Among-individual diet variation within a lake trout ecotype: Lack of stability of niche use.

In a polyphenic species, differences in resource use are expected among ecotypes, and homogeneity in resource use is expected within an ecotype. Yet, using a broad resource spectrum has been identified as a strategy for fishes living in unproductive northern environments, where food is patchily distributed and ephemeral. We investigated whether specialization of trophic resources by individuals occurred within the generalist piscivore ecotype of lake trout from Great Bear Lake, Canada, reflective of a form of diversity. Four distinct dietary patterns of resource use within this lake trout ecotype were detected from fatty acid composition, with some variation linked to spatial patterns within Great Bear Lake. Feeding habits of different groups within the ecotype were not associated with detectable morphological or genetic differentiation, suggesting that behavioral plasticity caused the trophic differences. A low level of genetic differentiation was detected between exceptionally large-sized individuals and other piscivore individuals. We demonstrated that individual trophic specialization can occur within an ecotype inhabiting a geologically young system (8,000-10,000 yr BP), a lake that sustains high levels of phenotypic diversity of lake trout overall. The characterization of niche use among individuals, as done in this study, is necessary to understand the role that individual variation can play at the beginning of differentiation processes.

From top to bottom: Do Lake Trout diversify along a depth gradient in Great Bear Lake, NT, Canada?

Depth is usually considered the main driver of Lake Trout intraspecific diversity across lakes in North America. Given that Great Bear Lake is one of the largest and deepest freshwater systems in North America, we predicted that Lake Trout intraspecific diversity to be organized along a depth axis within this system. Thus, we investigated whether a deep-water morph of Lake Trout co-existed with four shallow-water morphs previously described in Great Bear Lake. Morphology, neutral genetic variation, isotopic niches, and life-history traits of Lake Trout across depths (0-150 m) were compared among morphs. Due to the propensity of Lake Trout with high levels of morphological diversity to occupy multiple habitat niches, a novel multivariate grouping method using a suite of composite variables was applied in addition to two other commonly used grouping methods to classify individuals. Depth alone did not explain Lake Trout diversity in Great Bear Lake; a distinct fifth deep-water morph was not found. Rather, Lake Trout diversity followed an ecological continuum, with some evidence for adaptation to local conditions in deep-water habitat. Overall, trout caught from deep-water showed low levels of genetic and phenotypic differentiation from shallow-water trout, and displayed higher lipid content (C:N ratio) and occupied a higher trophic level that suggested an potential increase of piscivory (including cannibalism) than the previously described four morphs. Why phenotypic divergence between shallow- and deep-water Lake Trout was low is unknown, especially when the potential for phenotypic variation should be high in deep and large Great Bear Lake. Given that variation in complexity of freshwater environments has dramatic consequences for divergence, variation in the complexity in Great Bear Lake (i.e., shallow being more complex than deep), may explain the observed dichotomy in the expression of intraspecific phenotypic diversity between shallow- vs. deep-water habitats. The ambiguity surrounding mechanisms driving divergence of Lake Trout in Great Bear Lake should be seen as reflective of the highly variable nature of ecological opportunity and divergent natural selection itself.

Lessons Learned from an Industry, Government and University Collaboration to Restore Stream Habitats and Mitigate Effects.

Restoration ecologists conduct both basic and applied research using a diversity of funding and collaborative models. Over the last 17 years we have assessed the effectiveness of a stream compensation project in Canada's north, where an independent university-based research program was a condition of the regulatory approval process. This resulted in a non-traditional university-government-industry partnership. Here we share seven lessons that we learned from our collective experiences with the research partnership and use the Ekati diamond mine as a case study to illustrate and support lessons learned. Our advice includes opinions on the importance of: engaging collaborators early, defining roles and responsibilities, data sharing and standardization, the use of natural streams to set restoration targets, expect setbacks and surprises, treating restoration as an opportunity to experiment, and how to define success. Many of the lessons learned are broadly applicable to those whom embark on research collaborations among industry, universities, and consulting companies within a regulatory framework and may be of particular value to collaborators in early stages of their career.

Multiple generalist morphs of Lake Trout: Avoiding constraints on the evolution of intraspecific divergence?

A generalist strategy, as an adaptation to environmental heterogeneity, is common in Arctic freshwater systems, often accompanied, however, by intraspecific divergence that promotes specialization in niche use. To better understand how resources may be partitioned in a northern system that supports intraspecific diversity of Lake Trout, trophic niches were compared among four shallow-water morphotypes in Great Bear Lake (N65° 56' 39″, W120° 50' 59″). Bayesian mixing model analyses of stable isotopes of carbon and nitrogen were conducted on adult Lake Trout. Major niche overlap in resource use among four Lake Trout morphotypes was found within littoral and pelagic zones, which raises the question of how such polymorphism can be sustained among opportunistic generalist morphotypes. Covariances of our morphological datasets were tested against δ13C and δ15N values. Patterns among morphotypes were mainly observed for δ15N. This link between ecological and morphological differentiation suggested that selection pressure(s) operate at the trophic level (δ15N), independent of habitat, rather than along the habitat-foraging opportunity axis (δ13C). The spatial and temporal variability of resources in Arctic lakes, such as Great Bear Lake, may have favored the presence of multiple generalists showing different degrees of omnivory along a weak benthic-pelagic gradient. Morphs 1-3 had more generalist feeding habits using both benthic and pelagic habitats than Morph 4, which was a top-predator specialist in the pelagic habitat. Evidence for frequent cannibalism in Great Bear Lake was found across all four morphotypes and may also contribute to polymorphism. We suggest that the multiple generalist morphs described here from Great Bear Lake are a unique expression of diversity due to the presumed constraints on the evolution of generalists and contrast with the development of multiple specialists, the standard response to intraspecific divergence.

Assessing the effectiveness of a constructed Arctic stream using multiple biological attributes.

Objective assessment of habitat compensation is a central yet challenging issue for restoration ecologists. In 1997, a 3.4-km stream channel, designed to divert water around an open pit diamond mine, was excavated in the Barrenlands region of the Canadian Arctic to create productive stream habitat. We evaluated the initial success of this compensation program by comparing multiple biological attributes of the constructed stream during its first three years to those of natural reference streams in the area. The riparian zone of the constructed stream was largely devoid of vegetation throughout the period, in contrast to the densely vegetated zones of reference streams. The constructed stream also contained lower amounts of woody debris, coarse particulate organic matter (CPOM), and epilithon; had lower coverage by macrophytes and bryophytes; and processed leaf litter at a lower rate than reference streams. Species richness and densities of macroinvertebrates were consistently lower in the constructed stream compared to natural streams. This contributed to differences in macroinvertebrate assemblage structure throughout the period, although assemblages showed some convergence by year 3. The effectiveness of the constructed stream to emulate natural streams varied somewhat depending on the biological attribute being evaluated. Assessments based on individual attributes showed that minimal to moderate levels of similarity between the constructed stream and natural streams were achieved. A collective assessment of all biological and ecosystem attributes suggested that the constructed stream was not a good surrogate for natural streams during these first years. Additional time would be required before many characteristics of the constructed stream would resemble those of reference streams. Because initial efforts to improve fish habitat in the constructed stream focused on physical structures (e.g., weirs, vanes, rock, groins), ecological factors limiting fish growth were not considered and likely constrained success. We suggest that a greater focus on organic characteristics and vegetation within the stream and its riparian zone could have accelerated compensation. The addition of woody debris and CPOM, combined with planting of shrubs and herbs along the stream, should provide a source of allochthonous matter for the biotic community while large cobble and boulders should improve the physical stability of stream system, protecting its organic components.

Cascading life-history interactions: alternative density-dependent pathways drive recruitment dynamics in a freshwater fish.

Although density-dependent mechanisms in early life-history are important regulators of recruitment in many taxa, consequences of such mechanisms on other life-history stages are poorly understood. To examine interacting and cascading effects of mechanisms acting on different life-history stages, we stocked experimental ponds with fathead minnow (Pimephales promelas) at two different densities. We quantified growth and survival of the stocked fish, the eggs they produced, and the resulting offspring during their first season of life. Per-capita production and survival of eggs were inversely related to density of stocked fish; significant egg cannibalism by stocked minnows resulted in initial young-of-the-year (YOY) densities that were inversely related to adult densities. Subsequent growth and survival of YOY were then inversely related to these initial YOY densities, and survival of YOY was selective for larger fish. Because of these compensatory processes in the egg and YOY stages, treatments did not differ in YOY abundance and mean size at the end of the growing season. Because of differences in the intensity of size-selective mortality, however, variation in end-of season sizes of YOY was strongly (and inversely) related to densities of stocked fish. When mortality was severe in the egg stage (high densities of stocked fish), final YOY size distributions were more variable than when the dominant mortality was size-selective in the YOY stage (low stocked fish densities). These differences in size variation could have subsequent recruitment consequences, as overwinter survival is typically selective for YOY fish larger than a critical threshold size. Density-dependent effects on a given life stage are not independent, but will be influenced by earlier stages; alternative recruitment pathways can result when processes at earlier stages differ in magnitude or selectivity. Appreciation of these cascading effects should enhance our overall understanding of the dynamics of stage-structured populations.

Individual specialization and trophic adaptability of northern pike (Esox lucius): an isotope and dietary analysis.

Northern pike (Esox lucius) are often considered to be specialist piscivores, but under some circumstances will continue to eat invertebrates as adults. To examine effects of fish assemblage composition on the trophic ecology of pike, we combined stable isotope analysis (SIA) of carbon and nitrogen and stomach content analysis (SCA) on pike from five lakes in northern Alberta, three of which contain only pike ("pike-only") and two that also contain yellow perch (Perca flavescens) or white sucker (Catostomus commersoni) ("pike-other"). Fish were more important as prey and empty stomachs, which often characterize piscivores, were significantly more frequent in pike-other than in pike-only lakes. However, even though invertebrates were more important for pike in pike-only lakes, SIA and SCA indicated that invertebrates were also an important component of pike diets in pike-other lakes. SIA and SCA also revealed considerable intrapopulation variation in trophic ecology, with individuals in some populations differing by as much as two trophic levels. Comparisons of stomach contents and isotope signatures of the same fish suggested that within these variable populations, specialization on invertebrates or fish was a long-term trait of some individuals. SIA indicated that trophic position increased and diets shifted to a greater importance of littoral prey as pike grew in pike-only lakes, but not in lakes with other fish present. Trophic adaptability in northern pike is expressed at both the population level, where the trophic ecology is sensitive to differences in prey regimes, and at the organismal level, in the form of intrapopulation variation and individual specialization.

Competition in Umbra-Perca fish assemblages: experimental and field evidence.

One potentially important effect of interspecific competition in freshwater fish communities is to increase predation intensity from gape-limited piscivores by lowering growth rates of prey species. We investigated the operation and consequences of competition between central mudminnows (Umbra limi) and yearling yellow perch (Perca flavescens) in a system where size-limited predation on mudminnows by larger perch is a principle structuring mechanism. During laboratory experiments in which mudminnows foraged for patchily-presented food in the presence and absence of yearling perch, the food intake of mudminnows decreased at both the population and individual-fish levels when perch were present. Mudminnows were neither less active nor did they occupy lower-quality food patches in the presence of perch; exploitation competition, rather than interference or predator avoidance, appeared responsible for reduced feeding success. To assess effects of competition in the field, we examined size distributions and condition factors of yearling mudminnows and perch in small Wisconsin lakes having mudminnow-only and mudminnow-perch assemblages. Yearling mudminnows were smaller and in poorer condition in an assemblage composed (by mass) of 45% yearling perch than in two mudminnow-only assemblages. Conversely, yearling perch were larger and in better condition when the mudminnow-perch assemblage contained only 45% perch then when it contained 92% perch. Our experimental and field results indicated that an asymmetrical competitive relationship could contribute directly to the interspecific population dominance of perch over mudminnows in Umbra-Perca assemblages through reduced food intake, growth, and condition of mudminnows, and indirectly through increased vulnerability of mudminnows to size-limited predation.