instead, the influence of food and sediment as exposure routes should be considered as well, using comprehensive food chain modelling and/or laboratory studies.