Effects of population, family, and diet on craniofacial morphology of Icelandic Arctic charr (Salvelinus alpinus) (original) (raw)

Phenotypic plasticity in the morphology of small benthic Icelandic Arctic charr (Salvelinus alpinus)

Ecology of Freshwater Fish, 2017

Intraspecific phenotypic diversity is the raw material for evolution, so understanding its origin and maintenance is critically important for conservation of biodiversity. Intraspecific diversity in a trait or a suite of traits can result from genetic diversity and/or phenotypic plasticity. The two are, however, not independent as plasticity has been shown to evolve. In this study, we evaluated the importance of phenotypic plasticity in generating morphological diversity in populations of small benthic Arctic charr in Iceland, using a rearing experiment with contrasting modes of feeding. We also examined the association between phenotypic plasticity in offspring groups generated by the contrasting feeding modes and important ecological variables characterising the natural habitats of the respective populations. Although the level of plasticity could not be related to any of the ecological measurements, clear differences in morphological reaction norms among populations suggest that ...

Differences among families in craniofacial shape at early life-stages of Arctic charr (Salvelinus alpinus)

Background Organismal fitness can be determined at early life-stages, but phenotypic variation at these early life-stages has rarely been considered in studies on evolutionary diversification. The trophic apparatus has been shown to contribute to sympatric resource-mediated divergence in several taxa. However, processes underlying this diversification are poorly understood. Using a phenotypically variable morph of Icelandic Arctic charr (Salvelinus alpinus), we reared offspring from multiple families under standardized laboratory conditions and tested to what extent family (i.e. direct genetic and maternal effects) contributes to offspring morphology at hatching (H) and first feeding (FF). To understand the underlying mechanisms behind early life-stage variation in morphology, we examined how craniofacial shape varied according to family, egg size, offspring size and individual candidate genes related to craniofacial development. Finally, we assessed whether craniofacial shape and e...

Morphological variation over ontogeny and environments in resource polymorphic arctic charr (Salvelinus alpinus)

Evolution & Development, 2010

Natural selection requires genetically based phenotypic variation to facilitate its action and cause adaptive evolution. It has become increasingly recognized that morphological development can become canalized likely as a result of selection. However, it is largely unknown how selection may influence canalization over ontogeny and differing environments. Changes in environments or colonization of a novel one is expected to result in adaptive divergence from the ancestral population when selection favors a new phenotypic optimum. In turn, a novel environment may also expose variation previously hidden from natural selection. We tested for changes in phenotypic variation over ontogeny and environments among ecomorphs of Arctic charr (Salvelinus alpinus) from two Icelandic lakes. Populations represented varying degrees of ecological specialization, with one lake population possessing highly specialized ecomorphs exhibiting a large degree of phenotypic divergence, whereas the other displayed more subtle divergence with more ecological overlap. Here we show that ecomorphs hypothesized to be the most specialized in each lake possess significant reductions in shape variation over ontogeny regardless of environmental treatment suggesting canalized development. However, environments did change the amount of shape variation expressed in these ecomorphs, with novel environments slowing the rate at which variation was reduced over ontogeny. Thus, environmental conditions may play an important role in determining the type and amount of genetically based phenotypic variation exposed to natural selection.

Plastic resource polymorphism: effects of resource availability on Arctic char (Salvelinus alpinus) morphology

Biological Journal of the Linnean Society, 2005

Resource polymorphism has been suggested to be a platform for speciation. In some cases resource polymorphism depends on phenotypic plasticity but in other cases on genetic differences between morphotypes, which in turn has been suggested to be the ongoing development of a species pair. Here we study environmentally induced morphological differences in two age classes of Arctic char (Salvelinus alpinus) influencing char performance and diet in relation to resource availability. We found that structurally complex habitats with relatively lower zooplankton densities gave rise to individuals with a deeper body, and a downward positioned tip of the snout compared with individuals from structurally simple habitats with relatively higher zooplankton densities for both age classes. Environment also had an effect on foraging efficiency on zooplankton, with fish from structurally simple habitats had a higher foraging rate than fish from structurally complex habitats. Diet analyses showed that resource use in char mainly depends on the relative abundance of different resources. Therefore, to gain further understanding of resource polymorphism we suggest that future studies must include population dynamic feedbacks by the resources on the consumers.

Variation in the magnitude of morphological and dietary differences between individuals among populations of small benthic Arctic charr in relation to ecological factors

Ecology and Evolution

The early stages of intraspecific diversity are important for the evolution of diversification and speciation. Early stages of diversification can be seen in individual specialization, where individuals consume only a portion of the diet of the population as a whole, and how such specialization is related to phenotypic diversity within populations. Here, we study the strength of the relationship between morphological and dietary distances among individuals in eighteen populations of Icelandic small benthic charr. We furthermore studied if the strength of the relationship could be related to variation in local ecological factors these populations inhabit. In all the populations studied, there was a clear relationship between morphological and dietary distances, indicating that fish that had similar morphology were at the same time-consuming similar food items. Our findings show a systematic variation in the relationship between morphology and diet at early stages of diversification in a highly specialized small benthic charr morph. The results show the importance of fine scale comparisons within populations and furthermore the value that systematic comparisons among populations under parallel evolution can contribute toward our increased understanding of evolutionary and ecological processes.

Divergent functional traits in three sympatric Arctic charr Salvelinus alpinus morphs are not coupled with the age of the lineage divergence

Hydrobiologia, 2016

Three genetically discrete morphs of Arctic charr in Loch Rannoch, Scotland originated from a recent divergence within the lake (in situ) (piscivore and benthivore morphs) and from secondary contact of two older lineages (ex situ; a planktivore-piscivore/ benthivore divergence). To test if the expression of traits with strong functional roles was linked to the age of the divergence, fin and gill anatomy, and dentition were quantified and compared across morphs. Five additional working hypotheses suggesting a rank order of trait expression amongst morphs were also tested. The planktivorous morph had more rays in the dorsal and pectoral fins, longer gill rakers (but not more) as well as a smaller gill cavity than the other two morphs. The piscivorous morph had more palatine teeth and longer teeth on the mandible, pre-maxillary and glossohyal bones, and a larger buccal cavity. These differences indicate a differential response to selection in these functional anatomical features most likely related to morph foraging specialisms. Notably, between-morph divergences in the expression of these traits were not simply linked to the length of divergence between morphs and have arisen equally quickly in the recent (in situ) divergence as they have in older, ex situ divergences.

Natural selection for body shape in resource polymorphic Icelandic Arctic charr

Journal of evolutionary biology, 2018

Resource polymorphisms exhibit remarkable intraspecific diversity and in many cases are expected to be maintained by diversifying selection. Phenotypic trade-offs can constrain morphologically intermediate individuals from effectively exploiting both alternate resources, resulting in ecological barriers to gene flow. Determining if and how phenotypic trade-offs cause fitness variation in the wild is challenging because of phenotypic and environmental correlations associated with alternative resource strategies. We investigated multiple pathways through which morphology could affect organismal performance, as measured by growth rate, and whether these effects generate diversifying selection in polymorphic Icelandic Arctic charr (Salvelinus alpinus) populations. We considered direct effects of morphology on growth and indirect effects via trophic resource use, estimated by stable isotopic signatures, and via parasitism associated with trophic resources. We sampled over 3 years in (lak...

Relationships between lake ecology and morphological characters in Icelandic Arctic charr, Salvelinus alpinus

Biological Journal of the Linnean Society, 2011

The common occurrence of parallel phenotypic patterns suggests that a strong relationship exists between ecological dynamics and micro-evolution. Comparative studies from a large number of populations under varying sets of ecological drivers could contribute to a better understanding of this relationship. We used data on morphology of arctic charr (Salvelinus alpinus) and ecological factors from 35 Icelandic lakes to test the hypothesis that morphological patterns among monomorphic charr populations from different lakes are related to interlake variation in ecological characteristics. There is extensive phenotypic diversity among populations of Icelandic charr, and populations are easily distinguished based on overall body morphology. The results obtained in the present study showed that the morphological diversity of charr was related to large-scale diversity in lake ecology. Variation in charr morphology was related to water origin (e.g. spring fed versus run-off), bedrock age, and fish community structure. The present study shows how various ecological factors can shape the biological diversity that we observe.

The functional significance of inherited differences in feeding morphology in a sympatric polymorphic population of Arctic charr

Evolutionary Ecology, 2002

Artificially fertilised eggs from wild-caught Arctic charr parents of two sympatric morphs (benthivorous and planktivorous) from Loch Rannoch, Scotland were reared in the laboratory under identical conditions. During the subsequent 2 years, aspects of their trophic anatomy and feeding behaviour were compared. As previously described for wild-caught fish, charr derived from the benthivorous morph had an increasingly wider mouth gape for a given body length than those derived from the planktivorous morph. The functional significance of these differences in gape was tested by comparing the maximum size of prey that could be handled by each of the two morphs. In both forms, a larger gape enabled larger food particles to be eaten, but the elevation of the regression of maximum prey size on gape was higher in the benthivorous form, indicating the existence of additional morphological and/or behavioural differences influencing the size of prey consumed. When offered a choice between a typical benthic prey item and a typical pelagic food item, charr of benthivorous origin were more likely to feed on the former, whereas those of planktivorous origin were more likely to feed on the latter. Thus inherited differences in gape place constraints on foraging ability and are associated with inherited differences in dietary preference. We conclude that the functional significance of the foraging specialisations indicate a strong selection pressure for the evolution of the divergence and propose that heterochronic growth is the mechanism resulting in the divergence of tropic anatomy.

Gene expression in the phenotypically plastic Arctic charr (Salvelinus alpinus ): A focus on growth and ossification at early stages of development

Evolution & Development

Gene expression during development shapes the phenotypes of individuals. Although embryonic gene expression can have lasting effects on developmental trajectories, few studies consider the role of maternal effects, such as egg size, on gene expression. Using qPCR, we characterize relative expression of 14 growth and/or skeletal promoting genes across embryonic development in Arctic charr (Salvelinus alpinus). We test to what extent their relative expression is correlated with egg size and size at early life-stages within the study population. We predict smaller individuals to have higher expression of growth and skeletal promoting genes, due to less maternal resources (i.e., yolk) and prioritization of energy toward ossification. We found expression levels to vary across developmental stages and only three genes (Mmp9, Star, and Sgk1) correlated with individual size at a given developmental stage. Contrary to our hypothesis, expression of Mmp9 and Star showed a non-linear relationship with size (at post fertilization and hatching, respectively), whilst Sgk1 was higher in larger embryos at hatching. Interestingly, these genes are also associated with craniofacial divergence of Arctic charr morphs. Our results indicate that early life-stage variation in gene expression, concomitant to maternal effects, can influence developmental plasticity and potentially the evolution of resource polymorphism in fishes. 1 | INTRODUCTION The ability of individuals to respond to environmental heterogeneity largely depends upon changes in gene expression, framed by physiological, behavioral, and anatomic constraints (Taugbøl, Arntsen, Østbye, & Vøllestad, 2014). Such interactions between genes and environment are most often realized through development, thus providing pheno-typic variation upon which natural selection can act (e.g. West-Eberhard, 2005). During early stages of development, an organism's phenotype tends to be malleable (Hoverman & Relyea, 2007). The effects of such developmental plasticity often persist through to the adult phenotype (