The purpose of this review is to provide a global perspective on Oncorhynchus salmonine introductions and put-and-take fisheries based on modern stocking programs, with special emphasis on freshwater ecosystems. We survey the global introductions of nine selected salmonines of the genus Oncorhynchus: golden trout, cutthroat trout, pink salmon, chum salmon, coho salmon, masu/cherry salmon, rainbow trout/steelhead, sockeye salmon/kokanee, and chinook salmon. The information is organized on a geographical basis by continent, and then by species and chronology. Two different objectives and associated definitions of 'success' for introductions are distinguished: (a) seed introduction: release of individuals with the purpose of creating a wildreproducing, self-sustaining population; and (b) putand-take introduction: release of individuals with the purpose of maintaining some level of wild population abundance, regardless of wild reproduction. We identify four major phenomena regarding global salmonine introductions: (1) general inadequacy of documentation regarding introductions; (2) a fundamental disconnect between management actions and ecological consequences of introductions; (3) the importance of global climate change on success of previous and future introductions; and (4) the significance of aquaculture as a key uncertainty in accidental introductions. We conclude this review with a recognition of the need to terminate ongoing stocking programs for introduced salmonines worldwide.
Aquatic ecosystems support size structured food webs, wherein predator‐prey body sizes span orders of magnitude. As such, these food webs are replete with extremely generalized feeding strategies, especially among the larger bodied, higher trophic position taxa. The movement scale of aquatic organisms also generally increases with body size and trophic position. Together, these body size, mobility, and foraging relationships suggest that organisms lower in the food web generate relatively distinct energetic pathways by feeding over smaller spatial areas. Concurrently, the potential capacity for generalist foraging and spatial coupling of these pathways often increases, on average, moving up the food web toward higher trophic levels. We argue that these attributes make for a food web architecture that is inherently ‘adaptive’ in its response to environmental conditions. This is because variation in lower trophic level dynamics is dampened by the capacity of predators to flexibly alter their foraging behavior. We argue that empirical, theoretical, and applied research needs to embrace this inherently adaptive architecture if we are to understand the relationship between structure and function in the face of ongoing environmental change. Toward this goal, we discuss empirical patterns in the structure of lake food webs to suggest that ecosystems change consistently, from individual traits to the structure of whole food webs, under changing environmental conditions. We then explore an empirical example to reveal that explicitly unfolding the mechanisms that drive these adaptive responses offers insight into how human‐driven impacts, such as climate change, invasive species, and fisheries harvest, ought to influence ecosystem structure and function (e.g., stability, secondary productivity, maintenance of major energy pathways). We end by arguing that such a directed food web research program promises a powerful across‐scale framework for more effective ecosystem monitoring and management.
Teleost fishes are prominent vertebrate models of evolution, illustrated among old‐world radiations by the Cichlidae of East African Great Lakes and new‐world radiations by the circumpolar Arctic charr Salvelinus alpinus. Herein, we describe variation in lake charr S. namaycush morphology, life history, physiology and ecology, as another example of radiation. The lake charr is restricted to northern North America, where it originated from glacial refugia and diversified in large lakes. Shallow and deepwater morphs arose in multiple lakes, with a large‐bodied shallow‐water ‘lean’ morph in shallow inshore depths, a small‐bodied mid‐water ‘humper’ morph on offshore shoals or banks, a robust, large‐bodied moderate to deep‐water ‘redfin’; morph and a large‐bodied deep‐water ‘siscowet’ morph at depths > 100 m. Eye position, gape size, and gillraker length and spacing adapted for feeding on different‐sized prey, with piscivorous morphs (leans, siscowets and presumably redfins) reaching larger asymptotic size than invertivorous morphs (humpers). Lean morphs are light in colour, whereas deepwater morphs are drab and dark, although the pattern is reversed in dark tannic lakes. Morphs shift from benthic to pelagic feeding at a length of 400–490 mm. Phenotypic differences in locomotion, buoyancy and lipid metabolism evolved into different mechanisms for buoyancy regulation, with lean morphs relying on hydrodynamic lift and siscowet morphs relying on hydrostatic lift. We suggest that the Salvelinus genus, rather than the species S. alpinus, is a diverse genus that should be the subject of comparative studies of processes causing divergence and adaptation among member species that may lead to a more complete evolutionary conceptual model.
Four Lake Trout Salvelinus namaycush morphs were identified from Isle Royale, Lake Superior; the morphs differed in shape, traits linked to feeding and locomotion, buoyancy, and physical habitat use. Lean, humper, and siscowet Lake Trout generally conformed to previous descriptions, and we report, for the first time, quantitative evidence of a fourth morph, previously described anecdotally as the “redfin.” Jackknife classification of individuals to morphs based on body shape were 94% correct. High variation within and low variation among morphs led to moderately low percent agreement among visual identifications and high uncertainty in Bayesian model groupings of morphs. Eight linear measures of phenotypic traits linked to feeding (i.e., head and eyes) and locomotion (i.e., fin lengths and caudal peduncle shape) varied among morphs, consistent with specialized adaptations for trophic and physical resource use. Habitat differed among morphs with leans being most abundant in the 0–50‐m depth stratum and siscowets most abundant in two deeper strata (50–100 and 100–150 m). Differences in capture depth and percent buoyancy reflected physical habitat and known trophic resource partitioning among morphs. While the historical fingerprint of morphological and ecological diversity in Lake Superior Lake Trout persists, it is unknown whether the contemporary low level of differentiation is due to ecological release without subsequent reorganization or to a complete breakdown of differentiation. Received July 11, 2013; accepted February 7, 2014
The study of cisco diversity in inland lakes of North America has been plagued by taxonomic uncertainty linked to high phenotypic plasticity and an ongoing reliance on morphology to differentiate species. More recently, this uncertainty has hindered the development of conservation plans and status assessments of ciscoes. This study presents the first range-wide comparison of morphological and genetic variation between Cisco (Coregonus artedi) and Shortjaw Cisco (C. zenithicus). Using morphological and genetic data from 17 lakes, three sets of analyses were undertaken to evaluate alternate hypotheses explaining the pattern of cisco phenotypic diversity in inland lakes. Morphotypes (MTs) representing the two taxa were phenotypically distinct (largely reflective of differences in gill raker number and jaw morphology) within lakes but highly variable across lakes. Shortjaw Cisco was only recognizable when compared to sympatric Cisco and some populations were morphologically similar to Cisco from other lakes. Analysis of AFLP data revealed two genetic clusters that conformed to differences in geography (eastern and western groups), rather than hypothesized taxonomic boundaries. Genetic variation strongly suggests that each of these unique sympatric pairs of MTs originated recently and locally, in parallel, from the ancestral Cisco. Phenotypic and genetic distinctiveness between MTs were not related. MTs were sometimes clearly recognizable despite a lack of genetic differentiation, suggesting that the canalization of phenotypic plasticity is unevenly completed across lakes. These results provide evidence that the taxonbased approach is clearly inadequate for the protection of Shortjaw Cisco. In Canada, status assessment should aim to identify lake-specific designatable units (DU). Given the idiosyncratic nature of each instance of Shortjaw Cisco, it is expected that the strength of morphological, biological, ecological and genetic evidence for individual DUs will vary among lakes.
Understanding the emergence of species through the process of ecological speciation is a central question in evolutionary biology which also has implications for conservation and management. Lake trout (Salvelinus namaycush) is renowned for the occurrence of different ecotypes linked to resource and habitat use throughout North America. We aimed to unravel the fine genetic structure of the four lake trout ecotypes in Lake Superior. A total of 486 individuals from four sites were genotyped at 6822 filtered SNPs using RADseq technology. Our results revealed different extent of morphological and genetic differentiation within the different sites. Overall, genetic differentiation was weak but significant and was on average three times higher between sites (mean F = 0.016) than between ecotypes within sites (mean F = 0.005) indicating higher level of gene flow or a more recent shared ancestor between ecotypes within each site than between populations of the same ecotype. Evidence of divergent selection was also found between ecotypes and/or in association with morphological variation. Outlier loci found in genes related to lipid metabolism and visual acuity were of particular interest in this context of ecotypic divergence. However, we did not find clear indication of parallelism at the genomic level, despite the presence of phenotypic parallelism among some ecotypes from different sampling sites. Overall, the occurrence of different levels of both genomic and phenotypic differentiation between ecotypes within each site with several differentiated loci linked to relevant biological functions supports the presence of a continuum of divergence in lake trout.
We present digital imaging methods for geometric morphometric analysis of shape, and we describe issues associated with improper image acquisition by using lake trout Salvelinus namaycush as an example. The choice of imaging equipment, the configuration of that equipment, and the orientation of the specimens with respect to the camera lens can lead to inaccurate imaging and ultimately to error in landmark placement during morphometric analysis. Lake trout that were imaged at 15‐mm focal length and 0.5‐m focal distance (treatment 1) were distorted in comparison with fish that were imaged at 50‐mm focal length and 2‐m focal distance (treatment 2). Deformation grids showed dramatic variation in the horizontal plane along the length of the fish, especially midbody, suggesting that barrel distortion was occurring at the 15‐mm focal length. Partial warp scores resulting from geometric analysis of body shape differed for all fish on all 18 warps as a result of the different focal length and distance treatments for image capture. To minimize perspective (orientation) and distortion (equipment) errors, we recommend using a digital single‐lens reflex camera (>5 megapixels) with a lens that has a focal length exceeding 35 mm, a horizontal tripod to position the lens directly over the specimen, a mesh cradle to create a planar imaging surface, and dissection pins to display the fish in a standard orientation. The method presented herein will aid in reducing measurement error associated with landmark homology and will promote comparability of geometric shape data among studies.
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