Individual differences in migratory behavior shape population genetic structure and microhabitat choice in sympatric blackcaps (Sylvia atricapilla)

In migratory birds, traits such as orientation and distance are known to have a strong genetic background, and they often exhibit considerable within‐population variation. How this variation relates to evolutionary responses to ongoing selection is unknown because the underlying mechanisms that translate environmental changes into population genetic changes are unclear. We show that within‐population genetic structure in southern German blackcaps (Sylvia atricapilla) is related to individual differences in migratory behavior. Our 3‐year study revealed a positive correlation between individual migratory origins, denoted via isotope (δ²H) values, and genetic distances. Genetic diversity and admixture differed not only across a recently established migratory polymorphism with NW‐ and SW‐migrating birds but also across δ²H clusters within the same migratory route. Our results suggest assortment based on individual migratory origins which would facilitate evolutionary responses. We scrutinized arrival times and microhabitat choice as potential mechanisms mediating between individual variation in migratory behavior and assortment. We found significant support that microhabitat choice, rather than timing of arrival, is associated with individual variation in migratory origins. Moreover, examining genetic diversity across the migratory divide, we found migrants following the NW route to be genetically more distinct from each other compared with migrants following the traditional SW route. Our study suggests that migratory behavior shapes population genetic structure in blackcaps not only across the migratory divide but also on an individual level independent of the divide. Thus, within‐population variation in migratory behavior might play an important role in translating environmental change into genetic change.     

Habitat distribution of migratory and sedentary blackcaps Sylvia atricapilla wintering in southern Iberia: a morphological and biogeochemical approach

In migratory species, the way in which conspecifics from different breeding populations are distributed during the non‐breeding period is important from and ecological, evolutionary and conservation perspective, but such knowledge is still limited for most species. Migratory and sedentary blackcaps Sylvia atricapilla wintering in southern Spain can occupy two habitat types: forests and shrublands. According to earlier studies, blackcaps prefer forests over shrublands, and residents remain nearly restricted to forests. However, whether migrants with different breeding origin occupy the two habitats differently is unknown. We used morphological and biogeochemical data (hydrogen isotope ratios measured on feathers: δ²H_f), which show variation along the breeding range of the species, to answer this question. Isotope analyses supported the reliability of morphology as a method for distinguishing between migratory and sedentary blackcaps in sympatry, showing that sedentary individuals are rare in shrublands while migratory ones are abundant in both habitat types. However, migratory blackcaps scored similar δ²H_f values in forests and shrublands, and neither did vary in structural size or flight morphology between habitats. Our study suggests that migrants from a wide range of breeding origins end up mixing between forests and shrublands, which may explain the patterns of variation in space and time in the abundance of blackcaps in this area, and supports the view that inequalities may arise among migrants with the same origin but wintering in different habitats. Such inequalities might carry over into other stages of blackcaps’ life cycle contributing to the regulation of its migratory populations.     

Sex and age-specific differences in wing pointedness and wing length in blackcaps Sylvia atricapilla migrating through the southern Baltic coast

The blackcap Sylvia atricapilla shows a complex migratory pattern and is a suitable species for the studies of morphological migratory syndrome, including adaptations of wing shape to different migratory performance. Obligate migrants of this species that breed in northern, central, and Eastern Europe differ by migration distance and some cover shorter distance to the wintering grounds in the southern part of Europe/North Africa or the British Isles, although others migrate to sub-Saharan Africa. Based on ˃40 years of ringing data on blackcaps captured during autumn migration in the Southern Baltic region, we studied age- and sex-related correlations in wing pointedness and wing length of obligate blackcap migrants to understand the differences in migratory behavior of this species. Even though the recoveries of blackcaps were scarce, we reported some evidence that individuals which differ in migration distance differed also in wing length. We found that wing pointedness significantly increased with an increasing wing length of migrating birds, and adults had longer and more pointed wings than juvenile birds. This indicates stronger antipredator adaptation in juvenile blackcaps than selection on flight efficiency, which is particularly important during migration. Moreover, we documented more pronounced differences in wing length between adult and juvenile males and females. Such differences in wing length may enhance a faster speed of adult male blackcaps along the spring migration route and may be adaptive when taking into account climatic effects, which favor earlier arrival from migration to the breeding grounds.     

Age-related variation in wing shape of migratory and sedentary Blackcaps Sylvia atricapilla

In many passerines, juveniles have shorter and more rounded wings than adults. Given that (1) long and pointed wings improve endurance in migratory flights, (2) shorter and rounded wings improve manoeuvrability, and (3) juvenile birds are more vulnerable to predators than adults, it has been hypothesised that ontogenetic variation in wing shape results from a greater importance of predation avoidance relative to migration performance during the first year of life. If so, wing shape should not change with age in the absence of migration-related selection for longer and more pointed wings. We test this by studying the variation with respect to age in wing length and wing pointedness of migratory and sedentary Blackcaps wintering in southern Spain. Migratory Blackcaps had longer and more pointed wings than sedentary Blackcaps. Juveniles had shorter wings than adults in migratory populations, but not in sedentary populations. The variation with age in wing pointedness was less pronounced, and was found in migratory females only. These differences between the two traits could be related to a stronger selection for pointed wings than for longer wings with increasing distance of migration, and to an increased migratoriness of females in partially migratory Blackcap populations. We hypothesise that, in migratory Blackcaps, a shorter and more rounded wing in juveniles could be selected for if the decrease in predation rate compensated for the somewhat greater costs of the first migration attempt. On the other hand, there are no costs of migration in sedentary Blackcaps, which hence maintain a similar wing shape, giving high manoeuvrability, both as juveniles and as adults.     

A method for differentiating between sedentary and migratory Blackcaps Sylvia atricapilla in wintering areas of southern Iberia

We used the morphological differentiation of southern, sedentary Iberian Blackcaps Sylvia atricapilla to distinguish them from migratory Blackcaps during the winter. To do so, we studied three morphometric traits (wing length, wing shape and tail length) of three migratory Iberian populations and two sedentary ones during the breeding season. We obtained a discriminant function which allowed us to differentiate the two population types (>90% correct classifications). We discuss the utility of the method for differentiating between sedentary and migratory Blackcaps in their sympatric wintering areas, as well as the implications of our results for the study and conservation of sedentary, perhaps differentiated Blackcaps in southern Iberia.     

Fuel load and flight ranges northern Iberia during autumn of blackcaps Sylvia atricapilla in and spring migrations

Fuel accumulation, mainly as fatty acids, is one of the main characteristics of migratory, birds. Studying to what extent each population or species manages fuel load and how it varies along routes of migration or between seasons （autumn and spring migrations） is crucial to our understanding of bird migration strategies. Our aim here was to analyse whether migratory blackcaps Sylvia atrieapilla passing through northern Iberia differ in their mean fuel loads, rate of fuel accumulation and ＇ potential＇ flight ranges between migration seasons. Blackcaps were mist netted for 4 h-periods beginning at dawn from 16 September to 15 November 2003 - 2005, and from 1 March to 30 April 2004 - 2006 in a European Atlantic hedgerow at Loza, northern Iberia. Both fuel load and fuel deposition rate （this latter assessed with difference in body mass of within-season recaptured individuals） were higher in autumn than in spring. Possible hypotheses explaining these results could be seasonal-associated variations in food availability （likely lower during spring than during autumn）, the fact that a fraction of the migrants captured in spring could breed close to the study area and different selective pressures for breeding and wintering     

Genetic divergence among geographical populations of the migratory locust in China

The random amplified polymorphic DNA (RAPD) technique was used to examine genetic divergence and interrelations of 11 geographical populations of the migratory locust in China, and the role of spatial separation in the population differentiations. AMOVA analysis of genetic variations in all the populations indicated greater within- (79.55%) than among-population variability (20.45%), and that there were significant differentiations among the populations; 11 populations were divided into four regional groups, with significantly greater variability within (82.99%) than among the groups (17.01%), and there existed apparent regional differentiations. Paired comparisons showed significantly greater variability within-than between-groups, indicating significant differentiations between populations of different regional groups. Of all the pairwise comparisons, Hainan and Tibetan groups displayed the greatest differentiation, with the difference between the two groups being seven folds of that between populations within the groups; the least differentiations were exhibited between the groups of Hainan, Xinjiang, and Inner Mongolia, with the differences between groups being only half of the differences between populations within the groups. Mantel tests of the genetic and spatial distances showed that the two matrices were significantly correlated (p<0.01), indicating that the geographical isolation played an important role in the differentiations of the geographical populations of the migratory locusts. Cluster analysis divided all populations into four major groups: Xinjiang and Inner Mongolia group, the Great Plains of North China (the Yellow River and Huai River Plains) group, Hainan group, and Tibet group. Principal component analysis (PCA) supported the division of populations based on the cluster analysis. However, analysis of individuals clustered the locusts into five populations: Xinjiang and Inner Mongolia, Hami in Xinjiang, the Great Plains of North China, Hainan, and Tibet. The locust populations in eastern China displayed apparently continous and gradient variations; as such authors consider that there were no necessity and valid reasons for further division of subspecies. The subspecific status for the main geographical populations of the migratory locusts in China was discussed.     

Experimental evolution across different thermal regimes yields genetic divergence in recombination fraction but no divergence in temperature associated plastic recombination

Phenotypic plasticity is pervasive in nature. One mechanism underlying the evolution and maintenance of such plasticity is environmental heterogeneity. Indeed, theory indicates that both spatial and temporal variation in the environment should favor the evolution of phenotypic plasticity under a variety of conditions. Cyclical environmental conditions have also been shown to yield evolved increases in recombination frequency. Here, we use a panel of replicated experimental evolution populations of D. melanogaster to test whether variable environments favor enhanced plasticity in recombination rate and/or increased recombination rate in response to temperature. In contrast to expectation, we find no evidence for either enhanced plasticity in recombination or increased rates of recombination in the variable environment lines. Our data confirm a role of temperature in mediating recombination fraction in D. melanogaster, and indicate that recombination is genetically and plastically depressed under lower temperatures. Our data further suggest that the genetic architectures underlying plastic recombination and population‐level variation in recombination rate are likely to be distinct.     

Genetic divergence among geographical populations of the migratory locust in China

The migratory locust, Locusta migratoria L., which is distributed widely in the East Hemisphere, has long been regarded as the most important agricultural pest and a model insect. Its distribution ranges from thetropic and the subtropical zone, to the temperate zone, and to the cold-temperate zone. Because of the wide geographical distribution and adaptations, many geo- graphical populations of the migratory locust dis- played apparent variation in morphology, life history, physiology, and ot…     

Geographic variation and divergence of songs in the Olive Sparrow species complex

Phenotypic traits such as songs are important in species recognition. Variation in acoustic traits can form barriers to gene flow and promote speciation. Therefore, understanding song divergence is crucial in groups with controversial taxonomy such as Olive Sparrows (Arremonops rufivirgatus), a widespread Neotropical species of songbird with multiple allopatric populations. Taxonomic authorities disagree on the number of Olive Sparrow subspecies, placing them into either two or three main groups. These groups may represent separate species based on morphological traits, but trait divergence within the complex has not been examined. We studied geographic variation in the characteristics of the songs of Olive Sparrows at two geographical levels: among three proposed groups and among five allopatric populations. In a second analysis, we evaluated the strength of acoustic divergence within the complex by comparing acoustic distances among groups and allopatric populations of Olive Sparrows with the acoustic distance among three recognized species in the genus Arremonops. We analyzed 802 songs from 174 individuals across 81 locations and measured 12 variables to describe the fine structural characteristics of the songs of Olive Sparrows, Green-backed Sparrows (A. chloronotus), Black-striped Sparrows (A. conirostris), and Tocuyo Sparrows (A. tocuyensis). We found significant acoustic variation in the Olive Sparrow complex at both geographical levels. Our divergence analysis also revealed that vocal divergence within the complex is similar to or greater than that found between recognized species in the genus. Together, these results suggest that acoustic diversity within the Olive Sparrow complex probably originated by isolation in tandem with selective and/or non-selective factors.     

Genetic analysis of song dialect populations in Puget Sound white-crowned sparrows

The relationship between cultural variation and biological variationamong natural populations has been the subject of both theoreticaland empirical study. Zonotrichia leucophrys pugetensis is oneof three subspecies of white-crowned sparrow known to form geographicalsong dialects. We investigated whether these dialects correspondto genetic differences among Z. l. pugetensis populations. Wecompared allele frequencies at four microsatellite loci in malesfrom 11 sites spanning six dialects over the subspecies' rangein Oregon and Washington. Cluster analysis and genotype assignmenttests indicated no tendency for sample sites within dialectareas to be genetically more similar than are sites from differentdialect areas. AMOVA tests revealed high within-site variationand low but significant cross-site and cross-dialect-area variation.Finally, genetic distance between sites was not correlated withdialect differences when the effect of geographic distance wascontrolled statistically. We compare our finding of low geneticdifferentiation among Z. l. pugetensis dialect populations toresults of previous studies on Z. l. nuttalli and Z. l. oriantha.Because genetic structuring appears weaker than cultural (songdialect) structure in this species, we discuss the behavioralmechanisms underlying dialect maintenance in the presence ofapparent gene flow.     

Anatomical and histological changes in the alimentary tract of migrating blackcaps (Sylvia atricapilla): a comparison among fed, fasted, food-restricted, and refed birds

During northward migration, blackcaps that arrive to refuel at stopover sites in Israel's Negev Desert have reduced masses of organs that are important in food digestion and assimilation. We tested several predictions from the general hypothesis that smaller organs of digestion (small intestine and pancreas) and nutrient assimilation (liver) bring about a lower capacity to consume food and that the organs must be restored before blackcaps can feed and digest at a high rate. We used a fasting protocol to create a group of blackcaps with reduced intestine and liver mass (reduced by 45% and 36%, respectively) compared with controls fed ad lib. Because most of the small intestine's biochemical digestive capacity reside in enterocytes found on villi, we predicted and found that reduced intestinal mass in fasted blackcaps related mainly to changes in enterocytes rather than other cells and tissues such as nonabsorptive crypt cells or underlying muscle. Because migrating blackcaps that stop over to feed begin to increase in body mass only 2 d after arrival, we predicted and found a similar recovery period in blackcaps that were first fasted but then refed--the organ mass, structure, function, and ability to consume food was restored after 2 d of feeding. Another group of food-restricted blackcaps (fed at one-third ad lib. level) lost similar amounts of body mass as fasted blackcaps but had much greater capacity to consume food than fasted blackcaps, and so we predicted that they would exhibit little or no reduction in alimentary organs relative to controls fed ad lib. A surprising result was that, as in fasted blackcaps, in food-restricted blackcaps, the decreases in masses of small intestine, liver, and pancreas were proportionally greater than the decreases in body mass or in masses of nonalimentary organs (heart, pectoralis). Food restriction, like fasting, caused a decrease in amount of intestinal mucosa and an alteration in the phenotype of enterocytes. These results are thus not consistent with the general hypothesis, and although they can be rationalized by assuming that blackcaps fed ad lib. have excess digestive capacity, it may also be that the physiological process or processes limiting very high feeding rate lie elsewhere than in the digestive system.     

The probability of genetic parallelism and convergence in natural populations

Genomic and genetic methods allow investigation of how frequently the same genes are used by different populations during adaptive evolution, yielding insights into the predictability of evolution at the genetic level. We estimated the probability of gene reuse in parallel and convergent phenotypic evolution in nature using data from published studies. The estimates are surprisingly high, with mean probabilities of 0.32 for genetic mapping studies and 0.55 for candidate gene studies. The probability declines with increasing age of the common ancestor of compared taxa, from about 0.8 for young nodes to 0.1–0.4 for the oldest nodes in our study. Probability of gene reuse is higher when populations begin from the same ancestor (genetic parallelism) than when they begin from divergent ancestors (genetic convergence). Our estimates are broadly consistent with genomic estimates of gene reuse during repeated adaptation to similar environments, but most genomic studies lack data on phenotypic traits affected. Frequent reuse of the same genes during repeated phenotypic evolution suggests that strong biases and constraints affect adaptive evolution, resulting in changes at a relatively small subset of available genes. Declines in the probability of gene reuse with increasing age suggest that these biases diverge with time.     

Genetic Landscapes GIS Toolbox: tools to map patterns of genetic divergence and diversity

The Landscape Genetics GIS Toolbox contains tools that run in the Geographic Information System software, ArcGIS^®, to map genetic landscapes and to summarize multiple genetic landscapes as average and variance surfaces. These tools can be used to visualize the distribution of genetic diversity across geographic space and to study associations between patterns of genetic diversity and geographic features or other geo‐referenced environmental data sets. Together, these tools create genetic landscape surfaces directly from tables containing genetic distance or diversity data and sample location coordinates, greatly reducing the complexity of building and analyzing these raster surfaces in a Geographic Information System.     

Interactions of genetic and cultural evolution: Models and examples

This paper proposes models and examples of five principal modes of interaction between genes and culture in human evolution. Because genes and culture ultimately interact in the minds of individuals, the models are focused on individual level processes of constrained microevolution. The central hypotheses are (1) that cultural evolution as well as genetic evolution commonly proceeds by the differential transmission of alternative instructions among individuals, (2) that genetic and cultural processes directly interact through mutual influence on each other's differentials of transmission in a population, (3) that the cultural process is often self-selecting by its own criteria, and (4) that these criteria generally operate to enhance rather than oppose human adaptation. Evolutionary change at higher levels, which is particularly important in sociocultural evolution, is interpreted as restructuring the nature and extent of the variability available at the individual level. To clarify the conceptual differences of the models and hopefully to stimulate related analyses in other areas, I discuss selected examples of each of these interactions. I conclude with some remarks on the relative importance of the models to human ecology and evolution.     

Marked genetic divergence revealed by allozymes among populations of the guppy Poecilia reticulata (Poeciliidae), in Trinidad

Populations of the guppy Poecilia reticulata from six locations in N. Trinidad were examined by starch gel electrophoresis to estimate their degree of genetic divergence. Variability at seven enzyme-coding loci demonstrated that populations differed markedly in allele frequencies with some allelic substitution between sites (for 23 loci, Nei's mean genetic identity, Ī ranged from 0.869–1.00; the coefficient of gene differentiation, G _ST= 0.086; and the absolute differentiation between populations, _m= 0.044). There was a good correspondence between degree of physical isolation and extent of genetic differentiation, as exemplified by the partitioning of gene diversity (Nei's gene diversity analysis = 66% between river basins; 32% within river basins; 2% within rivers), though there was considerable variability in the contribution of individual loci. Most populations were in Hardy-Weinberg equilibrium, and any significant deviations were due exclusively to heterozygote deficiencies. The patterns of population differentiation are discussed in relation to probable past geological and historical events, and present-day evolutionary forces. Notable among these are previous continental land connections, periodic immigration from N.E. South America, and post-colonization events including differential predation, sexual selection, apparent restricted vagility and small effective population sizes.