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  • 1.
    Sunde, Johanna
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Adaptations of early development to local spawning temperature in anadromous populations of pike (Esox lucius)2019Ingår i: BMC Evolutionary Biology, ISSN 1471-2148, E-ISSN 1471-2148, Vol. 19, s. 1-13, artikel-id 148Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Background: In the wake of climate change many environments will be exposed to increased and more variable temperatures. Knowledge about how species and populations respond to altered temperature regimes is therefore important to improve projections of how ecosystems will be affected by global warming, and to aid management. We conducted a common garden, split-brood temperature gradient (4.5 degrees C, 9.7 degrees C and 12.3 degrees C) experiment to study the effects of temperature in two populations (10 families from each population) of anadromous pike (Esox lucius) that normally experience different temperatures during spawning. Four offspring performance measures (hatching success, day degrees until hatching, fry survival, and fry body length) were compared between populations and among families. Results: Temperature affected all performance measures in a population-specific manner. Low temperature had a positive effect on the Harfjarden population and a negative effect on the Lervik population. Further, the effects of temperature differed among families within populations. Conclusions: The population-specific responses to temperature indicate genetic differentiation in developmental plasticity between populations, and may reflect an adaptation to low temperature during early fry development in Harfjarden, where the stream leading up to the wetland dries out relatively early in the spring, forcing individuals to spawn early. The family-specific responses to temperature treatment indicate presence of genetic variation for developmental plasticity (G x E) within both populations. Protecting between- and within-population genetic variation for developmental plasticity and high temperature-related adaptive potential of early life history traits will be key to long-term viability and persistence in the face of continued climate change.

  • 2.
    Tamario, Carl
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Sunde, Johanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Petersson, Erik
    Swedish University of Agricultural Sciences, Sweden.
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Ecological and Evolutionary Consequences of Environmental Change and Management Actions for Migrating Fish2019Ingår i: Frontiers in Ecology and Evolution, E-ISSN 2296-701X, Vol. 7, s. 1-24, artikel-id 271Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    Migration strategies in fishes comprise a rich, ecologically important, and socioeconomically valuable example of biological diversity. The variation and flexibility in migration is evident between and within individuals, populations, and species, and thereby provides a useful model system that continues to inform how ecological and evolutionary processes mold biodiversity and how biological systems respond to environmental heterogeneity and change. Migrating fishes are targeted by commercial and recreational fishing and impact the functioning of aquatic ecosystems. Sadly, many species of migrating fish are under increasing threat by exploitation, pollution, habitat destruction, dispersal barriers, overfishing, and ongoing climate change that brings modified, novel, more variable and extreme conditions and selection regimes. All this calls for protection, sustainable utilization and adaptive management. However, the situation for migrating fishes is complicated further by actions aimed at mitigating the devastating effects of such threats. Changes in river connectivity associated with removal of dispersal barriers such as dams and construction of fishways, together with compensatory breeding, and supplemental stocking can impact on gene flow and selection. How this in turn affects the dynamics, genetic structure, genetic diversity, evolutionary potential, and viability of spawning migrating fish populations remains largely unknown. In this narrative review we describe and discuss patterns, causes, and consequences of variation and flexibility in fish migration that are scientifically interesting and concern key issues within the framework of evolution and maintenance of biological diversity. We showcase how the evolutionary solutions to key questions that define migrating fish-whether or not to migrate, why to migrate, where to migrate, and when to migrate-may depend on individual characteristics and ecological conditions. We explore links between environmental change and migration strategies, and discuss whether and how threats associated with overexploitation, environmental makeovers, and management actions may differently influence vulnerability of individuals, populations, and species depending on the variation and flexibility of their migration strategies. Our goal is to provide a broad overview of knowledge in this emerging area, spur future research, and development of informed management, and ultimately promote sustainable utilization and protection of migrating fish and their ecosystems.

  • 3.
    Tamario, Carl
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Sunde, Johanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Petersson, Erik
    Swedish University of Agricultural Sciences, Sweden.
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Ecological and Evolutionary Consequences of Environmental Change and Management Actions for Migrating Fish2019Ingår i: Frontiers in Ecology and Evolution, E-ISSN 2296-701X, Vol. 7, s. 1-24, artikel-id 271Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    Migration strategies in fishes comprise a rich, ecologically important, and socioeconomically valuable example of biological diversity. The variation and flexibility in migration is evident between and within individuals, populations, and species, and thereby provides a useful model system that continues to inform how ecological and evolutionary processes mold biodiversity and how biological systems respond to environmental heterogeneity and change. Migrating fishes are targeted by commercial and recreational fishing and impact the functioning of aquatic ecosystems. Sadly, many species of migrating fish are under increasing threat by exploitation, pollution, habitat destruction, dispersal barriers, overfishing, and ongoing climate change that brings modified, novel, more variable and extreme conditions and selection regimes. All this calls for protection, sustainable utilization and adaptive management. However, the situation for migrating fishes is complicated further by actions aimed at mitigating the devastating effects of such threats. Changes in river connectivity associated with removal of dispersal barriers such as dams and construction of fishways, together with compensatory breeding, and supplemental stocking can impact on gene flow and selection. How this in turn affects the dynamics, genetic structure, genetic diversity, evolutionary potential, and viability of spawning migrating fish populations remains largely unknown. In this narrative review we describe and discuss patterns, causes, and consequences of variation and flexibility in fish migration that are scientifically interesting and concern key issues within the framework of evolution and maintenance of biological diversity. We showcase how the evolutionary solutions to key questions that define migrating fish-whether or not to migrate, why to migrate, where to migrate, and when to migrate-may depend on individual characteristics and ecological conditions. We explore links between environmental change and migration strategies, and discuss whether and how threats associated with overexploitation, environmental makeovers, and management actions may differently influence vulnerability of individuals, populations, and species depending on the variation and flexibility of their migration strategies. Our goal is to provide a broad overview of knowledge in this emerging area, spur future research, and development of informed management, and ultimately promote sustainable utilization and protection of migrating fish and their ecosystems.

  • 4.
    Nordahl, Oscar
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Koch-Schmidt, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Sunde, Johanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Yildirim, Yeserin
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Genetic differentiation between and within ecotypes of pike (Esox lucius) in the Baltic Sea2019Ingår i: Aquatic conservation, ISSN 1052-7613, E-ISSN 1099-0755Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Aquatic systems often lack physical boundaries for gene flow, but ecological and behavioural barriers can form surprisingly fine spatial scale genetic patterns that challenge traditional, large scale management. To detect fine spatial scale structures, understand sources of intraspecific diversity, and design appropriate management plans requires identification of reproductively isolated units. This study reports on genetic differentiation in pike (Esox lucius) within a coastal area stretching 55 km from south to north in the central Baltic Sea. Pike is here an economically and ecologically important top predator that has declined in abundance. However, population structures have mostly been studied on large spatial scales, and without considering the potential for genetic divergence between the sympatric anadromous fresh water and the resident brackish water spawning ecotypes. To this end, 487 individuals from the east coast of Sweden and the island of oland, representing sympatric anadromous and resident spawning individuals, categorized to ecotype based on spawning location or otolith microchemistry, were genotyped for 10 microsatellites and used to test for divergence between ecotypes. Furthermore, divergence between regions (island/mainland), neighbouring spawning locations (n = 13) and isolation by distance within and between regions were evaluated for the anadromous ecotype. The results revealed strong genetic differences between regions, between spawning locations separated by as little as 5 km and the first evidence of genetic differentiation between resident and anadromous ecotypes; despite a high dispersal capacity of pike and a high connectivity within the study area. The signatures of isolation by distance indicated that connectivity among populations differed between regions, probably reflecting availability of spawning habitats. To safeguard against the challenges and uncertainties associated with environmental change, adaptive conservation management should aim to promote high intra-population functional genetic diversity without compromising the continued integrity and coexistence of the different ecotypes and of locally adapted sub-populations.

  • 5.
    Franzén, Markus
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Betzholtz, Per-Eric
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Variable color patterns influence continental range size and species-area relationships on islands2019Ingår i: Ecosphere, ISSN 2150-8925, E-ISSN 2150-8925, Vol. 10, nr 1, artikel-id e02577Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    It has been hypothesized that species with more variable color patterns should have higher establishment success and be less sensitive to environmental changes and local extinction compared with species that do not vary in color. This difference in colonization/extinction balance should manifest as larger continental range distributions and modulate the species-area relationship on true islands. We evaluated these predictions using data for 1216 species of butterflies and moths that differed with regard to inter-individual variation in color pattern. We show that species with more variable color patterns have larger continental range sizes in Europe compared with non-variable species. We also provide rare evidence that the slope of the species-area relationship on islands is steeper for species having non-variable color patterns, suggesting that to preserve 60% of non-variable species would require an area twice as large compared to what would be needed to preserve 60% of variable species. Our findings suggest that combining information on ecological characteristics with presence/absence data from small and medium sized islands can help identify traits that drive species range patterns at the continental scale, and that individual variation in color pattern can be used as a proxy for ecological generalization and the ability to cope with environmental change.

  • 6.
    Zverev, Vitali
    et al.
    University of Turku, Finland.
    Kozlov, Mikhail
    University of Turku, Finland.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Zvereva, Elena
    University fo Turku, Finland.
    Ambient temperatures differently influence colour morphs of the leaf beetle Chrysomela lapponica: roles of thermal melanism and developmental plasticity2018Ingår i: Journal of Thermal Biology, ISSN 0306-4565, E-ISSN 1879-0992, Vol. 74, s. 100-109Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    We asked whether ambient temperatures can affect morph frequencies within a subarctic population of thepolymorphic leaf beetle Chrysomela lapponica through thermal melanism and/or developmental plasticity. Bodytemperature increased faster in beetles of dark morph than in beetles of light morph under exposure to artificialirradiation. Dark males ran faster than light males in both field and laboratory experiments, and this differencedecreased with increasing ambient air temperature, from significant at 10 °C to non-significant at 20 °C and26 °C. On cold days (6–14 °C), significantly more dark males than light males were found on their host plants incopula (40.8% and 27.3% respectively); on warm days (15–22 °C) this difference disappeared. Light femalesproduced twice as many eggs as dark females; this difference did not depend on the ambient temperature. Theproportion of dark morphs in the progenies of pairs with one dark parent was twice as high as that in theprogenies of pairs in which both parents were light, and this proportion was greater when larvae developed atlow (10 and 15 °C) than at high (20 and 25 °C) temperatures. We conclude that low temperatures may increasethe frequencies of dark morphs in C. lapponica populations due to both the mating advantages of dark males overlight males and developmental plasticity. Variation in frequencies of low-fecund dark morphs in the population,caused by among-year differences in temperature together with density-dependent selection, may contribute tothe evolutionary dynamics of the colour polymorphism and may influence abundance fluctuations in these leafbeetle populations.

  • 7.
    Yildirim, Yeserin
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Tinnert, Jon
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Contrasting patterns of neutral and functional genetic diversity in stable and disturbed environments2018Ingår i: Ecology and Evolution, ISSN 2045-7758, E-ISSN 2045-7758, Vol. 8, nr 23, s. 12073-12089Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Genetic structure among and diversity within natural populations is influenced by acombination of ecological and evolutionary processes. These processes can differentlyinfluence neutral and functional genetic diversity and also vary according toenvironmental settings. To investigate the roles of interacting processes as drivers ofpopulation‐level genetic diversity in the wild, we compared neutral and functionalstructure and diversity between 20 Tetrix undulata pygmy grasshopper populations indisturbed and stable habitats. Genetic differentiation was evident among the differentpopulations, but there was no genetic separation between stable and disturbedenvironments. The incidence of long‐winged phenotypes was higher in disturbedhabitats, indicating that these populations were recently established by flight‐capablecolonizers. Color morph diversity and dispersion of outlier genetic diversity, estimatedusing AFLP markers, were higher in disturbed than in stable environments,likely reflecting that color polymorphism and variation in other functionally importanttraits increase establishment success. Neutral genetic diversity estimated usingAFLP markers was lower in disturbed habitats, indicating stronger eroding effects onneutral diversity of genetic drift associated with founding events in disturbed comparedto stable habitats. Functional diversity and neutral diversity were negativelycorrelated across populations, highlighting the utility of outlier loci in genetics studiesand reinforcing that estimates of genetic diversity based on neutral markers donot infer evolutionary potential and the ability of populations and species to copewith environmental change.

  • 8.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    On the role of sex differences for evolution in heterogeneous and changing fitness landscapes: insights from pygmy grasshoppers2018Ingår i: Philosophical Transactions of the Royal Society of London. Biological Sciences, ISSN 0962-8436, E-ISSN 1471-2970, Vol. 373, nr 1757, artikel-id 20170429Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    Much research has been devoted to study evolution of local adaptations by natural selection, and to explore the roles of neutral processes and developmental plasticity for patterns of diversity among individuals, populations and species. Some aspects, such as evolution of adaptive variation in phenotypic traits in stable environments, and the role of plasticity in predictable changing environments, are well understood. Other aspects, such as the role of sex differences for evolution in spatially heterogeneous and temporally changing environments and dynamic fitness landscapes, remain elusive. An increased understanding of evolution requires that sex differences in development, physiology, morphology, life-history and behaviours are more broadly considered. Studies of selection should take into consideration that the relationships linking phenotypes to fitness may vary not only according to environmental conditions but also differ between males and females. Such opposing selection, sex-by-environment interaction effects of selection and sex-specific developmental plasticity can have consequences for population differentiation, local adaptations and for the dynamics of polymorphisms. Integrating sex differences in analytical frameworks and population comparisons can therefore illuminate neglected evolutionary drivers and reconcile unexpected patterns. Here, I illustrate these issues using empirical examples from over 20 years of research on colour polymorphic Tetrix subulata and Tetrix undulata pygmy grasshoppers, and summarize findings from observational field studies, manipulation experiments, common garden breeding experiments and population genetics studies. This article is part of the theme issue 'Linking local adaptation with the evolution of sex differences'.

  • 9.
    Sunde, Johanna
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Sex-specific effects of outbreeding on offspring quality in pike (Esox lucius)2018Ingår i: Ecology and Evolution, ISSN 2045-7758, E-ISSN 2045-7758, Vol. 8, nr 21, s. 10448-10459Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Intraspecific genetic admixture occurs when previously separated populations withina species start interbreeding, and it can have either positive, negative, or neutral effectson reproductive performance. As there currently is no reliable predictor for theoutcome of admixture, an increased knowledge about admixture effects in differentspecies and populations is important to increase the understanding about what determinesthe response to admixture. We tested for effects of admixture on F1 offspringquality in three subpopulations of pike (Esox lucius). Gametes were collected inthe field, and eggs from each female were experimentally fertilized with milt from amale from each population (one “pure” and two “admixed” treatments). Three offspringquality measures (hatching success, fry survival, and fry length) were determinedand compared between (a) pure and admixed population combinations and (b)the sex-specifictreatments within each admixed population combination (based onthe origin of the male and female, respectively). The results suggested that althoughthere were no overall effects of admixture on offspring quality, the consequences fora given population combination could be sex-specificand thus differ depending onwhich of the parents originated from one or the other population. All offspring qualitytraits were influenced by both maternal ID and paternal ID. Sex-andindividual-specificeffects can have implications for dispersal behavior and gene flow betweennatural populations, and are important to consider in conservation efforts.

  • 10.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Merilaita, Sami
    Åbo Akad Univ, Finland;Univ Turku, Finland.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Size variability effects on visual detection are influenced by colour pattern and perceived size2018Ingår i: Animal Behaviour, ISSN 0003-3472, E-ISSN 1095-8282, Vol. 143, s. 131-138Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Most animals including humans use vision to detect, identify, evaluate and respond to potential prey items in complex environments. Theories predict that predators' visual search performance is better when targets are similar than when targets are dissimilar and require divided attention, and this may contribute to colour pattern polymorphism in prey. Most prey also vary in size, but how size variation influences detectability and search performance of predators that utilize polymorphic prey has received little attention. To evaluate the effect of size variability on prey detection we asked human subjects to search for images of black, grey and striped pygmy grasshoppers presented on computer screens in size-variable (large, medium and small) or in size-invariable (all medium) sequences (populations) against photographs of natural grasshopper habitat. Results showed that size variability either increased or reduced detection of medium-sized targets depending on colour morph. To evaluate whether bias in perceived size varies depending on colour pattern, subjects were asked to discriminate between two grasshopper images of identical size that were presented in pairs against a monochromatic background. Subjects more often incorrectly classified one of the two identical-sized targets as being larger than the other in colour-dimorphic than in monomorphic presentations. The distinctly patterned (striped) morph elicited stronger size perception biases than the dorsally grey or black morphs, and striped grasshoppers were incorrectly classified more often as smaller than grey grasshoppers. The direction of the effect of size variability on detection changed across colour patterns as the bias in perceived size increased. Such joint effects of variation in size and colour pattern on detection and perception can impact the outcome of behavioural and evolutionary interactions between visually oriented predators and their camouflaged prey. This may have consequences for population dynamics, evolution of polymorphisms, community species composition and ecosystem functioning. (C) 2018 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

  • 11.
    Nordahl, Oscar
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Koch-Schmidt, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Berggren, Hanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Sun-basking fish benefit from body temperatures that are higher than ambient water2018Ingår i: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 285, nr 1879, artikel-id 20180639Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    In terrestrial environments, cold-blooded animals can attain higher bodytemperatures by sun basking, and thereby potentially benefit from broaderniches, improved performance and higher fitness. The higher heat capacityand thermal conductivity of water compared with air have been universallyassumed to render heat gain from sun basking impossible for aquaticectotherms, such that their opportunities to behaviourally regulate body temperatureare largely limited to choosing warmer or colder habitats. Here wechallenge this paradigm. Using physical modelswe first showthat submergedobjects exposed to natural sunlight attain temperatures in excess of ambientwater. We next demonstrate that free-ranging carp (Cyprinus carpio) canincrease their body temperature during aquatic sun basking close to thesurface. The temperature excess gained by basking was larger in dark thanin pale individuals, increased with behavioural boldness, and was associatedwith faster growth. Overall, our results establish aquatic sun basking as a novelecologically significant mechanism for thermoregulation in fish. The discoveryof this previously overlooked process has practical implications for aquaculture,offers alternative explanations for behavioural and phenotypicadaptations, will spur future research in fish ecology, and calls for modificationsof models concerning climate change impacts on biodiversity inmarine and freshwater environments.

  • 12.
    Sunde, Johanna
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Tamario, Carl
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM). Swedish University of Agricultural Sciences.
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Variation in salinity tolerance between and within anadromous subpopulations of pike (Esox lucius)2018Ingår i: Scientific Reports, ISSN 2045-2322, E-ISSN 2045-2322, Vol. 8, artikel-id 22Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Environmental heterogeneity is a key determinant of genetic and phenotypic diversity. Stable andhomogenous environments tends to result in evolution of specialism and local adaptations, whiletemporally unpredictable environments may maintain a diversity of specialists, promote generaliststrategies, or favour diversified bet hedging strategies. We compared salinity tolerance between twoanadromous subpopulations of pike (Esox Lucius) that utilize freshwater spawning sites with differentsalinity regimes. Eggs from each population were artificially fertilized and incubated in a salinitygradient (0, 3, 5, 7, and 9 psu) using a split-brood design. Effects on embryonic development, hatchingsuccess, survival of larvae, and fry body length were compared between populations and families.The population naturally spawning in the stable freshwater habitat showed signs of specialization forfreshwater spawning. The population exposed to fluctuating selective pressure in a spawning area withoccasional brackish water intrusions tolerated higher salinities and displayed considerable variation inreaction norms. Genetic differences and plasticity of salinity tolerance may enable populations to copewith changes in salinity regimes associated with future climate change. That geographically adjacentsubpopulations can constitute separate units with different genetic characteristics must be consideredin management and conservation efforts to avoid potentially negative effects of genetic admixture onpopulation fitness and persistence.

  • 13.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Berggren, Hanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Can spatial sorting associated with spawning migration explain evolution of body size and vertebral number in Anguilla eels?2017Ingår i: Ecology and Evolution, ISSN 2045-7758, E-ISSN 2045-7758, Vol. 7, nr 2, s. 751-761Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Spatial sorting is a process that can contribute to microevolutionary change by assemblingphenotypes through space, owing to nonrandom dispersal. Here we first buildupon and develop the “neutral” version of the spatial sorting hypothesis by arguingthat in systems that are not characterized by repeated range expansions, the evolutionaryeffects of variation in dispersal capacity and assortative mating might not beindependent of but interact with natural selection. In addition to generating assortativemating, variation in dispersal capacity together with spatial and temporal variationin quality of spawning area is likely to influence both reproductive success and survivalof spawning migrating individuals, and this will contribute to the evolution of dispersal-enhancingtraits. Next, we use a comparative approach to examine whether differencesin spawning migration distance among 18 species of freshwater Anguilla eelshave evolved in tandem with two dispersal-favoringtraits. In our analyses, we use informationon spawning migration distance, body length, and vertebral number thatwas obtained from the literature, and a published whole mitochondrial DNA-basedphylogeny. Results from comparative analysis of independent contrasts showed thatmacroevolutionary shifts in body length throughout the phylogeny have been associatedwith concomitant shifts in spawning migration. Shifts in migration distance werenot associated with shifts in number of vertebrae. These findings are consistent withthe hypothesis that spatial sorting has contributed to the evolution of more elongatedbodies in species with longer spawning migration distances, or resulted in evolution oflonger migration distances in species with larger body size. This novel demonstrationis important in that it expands the list of ecological settings and hierarchical levels ofbiological organization for which the spatial sorting hypothesis seems to have predictivepower.

  • 14.
    Betzholtz, Per-Eric
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Franzén, Markus
    Helmholtz Centre for Environmental Research, Germany.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Colour pattern variation can inform about extinction risk in moths2017Ingår i: Animal Conservation, ISSN 1367-9430, E-ISSN 1469-1795, Vol. 20, nr 1, s. 72-79Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Theory posits that species with inter-individual variation in colour patterns should beless vulnerable to extinction, compared with species that do not vary in colour. Toevaluate this prediction, we explored whether differences in colour pattern diversitywas associated with extinction risk, using red-list status for more than 350 species ofnoctuid moths in Sweden. We also evaluated six other species characteristics thathave been proposed to influence extinction risk namely: host plant niche breadth,habitat type, area of occupancy, body size, overwintering life-history stage and lengthof flight activity period. We found that species with variable colour patterns hadreduced extinction risk overall compared with species having non-variable colourpatterns, and that this difference was pronounced more strongly among species havingsmaller areas of occupancy. There were also significant associations with hostplant niche breadth and habitat type, extinction risk being lower on average in polyphagousspecies and in generalist species that occupied different habitat types. Thesefindings represent the first evidence for insects that variable colouration is associatedwith reduced extinction risks. Information on colour pattern variation is readily availablefor many taxa and may be used as a cost-effective proxy for endangerment inthe work of halting national and global biodiversity loss.

  • 15.
    Tinnert, Jon
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    The role of dispersal for genetic and phenotypic variation: insights from comparisons of sympatric pygmy grasshoppers2017Ingår i: Biological Journal of the Linnean Society, ISSN 0024-4066, E-ISSN 1095-8312, Vol. 122, nr 1, s. 84-97Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Patterns of genetic and phenotypic variation within and among populations are influenced by a complex interplay of ecological and evolutionary processes. Theory posits that gene flow should increase diversity within and reduce differentiation between populations. Evaluating these predictions is potentially complicated by selection, population dynamics and plasticity that may also affect genetic and phenotypic variation. Here, we compare genetic and morphological variation between sympatric populations of two pygmy grasshopper species, Tetrix subulata and T. undulata, that differ in dispersal capacity. We found that genetic differentiation between populations is lower on average in the generally dispersive T. subulata compared with the mostly sedentary T. undulata, suggesting that genetic structure in the latter species has been less influenced by the homogenizing effects of migration. Our results also provided weak support for the hypothesis that neutral genetic diversity within populations should be higher in T. subulata than in T. undulata. We further found that body size varied among populations in both species, but the differences seen in T. subulata did not parallel those seen in T. undulata, indicating that the two species have unique plasticity responses or that they have responded differently to shared selective regimes. Our findings illustrate the utility of the pairwise comparative approach and further highlight that results and conclusions may not be transferrable even between closely related species.

  • 16.
    Tibblin, Petter
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Berggren, Hanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Nordahl, Oscar
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Causes and consequences of intra-specific variation in vertebral number2016Ingår i: Scientific Reports, ISSN 2045-2322, E-ISSN 2045-2322, Vol. 6, artikel-id 26372Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Intraspecific variation in vertebral number is taxonomically widespread. Much scientific attention hasbeen directed towards understanding patterns of variation in vertebral number among individualsand between populations, particularly across large spatial scales and in structured environments.However, the relative role of genes, plasticity, selection, and drift as drivers of individual variation andpopulation differentiation remains unknown for most systems. Here, we report on patterns, causesand consequences of variation in vertebral number among and within sympatric subpopulations ofpike (Esox lucius). Vertebral number differed among subpopulations, and common garden experimentsindicated that this reflected genetic differences. A QST-FST comparison suggested that populationdifferences represented local adaptations driven by divergent selection. Associations with fitness traitsfurther indicated that vertebral counts were influenced both by stabilizing and directional selectionwithin populations. Overall, our study enhances the understanding of adaptive variation, which iscritical for the maintenance of intraspecific diversity and species conservation.

  • 17.
    Tibblin, Petter
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Borger, Tobias
    Kalmar County Council.
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Causes and consequences of repeatability, flexibility and individual fine tuning of of migratory timing in pike2016Ingår i: Journal of Animal Ecology, ISSN 0021-8790, E-ISSN 1365-2656, Vol. 85, nr 1, s. 136-145Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    1. Many organisms undertake migrations between foraging and breeding habitats and while it is assumed that reproductive timing affects fitness, little is known about the degree of individual consistency, and about the causes and consequences of individual variation in migratory timing in organisms other than birds. 2. Here, we report on a 6-year mark-recapture study, including 2048 individuals, of breeding migration in anadromous pike (Esox lucius), an iteroparous top-predatory fish that displays homing behaviour. By repeated sampling across years at a breeding site, we first quantify individual variation both within and between breeding events and then investigate phenotypic correlates and fitness consequences of arrival timing to the breeding site. 3. Our data demonstrate that males arrive before females, that large males arrive later than small males, that the timing of breeding migration varies among years and that individuals are consistent in their timing across years relative to other individuals in the population. 4. Furthermore, data on return rates indicate that arrival time is under stabilizing viability selection, and that individuals who are more flexible in their timing of arrival during the first reproductive years survive longer compared with less flexible individuals. Finally, longitudinal data demonstrate that individuals consistently fine-tune their arrival timing across years, showing that the timing of arrival to breeding sites is influenced by experience. 5. These findings represent rare evidence of how between-and within-individual variations in migratory timing across breeding events are correlated with phenotypic and fitness traits in an ecologically important keystone species. Our results emphasize the importance of considering variation in migratory timing both between and within individuals in studies investigating the fitness consequences of migratory behaviour and have implications for future management.

  • 18.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Merilaita, Sami
    Åbo Akademi University, Finland.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Colour polymorphism protects prey individuals and populations against predation2016Ingår i: Scientific Reports, ISSN 2045-2322, E-ISSN 2045-2322, Vol. 6, artikel-id 22122Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Colour pattern polymorphism in animals can influence and be influenced by interactions between predators and prey. However, few studies have examined whether polymorphism is adaptive, and there is no evidence that the co-occurrence of two or more natural prey colour variants can increase survival of populations. Here we show that visual predators that exploit polymorphic prey suffer from reduced performance, and further provide rare evidence in support of the hypothesis that prey colour polymorphism may afford protection against predators for both individuals and populations. This protective effect provides a probable explanation for the longstanding, evolutionary puzzle of the existence of colour polymorphisms. We also propose that this protective effect can provide an adaptive explanation for search image formation in predators rather than search image formation explaining polymorphism.

  • 19.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Betzholtz, Per-Eric
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Franzén, Markus
    Helmholtz Centre for Environmental Research, Germany.
    Faster poleward range shifts in moths with more variable colour patterns2016Ingår i: Scientific Reports, ISSN 2045-2322, E-ISSN 2045-2322, Vol. 6, artikel-id 36265Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Range shifts have been documented in many organisms, and climate change has been implicated asa contributing driver of latitudinal and altitudinal range modifications. However, little is known aboutwhat species trait(s) allow for faster environmental tracking and improved capacity for distributionexpansions. We used data for 416 species of moths, and show that range limits in Sweden have shifted tothe north by on average 52.4 km per decade between 1973 and 2014. When also including non-expandingspecies, average expansion rate was 23.2 km per decade. The rate of boundary shifts increased withincreasing levels of inter-individual variation in colour patterns and decreased with increasing latitude. Theassociation with colour patterns indicate that variation in this functionally important trait enables speciesto cope with novel and changing conditions. Northern range limits also increased with average abundanceand decreased with increasing year-to-year abundance fluctuations, implicating production of dispersersas a driver of range dynamics. Studies of terrestrial animals show that rates of poleward shifts differbetween taxonomic groups, increase over time, and depend on study duration and latitude. Knowledge ofhow distribution shifts change with time, location, and species characteristics may improve projections ofresponses to climate change and aid the protection of biodiversity

  • 20.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Wennersten, Lena
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Inter-individual variation promotes ecological success of populations and species: evidence from experimental and comparative studies2016Ingår i: Ecography, ISSN 0906-7590, E-ISSN 1600-0587, Vol. 39, nr 7, s. 630-648Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    Biological diversity is threatened by exploitation, fragmentation of natural habitats, pollution, climatechange, and anthropogenic spread of species. The question of how among-individual variation influencesthe performance of populations and species is a poorly explored but currently growing field of research.Here, we review 31 experimental and 14 comparative studies and first investigate whether there is empiricalsupport for the propositions that higher levels of among-individual phenotypic and genetic variationpromote the ecological and evolutionary success of populations and species in the face of environmentalchange. Next, we examine whether and how the effect of diversity depends on environmental conditions.Finally, we explore whether the relationship linking population fitness to diversity is typically linear,asymptotic, or whether the benefits peak at intermediate diversity. The reviewed studies provide strong,almost invariable, evidence that more variable populations are less vulnerable to environmental changes,show decreased fluctuations in population size, have superior establishment success, larger distributionranges, and are less extinction prone, compared with less variable populations or species. Given theoverwhelming evidence that variation promotes population performance, it is important to identifyconditions when increased variation does not have the theoretically expected effect, a question ofconsiderable importance in biodiversity management, where there are many other practical constraints. Wefind that experimental outcomes generally support the notion that genetic and phenotypic variation is ofgreater importance under more stressful than under benign conditions. Finally, population performanceincreased linearly with increasing diversity in the majority (10 of 12) of manipulation studies that includedfour or more diversity levels; only two experiments detected curvilinear relationships.

  • 21.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Is colour polymorphism advantageous to populations and species?2016Ingår i: Molecular Ecology, ISSN 0962-1083, E-ISSN 1365-294X, Vol. 25, nr 12, s. 2693-2698Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    I am writing in response to an article by Bolton, Rollinsand Griffith (2015) entitled ‘The danger within: the roleof genetic, behavioural and ecological factors in populationpersistence of colour polymorphic species’ that wasrecently published as an Opinion under the NEWS ANDVIEWS section in Molecular Ecology. Bolton et al.(Molecular Ecology, 2015, 24, 2907) argue that colour polymorphismmay reduce population fitness and increaseextinction risk and emphasize that this is contrary to predictionsput forward by Forsman et al. (Ecology, 89, 2008,34) and Wennersten & Forsman (Biological Reviews 87,2012, 756) that the existence of multiple colour morphswith co-adapted gene complexes and associated trait valuesmay increase the ecological and evolutionary successof polymorphic populations and species. Bolton et al.(Molecular Ecology, 2015, 24, 2907) further state that thereis no clear evidence from studies of ‘true polymorphicspecies’ that polymorphism promotes population persistence.In response, I (i) challenge their classifications ofpolymorphisms and revisit the traditional definitions recognizingthe dynamic nature of polymorphisms, (ii)review empirical studies that have examined whetherand how polymorphism is associated with extinction risk,(iii) discuss the roles of trait correlations between colourpattern and other phenotypic dimensions for populationfitness and (iv) highlight that the causes and mechanismsthat influence the composition and maintenance of polymorphismsare different from the consequences of thepolymorphic condition and how it may impact on aspectsof ecological success and long-term persistence of populationsand species.

  • 22.
    Sunde, Johanna
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Mate choice is not consistent with short-term effects of intraspecific admixture in the common rough woodlouse (Porcellio scaber)2016Ingår i: Biological Journal of the Linnean Society, ISSN 0024-4066, E-ISSN 1095-8312, Vol. 119, nr 2, s. 359-369Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Theory posits that individuals should exhibit mate preferences in part based on genetic relatedness such that fitness is maximized. Intraspecific genetic admixture can have different effects depending on the genetic characteristics and evolutionary history of the individuals and populations involved. We investigated whether female mate choice behavior in the common rough woodlouse (Porcellio scaber) matched the fitness consequences of genetic admixture. We found that most females from two populations that were in sequence introduced to one male from each population mated with both males, and further that monandrous females (females that only mated with one male) predominantly mated with males from their own population. To test for effects of genetic admixture, females from four populations were divided into two replicate pairs and assigned to mate either with a male from the same population as the female (pure) or with a male from the other population (admixed). The effect of mating treatment on the proportion of females that produced eggs and hatched young, as well as on the number and viability of offspring depended on female source population. Mating treatment had opposing effects in two of the populations, whereas there were no detectable effects in the other two populations. Contrary to what was expected, the mating patterns did not match the observed effects of genetic admixture. We discuss alternative adaptive and non-adaptive explanations for the observed patterns.

  • 23.
    Tinnert, Jon
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Hellgren, Olof
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM). Lund University.
    Lindberg, Jenny
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM). Naturbruksskolan Sotasen.
    Koch-Schmidt, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Population genetic structure, differentiation, and diversity in Tetrix subulata pygmy grasshoppers: roles of population size and immigration2016Ingår i: Ecology and Evolution, ISSN 2045-7758, E-ISSN 2045-7758, Vol. 6, nr 21, s. 7831-7846Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Genetic diversity within and among populations and species is influenced by complex demographic and evolutionary processes. Despite extensive research, there is no consensus regarding how landscape structure, spatial distribution, gene flow, and population dynamics impact genetic composition of natural populations. Here, we used amplified fragment length polymorphisms (AFLPs) to investigate effects of population size, geographic isolation, immigration, and gene flow on genetic structure, divergence, and diversity in populations of Tetrix subulata pygmy grasshoppers (Orthoptera: Tetrigidae) from 20 sampling locations in southern Sweden. Analyses of 1564 AFLP markers revealed low to moderate levels of genetic diversity (PPL=59.5-90.1; Hj=0.23-0.32) within and significant divergence among sampling localities. This suggests that evolution of functional traits in response to divergent selection is possible and that gene flow is restricted. Genetic diversity increased with population size and with increasing proportion of long-winged phenotypes (a proxy of recent immigration) across populations on the island of oland, but not on the mainland. Our data further suggested that the open water separating oland from the mainland acts as a dispersal barrier that restricts migration and leads to genetic divergence among regions. Isolation by distance was evident for short interpopulation distances on the mainland, but gradually disappeared as populations separated by longer distances were included. Results illustrate that integrating ecological and molecular data is key to identifying drivers of population genetic structure in natural populations. Our findings also underscore the importance of landscape structure and spatial sampling scheme for conclusions regarding the role of gene flow and isolation by distance.

  • 24.
    Tinnert, Jon
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Berggren, Hanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Population-specific effects of interbreeding and admixture on reproductive decisions and offspring quality2016Ingår i: Annales Zoologici Fennici, ISSN 0003-455X, E-ISSN 1797-2450, Vol. 53, nr 1-2, s. 55-68Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    We investigated interbreeding and admixture in Tetrix subulata grasshoppers from two maternal origin populations that differed in life-history and dispersal traits. We compared reproductive output of females that had been experimentally mated with males from the same or from a different population. Interbreeding affected clutch size and number of clutches; in one population females in the admixed treatment produced smaller clutches, in the other population females in the admixed treatment produced more clutches. Behavioral observations indicated that individuals can discriminate scents emitted by individuals from different populations; such that females might adjust reproductive allocation depending on male origin. However, hatchability of eggs and survival of nymphs were not affected by the mating treatment. Admixture influenced the production of viable offspring in the F2 generation, but the effect was opposite in the two populations of maternal origin. Results suggested that responses to interbreeding and admixture can differ between populations within a species.

  • 25.
    Berggren, Hanna
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Nordahl, Oscar
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Testing for local adaptation to spawning habitat in sympatric subpopulations of northern pike by reciprocal translocation of embryos2016Ingår i: PLoS ONE, ISSN 1932-6203, E-ISSN 1932-6203, Vol. 11, nr 5, artikel-id e0154488Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    We tested for local adaption in early life-history traits by performing a reciprocal translocation experiment with approximately 2500 embryos of pike (Esox lucius) divided in paired split-family batches. The experiment indicated local adaptation in one of the two subpopulations manifested as enhanced hatching success of eggs in the native habitat, both when compared to siblings transferred to a non-native habitat, and when compared to immigrant genotypes from the other subpopulation. Gene-by-environment effects on viability of eggs and larvae were evident in both subpopulations, showing that there existed genetic variation allowing for evolutionary responses to divergent selection, and indicating a capacity for plastic responses to environmental change. Next, we tested for differences in female life-history traits. Results uncovered that females from one population invested more resources into reproduction and also produced more (but smaller) eggs in relation to their body size compared to females from the other population. We suggest that these females have adjusted their reproductive strategies as a counter-adaptation because a high amount of sedimentation on the eggs in that subpopulations spawning habitat might benefit smaller eggs. Collectively, our findings point to adaptive divergence among sympatric subpopulations that are physically separated only for a short period during reproduction and early development – which is rare. These results illustrate how combinations of translocation experiments and field studies of life-history traits might infer about local adaptation and evolutionary divergence among populations. Local adaptations in subdivided populations are important to consider in management and conservation of biodiversity, because they may otherwise be negatively affected by harvesting, supplementation, and reintroduction efforts targeted at endangered populations.

  • 26.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Berggren, Hanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Åström, Mats E.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    To what extent can existing research help project climate change impacts on biodiversity in aquatic environments?: A review of methodological approaches2016Ingår i: Journal of Marine Science and Engineering, E-ISSN 2077-1312, Vol. 4, nr 4, artikel-id 75Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    It is broadly accepted that continued global warming will pose a major threat to biodiversity in the 21st century. But how reliable are current projections regarding consequences of future climate change for biodiversity? To address this issue, we review the methodological approaches in published studies of how life in marine and freshwater environments responds to temperature shifts. We analyze and compare observational field surveys and experiments performed either in the laboratory or under natural conditions in the wild, the type of response variables considered, the number of species investigated, study duration, and the nature and magnitude of experimental temperature manipulations. The observed patterns indicate that, due to limitations of study design, ecological and evolutionary responses of individuals, populations, species, and ecosystems to temperature change were in many cases difficult to establish, and causal mechanism(s) often remained ambiguous. We also discovered that the thermal challenge in experimental studies was 10,000 times more severe than reconstructed estimates of past and projections of future warming of the oceans, and that temperature manipulations also tended to increase in magnitude in more recent studies. These findings raise some concerns regarding the extent to which existing research can increase our understanding of how higher temperatures associated with climate change will affect life in aquatic environments. In view of our review findings, we discuss the trade-off between realism and methodological tractability. We also propose a series of suggestions and directions towards developing a scientific agenda for improving the validity and inference space of future research efforts.

  • 27.
    Larsson, Per
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Koch-Schmidt, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Engstedt, Olof
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Nilsson, Jonas
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Nordahl, Oscar
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Ecology, evolution, and management strategies of northern pike populations in the Baltic Sea2015Ingår i: Ambio, ISSN 0044-7447, E-ISSN 1654-7209, Vol. 44, nr Supplement 3, s. S451-S461Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    Baltic Sea populations of the northern pike (Esox lucius) have declined since the 1990s, and they face additional challenges due to ongoing climate change. Pike in the Baltic Sea spawn either in coastal bays or in freshwater streams and wetlands. Pike recruited in freshwater have been found to make up about 50 % of coastal pike stocks and to show natal homing, thus limiting gene flow among closely located spawning sites. Due to natal homing, sub-populations appear to be locally adapted to their freshwater recruitment environments. Management actions should therefore not involve mixing of individuals originating from different sub-populations. We offer two suggestions complying with this advice: (i) productivity of extant freshwater spawning populations can be boosted by modifying wetlands such that they promote spawning and recruitment; and (ii) new sub-populations that spawn in brackish water can potentially be created by transferring fry and imprinting them on seemingly suitable spawning environments.

  • 28.
    Tibblin, Petter
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Koch-Schmidt, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Nordahl, Oscar
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Johannessen, Peter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Nilsson, Jonas
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Evolutionary divergence of adult body size and juvenile growth in sympatric subpopulations of a top predator in aquatic ecosystems2015Ingår i: American Naturalist, ISSN 0003-0147, E-ISSN 1537-5323, Vol. 186, nr 1, s. 98-110Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Evolutionary theory predicts that different selective regimes may contribute to divergent evolution of body size and growth rate among populations, but most studies have focused on allopatric populations. Here, we studied five sympatric subpopulations of anadromous northern pike (Esox lucius) in the Baltic Sea subjected to allopatric habitats for a short period of their life cycle due to homing behavior. We report differences in adult body size among subpopulations that were in part due to variation in growth rate. Body size of emigrating juveniles also differed among subpopulations, and differences remained when individuals were reared in a common environment, thus indicating evolutionary divergence among subpopulations. Furthermore, a QST-FST comparison indicated that differences had evolved due to divergent selection rather than genetic drift, possibly in response to differences in selective mortality among spawning habitats during the allopatric life stage. Adult and juvenile size were negatively correlated across subpopulations, and reconstruction of growth trajectories of adult fishes suggested that body size differences developed gradually and became accentuated throughout the first years of life. These results represent rare evidence that sympatric subpopulations can evolve differences in key life-history traits despite being subjected to allopatric habitats during only a very short fraction of their life.

  • 29.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Tibblin, Petter
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Berggren, Hanna
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Nordahl, Oscar
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Koch-Schmidt, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Larsson, Per
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Pike Esox lucius as an emerging model organism for studies in ecology and evolutionary biology: a review.2015Ingår i: Journal of Fish Biology, ISSN 0022-1112, E-ISSN 1095-8649, Vol. 87, nr 2, s. 472-479Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    The pikeEsox luciusis a large, long-lived, iteroparous, top- predator fish species with a circumpolardistribution that occupies a broad range of aquatic environments. This study reports on a literaturesearch and demonstrates that the publication rate ofE. luciusresearch increases both in absolute termsand relative to total scientific output, and that the focus of investigation has changed over time frombeing dominated by studies on physiology and disease to being gradually replaced by studies on ecol-ogy and evolution.Esox luciuscan be exploited as a model in future research for identifying causes andconsequences of phenotypic and genetic variation at the levels of individuals, populations and speciesas well as for investigating community processes.

  • 30.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Rethinking phenotypic plasticity and its consequences for individuals, populations and species.2015Ingår i: Heredity, ISSN 0018-067X, E-ISSN 1365-2540, Vol. 115, nr 4, s. 276-284Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    Much research has been devoted to identify the conditions under which selection favours flexible individuals or genotypes that are able to modify their growth, development and behaviour in response to environmental cues, to unravel the mechanisms of plasticity, and to explore its influence on patterns of diversity among individuals, populations, and species. The consequences of developmental plasticity and phenotypic flexibility for the performance and ecological success of populations and species have attracted a comparatively limited but currently growing interest. Here, I re-emphasize that an increased understanding of the roles of plasticity in these contexts requires a ‘whole organism’ (rather than ‘single trait’) approach, taking into consideration that organisms are integrated complex phenotypes. I further argue that plasticity and genetic polymorphism should be analysed and discussed within a common framework. I summarize predictions from theory on how phenotypic variation stemming from developmental plasticity and phenotypic flexibility may affect different aspects of population-level performance. I argue that it is important to distinguish between effects associated with greater inter-individual phenotypic variation resulting from plasticity, and effects mediated by variation among individuals in the capacity to express plasticity and flexibility as such. Finally, I claim that rigorous testing of predictions requires methods that allow for quantifying and comparing whole organism plasticity, as well as the ability to experimentally manipulate the level of and capacity for developmental plasticity and phenotypic flexibility independent of genetic variation.

  • 31.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Betzholtz, Per-Eric
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Franzén, Markus
    Helmholtz Centre for Environmental Research, Germany.
    Variable coloration is associated with dampened population fluctuations in noctuid moths2015Ingår i: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 282, nr 1808, s. 1-9, artikel-id 20142922Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Theory and recent reviews state that greater genetic and phenotypic variation should be beneficial for population abundance and stability. Experimental evaluations of this prediction are rare, of short duration and conducted under controlled environmental settings. The question whether greater diversity in functionally important traits stabilizes populations under more complex ecological conditions in the wild has not been systematically evaluated. Moths are mainly nocturnal, with a large variation in colour patterns among species, and constitute an important food source for many types of organisms. Here, we report the results of a long-term (2003-2013) monitoring study of 115 100 noctuid moths from 246 species. Analysis of time-series data provide rare evidence that species with higher levels of inter-individual variation in colour pattern have higher average abundances and undergo smaller between-year fluctuations compared with species having less variable colour patterns. The signature of interspecific temporal synchronization of abundance fluctuations was weak, suggesting that the dynamics were driven by species-specific biotic interactions rather than by some common, density-independent factor(s). We condude that individual variation in colour patterns dampens population abundance fluctuations, and suggest that this may partly reflect that colour pattern polymorphism provides protection from visually oriented predators and parasitoids.

  • 32.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Merilaita, Sami
    Åbo Akademi University, Finland.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Body size influences differently the detectabilities of colour morphs of cryptic prey2014Ingår i: Biological Journal of the Linnean Society, ISSN 0024-4066, E-ISSN 1095-8312, Vol. 113, nr 1, s. 112-122Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Body size and coloration may contribute to variation in performance and fitness among individuals, for instance by influencing vulnerability to predators. Yet, the combined effect of size and colour pattern on susceptibility to visual predators has received little attention, particularly in camouflaged prey. In the colour polymorphic pygmy grasshopper Tetrix subulata (Linnaeus, 1758) females are larger than males although there is a size overlap between sexes. We investigated how body size and colour morph influenced detection of these grasshoppers, and whether differences in protective value among morphs change with size. We conducted a computer-based experiment and compared how human ‘predators’ detected images of large, intermediate or small grasshoppers belonging to black, grey or striped colour morphs when embedded in photographs of natural grasshopper habitats. We found that time to detection increased with decreasing size, that differences in time to detection of the black, grey and striped morphs depended differently on body size, and that no single morph provided superior or inferior protection in all three size classes. By comparing morph frequencies in samples of male and female grasshoppers from natural populations we also examined whether the joint effects of size and colour morph on detection could explain evolutionary dynamics in the wild. Morph frequency differences between sexes were largely in accordance with expectations from the results of the detection experiment. Our results demonstrate that body size and colour morph can interactively influence detection of camouflaged prey. This may contribute to the morph frequency differences between male and female pygmy grasshoppers in the wild. Such interactive effects may also influence the dynamics of colour polymorphisms, and contribute to the evolution of ontogenetic colour change and sexual dichromatism.

  • 33.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Effects of genotypic and phenotypic variation on establishment are important for conservation, invasion and infection biology.2014Ingår i: Proceedings of the National Academy of Sciences of the United States of America, ISSN 0027-8424, E-ISSN 1091-6490, Vol. 111, nr 1, s. 302-307Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    There is abundant evidence that the probability of successful establishment in novel environments increases with number of individuals in founder groups and with number of repeated introductions. Theory posits that the genotypic and phenotypic variation among individuals should also be important, but few studies have examined whether founder diversity influences establishment independent of propagule pressure, nor whether the effect is model or context dependent. I summarize the results of 18 experimental studies and report on a metaanalysis that provides strong evidence that higher levels of genotypic and phenotypic diversity in founder groups increase establishment success in plants and animals. The effect of diversity is stronger in experiments carried out under natural conditions in the wild than under seminatural or standardized laboratory conditions. The realization that genetic and phenotypic variation is key to successful establishment may improve the outcome of reintroduction and translocation programs used to vitalize or restore declining and extinct populations. Founder diversity may also improve the ability of invasive species to establish and subsequently spread in environments outside of their native community, and enhance the ability of pathogens and parasites to colonize and invade the environment constituted by their hosts. It is argued that exchange of ideas, methodological approaches, and insights of the role of diversity for establishment in different contexts may further our knowledge, vitalize future research, and improve management plans in different disciplines.

  • 34.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Merilaita, Sami
    Åbo Akademi University, Finland.
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Natural levels of colour polymorphism reduce performance of visual predators searching for camouflaged prey2014Ingår i: Biological Journal of the Linnean Society, ISSN 0024-4066, E-ISSN 1095-8312, Vol. 112, nr 3, s. 546-555Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Polymorphism, the coexistence of two or more variants within a population, has served as a classic model system to address questions about the evolution and maintenance of intra-specific variation. It has been hypothesized that natural level of colour polymorphism may impair the search efficiency of visually oriented predators. To test this polymorphism protects hypothesis, we asked human participants to search for images of natural black, striped or grey Tetrix subulata grasshopper colour morphs presented against photographs of their natural habitat on computer screens. Fewer grasshoppers were detected when morphs were presented in mixed than in uniform sequences. All three morphs benefitted to comparable degrees, in terms of reduced detection, from being presented in polymorphic sequences. Our findings demonstrate that natural levels of polymorphic variation can impede the efficiency of visually oriented predators and increase survival of prey. This protective effect supports the limited attention hypothesis, explains why predators develop ‘search images’, may account for the spread and establishment of novel colour variants, and contribute to maintenance of polymorphisms.

  • 35.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Merilaita, Sami
    Department of Biosciences, Åbo Akademi University, Turku 780 FI-20520, Finland..
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Detection experiments with humans implicate visual predation as a driver of colour polymorphism dynamics in pygmy grasshoppers2013Ingår i: BMC Ecology, ISSN 1472-6785, E-ISSN 1472-6785, Vol. 13, s. Article number 17-Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Background: Animal colour patterns offer good model systems for studies of biodiversity and evolution of local adaptations. An increasingly popular approach to study the role of selection for camouflage for evolutionary trajectories of animal colour patterns is to present images of prey on paper or computer screens to human 'predators'. Yet, few attempts have been made to confirm that rates of detection by humans can predict patterns of selection and evolutionary modifications of prey colour patterns in nature. In this study, we first analyzed encounters between human 'predators' and images of natural black, grey and striped colour morphs of the polymorphic Tetrix subulata pygmy grasshoppers presented on background images of unburnt, intermediate or completely burnt natural habitats. Next, we compared detection rates with estimates of capture probabilities and survival of free-ranging grasshoppers, and with estimates of relative morph frequencies in natural populations. Results: The proportion of grasshoppers that were detected and time to detection depended on both the colour pattern of the prey and on the type of visual background. Grasshoppers were detected more often and faster on unburnt backgrounds than on 50% and 100% burnt backgrounds. Striped prey were detected less often than grey or black prey on unburnt backgrounds; grey prey were detected more often than black or striped prey on 50% burnt backgrounds; and black prey were detected less often than grey prey on 100% burnt backgrounds. Rates of detection mirrored previously reported rates of capture by humans of free-ranging grasshoppers, as well as morph specific survival in the wild. Rates of detection were also correlated with frequencies of striped, black and grey morphs in samples of T. subulata from natural populations that occupied the three habitat types used for the detection experiment. Conclusions: Our findings demonstrate that crypsis is background-dependent, and implicate visual predation as an important driver of evolutionary modifications of colour polymorphism in pygmy grasshoppers. Our study provides the clearest evidence to date that using humans as 'predators' in detection experiments may provide reliable information on the protective values of prey colour patterns and of natural selection and microevolution of camouflage in the wild.

  • 36.
    Johansson, Jenny
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Caesar, Sofia
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Multiple paternity increases phenotypic diversity in Tetrix subulata pygmy grasshoppers2013Ingår i: Journal of Orthoptera Research, ISSN 1082-6467, Vol. 22, nr 2, s. 79-85Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Multiple paternity within clutches has been recorded among a variety of organisms. The degree of genetic similarity between parents may influence the number and viability of offspring. Females may therefore mate with several males as an insurance against sterile, low quality or genetically incompatible mates, but also to obtain half sibling offspring that are genetically and phenotypically more diverse. We examine the links between polyandry, multiple paternity and offspring phenotypic diversity in the color polymorphic pygmy grasshopper Tetrix subulata. By experimentally mating virgin females and genotyping the resulting offspring using microsatellite markers, we demonstrate that polyandrous females can produce offspring sired by different males. Analyses of microsatellite data and color patterns of captive reared families produced by wild caught females that were not mated in the laboratory, confirmed that multiple paternity occurs in the wild, and that it may increase color morph diversity among half-siblings. Polyandrous mating behavior may thus influence the evolutionary dynamics and maintenance of color polymorphism in this species.

  • 37.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Forsman, Anders
    Linnéuniversitetet, Fakulteten för Hälso- och livsvetenskap (FHL), Institutionen för biologi och miljö (BOM).
    Stable isotopes reveal dietary divergence between dispersal phenotypes in Tetrix subulata pygmy grasshoppers (Orthoptera: Tetrigidae)2013Ingår i: European Journal of Entomology, ISSN 1210-5759, E-ISSN 1802-8829, Vol. 110, nr 1, s. 65-70Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    In some species of insects, individuals with fully developed wings and capable of flying coexist with flightless individuals that lack functional wings. Their diets may differ if long-winged individuals are more mobile and therefore likely to be better at finding and utilizing high quality food resources, or if they have different food preferences or physiological requirements. Despite its potential importance, differences in the diet of dispersal phenotypes have not been unequivocally demonstrated under natural conditions. To test for dietary divergence, we compared natural abundances of carbon and nitrogen stable isotope ratios (d13C and d15N) in long- and short-winged free ranging Tetrix subulata pygmy grasshoppers collected as adults from two natural populations. Overall, this comparison of stable isotopes indicated long-term differences in the diet of the two wing morphs in both populations, but not between males and females of the same morph. We conclude that it is likely that the dietary niches of the long winged and flightless individuals differ under natural conditions. This may reduce intra-specific competition, offset the expected trade-off between flight capacity and reproduction and promote ecological speciation.

  • 38.
    Wennersten, Lena
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Johansson, Jenny
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Karpestam, Einat
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Higher establishment success in more diverse groups of pygmy grasshoppers under seminatural conditions2012Ingår i: Ecology, ISSN 0012-9658, E-ISSN 1939-9170, Vol. 93, nr 12, s. 2519-2525Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Large founder groups and habitat match have been shown to increase the establishment success of reintroduced populations. Theory posits that the diversity of founder groups should also be important, but this has rarely been investigated. Here, experimental introductions of color-polymorphic Tetrix subulata pygmy grasshoppers into outdoor enclosures were used to test whether higher phenotypic diversity promotes establishment success. We show that the number of individuals present one year after introduction increases with color morph diversity in founder groups. Variance in establishment success did not decrease with increasing founder diversity, arguing against an important contribution of sampling effects or evolutionary rescue. Color morphs in T. subulata covary with a suite of other functionally important traits and utilize different resources. The higher establishment success in more diverse founder groups may therefore result, in part, from niche complementarity. Variation in establishment among groups was not associated with differences among source populations in reproductive capacities.

  • 39.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Wennersten, Lena
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Matching habitat choice by experimentally mismatched phenotypes2012Ingår i: Evolutionary Ecology, ISSN 0269-7653, E-ISSN 1573-8477, Vol. 26, nr 4, s. 893-907Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Gene flow is often regarded a random process that homogenizes differencesbetween populations and constrains local adaptation. However, the matching habitat choicehypothesis posits that individuals actively choose those microhabitats that best match theirspecific phenotype to maximize fitness. Dispersal (and possibly gene flow) may thus bedirected. Many studies report associations between habitats and phenotypes, but they mayreflect selection, plasticity or adaptation rather than matching choice. Here, we test twopredictions from the matching habitat choice hypothesis by manipulating the dorsal colourof Tetrix subulata, a pygmy grasshopper. (1) Is microhabitat choice flexible such thatdifferently manipulated phenotypes distribute themselves differently in a microclimaticand solar radiation mosaic? (2) If they do, are their fitness prospects higher in the morepreferred microhabitat? We find that individuals painted white or black do distributethemselves differently, with black individuals residing in habitats with less radiation, onaverage, than white individuals, demonstrating that microhabitat choices are plastic. Furthermore,white females had more hatchlings than black ones in the increased radiationtreatment, and this was mainly due to increased mortality of black females under increasedradiation. These findings provide rare experimental evidence in line with predictions fromthe matching habitat choice hypothesis.

  • 40.
    Wennersten, Lena
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Karpestam, Einat
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Phenotype manipulation influences microhabitat choice in pygmy grasshoppers2012Ingår i: Current Zoology, ISSN 1674-5507, Vol. 58, nr 3, s. 392-400Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    The matching habitat choice hypothesis posits that individuals actively choose those microhabitats that best match their specificphenotype to maximize fitness. Despite the profound implications, matching habitat choice has not been unequivocally demonstrated. Weconducted two experiments to examine the impact of pigmentation pattern in the color polymorphic pygmy grasshopper Tetrix subulata onhabitat choice in a laboratory thermal mosaic arena. We found no behavioral differences in the thermal mosaic among pygmy grasshoppersbelonging to either pale, intermediate or dark natural color morphs. However, after manipulating the grasshoppers’ phenotype, the utilizationthrough time of warmer and colder parts of the arena was different for black-painted and white-painted individuals. White-paintedindividuals used warmer parts of the arena, at least during the initial stage of the experiment. We conclude that microhabitat choicerepresents a form of behavioural plasticity. Thus, even if the choice itself is flexible and not genetically determined, it can still lead to spatialgenetic structure in the population because the phenotypes themselves may be genetically mediated

  • 41.
    Wennersten, Lena
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Population-level consequences of polymorphism, plasticity and randomized phenotype switching: a review of predictions2012Ingår i: Biological Reviews, ISSN 1464-7931, E-ISSN 1469-185X, Vol. 87, nr 3, s. 756-767Artikel, forskningsöversikt (Refereegranskat)
    Abstract [en]

    The consequences of among-individual phenotypic variation for the performance and ecological success of populations and species has attracted growing interest in recent years. Earlier reviews of this field typically address the consequences for population processes of one specific source of variation (plasticity or polymorphism), or consider one specific aspect of population performance, such as rate of speciation. Here we take a broader approach and study earlier reviews in order to summarize and compare predictions regarding several population-level consequences of phenotypic variation stemming from genetic polymorphism, developmental plasticity or randomized phenotype switching. Unravelling cause-dependent consequences of variation may increase our ability to understand the ecological dynamics of natural populations and communities, develop more informed management plans for protection of biodiversity, suggest possible routes to increased productivity and yield in natural and managed biological systems, and resolve inconsistencies in patterns and results seen in studies of different model systems. We find an overall agreement regarding the effects of higher levels of phenotypic variation generated by different sources, but also some differences between fine-grained and coarse-grained environments, modular and unitary organisms, mobile and sessile organisms, and between flexible and fixed traits. We propose ways to test the predictions and identify issues where current knowledge is limited and future lines of investigation promise to provide important novel insights.

  • 42.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Merilaita, Sami
    Åbo Akademi University, Finland.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Reduced predation risk for melanistic pygmy grasshoppers in post-fire environments2012Ingår i: Ecology and Evolution, ISSN 2045-7758, E-ISSN 2045-7758, Vol. 2, nr 9, s. 2204-2212Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    The existence of melanistic (black) color forms in many species represents interesting model systems that have played important roles for our understanding of selective processes, evolution of adaptations, and the maintenance of variation. A recent study reported on rapid evolutionary shifts in frequencies of the melanistic forms in replicated populations of Tetrix subulata pygmy grasshoppers; the incidence of the melanistic form was higher in recently burned areas with backgrounds blackened by fire than in nonburned areas, and it declined over time in postfire environments. Here, we tested the hypothesis that the frequency shifts of the black color variant were driven, at least in part, by changes in the selective regime imposed by visual predators. To study detectability of the melanistic form, we presented human “predators” with images of black grasshoppers and samples of the natural habitat on computer screens. We demonstrate that the protective value of black coloration differs between burnt and nonburnt environments and gradually increases in habitats that have been more blackened by fire. These findings support the notion that a black color pattern provides improved protection from visually oriented predators against blackened backgrounds and implicate camouflage and predation as important drivers of fire melanism in pygmy grasshoppers.

  • 43.
    Berggren, Hanna
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Tinnert, Jon
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Spatial sorting may explain evolutionary dynamics of wing polymorphism in pygmy grasshoppers.2012Ingår i: Journal of Evolutionary Biology, ISSN 1010-061X, E-ISSN 1420-9101, Vol. 25, nr 10, s. 2126-2138Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Wing polymorphism in insects provides a good model system for investigating evolutionary dynamics and population divergence in dispersal-enhancing traits. This study investigates the contribution of divergent selection, trade-offs, behaviour and spatial sorting to the evolutionary dynamics of wing polymorphism in the pygmy grasshopper Tetrix subulata (Tetrigidae: Orthoptera). We use data for > 2800 wild-caught individuals from 13 populations and demonstrate that the incidence of the long-winged (macropterous) morph is higher and changes faster between years in disturbed habitats characterized by succession than in stable habitats. Common garden and mother-offspring resemblance studies indicate that variation among populations and families is genetically determined and not influenced to any important degree by developmental plasticity in response to maternal condition, rearing density or individual growth rate. Performance trials show that only the macropterous morph is capable of flight and that propensity to fly differs according to environment. Markrecapture data reveal no difference in the distance moved between free-ranging long- and short-winged individuals. There is no consistent difference across populations and years in number of hatchlings produced by long- and shorter-winged females. Our findings suggest that the variable frequency of the long-winged morph among and within pygmy grasshopper populations may reflect evolutionary modifications driven by spatial sorting due to phenotype- and habitat typedependent emigration and immigration.

  • 44.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Wennersten, Lena
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Karlsson, Magnus
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Caesar, Sofia
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Variation in founder groups promotes establishment success in the wild2012Ingår i: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 279, nr 1739, s. 2800-2806Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Environmental changes currently pose severe threats to biodiversity, and reintroductions and translocations are increasingly used to protect declining populations and species from extinction. Theory predicts that establishment success should be higher for more variable groups of dissimilar individuals. To test this ‘diversity promotes establishment’ hypothesis, we introduced colour polymorphic pygmy grasshoppers (Tetrix subulata) to different sites in the wild. The number of descendants found at the release sites the subsequent year increased with increasing number of colour morphs in the founder group, and variation in founder groups also positively affected colour morph diversity in the established populations. Since colour morphs differ in morphology, physiology, behaviour, reproductive life history and types of niche used, these findings demonstrate that variation among individuals in functionally important traits promotes establishment success under natural conditions, and further indicate that founder diversity may contribute to evolutionary rescue and increased population persistence.

  • 45.
    Karpestam, Einat
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Dietary differences among colour morphs of pygmy grasshoppers revealed by behavioural experiments and stable isotopes.2011Ingår i: Evolutionary Ecology Research, ISSN 1522-0613, E-ISSN 1937-3791, Vol. 13, s. 461-477Artikel i tidskrift (Refereegranskat)
  • 46.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Merilä, Juha
    University of Helsinki.
    Ebenhard, Torbjörn
    Swedish Biodiversity Centre.
    Phenotypic evolution of dispersal-enhancing traits in insular voles2011Ingår i: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 278, nr 1703, s. 225-232Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Evolutionary theory predicts that in metapopulations subject to rapid extinction-recolonization dynamics, natural selection should favour evolution of traits that enhance dispersal and recolonization ability. Metapopulations of field voles (Microtus agrestis) on islands in the Stockholm archipelago, Sweden, are characterized by frequent local extinction and recolonization of subpopulations. Here, we show that voles on the islands were larger and had longer feet than expected for their body size, compared with voles from the mainland; that body size and size-specific foot length increased with increasing geographical isolation and distance from mainland; and that the differences in body size and size-specific foot length were genetically based. These findings provide rare evidence for relatively recent (less than 1000 years) and rapid (corresponding to 100-250 darwins) evolution of traits facilitating dispersal and recolonization in island metapopulations.

  • 47.
    Forsman, Anders
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Karlsson, Magnus
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Wennersten, Lena
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Johansson, Jenny
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Karpestam, Einat
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Rapid evolution of fire melanism in replicated populations of pygmy grasshoppers2011Ingår i: Evolution, ISSN 0014-3820, E-ISSN 1558-5646, Vol. 65, nr 9, s. 2530-2540Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Evolutionary theory predicts an interactive process whereby spatiotemporal environmental heterogeneity will maintain genetic variation, while genetic and phenotypic diversity will buffer populations against stress and allow for fast adaptive evolution in rapidly changing environments. Here, we study color polymorphism patterns in pygmy grasshoppers (Tetrix subulata) and show that the frequency of the melanistic (black) color variant was higher in areas that had been ravaged by fires the previous year than in nonburned habitats, that, in burned areas, the frequency of melanistic grasshoppers dropped from ca. 50% one year after a fire to 30% after four years, and that the variation in frequencies of melanistic individuals among and within populations was genetically based on and represented evolutionary modifications. Dark coloration may confer a selective benefit mediated by enhanced camouflage in recently fire-ravaged areas characterized by blackened visual backgrounds before vegetation has recovered. These findings provide rare evidence for unusually large, extremely rapid adaptive contemporary evolution in replicated natural populations in response to divergent and fluctuating selection associated with spatiotemporal environmental changes.

  • 48.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Rethinking the thermal melanism hypothesis: rearing temperature and coloration in pygmy grasshoppers2011Ingår i: Evolutionary Ecology, ISSN 0269-7653, E-ISSN 1573-8477, Vol. 25, nr 6, s. 1247-1257Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Selection for efficient conversion of solar radiation to body heat has favored theevolution of dark coloration in many ectotherms. The thermal melanism hypothesis positsthat dark coloration is beneficial under conditions of low ambient temperatures because itresults in faster heating rates and higher body temperatures. Fast heating rates, however,may come at a cost of overheating unless compensated for by thermal physiology orbehaviour. Pygmy grasshopper (Orthoptera, Tetrigidae) populations that inhabit fire-ravagedareas characterized by blackened backgrounds and hot surface temperatures due tohigh absorbance of solar radiation show an increased frequency of black phenotypes. Iraised the progeny of wild-captured Tetrix undulata in cold and hot temperatures and useddata on color patterns and survival in a greenhouse to examine whether a cold thermalenvironment triggered the development of melanic coloration or differently affected survivalof melanic versus non-melanic individuals. My results indicate that melanism was notinfluenced by rearing temperature but by genes or epigenetic maternal effects. Temperaturealso did not affect survival. However, melanic individuals produced by melanic motherssurvived longer than melanic individuals produced by non- melanic mothers, whereas nonmelanicindividuals produced by non-black mothers survived longer than melanic individualsproduced by non-black mothers. This suggests a mismatch between color andphysiology in offspring belonging to a different color morph than their mother. Futureinvestigations into the evolution of melanism should consider conflicting selection pressureson thermal capacity and camouflage as well as the influence of correlated responsesto selection on traits associated with coloration.

  • 49.
    Caesar, Sofia
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Karlsson, Magnus
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Diversity and relatedness enhance survival in colour polymorphic grasshoppers2010Ingår i: PLoS ONE, ISSN 1932-6203, E-ISSN 1932-6203, Vol. 5, nr 5, artikel-id e10880Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Evolutionary theory predicts that different resource utilization and behaviour by alternative phenotypes may reduce competition and enhance productivity and individual performance in polymorphic, as compared with monomorphic, groups of individuals. However, firm evidence that members of more heterogeneous groups benefit from enhanced survival has been scarce or lacking. Furthermore, benefits associated with phenotypic diversity may be counterbalanced by costs mediated by reduced relatedness, since closely related individuals typically are more similar. Pygmy grasshoppers (Tetrix subulata) are characterized by extensive polymorphism in colour pattern, morphology, behaviour and physiology. We studied experimental groups founded by different numbers of mothers and found that survival was higher in low than in high density, that survival peaked at intermediate colour morph diversity in high density, and that survival was independent of diversity in low density where competition was less intense. We further demonstrate that survival was enhanced by relatedness, as expected if antagonistic and competitive interactions are discriminately directed towards non-siblings. We therefore also performed behavioural observations and staged encounters which confirmed that individuals recognized and responded differently to siblings than to non-siblings. We conclude that negative effects associated with competition are less manifest in diverse groups, that there is conflicting selection for and against genetic diversity occurring simultaneously, and that diversity and relatedness may facilitate the productivity and ecological success of groups of interacting individuals.

  • 50.
    Karlsson, Magnus
    et al.
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Forsman, Anders
    Linnéuniversitetet, Fakultetsnämnden för naturvetenskap och teknik, Institutionen för naturvetenskap, NV.
    Is melanism in pygmy grasshoppers induced by crowding?2010Ingår i: Evolutionary Ecology, ISSN 0269-7653, E-ISSN 1573-8477, Vol. 24, nr 5, s. 975-983Artikel i tidskrift (Refereegranskat)
    Abstract [en]

    Color polymorphisms in animals may result from plasticity of the developmental system in response to genetic cues in the form of allelic variation at polymorphic loci, environmental cues, or a combination of genetic and environmental cues. An increased understanding of the evolution of color polymorphisms requires better knowledge of when we should expect genetic and environmental cues respectively to influence phenotype determination. Theory posits that the developmental systems of organisms should evolve sensitivity to such cues that most accurately predict coming selective conditions. Pygmy grasshoppers (Orthoptera, Tetrigidae) vary in color pattern within and among populations and show fire melanism, i.e., an increased frequency of black and dark colored phenotypes in high density populations inhabiting fire-ravaged areas. We examined if the population density experienced by individuals during development influenced the phenotypic expression of color pattern in Tetrix subulata. Individuals were experimentally reared either in solitude, at intermediate density or under crowded conditions. We found that color patterns of experimental individuals were independent of rearing density but strongly influenced by maternal color pattern. High population density and crowding may not constitute reliable predictors of the selective regime that characterizes post-fire environments.

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