<p>In Sweden today, deciduous woodlands are often restricted to small isolated remnants of what it once was. Managing practices have changed concerning both cattle grazing and logging. Thus, dispersal, habitat requirements and their importance for forest species distributions become a relevant issue. Most of the species found in the deciduous flora are perennial herbs. I have examined population dynamics in the forest perennial <i>Sanicula europaea</i>, and its relation to environmental factors such as grazing, competition and spatial and temporal variation in such factors. Moreover I examined species distributions in relation to dispersal and habitat suitability in <i>S. europaea</i> and<i> Dentaria bulbifera</i>. To understand mechanisms behind the observed patterns I performed a number of experiments.</p><p>The results clearly demonstrated that the distribution and abundance of <i>S. europaea</i> was not dispersal limited, whereas the opposite held true for <i>D. bulbifera. Moreover</i>, mollusc exclusion increased recruitment thus influencing population dynamics in <i>D. bulbifera</i>. Leaf losses had negative effects on <i>S. europaea</i> individuals. These negative effects depended both on the extent, frequency and timing of the leaf losses and early losses were more severe than late. Population level effects of grazing were not negative as negative direct effects were counterblanced by positive indirect effects. The positive effects of grazing were mainly in terms of an increased recruitment. </p><p>In<i> S. europaea</i>, high variation in a life cycle transition was always coupled to low elasticity, and traits that varied much due to the examined environmental factors had little importance to population growth rate. A population level perspective is required to assess total effects of environmental factors. In perennial organisms such evaluations need to calculate integrated measures of the effects over the entire life cycle. Field studies spanning several years and demographic models are important to achieve these objectives.</p>

In Sweden today, deciduous woodlands are often restricted to small isolated remnants of what it once was. Managing practices have changed concerning both cattle grazing and logging. Thus, dispersal, habitat requirements and their importance for forest species distributions become a relevant issue. Most of the species found in the deciduous flora are perennial herbs. I have examined population dynamics in the forest perennial Sanicula europaea, and its relation to environmental factors such as grazing, competition and spatial and temporal variation in such factors. Moreover I examined species distributions in relation to dispersal and habitat suitability in S. europaea and Dentaria bulbifera. To understand mechanisms behind the observed patterns I performed a number of experiments. The results clearly demonstrated that the distribution and abundance of S. europaea was not dispersal limited, whereas the opposite held true for D. bulbifera. Moreover, mollusc exclusion increased recruitment thus influencing population dynamics in D. bulbifera. Leaf losses had negative effects on S. europaea individuals. These negative effects depended both on the extent, frequency and timing of the leaf losses and early losses were more severe than late. Population level effects of grazing were not negative as negative direct effects were counterblanced by positive indirect effects. The positive effects of grazing were mainly in terms of an increased recruitment. In S. europaea, high variation in a life cycle transition was always coupled to low elasticity, and traits that varied much due to the examined environmental factors had little importance to population growth rate. A population level perspective is required to assess total effects of environmental factors. In perennial organisms such evaluations need to calculate integrated measures of the effects over the entire life cycle. Field studies spanning several years and demographic models are important to achieve these objectives.

R??sum?? : La reproduction entra??ne des d??penses ??nerg??tiques importantes chez les femelles mammif??res. Ces d??penses sont suppos??es diminuer l'??nergie disponible pour d'autres traits positivement corr??l??s ?? l'aptitude ph??notypique et augmenter les besoins d'alimentation. Toutefois, des diff??rences individuelles dans la capacit?? d'acquisition et d'allocation peuvent masquer l'impact n??gatif de l'effort reproducteur. La manipulation exp??rimentale de l'effort reproducteur et le contr??le statistique des effets individuels sont deux approches puissantes et compl??mentaires mises en ??uvre dans le cadre de mon ??tude afin de contr??ler pour l'h??t??rog??n??it?? individuelle. Elles ont permis de montrer clairement un co??t de la reproduction chez le kangourou gris de l'Est (Macropus giganteus). Dans un premier temps, nous avons trouv?? que le co??t de la reproduction affectait le montant d'??nergie allou?? ?? certaines fonctions somatiques (CHAPITRE I). L'effort reproducteur diminuait le gain de masse et la croissance des jambes pour des intervalles de capture sup??rieurs ?? deux ans. Chez les femelles non manipul??es, un effet n??gatif sur le gain de masse ??tait aussi observable pour un intervalle inf??rieur ?? 3 ans. ?? l'??chelle de deux ??v??nements successifs de reproduction, le gain de masse et dans une moindre mesure la croissance des bras, mais non des jambes diminuaient avec un effort reproducteur plus important ?? la pr??c??dente reproduction. Ensuite, nous avons d??montr?? qu'il y avait un co??t de la reproduction en terme de futur succ??s reproducteur (CHAPITRE II). Les individus dont l'effort reproducteur avait ??t?? diminu?? exp??rimentalement produisaient davantage de jeunes qui survivaient au stade 'LPY', ??g??s approximativement de 7 mois, que les femelles contr??les. Ils diminuaient ??galement davantage leur taux de reproduction en allongeant l'intervalle entre les naissances, mais la survie au sevrage n'??tait pas affect??e. Le CHAPITRE III montre que les femelles adaptaient leur comportement alimentaire en fonction de leur statut reproducteur. En comparaison avec les individus manipul??s ou ayant perdu leur jeune, les femelles allaitantes augmentaient leur temps pass?? ?? s'alimenter durant la journ??e, l'intensit?? de leurs bouch??es et de leur mastication sans impliquer de compromis avec la vigilance. Nous avons aussi d??couvert que la survie au sevrage du jeune pr??c??dent menait ?? une augmentation du taux de bouch??es pour l'ann??e en cours. Les CHAPITRES I et II ont mis en ??vidence l'effet non n??gligeable des diff??rences individuelles sur la d??tection des co??ts de la reproduction. En effet, en l'absence de manipulation exp??rimentale ou de contr??le statistique, aucun compromis n'??tait d??tect?? autant en terme de croissance que de prochaine reproduction. Au contraire, des corr??lations positives entre l'effort reproducteur et les autres traits ont ??t?? trouv??es. Le CHAPITRE I sugg??rait notamment que cette variabilit?? du succ??s reproducteur ??tait li??e ?? l'h??t??rog??n??it?? individuelle dans le gain de masse maternelle qui augmentait la survie du jeune. Dans le CHAPITRE II, une corr??lation positive entre les probabilit??s d'avoir un 'LPY' lors de deux ??v??nements successifs de reproduction sugg??rait que certaines femelles ??taient capables de mener ?? bien ou non leur reproduction, mais cela ind??pendamment de l'effort reproducteur pr??c??dent. Enfin dans le dernier CHAPITRE (III), l'effet al??atoire ??tait significatif dans l'analyse de diff??rents comportements d'alimentation, ce qui pourrait ??tre li?? aux diff??rences de gain de masse des femelles pr??sent??es dans le CHAPITRE I. Certaines contraintes individuelles affectant le co??t de la reproduction ont ??t?? identifi??es. La masse et la condition corporelle augmentaient le succ??s reproducteur et diminuaient l'intervalle entre deux naissances successives (CHAPITRES I et II), mais contrairement ?? de pr??c??dentes ??tudes le comportement d'alimentation des femelles n'??tait pas affect?? par leur masse (CHAPITRE III). L'??ge des individus avait ??galement une influence. Les jeunes femelles croissaient davantage, avaient aussi une prise alimentaire plus importante et subissaient un co??t de reproduction sup??rieur. Ce dernier se traduisait par un taux d'??chec plus ??lev?? ?? la reproduction suivante si les jeunes femelles avaient eu un jeune l'ann??e pr??c??dente (CHAPITRES I, II et III). Des contraintes environnementales fortes influen??aient la reproduction des femelles. La croissance, le succ??s reproducteur, l'intervalle entre les naissances et les comportements d'alimentation variaient suivant le site et l'ann??e d'??tude. Le site du Promontory et l'ann??e 2011 apparaissaient particuli??rement limitants. En effet en 2011, le gain de masse et le succ??s reproducteur ont diminu?? et l'intervalle de naissance et la prise de nourriture pour les femelles allaitantes ont augment?? (CHAPITRE I,II et III). Nous cherchions ??galement ?? mettre au jour une allocation diff??rentielle des m??res suivant le sexe de leur jeune. Si le co??t sup??rieur d'avoir un m??le par rapport ?? une femelle ??tait ??vident quant aux taux de bouch??es (CHAPITRE III), il s'est av??r?? plus difficile ?? d??tecter sur d'autres traits. ?? Anglesea, les jeunes m??res avaient moins de probabilit?? d'avoir un jeune qui atteigne le stade 'LPY' apr??s avoir eu un fils qu'une fille (CHAPITRE II). Toutefois, des r??sultats contraires ?? nos attentes ont ??t?? trouv??s, du moins au premier abord, sur la croissance et le succ??s reproducteur subs??quent. Ainsi, les femelles qui avaient eu une fille perdaient davantage de masse (CHAPITRE I) et avaient g??n??ralement une probabilit?? moindre de produire un jeune qui atteigne le stade 'LPY' ou qui soit sevr?? par la suite (CHAPITRE II). En revanche, l'intervalle de naissance n'??tait pas diff??rent suivant le sexe du jeune alors qu'il ??tait fortement affect?? par le co??t de la reproduction d??montr?? gr??ce ?? la manipulation, invoquant une autre explication qu'un co??t sup??rieur des filles par rapport aux fils. En effet, les femelles qui ??taient en mauvaise condition corporelle gagnaient de la masse quand elles produisaient une fille, mais pas un fils (CHAPITRE I). De surcro??t, les jeunes m??res avaient moins de chances de sevrer un jeune ?? l'??v??nement de reproduction suivant si elles avaient eu un fils plut??t qu'une fille, et le succ??s reproducteur des m??res des fils n'??tait plus diff??rent de celui des m??res des filles dans les ann??es plus difficiles (CHAPITRE II). Enfin, les m??res des fils augmentaient la quantit?? de nourriture ing??r??e si elles avaient sevr?? un jeune l'ann??e pr??c??dente, mais les m??res des filles la diminuaient. Ces diff??rents r??sultats sugg??raient fortement qu'un ajustement du sexe-ratio ??tait utilis?? quand les ressources individuelles ou environnementales contraignaient davantage la reproduction. En conclusion, pour limiter le d??calage entre les besoins ??nerg??tiques et la disponibilit?? en nourriture, les femelles chez le kangourou gris de l'Est pourraient modifier l'allocation de leurs ressources ?? la reproduction en reportant la prochaine mise bas et en produisant un jeune du sexe le moins co??teux en accord avec les contraintes individuelles et environnementales. Ces r??sultats soulignent l'importance d'??tudes avec un suivi individuel sur plusieurs ann??es afin de pouvoir comprendre la variabilit?? des strat??gies de reproduction et leurs cons??quences sur la dynamique des populations. // Abstract : Reproduction in living beings, particularly in female mammals that produce milk, is costly, potentially involving trade-offs with life-history traits if resources are limited and an increase in foraging effort. Individual differences may, however, hide the negative effects of this cost on life-history traits. I used two powerful and complementary approaches, to deal with individual heterogeneity: experimental manipulation of reproductive effort and statistical control of individual effect. Using both approaches, I investigated the effect of presence, size and sex of young on growth, subsequent reproduction and individual foraging behaviours of females. I used data of tagged free-ranging eastern grey kangaroos (Macropus giganteus) collected over six years at five study sites in Victoria, Australia. There was a clear cost of reproduction. Reproductive effort decreased mass gain and limb growth for inter-capture intervals greater than two years. Over two successive reproductive events, mass gain and arm growth were reduced but leg growth was independent of reproductive effort (CHAPTER II).In addition, survival to Large Pouch Young ('LPY') stage, about 7 months of age, was higher and birth rate lower in manipulated compared to control females but survival to weaning was not affected (CHAPTER III). CHAPTER IV shows that lactating females cope with current reproductive costs by increasing ivtime spent foraging as well as bite and chewing rates without decreasing vigilance comparedto non lactating ones. Bite rate was also greater for females that weaned a young at the previous reproductive event. My study supports reproductive cost hypothesis while showing substantial individual differences. To limit mismatch between energetic needs and resource availability, females of eastern grey kangaroo could modify resource allocation to reproduction by delaying birth date of subsequent young and producing the less costly sex according to individual and environmental constrains. My thesis shows the importance of experimental approach and individual monitoring over multiple years to understand the diversity of reproductive strategies and their consequences in evolutionary ecology and population dynamic.

A central goal of life history theory is to understand the selective factors that generate the diversity of reproductive patterns observed in nature. Within lifetime changes in reproductive investment will determine an organism's fitness; however, this area of life history theory has received less attention than comparisons among population that characterize life history traits as a single population mean. Reproductive allocation can be affected by multiple cues; the integration of these cues across an organism's lifetime generates the diversity in life history strategies observed in nature. Life history studies should examine the interacting effects of multiple cues on life history strategies to generate better predictions and generalizations of age-related changes in reproductive investment. An individual's life history strategy is inherently multivariate consisting of a coordinated suite of life history traits that, when combined across the organism's lifetime, determines its fitness. Life history strategies can therefore be described as a trajectory through multivariate space defined by life history traits. Here I describe life history trajectory analysis, a multivariate analytical approach for quantifying and comparing phenotypic change in life history strategies; this methodology is adapted from an analytical framework originally described for studies of morphological evolution. Life history trajectories have attributes (magnitude, direction, and shape) that can be quantified and statistically compared among taxa to determine if life history patterns are predictable. Using the life history trajectory analysis, I demonstrate the effect of prior experience on reproductive allocation in the burying beetle Nicrophorus orbicollis. The effect of prior experience resulted in a terminal investment or accentuated response to age-based cues, or resulted in a conservative investment strategy or reproductive restraint. In the livebearing fish Gambusia affinis, females adjust the level of reproductive investment to current reproduction based on age- or environment-based cues. Age-0 females decreased the level of reproductive investment to current reproduction in late summer prior to the onset of fall and winter months. Old females, on the other hand, increased the level of reproductive investment as the summer progressed. The reproductive restraint and terminal investment patterns exhibited by age-0 and age-1 females, respectively, were consistent with the predictions from the cost of reproduction hypothesis. These studies demonstrate how the life history trajectory analysis provides an analytical tool to test predictions of life history theory. Additionally, I provide evidence that organisms use multiple cues to determine the level of reproductive investment and that the strength of the effect of each cue will depend on the age of an individual.

<p>This thesis examines parental condition, how it is traded off against reproduction and how it is displayed in a secondary sexual trait. The studies were performed on nest-box breeding collared flycatchers Ficedula albicollis on the island of Gotland, in the Baltic Sea. Early breeding and high fitness were found to be associated with high levels of glycosylated haemoglobin possibly governed by migratory exertion and infectious disease. In order to test if immune function is expressed in secondary sexual traits and how it is traded off against reproductive effort a series of experiments were performed, in which birds were challenged with an antigen, via a vaccine containing neutralised paramyxovirus. The forehead patch of the male collared flycatcher serves as a badge of status and is under sexual selection. Good condition, as reflected in strong immune response and low levels of blood parasites was found to be associated with bigger patch size. Patch size was also found to vary in size within the same breeding season in a pattern predictable from immune response data. Immune response, in itself, was found to be costly in terms of reduced survival, confirming that trade-offs involving suppression of immune response may increase fitness. Mating effort was found to be traded off against immune function and moult. Experimental brood size manipulations revealed a trade-off females between number of offspring and immune function. Thus I suggest a set of parameters useful for condition estimation. I also show that immune response is costly and, second, that pathogen resistance probably plays an important role in the shaping of secondary sexual traits and life-history decisions.</p>

This thesis examines parental condition, how it is traded off against reproduction and how it is displayed in a secondary sexual trait. The studies were performed on nest-box breeding collared flycatchers Ficedula albicollis on the island of Gotland, in the Baltic Sea. Early breeding and high fitness were found to be associated with high levels of glycosylated haemoglobin possibly governed by migratory exertion and infectious disease. In order to test if immune function is expressed in secondary sexual traits and how it is traded off against reproductive effort a series of experiments were performed, in which birds were challenged with an antigen, via a vaccine containing neutralised paramyxovirus. The forehead patch of the male collared flycatcher serves as a badge of status and is under sexual selection. Good condition, as reflected in strong immune response and low levels of blood parasites was found to be associated with bigger patch size. Patch size was also found to vary in size within the same breeding season in a pattern predictable from immune response data. Immune response, in itself, was found to be costly in terms of reduced survival, confirming that trade-offs involving suppression of immune response may increase fitness. Mating effort was found to be traded off against immune function and moult. Experimental brood size manipulations revealed a trade-off females between number of offspring and immune function. Thus I suggest a set of parameters useful for condition estimation. I also show that immune response is costly and, second, that pathogen resistance probably plays an important role in the shaping of secondary sexual traits and life-history decisions.

Altitudinal gradients offer attractive opportunities for studies of population differentiation in response to environmental heterogeneity. In this thesis, I examined population differentiation along altitudinal gradients by combining common-garden experiments with field studies and experiments in alpine, subalpine and boreal populations of the perennial herb Solidago virgaurea. More specifically, I determined whether leaf physiology in terms of nitrogen concentration and resorption, flowering phenology, flower production and reproductive effort vary along altitudinal gradients. Nitrogen concentration in green leaves were higher in alpine than in subalpine and boreal populations. These differences persisted when plants were grown from seeds in a common-garden experiment at two sites, suggesting that the differences have a genetic component. There was mixed support for a trade-off between maximized carbon gain through the maintenance of high nitrogen concentration, and minimized nitrogen loss through high resorption. In their natural habitats alpine populations began flowering later than subalpine populations, but this difference was reversed when plants were grown in a common environment. This suggests that genetic differences among populations counteract environmental effects and reduce phenotypic variation in flowering time among populations. Flowering time thus shows countergradient genetic variation in S. virgaurea. In a common-garden experiment, boreal populations produced more flowers and had a higher reproductive effort than subalpine and alpine populations indicating habitat-specific genetic differences in reproductive allocation. In a field study, which included three populations, seed set was close to zero in the alpine population, intermediate in the subalpine population, and high in the boreal population. Experimental flower removal showed that seed production was associated with a considerable cost in terms of reduced flowering propensity the following year, but did not support the hypothesis that a large floral display is important for pollination success.

Rising archipelagos provide unique settings for the study of the temporal and spatial dynamics of their biota. This offers the possibility to study the ecology and genetics of early successional processes; both between islands that differ in age and within islands when already established organisms have to keep pace with the changing environment. I have worked in the Skeppsvik Archipelago housing about 100 islands that due to land uplift vary in age, thus representing various stages of primary succession. I have utilized a naturally created metapopulation of Silene dioica, which in this archipelago is a dominant plant of the deciduous border, offering the possibility to study subpopulations on islands of different ages and in different phases of primary succession. Many plant species exist as metapopulations, which consists of many local populations which may differ in size and degree of connectivity. Metapopulations are further characterized by recurrent colorizations and extinctions of local populations, meaning that a species continually must disperse and relocate to allow for persistence in this system. For a dioecious plant species, gene flow is in the shape of seeds and pollen and to allow for the persistence of populations, it is necessary that levels of seed dispersal and pollen gene flow are enough to ensure both colonisation, establishment and subsequent population growth. Levels of seed dispersal and pollen gene flow is in turn influenced by how the two sexes partition resources between reproduction, growth and survival. In paper I, I combined a field survey, a common garden experiment and a nine-year demographic study to assess the demographic consequences of sex-specific resource allocation and to investigate if differential costs of reproduction may be a driver in the evolution of sexual dimorphism in dioecious Silene dioica. Significant somatic intersexual dimorphism was found with females being the larger sex, both in terms of above – and belowground biomass. Furthermore, the reproductive effort of females exceeds that of males across a growing season which largely confirms what has been observed earlier in dioecious, herbaceous plant species. According to the cost of reproduction hypothesis, high reproductive investment should result in trade-offs with somatic and/or life-history traits. Somatic trade-offs were not observed, and instead I found strong, positive associations between reproductive investment and vegetative growth in both males and females. Compensation mechanisms were found in both sexes although females are generally more efficient at compensating their reproductive costs. At the end of a flowering season, after having paid the current costs of reproduction, females are better than males at provisioning perennial roots and rosettes potentially influencing the ability to set future flower buds and winter survival. Trade-offs were found between current and future reproduction and survival, but this is condition dependent and compensation through frequency of flowering plays an important role. The cost of reproduction hypothesis appears to play some role in driving the somatic and demographic sexual dimorphisms observed in this system but sexual selection acting on males will be a fruitful avenue for future research. In paper II, I investigated the population genetic consequences of metapopulation dynamics in Silene dioica. The occurrence of islands in different phases of primary succession together with successional gradients across islands, makes it possible to investigate the genetic dynamics occurring in an age-structured metapopulation across several hierarchical levels. Genetic diversity and differentiation were estimated in eight young, recently colonised populations and in ten populations of an intermediate successional stage. Young populations were less genetically diverse compared to older populations, indicating that bottlenecks, created by small founding groups derived from a limited number of source populations, reduce the genetic diversity within newly founded populations. The observation of strong genetic structure both between islands and between patches with islands, indicates that gene flow is restricted across several spatial levels in this system. However, the lack of statistically significant differences in genetic differentiation between young and intermediate populations, indicates that levels of gene flow may not be high enough to reduce the genetic differentiation that arise from the initial founder event. The patterns of sexual dimorphism and the roles of males and females in Silene dioica have evolved to allow persistence in an ecological and population context of this species. The nature of this habitat, where islands rise up from the sea creating new environments for colonisation while at the same time, autogenic primary succession processes eventually leads to extinction, means that S. dioica continuously must relocate within successional phases for its persistence. The obvious success of this dioecious plant is apparent as it is one of the few dominant species in the deciduous border. This suggests that levels of seed dispersal and gene flow are sufficient enough to allow for establishment and persistence of island populations and that the sexual dimorphisms that have evolved in this metapopulation system act to increase levels of gene flow. The "live hard – die young" strategy, with extensive flowering bouts, which we find in the males may have evolved as a way of maintaining sufficient levels of genetic diversity in the metapopulation but will only be a possible strategy if there are continuous opportunities for re-establishments. Thus, the continuous land uplift that is occurring in the northern part of the Gulf of Bothnia may very well be a prerequisite for the long-term persistence of this dioecious, perennial plant species. / Landhöjningsprocesser i skärgårdsmiljöer skapar nya habitat som gör det möjligt att studera naturliga populationer i ett rumsligt och tidsmässigt sammanhang. Detta möjliggör studier av ekologi och genetik i tidiga successionsprocesser, både mellan öar som skiljer sig åt åldersmässigt och inom öar, där redan etablerade organismer måste anpassa sig till en föränderlig miljö. Jag har utfört studier i Skeppsviks skärgård som rymmer cirka 100 öar. På grund av landhöjningen så varierar dessa öar i ålder och de representerar således olika stadier i primärsuccession. Jag har använt mig av en naturlig Silene dioica metapopulation lokaliserad i Skeppsviks skärgård. Många växtarter existerar i metapopulationer, vilket består av ett antal lokala populationer som kan skilja sig åt i storlek och grad av anknytning. Metapopulationer kännetecknas även av återkommande koloniseringar och utrotningar av lokala populationer, vilket innebär att en art kontinuerligt måste sprida sig för att garantera sin fortlevnad i detta system. Genflöde inom dioika växtarter är i form av pollen och frön, och för att populationer skall kunna överleva så är det nödvändigt att nivåerna av fröspridning och pollen-genflöde är tillräckliga för att säkerhetsställa både kolonisering, etablering och efterföljande populationstillväxt. Nivåer av fröspridning och pollen-genflöde påverkas i sin tur av hur de två könen partitionerar resurser mellan reproduktion, tillväxt och överlevnad. I den första studien har jag kombinerat en fältundersökning, ett frilandsexperiment och en nioårig demografisk studie för att undersöka de demografiska konsekvenserna av könsspecifik resursallokering och för att utreda om könsspecifika skillnader i reproduktiv kostnad kan vara en drivkraft för evolutionen av sexuell dimorfism hos den dioika växten Silene dioica. Jag upptäckte signifikant somatisk intersexuell dimorfism där honor hade betydligt mer ovanjordisk och underjordisk biomassa jämfört med hanar. Över en växtsäsong så investerar honorna mer resurser i reproduktion, vilket i stor utsträckning bekräftar vad som tidigare har observerats i örtartade, dioika växter. Enligt hypotesen för reproduktiv kostnad så bör en hög investering i reproduktion leda till trade-offs med somatiska egenskaper, t.ex. tillväxt. Jag observerade inga somatiska trade-offs och istället fann jag positiva associationer mellan reproduktion och tillväxt hos både honor och hanar. Båda könen verkar ha utvecklat kompensationsmekanismer, även om honorna generellt är mer effektiva i hur de kompenserar för sina reproduktiva kostnader. Vid slutet av en växtsäsong, efter att ha betalat för de nuvarande reproduktiva investeringarna, så är honor bättre än hanar på att allokera resurser till fleråriga strukturer, såsom bladrosetter och rötter. Detta kan potentiellt påverka hur de anlägger sina knoppanlag för nästkommande år och hur väl de överlever vintern. Trade-offs hittades mellan nuvarande reproduktion och framtida reproduktionsmöjligheter och överlevnad men detta var habitat-specifikt och kompensation med hjälp av hur ofta en växt blommar under sin livstid spelar en viktig roll. Hypotesen för reproduktiv kostnad verkar vara en del av förklaringen till den somatiska och demografiska könsdimorfism som observerats i detta system men sexuell selektion, som verkar på hanar, kan vara ett möjligt område för framtida studier. I den andra studien undersökte jag populationsgenetiska konsekvenser av metapopulationsdynamik i Silene dioica. Förekomsten av öar i olika faser av primär succession tillsammans med olika grader av succession inom öar gör det möjligt att undersöka den genetiska dynamiken som uppträder i en åldersstrukturerad metapopulation över flera hierarkiska nivåer. Genetisk mångfald och differentiering uppskattades i åtta unga, nyligen koloniserade populationer och i tio populationer av ett intermediärt successionsstadium. Unga populationer hade lägre genetisk diversitet jämfört med äldre populationer, vilket indikerar att genetiska flaskhalsar, skapade av fåtal antal koloniserande individer, s.k. founders, som härrör från ett begränsat antal källpopulationer, minskar den genetiska diversiteten inom nybildade populationer. Observationen av stark genetisk strukturering, mellan och inom öar, indikerar att genflödet är begränsat över flera rumsliga nivåer i detta system. Bristen på statistiskt signifikanta skillnader i genetisk differentiering mellan unga och intermediära populationer indikerar emellertid att nivåer av genflöde kanske inte är tillräckligt höga för att minska den genetiska differentieringen som uppstår från den ursprungliga founder-händelsen. Mönstren av sexuell dimorfism och hanarnas och honornas roll har utvecklats för att möjliggöra fortlevnad i ett ekologisk och populationsmässigt sammanhang hos Silene dioica. I denna livsmiljö, där öar stiger upp ur havet och skapar nya miljöer för kolonisering samtidigt som autogena primära successionsprocesser leder till utrotning, måste S. dioica kontinuerligt sprida sig mellan olika successionsfaser för att överleva. Den uppenbara framgången för den här dioika växten är uppenbar eftersom den är en av de få dominerande arterna i lövkanten. Detta tyder på att nivåer av fröspridning och genflöde är tillräckliga för att möjliggöra etablering och beständighet av ö-populationer och att de sexuella dimorfismer som har utvecklats i detta metapopulationssystem verkar för att öka nivåerna av genflöde. "Lev hårt – dö ung" -strategin med omfattande blomningar som vi finner hos hanarna kan ha utvecklats som ett sätt att upprätthålla tillräckliga nivåer av genetisk diversitet i metapopulationen men den kommer endast att vara en möjlig strategi om det finns kontinuerliga möjligheter för re-etableringar. Således kan den kontinuerliga landupphöjningen som förekommer i norra delen av Bottniska viken mycket väl vara en förutsättning för den långsiktiga beständigheten av denna dioika, fleråriga växtart. / <p>Felaktigt angivet "Dissertation for PhD" i kolofon.</p>