Summary1. Stable isotope analyses coupled with mixing models are being used increasingly to evaluate ecological management issues and questions. Such applications of stable isotope analyses often require simultaneous carbon and nitrogen analyses from the same sample. Correction of the carbon isotope values to take account of the varying content of 13 C-depleted lipids is then frequently achieved by a lipid-normalization procedure using a model describing the relationship between change in δ 13 C following lipid removal and the original C:N ratio of a sample. 2. We evaluated the applicability of two widely used normalization models using empirical data for muscle tissue from a wide range of fish and for aquatic invertebrates. Neither normalization model proved satisfactory, and we present some modifications that greatly improve the fit of one of the models to the fish muscle data. For invertebrates we found no clear relationship between change in δ 13 C following lipid removal and the original C:N ratio. 3. We also examined the effect of lipid-normalization on the output of a mixing model designed to calculate the proportional contribution of prey items to the diet of a consumer. Mixing model output was greatly influenced by whether prey or consumer values alone or together were lipid-normalized and we urge caution in the interpretation of results from these models pending further experimental evidence. 4. Synthesis and applications . We describe a revised lipid-normalization model that should be applicable to a wide range of marine and freshwater fish species in studies applying stable isotope analyses to ecological management issues. However, we strongly advise against applying these kinds of lipid-normalization models to aquatic invertebrate data. The interpretation of outputs from mixing models is greatly influenced by whether the carbon isotope data have been lipid-normalized or not.

Summary1. The adaptive radiation of fishes into benthic (littoral) and pelagic (lentic) morphs in post-glacial lakes has become an important model system for speciation. Although these systems are well studied, there is little evidence of the existence of morphs that have diverged to utilize resources in the remaining principal lake habitat, the profundal zone.2. Here, we tested phenotype-environment correlations of three whitefish (Coregonus lavaretus) morphs that have radiated into littoral, pelagic and profundal niches in northern Scandinavian lakes. We hypothesized that morphs in such trimorphic systems would have a morphology adapted to one of the principal lake habitats (littoral, pelagic or profundal niches). Most whitefish populations in the study area are formed by a single (monomorphic) whitefish morph, and we further hypothesized that these populations should display intermediate morphotypes and niche utilization. We used a combination of traditional (stomach content, habitat use, gill raker counts) and more recently developed (stable isotopes, geometric morphometrics) techniques to evaluate phenotype-environment correlations in two lakes with trimorphic and two lakes with monomorphic whitefish. 3. Distinct phenotype-environment correlations were evident for each principal niche in whitefish morphs inhabiting trimorphic lakes. Monomorphic whitefish exploited multiple habitats, had intermediate morphology, displayed increased variance in gillraker-counts, and relied significantly on zooplankton, most likely due to relaxed resource competition. 4. We suggest that the ecological processes acting in the trimorphic lakes are similar to each other, and are driving the adaptive evolution of whitefish morphs, possibly leading to the formation of new species.

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Ecological speciation has been the subject of intense research in evolutionary biology but the genetic basis of the actual mechanism driving reproductive isolation has rarely been identified. The extreme polymorphism of the major histocompatibility complex (MHC), probably maintained by parasite-mediated selection, has been proposed as a potential driver of population divergence. We performed an integrative field and experimental study using three-spined stickleback river and lake ecotypes. We characterized their parasite load and variation at MHC class II loci. Fish from lakes and rivers harbor contrasting parasite communities and populations possess different MHC allele pools that could be the result of a combined action of genetic drift and parasite-mediated selection. We show that individual MHC class II diversity varies among populations and is lower in river ecotypes. Our results suggest the action of homogenizing selection within habitat type and diverging selection between habitat types. Finally, reproductive isolation was suggested by experimental evidence: in a flow channel design females preferred assortatively the odor of their sympatric male. This demonstrates the role of olfactory cues in maintaining reproductive isolation between diverging fish ecotypes

Recent climatic change has been recorded across the globe. Although environmental change is a characteristic feature of life on Earth and has played a major role in the evolution and global distribution of biodiversity, predicted future rates of climatic change, especially in temperature, are such that they will exceed any that has occurred over recent geological time. Climate change is considered as a key threat to biodiversity and to the structure and function of ecosystems that may already be subject to significant anthropogenic stress. The current understanding of climate change and its likely consequences for the fishes of Britain and Ireland and the surrounding seas are reviewed through a series of case studies detailing the likely response of several marine, diadromous and freshwater fishes to climate change. Changes in climate, and in particular, temperature have and will continue to affect fish at all levels of biological organization: cellular, individual, population, species, community and ecosystem, influencing physiological and ecological processes in a number of direct, indirect and complex ways. The response of fishes and of other aquatic taxa will vary according to their tolerances and life stage and are complex and difficult to predict. Fishes may respond directly to climate-change-related shifts in environmental processes or indirectly to other influences, such as community-level interactions with other taxa. However, the ability to adapt to the predicted changes in climate will vary between species and between habitats and there will be winners and losers. In marine habitats, recent changes in fish community structure will continue as fishes shift their distributions relative to their temperature preferences. This may lead to the loss of some economically important cold-adapted species such as Gadus morhua and Clupea harengus from some areas around Britain and Ireland, and the establishment of some new, warm-adapted species. Increased temperatures are likely to favour cool-adapted (e.g. Perca fluviatilis) and warm-adapted freshwater fishes (e.g. roach Rutilus rutilus and other cyprinids) whose distribution and reproductive success may currently be constrained by temperature rather than by cold-adapted species (e.g. salmonids). Species that occur in Britain and Ireland that are at the edge of their distribution will be most affected, both negatively and positively. Populations of conservation importance (e.g.Salvelinus alpinus and Coregonus spp.) may decline irreversibly. However, changes in food-web dynamics and physiological adaptation, for example because of climate change, may obscure or alter predicted responses. The residual inertia in climate systems is such that even a complete cessation in emissions would still leave fishes exposed to continued climate change for at least half a century. Hence, regardless of the success or failure of programmes aimed at curbing climate change, major changes in fish communities can be expected over the next 50 years with a concomitant need to adapt manag...

Freshwater populations of three-spined sticklebacks (Gasterosteus aculeatus) in northern Germany are found as distinct lake and river ecotypes. Adaptation to habitat-specific parasites might influence immune capabilities of stickleback ecotypes. Here, naive laboratory-bred sticklebacks from lake and river populations were exposed reciprocally to parasite environments in a lake and a river habitat. Sticklebacks exposed to lake conditions were infected with higher numbers of parasite species when compared with the river. River sticklebacks in the lake had higher parasite loads than lake sticklebacks in the same habitat. Respiratory burst, granulocyte counts and lymphocyte proliferation of head kidney leucocytes were increased in river sticklebacks exposed to lake when compared with river conditions. Although river sticklebacks exposed to lake conditions showed elevated activation of their immune system, parasites could not be diminished as effectively as by lake sticklebacks in their native habitat. River sticklebacks seem to have reduced their immune-competence potential due to lower parasite diversity in rivers.

BackgroundAfter a volcano erupts, a lake may form in the cooled crater and become an isolated aquatic ecosystem. This makes fishes in crater lakes informative for understanding sympatric evolution and ecological diversification in barren environments. From a geological and limnological perspective, such research offers insight about the process of crater lake ecosystem establishment and speciation. In the present study we use genetic and coalescence approaches to infer the colonization history of Midas cichlid fishes (Amphilophus cf. citrinellus) that inhabit a very young crater lake in Nicaragua-the ca. 1800 year-old Lake Apoyeque. This lake holds two sympatric, endemic morphs of Midas cichlid: one with large, hypertrophied lips (~20% of the total population) and another with thin lips. Here we test the associated ecological, morphological and genetic diversification of these two morphs and their potential to represent incipient speciation.ResultsGene coalescence analyses [11 microsatellite loci and mitochondrial DNA (mtDNA) sequences] suggest that crater lake Apoyeque was colonized in a single event from the large neighbouring great lake Managua only about 100 years ago. This founding in historic times is also reflected in the extremely low nuclear and mitochondrial genetic diversity in Apoyeque. We found that sympatric adult thin- and thick-lipped fishes occupy distinct ecological trophic niches. Diet, body shape, head width, pharyngeal jaw size and shape and stable isotope values all differ significantly between the two lip-morphs. The eco-morphological features pharyngeal jaw shape, body shape, stomach contents and stable isotopes (δ15N) all show a bimodal distribution of traits, which is compatible with the expectations of an initial stage of ecological speciation under disruptive selection. Genetic differentiation between the thin- and thick-lipped population is weak at mtDNA sequence (FST = 0.018) and absent at nuclear microsatellite loci (FST < 0.001).ConclusionsThis study provides empirical evidence of eco-morphological differentiation occurring very quickly after the colonization of a new and vacant habitat. Exceptionally low levels of neutral genetic diversity and inference from coalescence indicates that the Midas cichlid population in Apoyeque is much younger (ca. 100 years or generations old) than the crater itself (ca. 1 800 years old). This suggests either that the crater remained empty for many hundreds of years after its formation or that remnant volcanic activity prevented the establishment of a stable fish population during the early life of the crater lake. Based on our findings of eco-morphological variation in the Apoyeque Midas cichlids, and known patterns of adaptation in Midas cichlids in general, we suggest that this population may be in a very early stage of speciation (incipient species), promoted by disruptive selection and ecological diversification.

Introduction of the invasive Asian cyprinid fish Pseudorasbora parva into a 0.3 ha pond in England with a fish assemblage that included Cyprinus carpio, Rutilus rutilus and Scardinius erythrophthalmus resulted in their establishment of a numerically dominant population in only 2 years; density estimates exceeded 60 ind. m(-2) and they comprised > 99% of fish present. Stable isotope analysis (SIA) revealed significant trophic overlap between P. parva, R. rutilus and C. carpio, a shift associated with significantly depressed somatic growth in R. rutilus. Despite these changes, fish community composition remained similar between the ponds. Comparison with SIA values collected from an adjacent pond free of P. parva revealed a simplified food web in P. parva presence, but with an apparent trophic position shift for several fishes, including S. erythrophthalmus which appeared to assimilate energy at a higher trophic level, probably through P. parva consumption. The marked isotopic shifts shown in all taxa in the P. parva invaded pond (C-13-enriched, N-15 depleted) were indicative of a shift to a cyanobacteria-dominated phytoplankton community. These findings provide an increased understanding of the ecological consequences of the ongoing P. parva invasion of European freshwater ecosystems