Eradication of Nonnative Fish by Gill Netting from a Small Mountain Lake in California

Nearly all mountain lakes in the western United States were historically fishless, but most now contain introduced trout populations. As a result of the impacts of these introductions on ecosystem structure and function, there is increasing interest in restoring some lakes to a fishless condition. To date, however, the only effective method of fish eradication is the application of rotenone, a pesticide that is also toxic to nontarget native species. The objective of this study was to assess the effectiveness of intensive gill netting in eradicating the trout population from a small subalpine lake in the Sierra Nevada, California. We removed the resident trout population and a second trout population accidentally stocked into the study lake within 18 and 15 gill net sets, respectively. Adult trout were highly vulnerable to gill nets, but younger fish were not readily captured until they reached approximately 110 mm. To determine the utility of gill netting as a fish eradication technique in other Sierra Nevada lakes, we used morphometry data from 330 Sierra Nevada lakes to determine what proportion had characteristics similar to the study lake (i.e., small, isolated lakes with little spawning habitat). We estimated that gill netting would be a viable eradication method in 15–20% of the high mountain lakes in the Sierra Nevada. We conclude that although gill netting is likely to be more expensive and time consuming than rotenone application, it is a viable alternative under some conditions and should be the method of choice when sensitive native species are present.     

Invasive trout increase the climatic sensitivity of zooplankton communities in naturally fishless lakes

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Rapid evolution in response to introduced predators II: the contribution of adaptive plasticity

Background  

Introductions of non-native species can significantly alter the selective environment for populations of native species, which can respond through phenotypic plasticity or genetic adaptation. We examined phenotypic and genetic responses of Daphnia populations to recent introductions of non-native fish to assess the relative roles of phenotypic plasticity versus genetic change in causing the observed patterns. The Daphnia community in alpine lakes throughout the Sierra Nevada of California (USA) is ideally suited for investigation of rapid adaptive evolution because there are multiple lakes with and without introduced fish predators. We conducted common-garden experiments involving presence or absence of chemical cues produced by fish and measured morphological and life-history traits in Daphnia melanica populations collected from lakes with contrasting fish stocking histories. The experiment allowed us to assess the degree of population differentiation due to fish predation and examine the contribution of adaptive plasticity in the response to predator introduction.     

Stocked trout have minimal effects on littoral invertebrate assemblages of productive fish‐bearing lakes: a whole‐lake BACI study

1. Rainbow Trout (Oncorhynchus mykiss [Walbaum]) is commonly stocked as a sport fish throughout the world but can have serious negative effects on native species, especially in headwater systems. Productive fish‐bearing lakes represent a frequently stocked yet infrequently studied system, and effects of trout in these systems may differ from those in headwater lakes. 2. We used a Before‐After Control‐Impact (BACI) design to determine how stocked trout affected assemblage‐level and taxon‐level biomass, abundance and average length of littoral invertebrates in a stocked lake relative to three unstocked control lakes in the boreal foothills of Alberta, Canada. Lakes were studied 1 year before and for 2 years after stocking. Because characteristics of productive fish‐bearing lakes should buffer impacts of introduced fish, we predicted that trout would not affect assemblage‐level structure of littoral invertebrates but might reduce the abundance or average length of large‐bodied taxa frequently consumed by trout. 3. Relative to the unstocked control lakes, biomass, but not abundance, of the littoral invertebrate assemblage was affected indirectly by trout through increases of some taxa after trout stocking. At the individual taxon‐level, trout stocking did not affect most (23 of the 27) taxa, with four taxa increasing in abundance or biomass after stocking. Only one taxon, Chironomidae, showed evidence of size‐selective predation by trout, being consumed frequently by trout and decreasing significantly in average length after stocking. 4. Our results contrast with the strong negative effects of trout stocking on invertebrate assemblages commonly reported from headwater lakes. A combination of factors, including large and robust native populations of forage fish, the generalised diet of trout, overwinter aeration, relatively high productivity and dense macrophyte beds, likely works in concert to reduce potentially negative effects of stocked trout in these systems. As such, productive, fish‐bearing lakes may represent a suitable system for trout stocking, especially where native sport fish populations are lacking.     

Changes in aquatic insect emergence in response to whole-lake experimental manipulations of introduced trout

1. Insects emerging from mountain lakes provide an important food source for many terrestrial predators. The amount of insect subsidy that emerges from lakes is influenced by predator composition, but predator effects could be ameliorated by greater habitat complexity. We conducted a replicated whole‐lake experiment to test the effects of introduced fish predators on the abundance and composition of aquatic insects within and emerging from the littoral zone of 16 mountain lakes in the Trinity Alps Wilderness in northwestern California. 2. Study treatments matched the fisheries management options being implemented in California’s wilderness areas: (i) continued stocking with non‐native trout, (ii) suspension of stocking, and (iii) suspension of stocking and removal of fish. We also included four naturally fishless ‘reference’ lakes. We compared abundances and biomass of emerging aquatic insects before treatments were initiated and for 3 years following their establishment. Abundances of benthic insects were also compared in the third year post‐treatment. 3. Trout removal rapidly increased abundances of mayflies, caddisflies, and insect predators, and overall insect biomass emerging from lakes. No significant differences were found between the suspension of stocking lakes and continued stocking lakes. Fish density was a more important predictor of aquatic insect emergence than habitat complexity. 4. Mayfly larvae responded positively to fish removal and caddisfly larvae tended to be more abundant in lakes without fish, but we did not detect effects on abundance of predatory insects. However, we found large insect predators in shallower water in lakes with fish compared to fish removal or fish‐free reference lakes. 5. These results provide insights into the continuing effects of past and current fish stocking practices on the flow of insect prey from mountain lakes into the neighbouring terrestrial environment. We also show that these consequences can rapidly be reversed by removing non‐native fishes.     

Geography of Invasion in Mountain Streams: Consequences of Headwater Lake Fish Introductions

The introduction of fish into high-elevation lakes can provide a geographic and demographic boost to their invasion of stream networks, thereby further endangering the native stream fauna. Increasingly, remaining populations of native salmonids are concentrated in fragmented headwater refugia that are protected by physical or biological barriers from introduced fishes that originate in the pervasive source populations established at lower elevations. Although fish introduced near mainstem rivers frequently encounter obstacles to upstream dispersal, such as steep slopes or falls, we found that brook trout (Salvelinus fontinalis) dispersed downstream through channel slopes of 80% and 18-m-high falls. Thus, headwater lake stocking provides source populations that may be capable of invading most downstream habitats, including headwater refugia of native fishes. The extent of additional area invasible from lakes, beyond that invasible from downstream, depends on the geography of the stream network, particularly the density and distribution of headwater lakes and their location relative to barriers inhibiting upstream dispersal. In the thermal and trophic environments downstream of lakes, fish commonly grow faster and thus mature earlier and have higher fecundity-at-age than their counterparts in other high-elevation streams. The resulting higher rates of population growth facilitate invasion. Larger body sizes also potentially aid the fish in overcoming barriers to invasion. Trout introductions to high-elevation headwater lakes thus pose disproportionately large risks to native fishes—even when the place of introduction may appear to be spatially dissociated from populations of the native species. Mapping the potential invasible area can help to establish priorities in stocking and eradication efforts. Received 28 March 2000; accepted 9 February 2001.     

Fish introductions reveal the temperature dependence of species interactions

A major area of current research is to understand how climate change will impact species interactions and ultimately biodiversity. A variety of environmental conditions are rapidly changing owing to climate warming, and these conditions often affect both the strength and outcome of species interactions. We used fish distributions and replicated fish introductions to investigate environmental conditions influencing the coexistence of two fishes in Swedish lakes: brown trout (Salmo trutta) and pike (Esox lucius). A logistic regression model of brown trout and pike coexistence showed that these species coexist in large lakes (more than 4.5 km²), but not in small, warm lakes (annual air temperature more than 0.9–1.5°C). We then explored how climate change will alter coexistence by substituting climate scenarios for 2091–2100 into our model. The model predicts that brown trout will be extirpated from approximately half of the lakes where they presently coexist with pike and from nearly all 9100 lakes where pike are predicted to invade. Context dependency was critical for understanding pike–brown trout interactions, and, given the widespread occurrence of context-dependent species interactions, this aspect will probably be critical for accurately predicting climate impacts on biodiversity.     

Invasive aquarium fish transform ecosystem nutrient dynamics

Trade of ornamental aquatic species is a multi-billion dollar industry responsible for the introduction of myriad fishes into novel ecosystems. Although aquarium invaders have the potential to alter ecosystem function, regulation of the trade is minimal and little is known about the ecosystem-level consequences of invasion for all but a small number of aquarium species. Here, we demonstrate how ecological stoichiometry can be used as a framework to identify aquarium invaders with the potential to modify ecosystem processes. We show that explosive growth of an introduced population of stoichiometrically unique, phosphorus (P)-rich catfish in a river in southern Mexico significantly transformed stream nutrient dynamics by altering nutrient storage and remineralization rates. Notably, changes varied between elements; the P-rich fish acted as net sinks of P and net remineralizers of nitrogen. Results from this study suggest species-specific stoichiometry may be insightful for understanding how invasive species modify nutrient dynamics when their population densities and elemental composition differ substantially from native organisms. Risk analysis for potential aquarium imports should consider species traits such as body stoichiometry, which may increase the likelihood that an invasion will alter the structure and function of ecosystems.     

Changes in the Spatial Distribution of Fishes in Lakes Along a Residential Development Gradient

As the human demand for freshwater natural resources such as fish and drinking water increases, we may rely more heavily on models to predict the response of aquatic ecosystems to natural and anthropogenic disturbance. Theses models in turn implicitly depend on the underlying spatial distribution of organisms. In terrestrial ecosystems, increased natural resource utilization has transformed habitat and changed the spatial distribution of organisms, with subsequent negative effects on biota. Recent studies in lakes demonstrate that human development of lakeshores alters the physical habitat and nutrient cycles. The impact of such disturbance by humans on the spatial distribution of aquatic organisms, however, remains unknown. Here we quantify the effect of lakeshore development on the spatial distribution of fishes in 23 lakes in the US Pacific Northwest. We found a significant decrease in the spatial aggregation of fishes with increased shoreline development by humans, reflecting a loss of refugia and resource heterogeneity that favors aggregation among fishes. We also found that lakes with a high perimeter–surface-area ratio and a relatively shallow littoral zone had much higher levels of fish aggregation, suggesting the importance of terrestrial inputs to lakes. Finally, we found a marginally significant decrease in fish spatial aggregation with increased total phosphorus concentration, but no effect of chlorophyll concentration, water transparency, the predator–prey ratio, or number of species on fish spatial distributions. These results suggest that anthropogenic modification of shorelines is significantly altering the spatial distribution of important aquatic organisms, and that these changes may have important implications for predictive modeling of ecosystem dynamics.     

Interdecadal declines in flood frequency increase primary production in lakes of a northern river delta

Human activities and climate change have greatly altered flooding regimes in many of the world's river deltas, but the impact of such changes remains poorly quantified on decadal to multidecadal timescales. This study identified the response of delta lake primary production (measured as the concentration of sedimentary pigments) to variations in flood frequency using spatial surveys and paleolimnological analyses of lakes in the Peace‐Athabasca Delta (PAD), Canada. Surveys of 61 lakes spanning a range of hydrological conditions showed that those lakes that received flood waters less frequently were associated with elevated algal production (surface sedimentary pigments) and, in some lakes, increased growth of emergent macrophytes and epiphytic diatoms. Paleolimnological analyses of five lakes corroborated the contemporary spatial survey results by showing that production of pigments from most algal groups increased during recent periods of lower flood frequency in the 20th century as determined from increases in cellulose‐inferred lake‐water oxygen isotope composition and plant macrofossils, but remained stable in a ‘reference’ basin. In general, past periods of elevated algal production coincided with the increased abundance of submerged macrophytes or emergent vegetation that provide habitat for attached algae. These results suggest that interdecadal declines in river discharge arising from increased aridity, hydrologic regulation or consumptive water use will cause long‐term increases in primary production and alter ecosystem processes (carbon sequestration, biological diversity) in aquatic delta ecosystems similar to the PAD where lakes become nutrient‐rich in the absence of flooding.     

Genetic structure of brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis> L.) from the Hardangervidda mountain plateau (Norway) analyzed by microsatellite DNA: a basis for conservation guidelines

The Hardangervidda in southern Norway, the largest mountain plateau in Europe, has thousands of lakes and streams, mainly between 1000 and 1300 m above sea level, where brown trout is the only fish species. To describe the current genetic diversity of brown trout in this area, a total of 863 fish from 20 lakes were genotyped with eleven microsatellites. Most diversity is within lake populations, but diversity among geographical groups and populations within groups was significant, too. Neighbor-joining, principle coordinate analysis and Bayesian clustering show three major geographic groups in accordance with the river systems. Bias was caused by recent stocking in two lakes. Low/no genetic differentiation among some populations indicates that intermixing is common when lakes are well-connected, as was also shown by assignment test. We recommend preserving the genetic diversity of brown trout in this unique area by managing stocking in lake systems according to genetic structure.     

Effects of stocking-up freshwater food webs

The establishment of exotic game fishes to enhance recreational fisheries through authorized and unauthorized stocking into freshwater systems is a global phenomenon. Stocked fishes are often top predators that either replace native top predators or increase the species richness of top predators. Many direct effects of stocking have been documented, but the ecosystem consequences are seldom quantified. New studies increasingly document how species and community shifts influence ecosystem processes. We discuss here how predator stocking might increase top-down effects, alter nutrient cycles and decrease links between aquatic and surrounding terrestrial ecosystems. As fisheries management moves beyond species-specific utilitarian objectives to incorporate ecosystem and conservation goals, ecologists must address how common management practices alter food-web structure and subsequent ecosystem-level effects.     

Biotic feedbacks in Lake phosphorus cycles

Limnologists are now reconsidering the role of the biota in the phosphorus (P) cycles of lakes. Changes in lake communities can have significant consequences for ecosystem P cycles. At seasonal timescales, the relative importance of nitrogen (N) and Pas limiting factors for primary production depends in part on zooplankton species composition. Phosphorus storage and recycling by fish and zooplankton can be large components of P budgets, and mobile consumers can be important vectors in P transport. Stability, resilience and resistance of lake P cycles may depend heavily on fluxes to and from upper trophic levels.     

Patterns of Fish Growth along a Residential Development Gradient in North Temperate Lakes

Residential development of lakeshores is expected to change a variety of key lake features that include increased nutrient loading, increased invasion rate of nonnative species, increased exploitation rates of fishes by anglers, and alteration of littoral habitats. All of these factors may alter the capacity of lakes to support productive native fish populations. Fourteen north temperate lakes were surveyed to examine how growth rates of two common fish species (bluegill sunfish, Lepomis macrochirus; largemouth bass, Micropterus salmoides) varied along a residential development gradient. Size-specific growth rates for both species were negatively correlated with the degree of lakeshore residential development, although this trend was not statistically significant for largemouth bass. On average, annual growth rates for bluegill sunfish were 2.6 times lower in heavily developed lakes than in undeveloped lakes. This effect of lakeshore development on fish growth was not size specific for bluegills between 60 and 140 mm in total length. An index of population production rate that accounted for both the size-specific growth rate and the size distribution of fishes showed that bluegill populations were approximately 2.3 times less productive in highly developed lakes than in undeveloped lakes. Our results suggest that extensive residential development of lakeshores may reduce the fish production capacity of aquatic ecosystems. Received 29 April 1999; Accepted 26 October 1999.     

Iron Constraints on Planktonic Primary Production in Oligotrophic Lakes

Phototrophic primary production is a fundamental ecosystem process, and it is ultimately constrained by access to limiting nutrients. Whereas most research on nutrient limitation of lacustrine phytoplankton has focused on phosphorus (P) and nitrogen (N) limitation, there is growing evidence that iron (Fe) limitation may be more common than previously acknowledged. Here we show that P was the nutrient that stimulated phytoplankton primary production most strongly in seven out of nine bioassay experiments with natural lake water from oligotrophic clearwater lakes. However, Fe put constraints on phytoplankton production in eight lakes. In one of these lakes, Fe was the nutrient that stimulated primary production most, and concurrent P and Fe limitation was observed in seven lakes. The effect of Fe addition increased with decreasing lake water concentrations of total phosphorus and dissolved organic matter. Possible mechanisms are low import rates and low bioavailability of Fe in the absence of organic chelators. The experimental results were used to predict the relative strength of Fe, N, and P limitation in 659 oligotrophic clearwater lakes (with total phosphorus ≤ 0.2 μM P and total organic carbon < 6 mg C l⁻¹) from a national lake survey. Fe was predicted to have a positive effect in 88% of these lakes, and to be the nutrient with the strongest effect in 30% of the lakes. In conclusion, Fe, along with P and N, is an important factor constraining primary production in oligotrophic clearwater lakes, which is a common lake-type throughout the northern biomes. This paper is dedicated to the memory of Prof. Peter Blomqvist (deceased 2004).     

Wilderness Fish Stocking: History and Perspective

The stocking of trout in wilderness lakes of the western United States began in the 1800s. This practice was followed for nearly a century with the singular goal of creating and enhancing sport fishing and without any consideration of its ecological ramifications. Following the advent of a new environmental awareness in the 1960s, and thanks to new research that revealed negative impacts on the biota attributable to introduced fishes, traditional fish-stocking practices came under question first at federal land management agencies and later at their counterparts within the states. The highly utilitarian ethic that drove resource management until well into the 1960s was gradually replaced by one that acknowledges the value of all life forms and their ecological complexity, a view currently supported even by many anglers. The necessity for wilderness fish stocking is now the subject of widespread debate, especially in view of changing social values and priorities. Options for future generations cannot be preserved if introductions continue to erode the biodiversity of mountain lake ecosystems. Received 28 March 2000; accepted 30 May 2000.     

Unimodal response of fish yield to dissolved organic carbon

Here, we demonstrate a contrasting effect of terrestrial coloured dissolved organic material on the secondary production of boreal nutrient poor lakes. Using fish yield from standardised brown trout gill‐net catches as a proxy, we show a unimodal response of lake secondary productivity to dissolved organic carbon (DOC). This suggests a trade‐off between positive and negative effects, where the initial increase may hinge upon several factors such as energy subsidising, screening of UV‐radiation or P and N load being associated with organic carbon. The subsequent decline in production with further increase in DOC is likely associated with light limitations of primary production. We also show that shallow lakes switch from positive to negative effects at higher carbon loads than deeper lakes. These results underpin the major role of organic carbon for structuring productivity of boreal lake ecosystems.     

Sub-alpine amphibian distributions related to species palatability to non-native salmonids in the Klamath mountains of northern California

The goal of this study was to examine how introduced trout influence the distributions and abundances of a sub‐alpine amphibian assemblage whose members display a variety of different life‐history and defence strategies. Our study was conducted in the sub‐alpine lentic habitats of three wilderness areas that form the core of the Klamath‐Siskiyou Bioregion of northern California, a biodiversity ‘hotspot’ that supports the highest diversity of sub‐alpine, lentic‐breeding amphibians in the western USA. These wilderness areas contain no native fishes, but all have been populated with non‐native trout for recreational fishing. Five of the eight amphibian species that occur in this region were sufficiently common to use in our study; these included one that breeds in both temporary and permanent waters and is palatable to fish (Pacific treefrog, Pseudacris regilla), two that breed primarily in permanent waters and are unpalatable to fish (western toad, Bufo boreas, and rough‐skinned newt, Taricha granulosa), and two that breed primarily in permanent waters and are palatable to fish (Cascades frog, Rana cascadae, and long‐toed salamander, Ambystoma macrodactylum). Based on life histories and predator defence strategies (i.e. palatable or not), we predicted that the three palatable species would likely be negatively correlated with introduced trout, but with P. regilla less impacted because of its use of both temporary and permanent waters. We predicted that B. boreas and T. granulosa would not be significantly correlated with introduced trout due to the lack of any predator/prey interactions between them. We surveyed 728 pond, lake, or wet meadow sites during the summers of 1999–2002, using timed gill‐net sets to measure trout occurrence and relative density, and visual encounter surveys to determine amphibian presence and abundance. We used semiparametric logistic regression models to quantify the effect of trout presence/absence and density on the probability of finding amphibian species in a water body while accounting for variation within and among the various lentic habitats sampled. The distributions of P. regilla, A. macrodactylum and R. cascadae were strongly negatively correlated with trout presence across all three wilderness areas. Ambystoma macrodactylum was 44 times more likely to be found in lakes without fish than in lakes with fish. Rana cascadae and P. regilla were 3.7 and 3.0 times more likely, respectively, to be found in fishless than fish‐containing waters. In contrast, the two unpalatable species were either uncorrelated (T. granulosa) or positively correlated (B. boreas) with fish presence. We found that the relative density of fish (catch per unit effort) was negatively correlated with the combined abundances of the three palatable amphibians, and also with both the length and the condition of the fish themselves. Our results are consistent with a compelling body of evidence that introduced fishes greatly alter the aquatic community structure of mountain lakes, ponds, and wet meadows.     

A bioenergetic assessment of the influence of stocking practices on rainbow trout (Oncorhynchus mykiss) growth and consumption in a New Zealand lake

Summary

• 1 To investigate the carrying capacity and factors affecting growth of rainbow trout in Lake Rotoiti, we employed a bioenergetics model to assess the influence of stocking rates, timing of releases and prey abundance on growth and prey consumption. We hypothesised that stocking rates and prey abundance would affect growth and prey consumption by influencing per‐capita prey availability, and that the environmental conditions encountered by fish at the time of stocking would affect growth and consumption.
• 2 Prey consumption of stocked rainbow trout was calculated with the Wisconsin bioenergetics model. We calculated growth trajectories of released trout based on data from stocked trout that were released in spring and autumn from 1993 to 2009 and then re‐captured by anglers. Diet, prey energy density, body mass lost during spawning and lake temperature were measured locally.
• 3 Stocking timing had no effect on return rates to anglers or length or weight of caught fish. Although trout released in autumn were smaller than those released in spring, autumn‐released trout grew at a faster rate and had similar lengths and weights to spring cohorts after 2 years of growth in the lake. Modelled consumption parameters were negatively correlated with trout population size, suggesting that stocking rates (347–809 fish ha^?1 year^?1) caused density‐dependent effects on growth. Although common smelt (Retropinna retropinna) accounted for 85% of total prey consumption, no significant relationship was found between prey consumption by individual trout and adult smelt abundance, possibly because trout are targeting smaller smelt that our abundance estimate did not account for.
• 4 Releasing trout in autumn appears to be advantageous for growth, possibly because (i) temperature is more suitable for growth in autumn–winter than in spring–summer and (ii) prey for small trout is abundant in autumn. Mild winter conditions appear to enhance overwinter survival and growth of rainbow trout in warm‐temperate lakes compared to higher latitudes. This implies that moderately productive warm‐temperate lake ecosystems are highly suitable for trout growth in winter, but less so in summer, when lake stratification and high nutrient levels may create conditions suitable for algal blooms and hypolimnetic deoxygenation. High growth rates of trout in warm‐temperate lakes can therefore be supported by timing releases to coincide with favourable winter conditions.

     

The history of the minnow Phoxinus phoxinus (L.) in Norway: from harmless species to pest

The causes, effects and extent of minnow Phoxinus phoxinus introductions in Norway are reviewed to assess why the introductions have had severe effects, especially where brown trout Salmo trutta is the only fish species present. The natural distribution of minnow in Norway was mainly restricted to low altitude localities in the south-eastern part of the country and in some northern areas. The distribution area expanded considerably throughout the 1900s, especially in mountain areas, due in part to the use of minnows as live bait for angling. Although minnow densities do not seem unusually high in the relatively complex fish communities of its native range, the species can achieve very high population densities when introduced to communities with few fish species, such as in the numerous recently invaded lakes where brown trout was the only fish species present. The dense minnow populations in these lakes appear to have led to reduced recruitment and growth rates in the brown trout, with abundances on average 35% lower in lakes where minnow has been introduced. The success of minnow in harsh habitats demonstrates their phenotypic and ecological plasticity, but also implies that their original distribution in Norway was restricted by early immigration history and not by environmental limitations. This suggests that human-assisted spread of the species could have strong adverse effects in Scandinavia lakes of low fish species richness.