Spatial Variation In Fitness Research Articles

Sex-specific spatial variation in fitness in the highly dimorphic Leucadendron rubrum.

Sexual dimorphism in plants may emerge as a result of sex-specific selection on traits enhancing access to nutritive resources and/or to sexual partners. Here we investigated sex-specific differences in selection of sexually dimorphic traits and in the spatial distribution of effective fecundity (our fitness proxy) in a highly dimorphic dioecious wind-pollinated shrub, Leucadendron rubrum. In particular, we tested for the effect of density on male and female effective fecundity. We used spatial and genotypic data of parent and offspring cohorts to jointly estimate individual male and female effective fecundity on the one hand and pollen and seed dispersal kernels on the other hand. This methodology was adapted to the case of dioecious species. Explicitly modelling dispersal avoids the confounding effects of heterogeneous spatial distribution of mates and sampled seedlings on the estimation of effective fecundity. We also estimated selection gradients on plant traits while modelling sex-specific spatial autocorrelation in fecundity. Males exhibited spatial autocorrelation in effective fecundity at a smaller scale than females. A higher local density of plants was associated with lower effective fecundity in males but was not related to female effective fecundity. These results suggest sex-specific sensitivities to environmental heterogeneity in L.rubrum. Despite these sexual differences, we found directional selection for wider canopies and smaller leaves in both sexes, and no sexually antagonistic selection on strongly dimorphic traits in L.rubrum. Many empirical studies in animals similarly failed to detect sexually antagonistic selection in species expressing strong sexual dimorphism, and we discuss reasons explaining this common pattern.

Fine-scale spatial variation in fitness is comparable to disturbance-induced fluctuations in a fire-adapted species.

The spatial scale at which demographic performance (e.g., net reproductive output) varies can profoundly influence landscape-level population growth and persistence, and many demographically pertinent processes such as species interactions and resource acquisition vary at fine scales. We compared the magnitude of demographic variation associated with fine-scale heterogeneity (<10m), with variation due to larger-scale (>1ha) fluctuations associated with fire disturbance. We used a spatially explicit model within an IPM modeling framework to evaluate the demographic importance of fine-scale variation. We used a measure of expected lifetime fruit production, EF , that is assumed to be proportional to lifetime fitness. Demographic differences and their effects on EF were assessed in a population of the herbaceous perennial Hypericum cumulicola (~2,600 individuals), within a patch of Florida rosemary scrub (400×80m). We compared demographic variation over fine spatial scales to demographic variation between years across 6 yr after a fire. Values of EF changed by orders of magnitude over<10m. This variation in fitness over fine spatial scales (<10m) is commensurate to postfire changes in fitness for this fire-adapted perennial. A life table response experiment indicated that fine-scale spatial variation in vital rates, especially survival, explains as much change in EF as demographic changes caused by time-since-fire, a key driver in this system. Our findings show that environmental changes over a few tens of meters can have ecologically meaningful implications for population growth and extinction.

Testing the potential mechanisms for the maintenance of a genetic color polymorphism in bluefin killifish populations.

The maintenance of genetic variation in the face of natural selection is a long-standing question in evolutionary biology. In the bluefin killifish Lucania goodei, male coloration is polymorphic. Males can produce either red or yellow coloration in their anal fins, and both color morphs are present in all springs. These 2 morphs are heritable and how they are maintained in nature is unknown. Here, we tested 2 mechanisms for the maintenance of the red/yellow color morphs. Negative frequency-dependent mating success predicts that rare males have a mating advantage over common males. Spatial variation in fitness predicts that different color morphs have an advantage in different microhabitat types. Using a breeding experiment, we tested these hypotheses by creating populations with different ratios of red to yellow males (5 red:1 yellow; 1 red:5 yellow) and determining male mating success on shallow and deep spawning substrates. We found no evidence of negative frequency-dependent mating success. Common morphs tended to have higher mating success, and this was particularly so on shallow spawning substrates. However, on deep substrates, red males enjoyed higher mating success than yellow males, particularly so when red males were rare. However, yellow males did not have an advantage at either depth nor when rare. We suggest that preference for red males is expressed in deeper water, possibly due to alterations in the lighting environment. Finally, male pigment levels were correlated with one another and predicted male mating success. Hence, pigmentation plays an important role in male mating success.

Evolution of dispersal in a multi‐trophic community context

Much is known about the evolution of dispersal when species interact with their resources or natural enemies, but very little is known about dispersal evolution when species interact with both resources and natural enemies. Here I investigate how the dispersal of an intermediate consumer evolves in response to its interactions with a basal resource and top predator. I find that dispersal evolution is possible even when the consumer species is not directly affected by environmental variability, but rather experiences the consequences that such variability has on its resource and predator. Spatial variation in the consumer's fitness is driven by spatial heterogeneity in resource productivity, which determines whether a predator can colonize a resource‐consumer community. Temporal variation in the consumer's fitness is driven by random disturbances that cause periodic local extinctions of the predator, followed by recolonizations that lead to transient fluctuations in consumer abundance. When spatial variation in resource productivity is low and the predator can colonize all patches in the landscape, there is no spatial variation in consumer fitness but temporal variation in fitness favors the evolution of a dispersal monomorphism. When spatial variation in resource productivity is high and the predator cannot colonize many patches in the landscape, spatial variation in fitness selects against dispersal. In this case, temporal variation can promote the evolution of a dispersal polymorphism with sedentary and mobile phenotypes, but only for certain types of tri‐trophic interactions. This finding underscores the importance of indirect interactions in shaping the evolution of dispersal. While the ecological community can provide a strong selective environment for the evolution of dispersal, the nature of interactions between trophic levels can also impose constraints on evolution.

中国大陆鸟类栖息地选择研究十年

Habitat selection can be regarded as a hierarchical process of different behavioral responses,which may lead to some being disproportionately used because of spatial variation in fitness and survival.In China,questions about selection have been the most important part of avian ecology research.Early research focused on characterizing the relationship between species diversity and type.In recent years,the number of studies about avian selection conducted by ornithologists in China has increased.To determine the status of bird selection research,we surveyed papers published by ornithologists in mainland China over the last 10 years(2001—2010).We searched titles,abstracts and keywords for the terms habitat selection and use,nest site selection and use,roost site selection and use,and foraging site selection and usein three Chinese journal databases and Google scholar.From each paper,we extracted:(1) the study species;(2) details about the methods;(3) whether the terms habitat selectionand habitat usewere used appropriately;(4) whether the results were placed in the context of fitness;and(5) any other relevant information.In total,we found 170 papers about bird selection or use.Of these,154 papers were published in journals of mainland China.The other 16 papers were published in international journals.The study species were 73 bird species belonging to 10 orders and 31 families.Only nine of the 59 endemic birds of mainland China have been studied in last ten years.The number of papers per year has continuously increased in the last ten years.The number of papers published in 2009—2010 was almost 3 times that in 2001—2002.However,the same questions have been consistently asked by Chinese bird selection studies.Furthermore,of the reviewed papers,33.3%(56 /168) suffered from semantic inconsistencies when usinghabitat selectionandhabitat useand this problem does not seem to vary among years.(x 2 = 2.94,P = 0.57).Some researchers appear to regard these two terms as synonymous.In total,93.2%(96 /103) of the papers examined selection by comparing used versus unused habitat;the other 6.8%(7 /103) compared used versus available habitat.However,many authors failed to clarify that not all is equally suitable for bird use.Population density,predators and scales may influence availability.Most(99.0%,101 /103) of the papers did not place their findings in a behavioral or fitness context.Habitat selection is a process in which birds decide to choose a suitable site for nest,roost or forage,and the choice can affect fitness.In most studies,the factors influencing selection were proposed to be food,water and coverage.However,the pathway through which these environmental factors influence selection was seldom discussed.Based on our findings,we propose ways in which bird selection research in mainland China can advance.First,there should be more study of the selection of bird species that are endemic to China.Second,bird selection should be studied at different spatial scales,and for more species and environments.Finally,bird selection should be studied by considering the behavior of individuals,and its fitness consequences.

Nonparametric spatial regression of survival probability: visualization of population sinks in Eurasian Woodcock

Both evolutionary ecologists and wildlife managers make inference based on how fitness and demography vary in space. Spatial variation in survival can be difficult to assess in the wild because (1) multisite study designs are not well suited to populations that are continuously distributed across a large area and (2) available statistical models accounting for detectability less than 1.0 do not easily cope with geographical coordinates. Here we use penalized splines within a Bayesian state-space modeling framework to estimate and visualize survival probability in two dimensions. The approach is flexible in that no parametric form for the relationship between survival and coordinates need be specified a priori. To illustrate our method, we study a game species, the Eurasian Woodcock Scolopax rusticola, based on band recovery data (5000 individuals) collected over a > 50 000-km2 area in west-central France with contrasted habitats and hunting pressures. We find that spatial variation in survival probability matches an index of hunting pressure and creates a mosaic of population sources and sinks. Such analyses could provide guidance concerning the spatial management of hunting intensity or could be used to identify pathways of spatial variation in fitness, for example, to study adaptation to changing landscape and climate.

The Fixation of Locally Beneficial Alleles in a Metapopulation

Extinction, recolonization, and local adaptation are common in natural spatially structured populations. Understanding their effect upon genetic variation is important for systems such as genetically modified organism management or avoidance of drug resistance. Theoretical studies on the effect of extinction and recolonization upon genetic variance started appearing in the 1970s, but the role of local adaptation still has no good theoretical basis. Here we develop a model of a haploid species in a metapopulation in which a locally adapted beneficial allele is introduced. We study the effect of different spatial patterns of local adaptation, and different metapopulation dynamics, upon the fixation probability of the beneficial allele. Controlling for the average selection pressure, we find that a small area of positive selection can significantly increase the global probability of fixation. However, local adaptation becomes less important as extinction rate increases. Deme extinction and recolonization have a spatial smoothing effect that effectively reduces spatial variation in fitness.

Spatio‐temporal shifts in gradients of habitat quality for an opportunistic avian predator

We used the conceptual framework of the theory of natural selection to study breeding habitat preferences by an opportunistic avian predator, the black kite Milvus migrans. In Europe, black kite populations are mostly found near large networks of aquatic habitats, usually considered optimal for foraging and breeding. We hypothesized that proximity to wetlands could vary among individuals and affect their fitness, and thus be subject to natural selection. We tested the hypothesis first on a population on Lake Lugano (Italian pre‐Alps) which has been monitored for nine years, and then on seven other populations, each studied for four–five years, located along a continuum of habitat from large water bodies to scarce aquatic habitat of any kind. In the Lake Lugano population, black kite abundance was negatively related to distance to the lake in all the nine years of study, consistent with long‐term natural selection. There was evidence of ongoing directional selection on strategic nest location in three of the years, and evidence of stabilizing selection in two years. In eight of the nine years the trend was for a linear increase in fitness with increasing proximity to the lake. At the population level, results were consistent with adaptive habitat choice in relation to the previous year's spatial variation in fitness: higher associations between fitness and distance to the lake (i.e. higher selection gradients) resulted in higher density variations in the following year, in turn related to the availability of fish, the main local prey. The progressive decline of inland pairs and increase in the density of lakeshore pairs caused a directional long‐term trend of declining mean distance to the lake. Breeding near aquatic habitats was associated with higher foraging success, and higher frequency and biomass of prey deliveries to offspring. There was weak evidence of selection in other populations. The inland‐wetland gradient of habitat quality may have been affected by predation risk, as estimated by density of a major predator of adults and nestlings, the eagle owl Bubo bubo. Behavioral decisions at the level of the individual probably translated into population effects on density and distribution at various spatial scales. Populations in optimal habitats showed higher density and produced six times as many young per unit space as those in sub‐optimal habitats.

Environmental heterogeneity and balancing selection in the acorn barnacle Semibalanus balanoides.

The northern acorn barnacle Semibalans banlanoides occupies several intertidal microhabitats which vary greatly in their degree of physical stress. This environmental heterogeneity creates distinct selection regimes which can maintain genetic variation in natural populations. Despite considerable attention placed on the link between spatial variation in fitness and balancing selection at specific loci, experimental manipulations and fitness estimates for molecular polymorphisms have rarely been conducted in the wild. The aim of this transplant experiment was to manipulate the level of physical stress experienced by a cohort of barnacles in the field and then investigate the spatial variation in fitness for genotypes at three loci: two candidate allozymes and the mitochondrial DNA control region. The viability of mannose-6-phosphate isomerase (Mpi) genotypes was dependent on the level of physical stress experienced in the various treatments; alternative homozygotes were favoured in alternative high stress-low stress environments. In contrast, the fitness of genotypes at other loci was equivalent among treatments and unaffected by the manipulation. Evaluated in the light of balancing selection models, these data indicate that the presence of multiple environmental niches is sufficient to promote a stable Mpi polymorphism in barnacle populations and that allelic variation at this locus reflects the process of adaptation to the heterogeneous intertidal landscape.

FINE-GRAINED SPATIAL AND TEMPORAL VARIATION IN SELECTION DOES NOT MAINTAIN GENETIC VARIATION IN ERIGERON ANNUUS.

Because interactions among plants are spatially local, the scale of environmental heterogeneity can have large effects on evolutionary dynamics. However, very little is known about the spatial patterns of variation in fitness and the relative magnitude of spatial and temporal variation in selection. Replicates of 12 genotypes of Erigeron annuus (Asteraceae) were planted in 288 locations within a field, separated by distances of 0.1 to 30.0 m, and replicated in two years. In a given year, most spatial variation in relative fitness (genotype-environment [G × E] interactions for fitness) occurred over distances of only 50 cm. Year effects were as large or larger than the spatial variation in fitness; in particular there was a large, three-way, genotype-year-environment interaction at the smallest spatial scale. The genetic correlation of fitness across years at a given location was near zero, 0.03. Thus, the relative fitness of genotypes is spatially unpredictable and a map of the selective environment has constantly shifting locations of peaks and valleys. Including measurements of soil nutrients as covariates in the analysis removed most of the spatial G × E interaction. Vegetation and microtopography had no effect on the G × E terms, suggesting that differential response to soil nutrients is the cause of spatial variation in fitness. However, the slope of response to NH4 and P04 was negative; therefore the soil nutrients are probably just indicators of other, unknown, environmental factors. We explored via simulation the evolutionary consequences of spatial and temporal variation in fitness and showed that, for this system, the spatial scale of variation was too fine grained (by a factor of 3 to 5) to be a powerful force maintaining genetic variation in the population. The inclusion of both spatial and temporal variation in fitness actually reduced the coexistence of genotypes compared to pure spatial models. Thus the presence of spatial or temporal variation in selection does not guarantee that it is an effective evolutionary force maintaining diversity. Instead the pattern of selection favors generalist genotypes.

Measuring spatial variation in natural selection using randomly‐sown seeds of Arabidopsis thaliana

AbstractWe describe a simple method of measuring spatial variation in fitness using randomly sown seeds. Given a random initial distribution of seeds, any spatial variation in the final distribution of genotypes must be the result of spatial variation in natural selection. Departures from spatial randomness are tested using a modification of join‐count statistics, based on the probability that two randomly chosen plants separated by a given distance carry the same genetic marker. Monte Carlo simulations showed that for the randomly sown seeds design the join‐count test had higher power to detect spatial variation in fitness than analysis of variance. The power was relatively independent of the number of genotypes used or the number of microsites sampled. The method was tested in the field using three standard lines of Arabidopsis thaliana. At the end of the life cycle, adjacent plants were significantly more likely to be the same line than randomly chosen pairs and the spatial dependence extended to distances of nearly 50 cm. Simple experiments like this may be especially useful for “comparative ecological genetics”, allowing systematic studies of natural selection among many species or habitats.