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  • 1. Alho, J. S.
    et al.
    Herczeg, G.
    Laugen, A. T.
    Raesaenen, K.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Merila, J.
    Allen's rule revisited: quantitative genetics of extremity length in the common frog along a latitudinal gradient2011In: Journal of Evolutionary Biology, ISSN 1010-061X, E-ISSN 1420-9101, Vol. 24, no 1, p. 59-70Article in journal (Refereed)
    Abstract [en]

    Ecogeographical rules linking climate to morphology have gained renewed interest because of climate change. Yet few studies have evaluated to what extent geographical trends ascribed to these rules have a genetic, rather than environmentally determined, basis. This applies especially to Allen's rule, which states that the relative extremity length decreases with increasing latitude. We studied leg length in the common frog (Rana temporaria) along a 1500 km latitudinal gradient utilizing wild and common garden data. In the wild, the body size-corrected femur and tibia lengths did not conform to Allen's rule but peaked at mid-latitudes. However, the ratio of femur to tibia length increased in the north, and the common garden data revealed a genetic cline consistent with Allen's rule in some trait and treatment combinations. While selection may have shortened the leg length in the north, the genetic trend seems to be partially masked by environmental effects.

  • 2. Almeida, Erika
    et al.
    Nunes, Ana
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Andrade, Pedro
    Alves, Susana
    Guerreiro, Catia
    Rebelo, Rui
    Antipredator responses of two anurans towards native and exotic predators2011In: Amphibia-Reptilia, ISSN 0173-5373, E-ISSN 1568-5381, Vol. 32, no 3, p. 341-350Article in journal (Refereed)
    Abstract [en]

    When faced with the risk of predation, tadpoles of many amphibian species are known to modify their phenotype. In this work we studied the effect of an exotic species, the red swamp crayfish (Procambarus clarkii), on the phenotype of two species of amphibians with different reproduction habitats: the Iberian painted frog, Discoglossus galganoi, that normally reproduces in temporary water bodies and the common toad, Bufo bufo, that reproduces in permanent water bodies. The responses were compared with the ones shown in the presence of a native predator, dragonfly (Aeshnidae) larvae. Behaviour, growth and morphology of tadpoles were monitored in a factorial experiment with five treatments. Our results showed that only the permanent habitat species altered its behaviour and life-history traits in the presence of P. clarkii; however, this was mediated by chemical cues from consumed conspecifics. Antipredator responses of B. bufo towards the exotic crayfish were similar to the ones towards the native predator, while D. galganoi responded to the dragonfly larvae but not to P. clarkii. This may be the result of infrequent colonization events of temporary habitats by the crayfish. Therefore, the consequences of the introduction of P. clarkii might be more serious for D. galganoi and other species living in temporary habitats. Species breeding in permanent habitats, more prone to having generalized antipredator responses, may be relatively protected against this exotic crayfish although the effectiveness of these responses still needs to be tested.

  • 3. Arce, Fernando
    et al.
    Orizaola, German
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Navedo, Juan G.
    Storm petrel's breeding skipping in response to oil-spill pollution: Raising concerns over Zabala et al. (2011) methodological approach2011In: Marine Pollution Bulletin, ISSN 0025-326X, E-ISSN 1879-3363, Vol. 62, no 11, p. 2576-2577Article in journal (Refereed)
  • 4.
    Barnaby, Jonathan
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Cooperative Breeding in the Southern Anteater-Chat: Sexual Disparity, Survival and Dispersal2012Doctoral thesis, comprehensive summary (Other academic)
    Abstract [en]

    Group-living sets the scene for complex social behaviours such as cooperative breeding, and exploring the factors that shape group-living is crucial in understanding these behaviours. This thesis explores the ecology of a population of the facultative cooperative breeding southern anteater-chat (Myrmecocichla formicivora), a group-living bird species endemic to southern Africa. It reveals a breeding system based around a breeding pair and up to three auxiliary males. Despite equal numbers of males and females produced as fledglings there was a surplus of adult males, which remained philopatric. Dispersal was strongly female biased. Females dispersed within their first year, they dispersed further than males, and they lost the benefits of the natal site. The sex skew in the population suggested that these factors drive differential mortality, with juvenile females having much lower annual survival than juvenile males. Adult survival was higher, with female survival only slightly lower than male survival. Dispersal distances suggested that males selected the breeding location, nearer to their natal site. There was no evidence of surplus non-breeding females. On the loss of a breeding female there was no replacement until new females entered the population, yet if a breeding male disappeared the female promptly re-paired with a male from another group. There was no indication of birds floating in the population, and if males were orphaned or widowed they joined other groups as unrelated helpers in preference to floating. There was no sign of inter-group or individual aggression among chats, and unrelated helpers were peacefully accepted into groups, suggesting mutual benefits. In fact all birds in a group helped raise offspring of the breeding pair, and groups with more helpers fledged more offspring, which implies that both direct and indirect fitness benefits can be gained through joining a group and helping. There was surprisingly little inheritance of breeding position by auxiliaries, and strikingly low levels of extra-pair paternity. This study suggests that the Southern anteater-chat group structure arises through male philopatry due to a shortage of breeding females, the benefits of remaining on the natal site and helping, and the potentially high costs of living alone.

    List of papers
    1. Group-living in the southern anteater-chat Myrmecocichla formicivora.
    Open this publication in new window or tab >>Group-living in the southern anteater-chat Myrmecocichla formicivora.
    (English)Manuscript (preprint) (Other academic)
    Abstract [en]

    Group-living sets the scene for complex social behaviours such as cooperative breeding, and exploring the factors that shape group-living is crucial in understanding these behaviours. Here we describe some aspects of the ecology of a population of the facultative cooperative breeding southern anteater-chat (Myrmecocichla formicivora), a group living bird species endemic to southern Africa. We used data from a four year study of individually marked birds, with pedigrees completed using microsatellite genotyping. Southern anteater-chats live in groups of 2-5 individuals - a breeding pair and up to three additional none-breeders. These auxiliary birds were either retained offspring or unrelated individuals, and all birds in a group assisted by feeding at the nest. Our population had a skewed sex ratio of approximately 58% males to 42% females, yet the sex ratio of fledglings was equal, suggesting sex-biased mortality. Helpers were predominantly retained male offspring; however 21% of helpers were unrelated to either of the breeding pair. Southern anteater-chats appear to be non-territorial, with an apparent lack of aggression both within and between groups. Our study confirms that the southern anteater-chat is a facultative cooperative breeder, with both pair breeders and groups with helpers capable of fledging youngsters. We provide evidence suggesting that the breeding system of the southern anteater-chat is based on prompt female dispersal, and male philopatry due to an apparent shortage of mates, potential benefits of the natal site and possible high costs of floating. It appears that ecological constraints promoting delayed dispersal are reinforced by benefits gained from remaining philopatric.

    Keywords
    Southern anteater-chat, cooperative breeding, social evolution, behavioural ecology, delayed dispersal, family living, Africa
    National Category
    Evolutionary Biology
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-179071 (URN)
    Available from: 2012-08-13 Created: 2012-08-07 Last updated: 2012-09-05
    2. Sex specific survival in the southern anteater-chat Mymecocichla formicivora.
    Open this publication in new window or tab >>Sex specific survival in the southern anteater-chat Mymecocichla formicivora.
    (English)Manuscript (preprint) (Other academic)
    Abstract [en]

    Survival is a key factor behind life-history variation both between and within species. It is also a major influence on sociality in species which delay dispersal and live in family groups. Knowledge of differential survival rates between males and females and juveniles and adults give insights into the costs and benefits of different behavioural and life-history strategies. Here we investigate patterns of survival in a population of the southern anteater-chat (Myrmecocichla formicivora), a facultative cooperatively breeding passerine of southern Africa. Using data from a 9-year mark-capture-recapture study, we examined juvenile and adult sex related variation in survival, and the role of environmental variables (rainfall, temperature) for survival patterns in the population. Annual adult survival probability (mean ± SE) was 0.71 ± 0.03 for males and 0.60 ± 0.04 for females. Juvenile survival was lower for each sex, with juvenile female survival (0.36 ± 0.04) being 35% lower than juvenile male survival (0.55 ± 0.04). Using these estimates we calculated the mean life span (MLS) in years for male southern anteater-chat to be 4.0 ± 0.7, considerably higher than for females at 2.0 ± 0.4. These figures closely matched the population-age structure of the study area, and could explain the high male biased sex skew of adult birds in this population. Higher annual mean temperature was associated with higher survival, whereas higher annual rainfall was associated with lower survival for both sex and age classes. Female survival, particularly female juvenile survival, may be reduced due to prompt dispersal and longer dispersal distances, and the additional costs of breeding early in life. Differential survival can promote male philopatry and this in turn could well encourage the cooperative breeding we see in the southern anteater-chat.

    Keywords
    Southern anteater-chat, cooperative breeding, social evolution, behavioural ecology, delayed dispersal, family living, Africa
    National Category
    Evolutionary Biology
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-179072 (URN)
    Available from: 2012-08-13 Created: 2012-08-07 Last updated: 2012-09-05
    3. The rarer sex - female natal dispersal and breeder replacement in the southern anteater-chat Myrmecocichla formicivora.
    Open this publication in new window or tab >>The rarer sex - female natal dispersal and breeder replacement in the southern anteater-chat Myrmecocichla formicivora.
    (English)Manuscript (preprint) (Other academic)
    Abstract [en]

    Sex biased dispersal is a crucial factor in understanding the mechanism of family dynamics in many cooperative breeders. Female biased dispersal occurs in many cooperatively breeding birds. It is often associated with females dispersing earlier and further, and a male biased sex skew in the population. Here we investigated female dispersal in the southern anteater-chat, a facultative cooperatively breeding passerine of southern Africa. Our study population had a male biased sex skew, and females had lower annual survival than males. Dispersal was strongly female biased, with females dispersing within their first year whereas many males remained philopatric beyond the next breeding season. Breeding females were on average younger than breeding males, and also dispersed further. Each breeding group contained only one female. No females were found floating in the population, and all females were associated with one or more males in a breeding group. It appears that all females disperse in their first year directly to a breeding position. If a female disappeared in the breeding season they were not replaced until new females matured and dispersed the following season, yet if a male breeder disappeared during the breeding season he was almost immediately replaced, indicating that there are no surplus females.

    Keywords
    Southern anteater-chat, cooperative breeding, social evolution, behavioural ecology, delayed dispersal, family living, Africa
    National Category
    Evolutionary Biology
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-179073 (URN)
    Available from: 2012-08-13 Created: 2012-08-07 Last updated: 2012-09-05
    4. From helping to breeding – male choice in the southern anteater-chats Myrmecocichla formicivora.
    Open this publication in new window or tab >>From helping to breeding – male choice in the southern anteater-chats Myrmecocichla formicivora.
    (English)Manuscript (preprint) (Other academic)
    Abstract [en]

    Dispersal choice is important in understanding population structure and dynamics. Here we examine male choice in the facultative cooperative breeding southern anteater-chat Myrmecocichla formicivora based on a four year study in South Africa. The sex ratio in our study population was male biased, with many males remaining philopatric. All groups consisted of one or more adult males associated with one adult female. We found a significant positive effect of auxiliary number on group productivity (both number of young fledged, and first year survival), while controlling for potentially confounding variables (territory and breeder identity). The majority of auxiliaries, 54%, were related to both birds in the breeding pair, with another 25% related to one member of the pair, and 21% related to neither of the breeders. There was no evidence of males floating within our study population, and it appears that if males lose their family due to mortality they join other groups as unrelated non-breeding auxiliaries rather than float. No aggression was observed between individual southern anteater-chats, and unrelated group members helped rear offspring in the group they had joined. Despite the presence of, and helping by unrelated group members there was very little evidence of breeding position inheritance (1/24 auxiliaries unrelated to the breeding female) or extra-pair paternity (2.4% of fledglings). This study suggests that the southern anteater-chat group structure arises through male philopatry due to a lack of breeding females and potentially high costs of living alone.

    Keywords
    Southern anteater-chat, cooperative breeding, social evolution, behavioural ecology, delayed dispersal, family living, Africa
    National Category
    Evolutionary Biology
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-179074 (URN)
    Available from: 2012-08-13 Created: 2012-08-07 Last updated: 2012-09-05
    5. Development of a suit of microsatellite markers for the southern anteater-chat Myrmecocichla formicivora.
    Open this publication in new window or tab >>Development of a suit of microsatellite markers for the southern anteater-chat Myrmecocichla formicivora.
    (English)Manuscript (preprint) (Other academic)
    Abstract [en]

    We tested the cross amplification of 37 microsatellite markers for their suitability in genotyping the southern anteater-chat Myrmecocichla formicivora, an opportunistic cooperatively breeding passerine bird endemic to southern Africa. Fourteen microsatellite markers were identified as having suitable characteristics, with minor deviation from Hardy-Weinberg equilibrium and little evidence of null alleles. These 14 Primer pairs were combined in 4 multiplexes and run on 183 individual samples from our study population of southern anteater-chat on Benfontein Nature Reserve, near Kimberley in central South Africa. The loci ranged from 3-34 alleles per locus, and observed heterozygosity ranged from 0.45 -0.93. We then tested these 14 microsatellites for their use in examining paternity in a population of southern anteater-chat being studied on Benfontein Nature Reserve, near Kimberley in South Africa. Of the population of 183 individuals (the 2011 population) 93% of the offspring could be allocated a mother, 97% a father, and 87% a parent pair with 95% confidence. The remainder could be allocated at the 80% confidence level. Where mothers could be assigned from observations this was in 100% agreement with the microsatellite results, giving us good support for the accurate assignment of parentage in our population.

    Keywords
    Southern anteater-chat, cooperative breeding, social evolution, microsatellites, genotyping, cross-amplification, behavioural ecology, delayed dispersal, family living, Africa
    National Category
    Evolutionary Biology
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-179075 (URN)
    Available from: 2012-08-13 Created: 2012-08-07 Last updated: 2012-09-05
  • 5.
    Barnaby, Jonathan
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Ekman, Jan
    From helping to breeding – male choice in the southern anteater-chats Myrmecocichla formicivora.Manuscript (preprint) (Other academic)
    Abstract [en]

    Dispersal choice is important in understanding population structure and dynamics. Here we examine male choice in the facultative cooperative breeding southern anteater-chat Myrmecocichla formicivora based on a four year study in South Africa. The sex ratio in our study population was male biased, with many males remaining philopatric. All groups consisted of one or more adult males associated with one adult female. We found a significant positive effect of auxiliary number on group productivity (both number of young fledged, and first year survival), while controlling for potentially confounding variables (territory and breeder identity). The majority of auxiliaries, 54%, were related to both birds in the breeding pair, with another 25% related to one member of the pair, and 21% related to neither of the breeders. There was no evidence of males floating within our study population, and it appears that if males lose their family due to mortality they join other groups as unrelated non-breeding auxiliaries rather than float. No aggression was observed between individual southern anteater-chats, and unrelated group members helped rear offspring in the group they had joined. Despite the presence of, and helping by unrelated group members there was very little evidence of breeding position inheritance (1/24 auxiliaries unrelated to the breeding female) or extra-pair paternity (2.4% of fledglings). This study suggests that the southern anteater-chat group structure arises through male philopatry due to a lack of breeding females and potentially high costs of living alone.

  • 6.
    Barnaby, Jonathan
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Ekman, Jan
    The rarer sex - female natal dispersal and breeder replacement in the southern anteater-chat Myrmecocichla formicivora.Manuscript (preprint) (Other academic)
    Abstract [en]

    Sex biased dispersal is a crucial factor in understanding the mechanism of family dynamics in many cooperative breeders. Female biased dispersal occurs in many cooperatively breeding birds. It is often associated with females dispersing earlier and further, and a male biased sex skew in the population. Here we investigated female dispersal in the southern anteater-chat, a facultative cooperatively breeding passerine of southern Africa. Our study population had a male biased sex skew, and females had lower annual survival than males. Dispersal was strongly female biased, with females dispersing within their first year whereas many males remained philopatric beyond the next breeding season. Breeding females were on average younger than breeding males, and also dispersed further. Each breeding group contained only one female. No females were found floating in the population, and all females were associated with one or more males in a breeding group. It appears that all females disperse in their first year directly to a breeding position. If a female disappeared in the breeding season they were not replaced until new females matured and dispersed the following season, yet if a male breeder disappeared during the breeding season he was almost immediately replaced, indicating that there are no surplus females.

  • 7.
    Barnaby, Jonathan
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Fletcher, Kevin
    Ekman, Jan
    Development of a suit of microsatellite markers for the southern anteater-chat Myrmecocichla formicivora.Manuscript (preprint) (Other academic)
    Abstract [en]

    We tested the cross amplification of 37 microsatellite markers for their suitability in genotyping the southern anteater-chat Myrmecocichla formicivora, an opportunistic cooperatively breeding passerine bird endemic to southern Africa. Fourteen microsatellite markers were identified as having suitable characteristics, with minor deviation from Hardy-Weinberg equilibrium and little evidence of null alleles. These 14 Primer pairs were combined in 4 multiplexes and run on 183 individual samples from our study population of southern anteater-chat on Benfontein Nature Reserve, near Kimberley in central South Africa. The loci ranged from 3-34 alleles per locus, and observed heterozygosity ranged from 0.45 -0.93. We then tested these 14 microsatellites for their use in examining paternity in a population of southern anteater-chat being studied on Benfontein Nature Reserve, near Kimberley in South Africa. Of the population of 183 individuals (the 2011 population) 93% of the offspring could be allocated a mother, 97% a father, and 87% a parent pair with 95% confidence. The remainder could be allocated at the 80% confidence level. Where mothers could be assigned from observations this was in 100% agreement with the microsatellite results, giving us good support for the accurate assignment of parentage in our population.

  • 8.
    Barnaby, Jonathan
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Fletcher, Kevin
    Ekman, Jan
    Group-living in the southern anteater-chat Myrmecocichla formicivora. Manuscript (preprint) (Other academic)
    Abstract [en]

    Group-living sets the scene for complex social behaviours such as cooperative breeding, and exploring the factors that shape group-living is crucial in understanding these behaviours. Here we describe some aspects of the ecology of a population of the facultative cooperative breeding southern anteater-chat (Myrmecocichla formicivora), a group living bird species endemic to southern Africa. We used data from a four year study of individually marked birds, with pedigrees completed using microsatellite genotyping. Southern anteater-chats live in groups of 2-5 individuals - a breeding pair and up to three additional none-breeders. These auxiliary birds were either retained offspring or unrelated individuals, and all birds in a group assisted by feeding at the nest. Our population had a skewed sex ratio of approximately 58% males to 42% females, yet the sex ratio of fledglings was equal, suggesting sex-biased mortality. Helpers were predominantly retained male offspring; however 21% of helpers were unrelated to either of the breeding pair. Southern anteater-chats appear to be non-territorial, with an apparent lack of aggression both within and between groups. Our study confirms that the southern anteater-chat is a facultative cooperative breeder, with both pair breeders and groups with helpers capable of fledging youngsters. We provide evidence suggesting that the breeding system of the southern anteater-chat is based on prompt female dispersal, and male philopatry due to an apparent shortage of mates, potential benefits of the natal site and possible high costs of floating. It appears that ecological constraints promoting delayed dispersal are reinforced by benefits gained from remaining philopatric.

  • 9.
    Barnaby, Jonathan
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Griesser, Michael
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Evolution, Population and Conservation Biology.
    The role of nepotism, cooperation, and competition in the avian families2010Book (Refereed)
    Abstract [en]

    A large number of bird species live in stable groups, and this sets the scene for complex social behaviours, such as cooperative breeding. The vast majority of groups consist of families which arise when young postpone dispersal and remain with their parents beyond independence. However, the factors selecting for the evolution of families and thus also cooperative breeding among birds, are still a challenging puzzle. The currently accepted key explanation for the evolution of families and cooperative breeding focuses on dispersal constraints. While constraints successfully explain within‐population dispersal decisions, they fail as an ultimate explanation because offspring in the majority of species face some sort of dispersal constraint, yet still disperse promptly. Recent alternative explanations focus on the role of philopatry and nepotism, and emphasise a key role of life‐history for the evolution of families. Phylogenetic analyses and field studies have indicated that living in family groups is far more widespread among long‐lived species than short‐lived ones. A long lifespan gives parents the opportunity to invest in their offspring for a prolonged period, while this option is less viable for short‐lived species. Thus, living with nepotistic parents provides offspring with direct fitness benefits that can select for the evolution of family living beyond independence. Nevertheless this generalisation is brought into question since many long‐lived bird species do not live in family groups. An alternative approach attempts to explain family living through the variation in territory quality. Here the incentive to remain with the parents is created by the availability of resources on the natal territory independent of parental nepotism. However, there is not only cooperation, conflicts are also common place in families. Living with independent, sexually mature offspring can lead to conflicts through a change in resource availability or the death of aparent. Therefore families can be expected to be dynamic societies where both parent and offspring decisions depend on each other, and family maintenance depends upon the current ecological conditions. Based on  this background, here we review recent studies that have investigated the processes that facilitate family formation, and which highlight both cooperation and conflict that arises from living in family groups. We examine the strengths of current models and explore ideas for a more coherent framework in which to understand prolonged family association in birds. We argue that two paths lead to family living, depending on the life-history. In medium-short lived species where the postponement of independent reproduction comes at a high cost, offspring can benefit from an association with their parents until the next breeding season. In longer-lived species, offspring actually benefit from postponing the onset of independent reproduction, making family living beyond the first year of life an adaptive strategy, and giving the option for cooperative breeding. These processes are illustrated by 5 species-specific case studies. We then finally suggest a number of key questions to developing a deeper understanding of the evolution of family living in birds.

  • 10.
    Barnaby, Jonathan
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Herrmann, Eric
    Ekman, Jan
    Sex specific survival in the southern anteater-chat Mymecocichla formicivora. Manuscript (preprint) (Other academic)
    Abstract [en]

    Survival is a key factor behind life-history variation both between and within species. It is also a major influence on sociality in species which delay dispersal and live in family groups. Knowledge of differential survival rates between males and females and juveniles and adults give insights into the costs and benefits of different behavioural and life-history strategies. Here we investigate patterns of survival in a population of the southern anteater-chat (Myrmecocichla formicivora), a facultative cooperatively breeding passerine of southern Africa. Using data from a 9-year mark-capture-recapture study, we examined juvenile and adult sex related variation in survival, and the role of environmental variables (rainfall, temperature) for survival patterns in the population. Annual adult survival probability (mean ± SE) was 0.71 ± 0.03 for males and 0.60 ± 0.04 for females. Juvenile survival was lower for each sex, with juvenile female survival (0.36 ± 0.04) being 35% lower than juvenile male survival (0.55 ± 0.04). Using these estimates we calculated the mean life span (MLS) in years for male southern anteater-chat to be 4.0 ± 0.7, considerably higher than for females at 2.0 ± 0.4. These figures closely matched the population-age structure of the study area, and could explain the high male biased sex skew of adult birds in this population. Higher annual mean temperature was associated with higher survival, whereas higher annual rainfall was associated with lower survival for both sex and age classes. Female survival, particularly female juvenile survival, may be reduced due to prompt dispersal and longer dispersal distances, and the additional costs of breeding early in life. Differential survival can promote male philopatry and this in turn could well encourage the cooperative breeding we see in the southern anteater-chat.

  • 11.
    Beier, Sara
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Limnology.
    Mohit, Vani
    Département de Biologie, Québec-Océan and Institut de biologie integrative et des systèmes, Université Laval.
    Ettema, Thijs J. G.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Cell and Molecular Biology, Molecular Evolution.
    Östman, Örjan
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Tranvik, Lars J.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Limnology.
    Bertilsson, Stefan
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Limnology.
    Pronounced seasonal dynamics of freshwater chitinase genes and chitin processing2012In: Environmental Microbiology, ISSN 1462-2912, E-ISSN 1462-2920, Vol. 14, no 9, p. 2467-2479Article in journal (Refereed)
    Abstract [en]

    Seasonal variation in activity of enzymes involved in polymer degradation, including chitinases, has been observed previously in freshwater environments. However, it is not known whether the seasonal dynamics are due to shifts in the activity of bacteria already present, or shifts in community structure towards emergence or disappearance of chitinolytic organisms. We traced seasonal shifts in the chitinase gene assemblage in a temperate lake and linked these communities to variation in chitinase activity. Chitinase genes from 20 samples collected over a full yearly cycle were characterized by pyrosequencing. Pronounced temporal shifts in composition of the chitinase gene pool (beta diversity) occurred along with distinct shifts in richness (alpha diversity) as well as chitin processing. Changes in the chitinase gene pool correlated mainly with temperature, abundance of crustacean zooplankton and phytoplankton blooms. Also changes in the physical structure of the lake, e.g. stratification and mixing were associated with changes in the chitinolytic community, while differences were minor between surface and suboxic hypolimnetic water. The lake characteristics influencing the chitinolytic community are all linked to changes in organic particles and we suggest that seasonal changes in particle quality and availability foster microbial communities adapted to efficiently degrade them.

  • 12. Bourdeau, Paul E.
    et al.
    Johansson, Frank
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Predator-induced morphological defences as by-products of prey behaviour: a review and prospectus2012In: Oikos, ISSN 0030-1299, E-ISSN 1600-0706, Vol. 121, no 8, p. 1175-1190Article, review/survey (Refereed)
    Abstract [en]

    Predator-induced morphological defences (PIMDs) are ubiquitous. Many PIMDs may be mediated by prey behaviour rather than directly cued by predators. A survey of 92 studies indicated 40 that quantified prey behaviour, all of which document positive associations between defence production and activity reduction. Thus, PIMDs are associated with changes in prey activity, which could have caused the morphological change. We propose two possible mechanisms: 1) decreased activity reduces feeding rate, resulting in lower growth and morphological change; and 2) activity reduction conserves energy, which is reallocated for growth, subsequently changing morphology. Resource availability also causes similar morphological change to predator presence, suggesting confounding effects of resources and predators with current methodology. Future studies should estimate food ingestion, assimilation efficiency, and growth rate in the presence and absence of predators, crossing predator presence with resource levels. Not all PIMDs will be behaviourally-mediated, but consideration of causal linkages between prey behaviour and PIMDs is warranted.

  • 13. Brodin, Tomas
    et al.
    Lind, Martin I.
    Wiberg, Miria Kaltiala
    Johansson, Frank
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology. Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Animal ecology.
    Personality trait differences between mainland and island populations in the common frog (Rana temporaria)2013In: Behavioral Ecology and Sociobiology, ISSN 0340-5443, E-ISSN 1432-0762, Vol. 67, no 1, p. 135-143Article in journal (Refereed)
    Abstract [en]

    Understanding and predicting species range expansions is an important challenge in modern ecology because of rapidly changing environments. Recent studies have revealed that consistent within-species variation in behavior (i.e., animal personality) can be imperative for dispersal success, a key process in range expansion. Here we investigate how habitat isolation can mediate differentiation of personality traits between recently founded island populations and the main population. We performed laboratory studies of boldness and exploration across life stages (tadpoles and froglets) using four isolated island populations and four mainland populations of the common frog (Rana temporaria). Both tadpoles and froglets from isolated populations were bolder and more exploratory than conspecifics from the mainland. Although the pattern can be influenced by possible differences in predation pressure, we suggest that this behavioral differentiation might be the result of a disperser-dependent founder effect brought on by an isolation-driven environmental filtering of animal personalities. These findings can have important implications for both species persistence in the face of climate change (i.e., range expansions) and ecological invasions as well as for explaining rapid speciation in isolated patches.

  • 14. Cano, J. M.
    et al.
    Li, M-H
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Vilkki, J.
    Merila, J.
    First-generation linkage map for the common frog Rana temporaria reveals sex-linkage group2011In: Heredity, ISSN 0018-067X, E-ISSN 1365-2540, Vol. 107, no 6, p. 530-536Article in journal (Refereed)
    Abstract [en]

    The common frog (Rana temporaria) has become a model species in the fields of ecology and evolutionary biology. However, lack of genomic resources has been limiting utility of this species for detailed evolutionary genetic studies. Using a set of 107 informative microsatellite markers genotyped in a large full-sib family (800 F1 offspring), we created the first linkage map for this species. This partial map-distributed over 15 linkage groups-has a total length of 1698.8 cM. In line with the fact that males are the heterogametic sex in this species and a reduction of recombination is expected, we observed a lower recombination rate in the males (map length: 1371.5 cM) as compared with females (2089.8 cM). Furthermore, three loci previously documented to be sex-linked (that is, carrying male-specific alleles) in adults from the wild mapped to the same linkage group. The linkage map described in this study is one of the densest ones available for amphibians. The discovery of a sex linkage group in Rana temporaria, as well as other regions with strongly reduced male recombination rates, should help to uncover the genetic underpinnings of the sex-determination system in this species. As the number of linkage groups found (n = 15) is quite close to the actual number of chromosomes (n = 13), the map should provide a useful resource for further evolutionary, ecological and conservation genetic work in this and other closely related species.

  • 15.
    Corrales, Carolina
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Höglund, Jacob
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Maintenance of gene flow by female-biased dispersal of black grouse, Tetrao tetrix in northern Sweden2012In: Journal of Ornithology = Journal fur Ornithologie, ISSN 0021-8375, E-ISSN 1439-0361, Vol. 153, no 4, p. 1127-1139Article in journal (Refereed)
    Abstract [en]

    Sex-biased dispersal is a common phenomenon in most birds. In general, males breed at or near their site of birth while most of the females disperse. We investigated the dispersal patterns and genetic structure of lekking Black Grouse Tetrao tetrix based on ten microsatellite loci. Data for 469 individuals from 25 localities spaced from 45 to 558 km apart revealed low levels of genetic differentiation and high connectivity among studied sites due to female-biased dispersal. The spatial distribution of the genetic variation did not follow an isolation by distance pattern neither for females nor for males. STRUCTURE identified three clusters of male individuals but without any geographical pattern. Only one cluster was identified for females. Several tests of sex-biased dispersal were executed. Most of them showed no difference between sexes, but the mean assignment index and F IS showed a statistically significant female-biased dispersal. Therefore, we consider that the northern Swedish Black Grouse population is a panmictic population. The amount of gene flow throughout time has been consistent with dispersal and with no strong effect of forest fragmentation in the region.

  • 16.
    Corrales, Carolina
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Pavlovska, Mariia
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Hoglund, Jacob
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Phylogeography and subspecies status of Black Grouse2014In: Journal of Ornithology = Journal fur Ornithologie, ISSN 0021-8375, E-ISSN 1439-0361, Vol. 155, no 1, p. 13-25Article in journal (Refereed)
    Abstract [en]

    The cold periods of the Pleistocene have had a striking impact on the diversification of most organisms in temperate regions. Phylogeographic patterns and postglacial expansion are poorly understood in the Black Grouse (Tetrao tetrix). This species is widely distributed across Eurasia, and has been divided into a number of subspecies on the basis of morphological differences and geographic isolation. To investigate the evolutionary history of the species, 143 samples from different regions were examined for a mtDNA control region fragment. Overall, analyses of mtDNA gave support for the divergence between Tetrao tetrix tetrix, T. t. ussuriensis and T. t. mongolicus. The analyses reveal the effects of colonisation out of glacial refugia on the genetic diversity and genetic structure of Black Grouse. The phylogeographical results are consistent with a demographic population expansion following a bell-shaped mismatch distribution, a star-shaped phylogeny and low nucleotide diversity. Patterns of postglacial dispersal imply that Black Grouse from southern Europe have been restricted to this area, and did not contribute to the genetic diversity of northern Europe. Instead, Black Grouse spread out to northern Europe from a refugium in the east and a possible one in western Europe, following the retreat of glacial ice sheets, although both refugia remain unidentified. We suggest that T. t. britannicus and T. t. viridanus correspond to northern T. t. tetrix, and that this lineage has diverged from the other subspecies. This division is tentative due to limited sampling, but it will facilitate the management of different evolutionary significant units of the species.

  • 17.
    Corrales Duque, Carolina
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics. Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Population Genetic Structure of Black Grouse (Tetrao tetrix): From a Large to a Fine Scale Perspective2011Doctoral thesis, comprehensive summary (Other academic)
    Abstract [en]

    Black grouse (Tetrao tetrix) is a bird species with a lek mating system found in the Palearctic boreal taiga. It is assumed that it has a continuous distribution along Scandinavia and Siberia, whereas in Central Europe it has declined during the last decades. The primary objective of this thesis was to obtain a deeper understanding of the history, systematic classification and the genetic structure of black grouse on different geographical scales using microsatellites and control region mtDNA sequences (CR). I determined how much the mating system, habitat fragmentation and historical population processes have influenced the partitioning of genetic diversity in this species. Phylogeographical results are consistent with a demographic population expansion, and the patterns of postglacial dispersal suggest that a glacial refugium was located somewhere in central Asia, and from there black grouse spread out to Europe following the retreat of glacial ice sheets. I suggest that the two European black grouse subspecies, T. t. Tetrix and T. t. britannicus correspond to only one subspecies: T. t. tetrix, and that this lineage has diverged from T.t. viridanus, a subspecies found in Kazakhstan. The British population is significantly divergent from the remaining Eurasian samples for microsatellites but it is not for mtDNA. Therefore, they should regard as a separate Management Unit and not as a subspecies. Furthermore, British black grouse occur in three independent genetic units, corresponding to Wales, northern England/southern Scotland and northern Scotland. There was also genetic structure within Sweden. Habitat fragmentation is the main cause of population genetic structure in southern Swedish black grouse. In contrast, low levels of genetic differentiation and high connectivity were found in northern Sweden due to female-biased dispersal. On a finer geographical scale, I found genetic differences between leks due to a mixture of related and unrelated individuals within leks. However, mean relatedness values hardly differed from zero. Some leks were similar to one another and I interpret this as a result of variation in local reproductive success and philopatry. These factors would cause genetic structuring but this by itself would not reveal that kin selection is operating within black grouse leks.

    List of papers
    1. Population history and subspecies status of black grouse
    Open this publication in new window or tab >>Population history and subspecies status of black grouse
    (English)Manuscript (preprint) (Other academic)
    National Category
    Biological Systematics
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-150115 (URN)
    Available from: 2011-03-25 Created: 2011-03-25 Last updated: 2011-05-05
    2. Genetic structure among black grouse in Britain: implications for designing conservation units
    Open this publication in new window or tab >>Genetic structure among black grouse in Britain: implications for designing conservation units
    Show others...
    2011 (English)In: Animal Conservation, ISSN 1367-9430, E-ISSN 1469-1795, Vol. 14, no 4, p. 400-408Article in journal (Refereed) Published
    Abstract [en]

    Black grouse in Britain have faced contraction of their range and have declined in numbers during the recent decades. As such, the species is a conservation concern in the UK. In order to aid conservation decisions, the terms Evolutionary Significant Unit (ESU) and Management Unit (MU) have been proposed. An ESU is an independently evolving evolutionary lineage defined by being reciprocally monophyletic for mitochondrial alleles, and which is significantly different from other lineages with regard to nuclear alleles, whereas an MU is operationally defined by only the latter criterion. Using mitochondrial sequences and nuclear microsatellite loci, we failed to find evidence that British black grouse is an ESU. However, British black grouse are sufficiently different from continental black grouse both with respect to mitochondrial and nuclear data to regard them as a separate MU. Furthermore, we present genetic data which suggest that British black grouse presently occur in what are probably three demographically independent units (roughly corresponding to Wales, northern England/southern Scotland and northern Scotland), which are genetically differentiated. The two southern units (Wales and northern England/southern Scotland) have lower genetic diversity and show signs of having lost genetic variability

    Keywords
    Management Unit, Evolutionary Significant Unit, population structure
    National Category
    Biological Systematics Biological Sciences
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-150110 (URN)10.1111/j.1469-1795.2011.00436.x (DOI)000293173900013 ()
    Available from: 2011-03-25 Created: 2011-03-25 Last updated: 2017-12-11Bibliographically approved
    3. Genetic structure of black grouse in Sweden: consequence of historic or contemporary patterns
    Open this publication in new window or tab >>Genetic structure of black grouse in Sweden: consequence of historic or contemporary patterns
    (English)Manuscript (preprint) (Other academic)
    National Category
    Biological Sciences
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-150116 (URN)
    Available from: 2011-03-25 Created: 2011-03-25 Last updated: 2011-05-05
    4. Maintenance of gene flow by female-biased dispersal of black grouse, Tetrao tetrix in northern Sweden
    Open this publication in new window or tab >>Maintenance of gene flow by female-biased dispersal of black grouse, Tetrao tetrix in northern Sweden
    2012 (English)In: Journal of Ornithology = Journal fur Ornithologie, ISSN 0021-8375, E-ISSN 1439-0361, Vol. 153, no 4, p. 1127-1139Article in journal (Refereed) Published
    Abstract [en]

    Sex-biased dispersal is a common phenomenon in most birds. In general, males breed at or near their site of birth while most of the females disperse. We investigated the dispersal patterns and genetic structure of lekking Black Grouse Tetrao tetrix based on ten microsatellite loci. Data for 469 individuals from 25 localities spaced from 45 to 558 km apart revealed low levels of genetic differentiation and high connectivity among studied sites due to female-biased dispersal. The spatial distribution of the genetic variation did not follow an isolation by distance pattern neither for females nor for males. STRUCTURE identified three clusters of male individuals but without any geographical pattern. Only one cluster was identified for females. Several tests of sex-biased dispersal were executed. Most of them showed no difference between sexes, but the mean assignment index and F IS showed a statistically significant female-biased dispersal. Therefore, we consider that the northern Swedish Black Grouse population is a panmictic population. The amount of gene flow throughout time has been consistent with dispersal and with no strong effect of forest fragmentation in the region.

    Abstract [de]

    Geschlechterunterschiede bei der Auswanderung aus dem Heimatterritorium und Ausbreitung gibt es bei den meisten Vogelarten. Üblicherweise brüten Männchen in der Nähe ihres Geburtsorts, während Weibchen vermehrt abwandern. Wir untersuchten Muster in der Verbreitung und die genetische Struktur im Lek balzender Birkhühner Tetrao tetrix anhand von zehn Mikrosatelliten-Loci. Daten von 469 Individuen aus 25 Gebieten, die zwischen 45 und 558 km voneinander entfernt lagen, zeigten geringe genetische Differenzierung und hohe Konnektivität zwischen den Gebieten, gewährleistet durch die Ausbreitung der Weibchen. Die räumliche Verteilung der genetischen Variation zeigte keine Isolation durch hohe Entfernungen, weder für Männchen noch für Weibchen. Das Programm STRUCTURE fand drei Cluster für männliche Individuen, die nicht mit geographischen Mustern übereinstimmten. Bei Weibchen konnte nur ein Cluster identifiziert werden. Wir führten verschiedene Tests für geschlechtsabhängige Verbreitung durch. Die meisten fanden keinen Unterschied zwischen den Geschlechtern, aber der ‘mean assignment index’ und FIS zeigten signifikant stärkere Ausbreitung der Weibchen. Daraus schließen wir, dass die nordschwedische Birkhuhn Population panmiktisch ist. Der genetische Austausch der Populationen konnte stets durch Verteilung weiblicher Individuen erklärt werden, während die Fragmentierung der Wälder in der Region keinen großen Einfluss gehabt zu haben scheint.

    Keywords
    microsatellites, sex-biased dispersal, philopatry
    National Category
    Biological Sciences
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-150111 (URN)10.1007/s10336-012-0844-0 (DOI)000308823100013 ()
    Available from: 2011-03-25 Created: 2011-03-25 Last updated: 2017-12-11Bibliographically approved
    5. Fine scale genetic structure in the lek-breeding black grouse Tetrao tetrix
    Open this publication in new window or tab >>Fine scale genetic structure in the lek-breeding black grouse Tetrao tetrix
    (English)Article in journal (Refereed) Submitted
    National Category
    Biological Sciences
    Research subject
    Biology with specialization in Population Biology
    Identifiers
    urn:nbn:se:uu:diva-150113 (URN)
    Available from: 2011-03-25 Created: 2011-03-25 Last updated: 2011-05-18Bibliographically approved
  • 18.
    Dahl, Emma
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Backström, T.
    Winberg, Svante
    Uppsala University, Disciplinary Domain of Medicine and Pharmacy, Faculty of Medicine, Department of Neuroscience, Physiology.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Is growth hormone expression correlated with variation in growth rate along a latitudinal gradient in Rana temporaria?2011In: Journal of Zoology, ISSN 0952-8369, E-ISSN 1469-7998, Vol. 285, no 2, p. 85-92Article in journal (Refereed)
    Abstract [en]

    In ectotherms, decreasing season length and lower temperature towards higher latitudes often favour higher growth and developmental rates. However, the underlying physiological mechanisms and particularly the hormonal correlates of clinal variation remain unexplored. The growth hormone (GH) plays a crucial role in growth of all vertebrates and high expression of GH is associated with rapid growth in many species. We tested the hypothesis that GH expression is correlated with a latitudinal gradient in growth in Scandinavian Rana temporaria tadpoles. Using quantitative polymerase chain reaction, we measured GH and growth hormone receptor (GHR) expression at two time points from laboratory-raised tadpoles originating from eight populations collected along the latitudinal gradient. To explore latitudinal differences in stress-induced changes in GH expression, we also compared GH expression in tadpoles raised with and without predators. In accordance with previous studies we found a clear latitudinal gradient in growth. There were no latitudinal effects, or predator-induced effects on GH or GHR expression. However, there was some indication for among-population variation in GH expression. The lack of a latitudinal pattern in GH and GHR expression may be due to that the growth promoting effects of GH is dependent on other factors including insulin-like growth factor-I (IGF), IGF-binding proteins or prolactin. Further studies on these factors may provide insight on the proximal mechanisms of differences in growth in R. temporaria tadpoles.

  • 19.
    Dahl, Emma
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Orizaola, German
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Nicieza, A. G.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Time constraints and flexibility of growth strategies: Geographic variation in catch-up growth responses in amphibian larvae2012In: Journal of Animal Ecology, ISSN 0021-8790, E-ISSN 1365-2656, Vol. 81, no 6, p. 1233-1243Article in journal (Refereed)
    Abstract [en]

    1. As size is tightly associated with fitness, compensatory strategies for growth loss can be vital for restoring individual fitness. However, immediate and delayed costs of compensatory responses may prevent their generalization, and the optimal strategy may depend on environmental conditions. Compensatory responses may be particularly important in high-latitude habitats with short growing seasons, and thus, high-latitude organisms might be more efficient at compensating after periods of unfavourable growth conditions than low-latitude organisms. 2. We investigated geographical differences in catch-up growth strategies of populations of the common frog (Rana temporaria) from southern and northern Sweden in two factorial common garden experiments involving predation risk and two different causes of growth arrest (nutritional stress and low temperatures) to evaluate how the compensatory strategies can be affected by context-dependent costs of compensation. Larval and metamorphic traits, and post-metamorphic performance were used as response variables. 3. Only northern tadpoles exposed to low food completely caught up in terms of metamorphic size, mainly by extending the larval period. Low food decreased survival and post-metamorphic jumping performance in southern, but not in northern tadpoles, suggesting that northern tadpoles have a better ability to compensate after periods of restricted food. 4. Both northern and southern tadpoles were able to metamorphose at the same size as control tadpoles after being exposed to low temperatures, indicating that consequences of variation in temperature and food availability differed for tadpoles. However, the combination of low temperatures and predation risk reduced survival in both southern and northern tadpoles. Also, predation risk decreased energy storage in both experiments. 5. Our results highlight the influence of climatic variation and the type of stressor as selective factors shaping compensatory strategies.

  • 20.
    Dahl, Emma
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Orizaola, German
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Winberg, Svante
    Uppsala University, Disciplinary Domain of Medicine and Pharmacy, Faculty of Medicine, Department of Neuroscience, Physiology.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Geographic variation in corticosterone response to chronic predator stress in tadpoles2012In: Journal of Evolutionary Biology, ISSN 1010-061X, E-ISSN 1420-9101, Vol. 25, no 6, p. 1066-1076Article in journal (Refereed)
    Abstract [en]

    Chronic stress often affects growth and development negatively, and these effects are often mediated via glucocorticoid hormones, which elevate during stress. We investigated latitudinal variation in corticosterone (CORT) response to chronic predator stress in Rana temporaria tadpoles along a 1500-km latitudinal cline in Sweden tadpoles, in a laboratory experiment. We hypothesized that more time-constrained high-latitude populations have evolved a lower CORT response to chronic stress to maintain higher growth under stressful conditions. Southern tadpoles had higher CORT content in response to predators after 1 day of exposure, whereas there was no increase in CORT in the northern populations. Two weeks later, there were no predator-induced CORT elevations. Artificially elevated CORT levels strongly decreased growth, development and survival in both northern and southern tadpoles. We suggest that the lower CORT response in high-latitude populations can be connected with avoidance of CORT-mediated reduction in growth and development, but also discuss other possible explanations.

  • 21. Dedukh, Dmitry
    et al.
    Mazepa, Glib
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Shabanov, Dmitry
    Rosanov, Juriy
    Litvinchuk, Spartak
    Borkin, Leo
    Saifitdinova, Alsu
    Krasikova, Alla
    Cytological maps of lampbrush chromosomes of European water frogs (Pelophylax esculentus complex) from the Eastern Ukraine2013In: BMC Genetics, ISSN 1471-2156, E-ISSN 1471-2156, Vol. 14, p. 26-Article in journal (Refereed)
    Abstract [en]

    Background: Hybridogenesis (hemiclonal inheritance) is a kind of clonal reproduction in which hybrids between parental species are reproduced by crossing with one of the parental species. European water frogs (Pelophylax esculentus complex) represent an appropriate model for studying interspecies hybridization, processes of hemiclonal inheritance and polyploidization. P. esculentus complex consists of two parental species, P. ridibundus (the lake frog) and P. lessonae (the pool frog), and their hybridogenetic hybrid - P. esculentus (the edible frog). Parental and hybrid frogs can reproduce syntopically and form hemiclonal population systems. For studying mechanisms underlying the maintenance of water frog population systems it is required to characterize the karyotypes transmitted in gametes of parental and different hybrid animals of both sexes. Results: In order to obtain an instrument for characterization of oocyte karyotypes in hybrid female frogs, we constructed cytological maps of lampbrush chromosomes from oocytes of both parental species originating in Eastern Ukraine. We further identified certain molecular components of chromosomal marker structures and mapped coilin-rich spheres and granules, chromosome associated nucleoli and special loops accumulating splicing factors. We recorded the dissimilarities between P. ridibundus and P. lessonae lampbrush chromosomes in the length of orthologous chromosomes, number and location of marker structures and interstitial (TTAGGG)(n)-repeat sites as well as activity of nucleolus organizer. Satellite repeat RrS1 was mapped in centromere regions of lampbrush chromosomes of the both species. Additionally, we discovered transcripts of RrS1 repeat in oocytes of P. ridibundus and P. lessonae. Moreover, G-rich transcripts of telomere repeat were revealed in association with terminal regions of P. ridibundus and P. lessonae lampbrush chromosomes. Conclusions: The constructed cytological maps of lampbrush chromosomes of P. ridibundus and P. lessonae provide basis to define the type of genome transmitted within individual oocytes of P. esculentus females with different ploidy and from various population systems.

  • 22. Duarte, Helder
    et al.
    Tejedo, Miguel
    Katzenberger, Marco
    Marangoni, Federico
    Baldo, Diego
    Francisco Beltran, Juan
    Andrea Marti, Dardo
    Richter Boix, Alex
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Gonzalez-Voyer, Alejandro
    Can amphibians take the heat?: Vulnerability to climate warming in subtropical and temperate larval amphibian communities2012In: Global Change Biology, ISSN 1354-1013, E-ISSN 1365-2486, Vol. 18, no 2, p. 412-421Article in journal (Refereed)
    Abstract [en]

    Predicting the biodiversity impacts of global warming implies that we know where and with what magnitude these impacts will be encountered. Amphibians are currently the most threatened vertebrates, mainly due to habitat loss and to emerging infectious diseases. Global warming may further exacerbate their decline in the near future, although the impact might vary geographically. We predicted that subtropical amphibians should be relatively susceptible to warming-induced extinctions because their upper critical thermal limits (CTmax) might be only slightly higher than maximum pond temperatures (Tmax). We tested this prediction by measuring CTmax and Tmax for 47 larval amphibian species from two thermally distinct subtropical communities (the warm community of the Gran Chaco and the cool community of Atlantic Forest, northern Argentina), as well as from one European temperate community. Upper thermal tolerances of tadpoles were positively correlated (controlling for phylogeny) with maximum pond temperatures, although the slope was steeper in subtropical than in temperate species. CTmax values were lowest in temperate species and highest in the subtropical warm community, which paradoxically, had very low warming tolerance (CTmaxTmax) and therefore may be prone to future local extinction from acute thermal stress if rising pond Tmax soon exceeds their CTmax. Canopy-protected subtropical cool species have larger warming tolerance and thus should be less impacted by peak temperatures. Temperate species are relatively secure to warming impacts, except for late breeders with low thermal tolerance, which may be exposed to physiological thermal stress in the coming years.

  • 23.
    Eggers, Sönke
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Low, Matthew
    Differential demographic responses of sympatric Parids to vegetation management in boreal forest2014In: Forest Ecology and Management, ISSN 0378-1127, E-ISSN 1872-7042, Vol. 319, p. 169-175Article in journal (Refereed)
    Abstract [en]

    Large-scale removal of small-diameter trees (i.e. thinning from below) in boreal forest can diminish niche diversity for birds that rely on a well-developed understory for nesting and foraging. Yet, few studies have examined how reduced niche diversity in managed forests affects fitness measures in closely-related species and the ability of competing species to co-exist. We related forest thinning to population trends of the willow tit Poecile montana (declining) and its dominant competitor the crested tit Lophophanes cristatus (stable), and conducted a 3-year comparative study to determine how variation in understory spruce density differentially influences survival and reproduction in these species. In line with our prediction that crested tits would gain resource priority under conditions of reduced forest understory complexity, willow tits and their nestlings suffered a disproportionate decline in both nest and adult survival prospects relative to crested tits as understoiy spruce density declined. Willow tits also had increased numbers of tail feather fault bars with decreasing understory complexity, further supporting the idea that willow tits suffer from food shortage and increased predation risk in areas of reduced understory. The long-term population declines of willow tits in boreal forest appears linked to large scale harvest of small-diameter spruce trees that provide important understory vegetation. A patchy arrangement of different thinning treatments through 'Understory Retention Thinning' (URT) may provide a cost-effective way to restore long-term structural complexity and biodiversity in densely stocked conifer stands. (C) 2014 Elsevier B.V. All rights reserved.

  • 24.
    Ekblom, Robert
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Evaluation of the analysis of distance sampling data: a simulation study2010In: Ornis Svecica, ISSN 1102-6812, Vol. 20, p. 43-53Article in journal (Refereed)
    Abstract [en]

    Distance sampling is used to estimate number of indi- viduals in an area of interest. The idea is that with known distances to the observed individuals, one can model the probability of detection in relation to distance and thereby account for individuals that were not detected. Distances can be recorded either exactly or in discrete categories. In this study I validated the method using simulated dis- tance sampling data for two hypothetical bird species and compared the estimated density values to the known true densities. Generally the true densities and numbers of individuals were very similar to (and always within the 95% confidence interval of) the parameter estimates from the analysis of the simulated data. The analyses were also robust to modifications of the data such as truncation and

    grouping of the distances into discrete categories. The confidence intervals increased, however, when using only two distance groups. Given that critical assumptions of the model can be met in the field situation, distance data can thus be used in a wide range of bird studies to calculate reliable density estimates.

  • 25.
    Ekblom, Robert
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Balakrishnan, Christopher N.
    Burke, Terry
    Slate, Jon
    Digital gene expression analysis of the zebra finch genome2010In: BMC Genomics, ISSN 1471-2164, E-ISSN 1471-2164, Vol. 11, p. 219-Article in journal (Refereed)
    Abstract [en]

    Background: In order to understand patterns of adaptation and molecular evolution it is important to quantify both variation in gene expression and nucleotide sequence divergence. Gene expression profiling in non-model organisms has recently been facilitated by the advent of massively parallel sequencing technology. Here we investigate tissue specific gene expression patterns in the zebra finch (Taeniopygia guttata) with special emphasis on the genes of the major histocompatibility complex (MHC). Results: Almost 2 million 454-sequencing reads from cDNA of six different tissues were assembled and analysed. A total of 11,793 zebra finch transcripts were represented in this EST data, indicating a transcriptome coverage of about 65%. There was a positive correlation between the tissue specificity of gene expression and non-synonymous to synonymous nucleotide substitution ratio of genes, suggesting that genes with a specialised function are evolving at a higher rate (or with less constraint) than genes with a more general function. In line with this, there was also a negative correlation between overall expression levels and expression specificity of contigs. We found evidence for expression of 10 different genes related to the MHC. MHC genes showed relatively tissue specific expression levels and were in general primarily expressed in spleen. Several MHC genes, including MHC class I also showed expression in brain. Furthermore, for all genes with highest levels of expression in spleen there was an overrepresentation of several gene ontology terms related to immune function. Conclusions: Our study highlights the usefulness of next-generation sequence data for quantifying gene expression in the genome as a whole as well as in specific candidate genes. Overall, the data show predicted patterns of gene expression profiles and molecular evolution in the zebra finch genome. Expression of MHC genes in particular, corresponds well with expression patterns in other vertebrates.

  • 26.
    Ekblom, Robert
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    French, Lisa
    Slate, Jon
    Burke, Terry
    Evolutionary Analysis and Expression Profiling of Zebra Finch Immune Genes2010In: Genome Biology and Evolution, ISSN 1759-6653, Vol. 2, p. 781-790Article in journal (Refereed)
    Abstract [en]

    Genes of the immune system are generally considered to evolve rapidly due to host-parasite coevolution. They are therefore of great interest in evolutionary biology and molecular ecology. In this study, we manually annotated 144 avian immune genes from the zebra finch (Taeniopygia guttata) genome and conducted evolutionary analyses of these by comparing them with their orthologs in the chicken (Gallus gallus). Genes classified as immune receptors showed elevated d(N)/d(S) ratios compared with other classes of immune genes. Immune genes in general also appear to be evolving more rapidly than other genes, as inferred from a higher d(N)/d(S) ratio compared with the rest of the genome. Furthermore, ten genes (of 27) for which sequence data were available from at least three bird species showed evidence of positive selection acting on specific codons. From transcriptome data of eight different tissues, we found evidence for expression of 106 of the studied immune genes, with primary expression of most of these in bursa, blood, and spleen. These immune-related genes showed a more tissue-specific expression pattern than other genes in the zebra finch genome. Several of the avian immune genes investigated here provide strong candidates for in-depth studies of molecular adaptation in birds.

  • 27.
    Ekblom, Robert
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Galindo, Juan
    Applications of next generation sequencing in molecular ecology of non-model organisms2011In: Heredity, ISSN 0018-067X, E-ISSN 1365-2540, Vol. 107, no 1, p. 1-15Article, review/survey (Refereed)
    Abstract [en]

    As most biologists are probably aware, technological advances in molecular biology during the last few years have opened up possibilities to rapidly generate large-scale sequencing data from non-model organisms at a reasonable cost. In an era when virtually any study organism can 'go genomic', it is worthwhile to review how this may impact molecular ecology. The first studies to put the next generation sequencing (NGS) to the test in ecologically well-characterized species without previous genome information were published in 2007 and the beginning of 2008. Since then several studies have followed in their footsteps, and a large number are undoubtedly under way. This review focuses on how NGS has been, and can be, applied to ecological, population genetic and conservation genetic studies of non-model species, in which there is no (or very limited) genomic resources. Our aim is to draw attention to the various possibilities that are opening up using the new technologies, but we also highlight some of the pitfalls and drawbacks with these methods. We will try to provide a snapshot of the current state of the art for this rapidly advancing and expanding field of research and give some likely directions for future developments.

  • 28.
    Ekblom, Robert
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Stapley, Jessica
    Ball, Alex D.
    Birkhead, Tim
    Burke, Terry
    Slate, Jon
    Genetic mapping of the major histocompatibility complex in the zebra finch (Taeniopygia guttata)2011In: Immunogenetics, ISSN 0093-7711, E-ISSN 1432-1211, Vol. 63, no 8, p. 523-530Article in journal (Refereed)
    Abstract [en]

    Genes of the major histocompatibility complex (MHC) have received much attention in immunology, genetics, and ecology because they are highly polymorphic and play important roles in parasite resistance and mate choice. Until recently, the MHC of passerine birds was not well-described. However, the genome sequencing of the zebra finch (Taeniopygia guttata) has partially redressed this gap in our knowledge of avian MHC genes. Here, we contribute further to the understanding of the zebra finch MHC organization by mapping SNPs within or close to known MHC genes in the zebra finch genome. MHC class I and IIB genes were both mapped to zebra finch chromosome 16, and there was no evidence that MHC class I genes are located on chromosome 22 (as suggested by the genome assembly). We confirm the location in the MHC region on chromosome 16 for several other genes (BRD2, FLOT1, TRIM7.2, GNB2L1, and CSNK2B). Two of these (CSNK2B and FLOT1) have not previously been mapped in any other bird species. In line with previous results, we also find that orthologs to the immune-related genes B-NK and CLEC2D, which are part of the MHC region in chicken, are situated on zebra finch chromosome Z and not among other MHC genes in the zebra finch.

  • 29. Gabriel, Sofia I.
    et al.
    Jóhannesdóttir, Fríða
    Jones, Eleanor P.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Searle, Jeremy B.
    Colonization, mouse-style2010In: BMC Biology, ISSN 1741-7007, E-ISSN 1741-7007, Vol. 8, p. 131-Article in journal (Other academic)
    Abstract [en]

    Several recent papers, including one in BMC Evolutionary Biology, examine the colonization history of house mice. As well as background for the analysis of mouse adaptation, such studies offer a perspective on the history of movements of the humans that accidentally transported the mice.

  • 30. Garriga, Nuria
    et al.
    Santos, Xavier
    Montori, Albert
    Richter-Boix, Alex
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Franch, Marc
    Llorente, Gustavo A.
    Are protected areas truly protected?: The impact of road traffic on vertebrate fauna2012In: Biodiversity and Conservation, ISSN 0960-3115, E-ISSN 1572-9710, Vol. 21, no 11, p. 2761-2774Article in journal (Refereed)
    Abstract [en]

    The extension of road networks is considered one of the major factors affecting fauna survival. Roadkill has been documented widely and affects all taxonomic groups. Although roadkill is associated mainly with traffic density, some life-history traits of species and the area surrounding roads are expected to modify number of roadkills both taxonomically and geographically. Here we studied the number of roadkills of vertebrates in an extensive region in the northeastern Iberian Peninsula. We surveyed 820 km of 41 roads in two different seasons (spring and autumn), that differ in traffic intensity. In addition, we covered zones with distinct climatic characteristics and levels of protection of the surrounding habitats. Amphibians showed the highest number of roadkills whereas reptiles, birds and mammals had similar rates. General Linear Model tests showed no differences in roadkills by climatic region; however, differences in number of roadkills were linked to protection status, with the highest number of casualties in highly protected areas. Redundancy Analysis demonstrated that the number of amphibians and reptiles killed was associated with roads in highly protected areas whereas that of mammals and birds was linked to unprotected areas. Protected areas often receive many visitors, which in turn may increase wildlife casualties as a result of greater traffic density. We recommend that correction measures be taken to reduce the high number of vertebrate fauna killed along roads that cross protected areas.

  • 31. Givskov Sørensen, Jesper
    et al.
    Loeschcke, Volker
    Merilä, Juha
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Effects of predator exposure on Hsp70 expression and survival in tadpoles of the Common Frog (Rana temporaria)2011In: Canadian Journal of Zoology, ISSN 0008-4301, E-ISSN 1480-3283, Vol. 89, no 12, p. 1249-1255Article in journal (Refereed)
    Abstract [en]

    Predator-induced changes in prey behavior and morphology are widespread, but little is known about physiological and cellular-level responses in prey in response to predation risk. We investigated whether predator (larvae of the dragonfly Aeshna Fabricius, 1775) presence elevated the expression level of heat-shock protein 70 (Hsp70)-a commonly found response to stress-in tadpoles of the Common Frog (Rana temporaria L., 1758). In another experiment, we tested the survival of tadpoles in the presence of a free-ranging predator. Prior to this encounter, the tadpoles were exposed to either an Hsp-inducing environmental stress in the form of heat (31 degrees C) or to predator cues from a caged predator. We found no evidence for increased Hsp70 expression in tadpoles either in the presence of fed or starved predators. We did not find any effects of prior exposure to neither heat nor predator presence on survival at the end of experiment. Our results do not point to either Hsp70-mediated effect of predator-induced responses or to beneficial effects of the stress response on survival under predation risk.

  • 32.
    Griesser, Michael
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Lagerberg, Sarah
    Long-term effects of forest management on territory occupancy and breeding success of an open-nesting boreal bird species, the Siberian jay2012In: Forest Ecology and Management, ISSN 0378-1127, E-ISSN 1872-7042, Vol. 271, p. 58-64Article in journal (Refereed)
    Abstract [en]

    In large parts of the world forests are intensively managed, affecting stand structure and biodiversity. Since re-growth of trees can be slow and management cycles can span over 100 years, long-term longitudinal data are needed to assess the effects of forestry on organisms living in these habitats. Here, we draw upon 50 years of population data of an open-nesting, sedentary bird species, typically of boreal forests, the Siberian jay (Perisoreus infaustus). Forests at the study site were managed by clear-cutting and re-plantation, and a management cycle spanning over 120 years. We tracked changes in forest structure in the nest vicinity and the whole territory with the help of aerial images, and linked them to territory occupancy and offspring production in 13 territories. While territories were occupied in 77.2% of all instances, reproduction was only successful in 19.6% of all instances. Both parameters were positively influenced by a large proportion of unthinned forests older than 50 years, which provide visual protection from nest and adult predators. Thus, it is crucial to assess both territory occupancy and reproductive output to understand the effects of forestry on bird populations. Moreover, our results suggest that in forests with slow turnover cycles, management which avoids thinning (such as green tree retention forestry) can promote population persistence of open-nesting bird species.

  • 33.
    Griesser, Michael
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Ma, Qi
    Uppsala University, Disciplinary Domain of Science and Technology, Mathematics and Computer Science, Department of Mathematics, Analysis and Applied Mathematics.
    Webber, Simone
    Bowgen, Katharine
    Sumpter, David J. T.
    Uppsala University, Disciplinary Domain of Science and Technology, Mathematics and Computer Science, Department of Mathematics, Analysis and Applied Mathematics.
    Understanding Animal Group-Size Distributions2011In: PLoS ONE, ISSN 1932-6203, E-ISSN 1932-6203, Vol. 6, no 8, p. e23438-Article in journal (Refereed)
    Abstract [en]

    One of the most striking aspects of animal groups is their remarkable variation in size, both within and between species. While a number of mechanistic models have been proposed to explain this variation, there are few comprehensive datasets against which these models have been tested. In particular, we only vaguely understand how environmental factors and behavioral activities affect group-size distributions. Here we use observations of House sparrows (Passer domesticus) to investigate the factors determining group-size distribution. Over a wide range of conditions, we observed that animal group sizes followed a single parameter distribution known as the logarithmic distribution. This single parameter is the mean group size experienced by a randomly chosen individual (including the individual itself). For sparrows, the experienced mean group size, and hence the distribution, was affected by four factors: morning temperature, place, behavior and the degree of food spillage. Our results further indicate that the sparrows regulate the mean group size they experience, either by groups splitting more or merging less when local densities are high. We suggest that the mean experienced group size provides a simple but general tool for assessing the ecology and evolution of grouping.

  • 34. Hangartner, S.
    et al.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Raesaenen, K.
    The quantitative genetic basis of adaptive divergence in the moor frog (Rana arvalis) and its implications for gene flow2012In: Journal of Evolutionary Biology, ISSN 1010-061X, E-ISSN 1420-9101, Vol. 25, no 8, p. 1587-1599Article in journal (Refereed)
    Abstract [en]

    Knowledge on the relative contribution of direct genetic, maternal and environmental effects to adaptive divergence is important for understanding the drivers of biological diversification. The moor frog (Rana arvalis) shows adaptive divergence in embryonic and larval fitness traits along an acidification gradient in south-western Sweden. To understand the quantitative genetic basis of this divergence, we performed reciprocal crosses between three divergent population pairs and reared embryos and larvae at acid and neutral pH in the laboratory. Divergence in embryonic acid tolerance (survival) was mainly determined by maternal effects, whereas the relative contributions of maternal, additive and nonadditive genetic effects in larval life-history traits differed between traits, population pairs and rearing environments. These results emphasize the need to investigate the quantitative genetic basis of adaptive divergence in multiple populations and traits, as well as different environments. We discuss the implications of our findings for maintenance of local adaptation in the context of migrant and hybrid fitness.

  • 35. Hangartner, Sandra
    et al.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Effects of the disinfectant Virkon S on early life-stages of the moor frog (Rana arvalis)2012In: Amphibia-Reptilia, ISSN 0173-5373, E-ISSN 1568-5381, Vol. 33, no 3-4, p. 349-353Article in journal (Refereed)
    Abstract [en]

    Emerging diseases, such as the chytrid fungus Batrachochytrium dendrobatidis, contribute to global population declines of amphibians. Virkon S is one of the most commonly used disinfectants to reduce risk of spreading such pathogens. Virkon S is classified as harmful to aquatic organisms, but until today no negative effects on tadpoles have been reported. We studied the effects of three concentrations of Virkon S on early life-stages (embryos and hatchlings) of the moor frog Rana anvils. Overall, Virkon S had no significant effects. However, hatching success was highest in the control treatment, suggesting that Virkon S may have weak negative effects on amphibian embryos. We suggest that further studies are needed to assess the negative effect of Virkon S on amphibians, and recommend that Virkon S is used with care and a minimized run-off into natural wetlands.

  • 36. Hangartner, Sandra
    et al.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Räsänen, Katja
    Adaptive Divergence in Moor Frog (Rana Arvalis) Populations Along an Acidification Gradient: Inferences from QST-FST Correlations2012In: Evolution, ISSN 0014-3820, E-ISSN 1558-5646, Vol. 66, no 3, p. 867-881Article in journal (Refereed)
    Abstract [en]

    Microevolutionary responses to spatial variation in the environment seem ubiquitous, but the relative role of selection and neutral processes in driving phenotypic diversification remain often unknown. The moor frog (Rana arvalis) shows strong phenotypic divergence along an acidification gradient in Sweden. We here used correlations among population pairwise estimates of quantitative trait (PST or QST from common garden estimates of embryonic acid tolerance and larval life-history traits) and neutral genetic divergence (FST from neutral microsatellite markers), as well as environmental differences (pond pH, predator density, and latitude), to test whether this phenotypic divergence is more likely due to divergent selection or neutral processes. We found that trait divergence was more strongly correlated with environmental differences than the neutral marker divergence, suggesting that divergent natural selection has driven phenotypic divergence along the acidification gradient. Moreover, pairwise PSTs of embryonic acid tolerance and QSTs of metamorphic size were strongly correlated with breeding pond pH, whereas pairwise QSTs of larval period and growth rate were more strongly correlated with geographic distance/latitude and predator density, respectively. We suggest that incorporating measurements of environmental variation into QSTFST studies can improve our inferential power about the agents of natural selection in natural populations.

  • 37. Hangartner, Sandra
    et al.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Räsänen, Katja
    Adaptive divergence of the moor frog (Rana arvalis) along an acidification gradient2011In: BMC Evolutionary Biology, ISSN 1471-2148, E-ISSN 1471-2148, Vol. 11, p. 366-Article in journal (Refereed)
    Abstract [en]

    Background: Environmental stress can result in strong ecological and evolutionary effects on natural populations, but to what extent it drives adaptive divergence of natural populations is little explored. We used common garden experiments to study adaptive divergence in embryonic and larval fitness traits (embryonic survival, larval growth, and age and size at metamorphosis) in eight moor frog, Rana arvalis, populations inhabiting an acidification gradient (breeding pond pH 4.0 to 7.5) in southwestern Sweden. Embryos were raised until hatching at three (pH 4.0, 4.3 and 7.5) and larvae until metamorphosis at two (pH 4.3 and 7.5) pH treatments. To get insight into the putative selective agents along this environmental gradient, we measured relevant abiotic and biotic environmental variables from each breeding pond, and used linear models to test for phenotype-environment correlations.

    Results: We found that acid origin populations had higher embryonic and larval acid tolerance (survival and larval period were less negatively affected by low pH), higher larval growth but slower larval development rates, and metamorphosed at a larger size. The phenotype-environment correlations revealed that divergence in embryonic acid tolerance and metamorphic size correlated most strongly with breeding pond pH, whereas divergence in larval period and larval growth correlated most strongly with latitude and predator density, respectively.

    Conclusion: Our results suggest that R. arvalis has diverged in response to pH mediated selection along this acidification gradient. However, as latitude and pH were closely spatially correlated in this study, further studies are needed to disentangle the specific agents of natural selection along acidification gradients. Our study highlights the need to consider the multiple interacting selective forces that drive adaptive divergence of natural populations along environmental stress gradients.

  • 38. Hartman, Goran
    et al.
    Kolzsch, Andrea
    Larsson, Karl
    Nordberg, Marcus
    Höglund, Jacob
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Trends and population dynamics of a Velvet Scoter (Melanitta fusca) population: influence of density dependence and winter climate2013In: Journal of Ornithology = Journal fur Ornithologie, ISSN 0021-8375, E-ISSN 1439-0361, Vol. 154, no 3, p. 837-847Article in journal (Refereed)
    Abstract [en]

    As many seaduck populations around the world have been reported to be in decline, there is an increasing demand for knowledge about intrinsic and extrinsic factors determining population dynamics of these species. In this study, we analyzed long-term dynamics of the summer population of Velvet Scoters (Melanitta fusca) breeding in the Aland archipelago in the Baltic Sea; in particular, we examined the influence of winter weather and density dependence on population change. The studied population exhibited substantial fluctuations but only a weak negative trend during the total period of 58 years (1949-2007), and no significant trend at all during the latter three decades of the study (1977-2007). We tested for density dependence and incorporated the winter North Atlantic Oscillation index into the model to test for effects of winter conditions. Our final model explained 56.3 % of the variance of population growth of the studied population. Delayed density dependence explained 29.7 %, pre-breeding climate 8.3 %, and post-breeding climate 18.3 % of the variance. That breeding success is density dependent in a delayed manner is in accordance with the apprehension that Velvet Scoters breed at the age of 2 years. We conclude that density dependence and winter conditions must be taken into consideration when discussing population changes in seaducks in general and the Velvet Scoter in particular.

  • 39.
    Hesson, Jenny C.
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Animal ecology.
    Rettich, Frantisek
    Merdic, Enrih
    Vignjevic, Goran
    Östman, Örjan
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Animal ecology.
    Schäfer, Martina
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Schaffner, Francis
    Foussadier, Remi
    Besnard, Gilles
    Medlock, Jolyon
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics.
    Scholte, Ernst-Jan
    Lundström, Jan O.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Animal ecology.
    The arbovirus vector Culex torrentium is more prevalent than Culex pipiens in northern and central Europe2014In: Medical and Veterinary Entomology, ISSN 0269-283X, E-ISSN 1365-2915, Vol. 28, no 2, p. 179-186Article in journal (Refereed)
    Abstract [en]

    Two species of arbovirus vector, Culex torrentium and Culex pipiens (Diptera: Culicidae), occur in several European countries, but difficulties in their accurate identification and discrimination have hampered both detailed and large-scale distribution and abundance studies. Using a molecular identification method, we identified to species 2559 larvae of Cx. pipiens/torrentium collected from 138 sites in 13 European countries ranging from Scandinavia to the Mediterranean coast. In addition, samples of 1712 males of Cx. pipiens/torrentium collected at several sites in the Czech Republic were identified to species based on the morphology of their hypopygia. We found that the two species occur together in large areas of Europe, and that Cx. torrentium dominates in northern Europe and Cx. pipiens dominates south of the Alps. The transition in dominance occurs in central Europe, where both species are roughly equally common. There was a strong correlation between the length of the growing season at different sites and occurrences of the two species. As the growing season increases, the proportion and detection of Cx. torrentium decrease, whereas those of Cx. pipiens increase. The present findings have important consequences for the interpretation of the results of studies on major enzootic and link-vectors of mosquito-borne bird-associated viruses (i.e. Sindbis, West Nile and Usutu viruses), especially in central Europe and Scandinavia.

  • 40.
    Hesson, Jenny C.
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Östman, Örjan
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Schäfer, Martina
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Lundström, Jan O.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Geographic distribution and relative abundance of the sibling vector species Culex torrentium and Culex pipiens in Sweden2011In: Vector Borne and Zoonotic Diseases, ISSN 1530-3667, E-ISSN 1557-7759, Vol. 11, no 10, p. 1383-1389Article in journal (Refereed)
    Abstract [en]

    Culex torrentium and Culex pipiens are sibling species and potential viral vectors that coexist in Europe. Larvae and females of the two species are morphologically almost identical, and reliable identification can only be done on males. To investigate the distribution and relative abundance of the two species in Sweden, we collected Culex larvae from sites spread over the country, identified them as Culex pipiens/torrentium based on morphology, and identified them to species using a recently developed restriction enzyme method. Cx. torrentium was the dominant species (89%, n = 1012) and it occurred in 48 of the 49 sites investigated, and also dominated in most of the study sites. The proportion of Cx. pipiens larvae in relation to Cx. torrentium collected at each site decreased with both increasing latitude and altitude, and the presence of Cx. pipiens decreased with latitude. In addition, Cx. pipiens/torrentium females were sampled with Centres for Disease Control light traps baited with carbon dioxide. The overall country mean was 4.0 Cx. pipiens/torrentium caught per trap night, with decreasing numbers of Cx. pipiens/torrentium caught per trap night with increasing latitude. Thus, the abundance of Cx. pipiens/torrentium decreased, but the proportion Cx. torrentium increased, with increasing latitude. This is the first study that shows the vast dominance of Cx. torrentium over Cx. pipiens in Sweden. The unexpected dominance of Cx. torrentium highlights the importance of distinguishing between the two species in studies of Culex-borne arboviruses in Europe.

  • 41. Hettyey, A.
    et al.
    Vincze, K.
    Zsarnoczai, S.
    Hoi, H.
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Costs and benefits of defences induced by predators differing in dangerousness2011In: Journal of Evolutionary Biology, ISSN 1010-061X, E-ISSN 1420-9101, Vol. 24, no 5, p. 1007-1019Article in journal (Refereed)
    Abstract [en]

    While theoretical studies predict that inducible defences should be fine-tuned according to the qualities of the predator, very few studies have investigated how dangerousness of predators, i.e. the rate at which predators kill prey individuals, affects the strength of phenotypic responses and resulting benefits and costs of induced defences. We performed a comprehensive study on fitness consequences of predator-induced responses by involving four predators (leech, water scorpion, dragonfly larva and newt), evaluating costs and benefits of responses, testing differences in dangerousness between predators and measuring responses in several life history traits of prey. We raised Rana dalmatina tadpoles in the presence of free-ranging predators, in the presence of caged predators, and exposed naive and experienced tadpoles to free-ranging predators. Tadpoles adjusted the intensities of their behavioural and morphological defences to predator dangerousness. Survival was lower in the nonlethal presence of the most dangerous predator, while we could not detect costs of induced defences at or after metamorphosis. When exposed to free-ranging predators, small, but not large, tadpoles benefited from exhibiting an induced phenotype in terms of elevated survival when compared to naive tadpoles, but we did not observe higher survival either in tadpoles exhibiting more extreme phenotypes or in tadpoles exposed to the type of predator they were raised with. These results indicate that while predator-induced defences can mirror dangerousness of predators, costs and benefits do not necessarily scale to the magnitude of plastic responses.

  • 42. Hettyey, Attila
    et al.
    Zsarnoczai, Szilvia
    Vincze, Krisztina
    Hoi, Herbert
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Interactions between the information content of different chemical cues affect induced defences in tadpoles2010In: Oikos, ISSN 0030-1299, E-ISSN 1600-0706, Vol. 119, no 11, p. 1814-1822Article in journal (Refereed)
    Abstract [en]

    Animals often alter their behaviour, morphology and physiology in the presence of predators. These induced defences can be fine-tuned by a variety of environmental factors such as predator species, acute predation risk or food availability. It has, however, remained unclear what cues influence the extent and quality of induced defences and how the information content of these cues interact to determine the development of antipredator defences. We performed an experiment to study the significance of direct chemical cues, originating from the predators themselves, and indirect cues, released by attacked or consumed prey, for phenotypic responses in Rana dalmatina tadpoles. We reared tadpoles in the presence of caged predators (Triturus vulgaris, Aeshna cyanea) fed either one or three tadpoles every other day outside the tadpole-rearing tanks. Fifteen hours after food provisioning, predators were put back into the tanks containing focal tadpoles either after washing (direct + digestion-released cues) or with the water containing remnants of the prey (direct + all types of indirect cues). Our results suggest that direct cues together with digestion-released cues can be sufficient to induce strong antipredator responses. Induced defences depended on both direct cues, affecting predator-specific responses, and the quantity of indirect cues, resulting in graded responses to differences in predation threat. Moreover, direct and indirect cues interacted in behaviour, resulting in predator-specific graded responses. We also observed a decrease in the extent of predator-induced responses in large tadpoles as compared to small ones. Our results, thus, suggest that prey integrate multiple cues about predators to optimize induced defences and that this process changes during ontogeny.

  • 43.
    Hjernquist, Mårten B.
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Söderman, Fredrik
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Jonsson, K. Ingemar
    Herczeg, Gabor
    Laurila, Anssi
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Merila, Juha
    Seasonality determines patterns of growth and age structure over a geographic gradient in an ectothermic vertebrate2012In: Oecologia, ISSN 0029-8549, E-ISSN 1432-1939, Vol. 170, no 3, p. 641-649Article in journal (Refereed)
    Abstract [en]

    Environmental variation connected with seasonality is likely to affect the evolution of life-history strategies in ectotherms, but there is no consensus as to how important life-history traits like body size are influenced by environmental variation along seasonal gradients. We compared adult body size, skeletal growth, mean age, age at first reproduction and longevity among 11 common frog (Rana temporaria) populations sampled along a 1,600-km-long latitudinal gradient across Scandinavia. Mean age, age at first reproduction and longevity increased linearly with decreasing growth season length. Lifetime activity (i.e. the estimated number of active days during life-time) was highest at mid-latitudes and females had on average more active days throughout their lives than males. Variation in body size was due to differences in lifetime activity among populations-individuals (especially females) were largest where they had the longest cumulative activity period-as well as to differences between populations in skeletal growth rate as determined by skeletochronological analyses. Especially, males grew faster at intermediate latitudes. While life-history trait variation was strongly associated with latitude, the direction and shape of these relationships were sex- and trait-specific. These context-dependent relationships may be the result of life-history trade-offs enforced by differences in future reproductive opportunities and time constraints among the populations. Thus, seasonality appears to be an important environmental factor shaping life-history trait variation in common frogs.

  • 44.
    Husby, Arild
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Animal ecology.
    Ekblom, Robert
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology. Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Evolutionary Biology.
    Qvarnström, Anna
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Animal ecology.
    Let's talk turkey: immune competence in domestic and wild fowl2011In: Heredity, ISSN 0018-067X, E-ISSN 1365-2540, Vol. 107, no 2, p. 103-104Article in journal (Other academic)
  • 45.
    Höglund, Jacob
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Cortazar-Chinarro, Maria
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Jarnemo, Anders
    Thulin, Carl-Gustaf
    Genetic variation and structure in Scandinavian red deer (Cervus elaphus): influence of ancestry, past hunting, and restoration management2013In: Biological Journal of the Linnean Society, ISSN 0024-4066, E-ISSN 1095-8312, Vol. 109, no 1, p. 43-53Article in journal (Refereed)
    Abstract [en]

    In the 19th century, the red deer (Cervus elaphus) population in Sweden experienced a rapid decline in numbers and distribution. A small population was, however, remnant in the southernmost province (Skane) of the country, presumably corresponding to the nominate form of red deer (Cervus elaphus elaphusLinnaeus, 1758). After management, reintroductions, and supplementary release during the 20th century the Swedish C.elaphus population recovered. The recovery was partially uncontrolled, and included introductions of C.elaphus of continental origin. In northern central Sweden (Jamtland) the current C.elaphus population may stem from natural colonization from Norway and/or from specimens of Swedish origin that have escaped from enclosures. To evaluate the status of the current, partially separated populations, we investigated variation at microsatellite markers in 157 C.elaphus specimens from ten locations in Sweden and Norway. Analyses suggest that the highest-likelihood phylogenetic structure among the individuals sampled is described four distinct genetic clusters: (1) animals from the province of Vastergotland in south-western Sweden; (2) deer from the southernmost province of Skane; (3) deer from the provinces Jamtland, Blekinge, and Vastmanland; and (4) Norwegian deer. Cervus elaphus from a captive herd at the Skane Zoo cluster with deer from Skane or deer from Vastergotland, depending on the method of analysis. A number of populations in Sweden may genetically match the nominate form of red deer (C.e.elaphus). The recently established C.elaphus population in Jamtland seems to stem mainly from escapees from enclosures, with a mixed ancestry from the wild remnant population in Skane and continental deer, whereas the influx from Norway is minor, if any. Our results show the need for a detailed assessment of genetic differentiation, and emphasize the value of local management plans when planning and managing introductions.

  • 46.
    Höglund, Jacob
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Larsson, Jobs Karl
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Corrales, Carolina
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Santafé, Guzman
    Baines, David
    Segelbacher, Gernot
    Genetic structure among black grouse in Britain: implications for designing conservation units2011In: Animal Conservation, ISSN 1367-9430, E-ISSN 1469-1795, Vol. 14, no 4, p. 400-408Article in journal (Refereed)
    Abstract [en]

    Black grouse in Britain have faced contraction of their range and have declined in numbers during the recent decades. As such, the species is a conservation concern in the UK. In order to aid conservation decisions, the terms Evolutionary Significant Unit (ESU) and Management Unit (MU) have been proposed. An ESU is an independently evolving evolutionary lineage defined by being reciprocally monophyletic for mitochondrial alleles, and which is significantly different from other lineages with regard to nuclear alleles, whereas an MU is operationally defined by only the latter criterion. Using mitochondrial sequences and nuclear microsatellite loci, we failed to find evidence that British black grouse is an ESU. However, British black grouse are sufficiently different from continental black grouse both with respect to mitochondrial and nuclear data to regard them as a separate MU. Furthermore, we present genetic data which suggest that British black grouse presently occur in what are probably three demographically independent units (roughly corresponding to Wales, northern England/southern Scotland and northern Scotland), which are genetically differentiated. The two southern units (Wales and northern England/southern Scotland) have lower genetic diversity and show signs of having lost genetic variability

  • 47.
    Johansson, Frank
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Lind, Martin I.
    Ingvarsson, Par K.
    Bokma, Folmer
    Evolution of the G-matrix in life history traits in the common frog during a recent colonisation of an island system2012In: Evolutionary Ecology, ISSN 0269-7653, E-ISSN 1573-8477, Vol. 26, no 4, p. 863-878Article in journal (Refereed)
    Abstract [en]

    Studies of genetic correlations between traits that ostensibly channel the path of evolution away from the direction of natural selection require information on key aspects such as ancestral phenotypes, the duration of adaptive evolution, the direction of natural selection, and genetic covariances. In this study we provide such information in a frog population system. We studied adaptation in life history traits to pool drying in frog populations on islands of known age, which have been colonized from a mainland population. The island populations show strong local adaptation in development time and size. We found that the first eigenvector of the variance-covariance matrix (g (max)) had changed between ancestral mainland populations and newly established island populations. Interestingly, there was no divergence in g (max) among island populations that differed in their local adaptation in development time and size. Thus, a major change in the genetic covariance of life-history traits occurred in the colonization of the island system, but subsequent local adaptation in development time took place despite the constraints imposed by the genetic covariance structure.

  • 48. Johansson, Helena
    et al.
    Ingvarsson, Par K.
    Johansson, Frank
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Cross-species amplification and development of microsatellites for six species of European Coenagrionid damselflies2012In: Conservation Genetics Resources, ISSN 1877-7252, E-ISSN 1877-7260, Vol. 4, no 1, p. 191-196Article in journal (Refereed)
    Abstract [en]

    We describe the cross-amplification and development of new loci for six species of closely related European damselflies. First, twenty-nine published microsatellites for the damselflies Coenagrion puella and C. mercuriale were multiplexed using M13-tagged primers, tested on 23 individuals, and then cross-species amplified on 21-26 individuals of C. armatum, C. johanssoni, C. pulchellum and C. scitulum. Second, sixteen new primers were developed for use in C. armatum, C. johanssoni and C. scitulum, and screened on 21 individuals. Values for observed heterozygosities and number of alleles ranged between 0.00-0.87 and 2-19 respectively (over all loci and species). For all species the tested loci provide a minimum of 1-8 usable markers for population genetic studies.

  • 49.
    Johansson, Magnus P.
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Mcmahon, Barry J.
    Höglund, Jacob
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Segelbacher, Gernot
    Amplification success of multilocus genotypes from feathers found in the field compared with feathers obtained from shot birds2012In: Ibis, ISSN 0019-1019, E-ISSN 1474-919X, Vol. 154, no 1, p. 15-20Article in journal (Refereed)
    Abstract [en]

    Effective DNA extraction methods from bird feathers have facilitated non-invasive sampling, leading to the suggestion that feathers are a great source for genetic studies. However, few studies have assessed whether all feathers can be used or provide equal numbers of useful templates. In this study, feathers collected in various ways from Red Grouse Lagopus lagopus were examined to establish the quality of DNA extracted. Individual samples were classified into two categories according to whether they were collected from shot birds or found in the field. DNA was extracted from all samples and genotyped at 19 microsatellite loci. PCR products were analysed on a MegaBACE 1000. A total of 93% of the shot category produced a genotype that was considered successful (i.e. 15 of 18 loci) and 23% of the collected category produced successful genotypes under the same criteria. There was a significant difference between shot and collected samples in genotyping success and the observed number of missing loci. Recommendations and best practices are discussed along with the utility of bird feathers as a source of DNA for population and conservation biology.

  • 50.
    Jones, Eleanor P.
    et al.
    Uppsala University, Disciplinary Domain of Science and Technology, Biology, Department of Ecology and Genetics, Population and Conservation Biology.
    Eager, Heidi M.
    Gabriel, Sofia I.
    Johannesdottir, Frioa
    Searle, Jeremy B.
    Genetic tracking of mice and other bioproxies to infer human history2013In: Trends in Genetics, ISSN 0168-9525, E-ISSN 1362-4555, Vol. 29, no 5, p. 298-308Article, review/survey (Refereed)
    Abstract [en]

    The long-distance movements made by humans through history are quickly erased by time but can be reconstructed by studying the genetic make-up of organisms that travelled with them. The phylogeography of the western house mouse (Mus musculus domesticus), whose current widespread distribution around the world has been caused directly by the movements of (primarily) European people, has proved particularly informative in a series of recent studies. The geographic distributions of genetic lineages in this commensal have been linked to the Iron Age movements within the Mediterranean region and Western Europe, the extensive maritime activities of the Vikings in the 9th to 11th centuries, and the colonisation of distant landmasses and islands by the Western European nations starting in the 15th century. We review here recent insights into human history based on phylogeographic studies of mice and other species that have travelled with humans, and discuss how emerging genomic methodologies will increase the precision of these inferences.

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