Trophic polymorphism, habitat and diet segregation in Percichthys trucha (Pisces: Percichthyidae) in the Andes

Divergent natural selection affecting specific trait combinations that lead to greater efficiency in resource exploitation is believed to be a major mechanism leading to trophic polymorphism and adaptive radiation. We present evidence of trophic polymorphism involving two benthic morphs within Percichthys trucha , a fish endemic to temperate South America. In a series of lakes located in the southern Andes, we found two morphs of P. trucha that could be distinguished on the basis of gill raker length and five other morphological measures, most of which are likely associated with the use of food resources. The differences were consistent across all lakes examined, and were correlated with habitat use and diet. Individuals with longer gill rakers were more abundant in the littoral zone (littoral morph) while the short gill-raker morph was more abundant at 10 m depth and deeper (deep benthic morph). Both morphs fed primarily on benthic invertebrates, but the littoral morph fed more on larval Anisoptera than did the deep benthic morph. Phenotypic correlations among traits were high for the littoral morph, but low and non-significant for the deep-benthic morph. We suggest that gill raker length may influence the relative efficiency of suction feeding for the two morphs. This is the first evidence of trophic polymorphism in fishes from temperate South America.     

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

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.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 341–351.     

Intraspecific vicariant history and the evolution of adaptive morphological diversity in a fish species (Osmerus mordax)

Vicariant geographic isolation and resource partitioning have long been independently identified as processes contributing to the morphological divergence of closely-related species. However, little is known about the extent to which vicariant history influences the adaptive ecological divergence associated with resource partitioning and trophic specialization within species. The present study thus quantified the contribution of vicariant historical genetic divergence to the adaptive contemporary morphological divergence of intraspecific feeding specialists in the Rainbow smelt (Pisces: Osmerus mordax ). This species is characterized by the polyphyletic origin of two lacustrine feeding specialists originating in two intraspecific lineages associated with independent glacial refuges. The historical genetic segregation was initiated approximately 350 000 years ago, whereas the lacustrine trophic segregation arose within the past 10 000 years. Wild caught lacustrine smelt populations were grouped a priori based on known historical genetic identities (Acadian and Atlantic mitochondrial DNA clades) and contemporary feeding specializations (microphageous and macrophageous morphotypes). The present study demonstrated that independent suites of correlated morphological traits are associated with either vicariant history or contemporary feeding specializations. Second, functionally-similar feeding specialists exhibit distinct morphologies resulting largely from vicariant historical processes. Although, the evolutionary processes producing historical phenotypes remains unknown, the results obtained demonstrate how adaptive radiation associated with ecological resource partitioning and feeding specializations can be strongly influenced by intraspecific phenotypic diversification resulting from relatively recent vicariant histories.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 140–151.     

Morphological segregation of Icelandic threespine stickleback (Gasterosteus aculeatus L).

Icelandic threespine sticklebacks show parallel sympatric morphological differences related to different substrate habitats in four Icelandic lakes. The level of morphological diversification varies among the lakes, ranging from a population with a wide morphological distribution to a population with clear resource morphs, where morphological diversification was reflected in diet differences. These differences in morphological divergence are closely related to the differences in the ecological surroundings of each population. This appears to be resource polymorphism, which may lead to population differentiation and speciation. Trophically related sexual dimorphism was also common in these sticklebacks, which is possibly the result of sexual selection or habitat segregation by the sexes. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 247–257.     

Disjunctions in relict alpine plants: phylogeography of Androsace brevis and A. wulfeniana (Primulaceae)

Amplified Fragment Length Polymorphism (AFLP) was used to clarify the glacial history of the rare, disjunctly distributed, alpine cushion plant Androsace wulfeniana , which is endemic to the Eastern Alps (Austria and Italy). Disjunct populations in the Dolomites are genetically very distinct from those in the main distributional area. It is hypothesized that they are descendants of long-term isolated glacial survivors and are not a result of recent long-­distance dispersal. Within the main distributional area of the species in the central Eastern Alps, two groups of ­populations can be distinguished, which are congruent with hotspots of rare relictual vascular plant taxa. In the ­taxonomically closely related A. brevis growing in the Southern Alps (Italy, Switzerland), no genetic-geographical structure was found. Genetic variation is extremely low in disjunct populations of A. wulfeniana in the Dolomites and in A. brevis . In contrast, in the main distributional area of A. wulfeniana , genetic variation is similar to that of the colonizing widespread congener A. alpina .  © 2003 The Linnean Society of London, Botanical Journal of the ­Linnean Society , 2003, 141 , 437–446.     

Genetic and morphological divergence along the littoral–pelagic axis in two common and sympatric fishes: perch,Perca fluviatilis (Percidae) and roach,Rutilus rutilus (Cyprinidae)

Individuals are constantly in competition with one another and, on both ecological and evolutionary timescales, processes act to reduce this competition and promote the gain of fitness advantages via diversification. Here we have investigated the genetic (AFLP) and morphological (geometric morphometrics) aspects of the littoral–pelagic axis, a commonly observed resource polymorphism in freshwater fishes of postglacial lakes. We found a large degree of variation in the genetic and morphological divergence between littoral and pelagic perch and roach across Swedish lakes. Although there was evidence of assortative mating (elevated kinship values) in both species, we could not find any significant coupling of morphology and genetic divergence. Instead, there was evidence that the extent of resource polymorphism may be largely caused by phenotypic plasticity. These results suggest that assortative mating, which can lead to genetically determined adaptive divergence, does occur in these species, particularly perch, but not according to genetically fixed morphological traits. The behavioural mechanisms facilitating associative mating need to be investigated to explore the interaction between phenotypic plasticity and adaptive genetic divergence and their roles in diversification. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 929–940.     

The role of gill raker number variability in adaptive radiation of coregonid fish

Gill raker divergence is a general pattern in adaptive radiations of postglacial fish, but few studies have addressed the adaptive significance of this morphological trait in foraging and eco-evolutionary interactions among predator and prey. Here, a set of subarctic lakes along a diversifying gradient of coregonids was used as the natural setting to explore correlations between gill raker numbers and planktivory as well as the impact of coregonid radiation on zooplankton communities. Results from 19 populations covering most of the total gill raker number gradient of the genus Coregonus, confirm that the number of gill rakers has a central role in determining the foraging ability towards zooplankton prey. Both at the individual and population levels, gill raker number was correlated with pelagic niche use and the size of utilized zooplankton prey. Furthermore, the average body size and the abundance and diversity of the zooplankton community decreased with the increasing diversity of coregonids. We argue that zooplankton feeding leads to an eco-evolutionary feedback loop that may further shape the gill raker morphology since natural selection intensifies under resource competition for depleted prey communities. Eco-evolutionary interactions may thus have a central role creating and maintaining the divergence of coregonid morphs in postglacial lakes.     

Long term diet differences between morphs in trophically polymorphic Perdchthys trucha (Pisces: Percichthyidae) populations from the southern Andes

Divergent natural selection is often believed to be the driving force behind phenotypic differentiation in characters related to resource acquisition, leading to trophic polymorphism in fishes. Here we use variation in the fatty acid composition of adipose and muscle tissues to look at differences in resource use by two recently described sympatric morphs of Percichthys trucha , a common freshwater fish of the Andean and Patagonian regions of South America. Because dietary fatty acids are often stored in carnivorous animals with litde modification after consumption, they can be used to infer information about dietary habits of individuals. We found that the two morphs differed in the overall composition of fatty acids in both adipose and muscle tissue, but that there were some differences in how die morphs differed in lakes from the northern vs southern part of the range. Furthermore, we found that certain fatty acids were correlated with diet as determined by gut content analysis. Consumption of anisopteran larvae was highly correlated widi 14:0 in adipose and muscle tissue; and higher levels of longer chain unsaturated fatty acids (i.e. 20 and 22 carbons) were correlated with die presence offish and also amphipods in die diets. Taken togetiier, die results suggest that diere are marked differences in die foraging ecology of die two morphs of P. trucha inhabiting southern Andean lakes.     

Genetic divergence in adaptive characters between sympatric species of stickleback

This study explored the genetic basis of phenotypic differences between two sympatric species of ecologically and morphologically divergent sticklebacks (Gasterosteus aculeatus complex). The aim was to understand how many loci determine the differences and to what extent the differences are due to additive or nonadditive gene action. I reared the two parental species, F1 and F2 hybrids, and both backcrosses in the laboratory and measured the following quantitative characters: gill raker number and length (both involved in feeding), lateral plate number and pelvic spine length (both involved in predator defense), and growth (a fitness component). I then applied joint-scaling regression models to estimate composite additive, dominance and epistatic effects, and their contribution to divergence of parental lines. A simple additive model was sufficient for gill raker number and growth; additive and dominance effects contributed significantly to divergence in plate number and pelvic spine length; and additive, dominance, and epistatic effects contributed significantly to divergence in gill raker length. Wright's estimator for the number of loci for the four morphological characters ranged from 1 to 50. My results suggest that adaptive divergence between limnetic and benthic sticklebacks has taken place through a variety of genetic mechanisms specific to different traits. Though interspecific hybrids are completely fertile and viable in the laboratory, they are selected against in the wild. The pattern of inheritance for the traits examined here directly influences how well hybrids can exploit the two major resource environments in the wild.     

Incipient speciation driven by phenotypic plasticity? Evidence from sympatric populations of Arctic charr

Recent models suggest that the existence of environmentally induced polymorphisms within a single population (especially those related to foraging) facilitates the process of evolutionary divergence within a single gene pool by generating distinct phenotypic modes that are exposed to differential selection. In order to test a prediction of the phenotypic plasticity model of divergence, we used a well-documented polymorphism to disentangle the relative effects of morph and rearing environment in generating phenotypic variance. We reared first-generation offspring of two sympatric morphs of Arctic charr Salvelinus alpinus in the laboratory and compared their head morphology with that of their wild parents. Morphological characters with a known functional role in foraging were highly plastic. Rearing environment accounted for the largest component of the variation in expressed phenotype, but this environmental effect overlaid a clear (but small) genetic effect. We conclude that phenotypic plasticity has played a significant role in the evolution of this trophic polymorphism, but that the evolutionary process has progressed to the point that the gene pool is now segregated.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 611–618.     

Discordance in body size, colour pattern, and advertisement call across genetically distinct populations in a Neotropical anuran (Dendropsophus ebraccatus)

Patterns of intraspecific geographic variation in morphology and behaviour, when examined in a phylogenetic context, can provide insight into the microevolutionary processes driving population divergence and ultimately speciation. In the present study, we quantified behavioural and phenotypic variation among populations from genetically divergent regions in the Central American treefrog, Dendropsophus ebraccatus . Our fine-scale population comparisons demonstrated regional divergence in body size, colour pattern frequencies, and male advertisement call. None of the characters covaried with phylogenetic history or geographic proximity among sampled populations, indicating the importance of highly localized selection pressures and genetic drift in shaping character divergence among isolated regions. The study underscores how multiple phenotypic characters can evolve independently across relatively small spatial scales.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 298–313.     

Constraints on speciation suggested by comparing lake-stream stickleback divergence across two continents

Adaptation to ecologically distinct environments can coincide with the emergence of reproductive barriers. The outcome of this process is highly variable and can range along a continuum from weak population differentiation all the way to complete, genome-wide divergence. The factors determining how far diverging taxa will move along this continuum remain poorly understood but are most profitably investigated in taxa under replicate divergence. Here, we explore determinants of progress towards speciation by comparing phenotypic and molecular divergence within young (<150 years) lake-stream stickleback pairs from Central Europe to divergence in older (thousands of years) archetypal lake-stream pairs from Vancouver Island, Canada. We generally find relatively weak divergence in most aspects of foraging morphology (gill raker number, body shape) in the European pairs, although substantial adaptive divergence is seen in gill raker length. Combined with striking overall phenotypic differences between the continents, this argues for genetic and time constraints on adaptive divergence in the European pairs. The European lake-stream pairs also do not display the strong habitat-related differentiation in neutral (microsatellite) markers seen in the Canadian watersheds. This indicates either the lack of strong reproductive barriers owing to weak adaptive divergence, or alternatively that neutral markers are poorly suited for detecting reproductive barriers if these emerge rapidly. Overall, our comparative approach suggests constraints on speciation due to genetic architecture and limited time for divergence. The relative importance of these factors remains to be quantified by future investigation.     

Genetic differentiation in gill raker number and length in sympatric anadromous and nonanadromous morphs of sockeye salmon, Oncorhynchus nerka

The genetic and environmental basis for polymorphism in gill raker number and length in sympatric anadromous and nonanadromous morphs of sockeye salmon, Oncorhynchus nerka, was investigated. Analysis of 30 full sib families involving pure types and reciprocal hybrids revealed that the variation was partitioned significantly among families within cross types and among cross types in both traits. As in the wild, kokanee displayed more gill rakers than sockeye; reciprocal hybrids displayed intermediate counts. Gill raker length also varied markedly among cross types, with pure sockeye displaying 19% longer gill rakers than comparable sized kokanee. This difference was in the opposite direction predicted, given the common positive association between gill raker number and length in sympatric morphs of the same species in fishes. Gill raker number and length were generally not correlated within cross types, suggesting independent divergence of the traits. The results are discussed in relation to genetic and trophic divergence of the morphs and to factors selecting for differentiation in the two gill raker traits.     

Phylogenetic relationships of Atherina hepsetus and Atherina boyeri (Pisces: Atherinidae) populations from Greece, based on mtDNA sequences

The genetic divergence and the phylogenetic relationships of six Atherina boyeri (freshwater and marine origin) and five Atherina hepsetus populations from Greece were investigated using partial sequence analysis of 12s rRNA, 16s rRNA and control region mtDNA segments. Three different well divergent groups were revealed; the first one includes A. boyeri populations living in the sea, the second includes A. boyeri populations living in the lakes and lagoons whereas the third one includes all A. hepsetus populations. Fifty-seven different haplotypes were detected among the populations studied. In all three mtDNA segments examined, sequence analysis revealed the existence of fixed haplotypic differences discriminating A. boyeri populations inhabiting the lagoon and the lakes from both the coastal A. boyeri and the A. hepsetus populations. The genetic divergence values estimated between coastal (marine) A. boyeri populations and those living in the lagoon and the lakes are of the same order of magnitude as those observed among coastal A. boyeri and A. hepsetus populations. The results obtained by different phylogenetic methods were identical. The deep sequence divergence with the fixed different haplotypes observed suggests the occurrence of a cryptic or sibling species within A. boyeri complex.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 151–161.     

Experimental evidence for trait utility of gill raker number in adaptive radiation of a north temperate fish

North temperate fish in post‐glacial lakes are textbook examples for rapid parallel adaptive radiation into multiple trophic specialists within individual lakes. Speciation repeatedly proceeded along the benthic–limnetic habitat axis, and benthic–limnetic sister species diverge in the number of gill rakers. Yet, the utility of different numbers of gill rakers for consuming benthic vs. limnetic food has only very rarely been experimentally demonstrated. We bred and raised families of a benthic–limnetic species pair of whitefish under common garden conditions to test whether these species (i) show heritable differentiation in feeding efficiency on zooplankton, and (ii) whether variation in feeding efficiency is predicted by variation in gill raker numbers. We used zooplankton of three different size classes to investigate prey size dependency of divergence in feeding efficiency and to investigate the effect strength of variation in the number of gill rakers. Our results show strong interspecific differences in feeding efficiency. These differences are largest when fish were tested with the smallest zooplankton. Importantly, feeding efficiency is significantly positively correlated with the number of gill rakers when using small zooplankton, also when species identity is statistically controlled for. Our results support the hypothesis that a larger number of gill rakers are of adaptive significance for feeding on zooplankton and provide one of the first experimental demonstrations of trait utility of gill raker number when fish feed on zooplankton. These results are consistent with the suggested importance of divergent selection driven feeding adaptation during adaptive radiation of fish in post‐glacial lakes.     

Morphotypes,varieties, or subspecies?: genetic diversity and differentiation of four Saintpaulia (Gesneriaceae) morphotypes from the East Usambara Mountains,Tanzania

Saintpaulia (H. Wendl.) is a forest-dependent, endangered plant genus endemic to Tanzania and Kenya. The taxonomy of Saintpaulia from the Usambara Mountains and adjacent lowlands in north-eastern Tanzania is problematic because of the morphological similarity of the species and the presence of considerable intraspecific variation. Conventional molecular phylogenetic methods have failed to reveal the genetic structure of this Saintpaulia complex. In this study, we assessed the genetic composition of 12 Saintpaulia populations, representing four different morphotypes, from the East Usambara Mountains using inter-simple sequence repeat (ISSR) markers. Relatively high genetic diversities were observed within populations (mean h  = 0.320), indicating their adaptive potential. Little genetic differentiation amongst populations (mean F _st  = 0.063) and the genetic divergence of the rosulate and trailing morphotypes support the hypothesis of ongoing divergent evolution within the East Usambara metapopulation(s) of Saintpaulia .  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 347–355.     

Description, biology and conservation of a new species of Australian tree frog (Amphibia: Anura: Hylidae: Litoria) and an assessment of the remaining populations of Litoria genimaculata Horst, 1883: systematic and conservation implications of an unusual speciation event

The Australian populations of the green-eyed tree frog Litoria genimaculata consist of a northern and southern genetic lineage that meet in a mosaic contact zone comprising two independent areas of contact: one where the main ranges of the lineages overlap, and the second where a population of the southern lineage is isolated within the range of the northern lineage. A recent study failed to find significant reproductive isolation between the main ranges of the two lineages, despite deep genetic divergence, partial postzygotic isolation, and call differences. The study did, however, demonstrate rapid phenotypic divergence and speciation of the isolated population of the southern lineage from both the parapatric northern lineage and from the allopatric, but genetically similar, main range of the southern lineage. Herein, the isolated population of the southern lineage is described as a distinct species, Litoria myola sp. nov. , whereas the remainder of the southern lineage and the northern lineage are retained as a single, paraphyletic species, Litoria genimaculata . Resolving this unusual systematic situation demonstrates the value of using multiple lines of evidence in delimiting species. Litoria myola sp. nov. has a very small distribution and population size and warrants a Critically Endangered listing (B1, 2) under IUCN criteria. Threats and management recommendations are outlined, and the conservation of hybrid zones as areas of evolutionary novelty is discussed.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 549–563.     

Diversifying selection drives parallel evolution of gill raker number and body size along the speciation continuum of European whitefish

Adaptive radiation is the evolution of ecological and phenotypical diversity. It arises via ecological opportunity that promotes the exploration of underutilized or novel niches mediating specialization and reproductive isolation. The assumed precondition for rapid local adaptation is diversifying natural selection, but random genetic drift could also be a major driver of this process. We used 27 populations of European whitefish (Coregonus lavaretus) from nine lakes distributed in three neighboring subarctic watercourses in northern Fennoscandia as a model to test the importance of random drift versus diversifying natural selection for parallel evolution of adaptive phenotypic traits. We contrasted variation for two key adaptive phenotypic traits correlated with resource utilization of polymorphic fish; the number of gill rakers and the total length of fish, with the posterior distribution of neutral genetic differentiation from 13 microsatellite loci, to test whether the observed phenotypic divergence could be achieved by random genetic drift alone. Our results show that both traits have been under diversifying selection and that the evolution of these morphs has been driven by isolation through habitat adaptations. We conclude that diversifying selection acting on gill raker number and body size has played a significant role in the ongoing adaptive radiation of European whitefish morphs in this region.     

Empirical evaluation of phenotype–environment correlation and trait utility with allopatric and sympatric whitefish, Coregonus lavaretus (L.), populations in subarctic lakes

Adaptive phenotypic divergence of sympatric morphs in a single species may have significant evolutionary consequences. In the present study, phenotypic impacts of predator on zooplankton prey populations were compared in two northern Finnish lakes; one with an allopatric whitefish, Coregonus lavaretus (L.), population and the other with three sympatric whitefish populations. First, we examined whether there were phenotypic associations with specific niches in allopatric and sympatric whitefish. Second, trait utility (i.e. number of gillrakers) of allopatric and sympatric whitefish in utilizing a pelagic resource was explored by comparing predator avoidance of prey, prey size in environment, and prey size in predator diet. The allopatric living large sparsely rakered (LSR) whitefish morph, was a generalist using both pelagic and benthic niches. In contrast, sympatric living whitefish morphs were specialized: LSR whitefish was a littoral benthivore, small sparsely rakered whitefish was a profundal benthivore and densely rakered (DR) whitefish was a pelagic planktivore. In the lake with allopatric whitefish, zooplankton prey did not migrate vertically to avoid predation whereas, in the lake with sympatric whitefish, all important prey taxa migrated significantly. Trait utility was observed as significantly smaller size of prey in environment and predator diet in the lake with DR whitefish than in the lake with only LSR whitefish.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 561–572.