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.     

Sizing up your enemy: individual predation vulnerability predicts migratory probability

Partial migration, in which a fraction of a population migrate and the rest remain resident, occurs in an extensive range of species and can have powerful ecological consequences. The question of what drives differences in individual migratory tendency is a contentious one. It has been shown that the timing of partial migration is based upon a trade-off between seasonal fluctuations in predation risk and growth potential. Phenotypic variation in either individual predation risk or growth potential should thus mediate the strength of the trade-off and ultimately predict patterns of partial migration at the individual level (i.e. which individuals migrate and which remain resident). We provide cross-population empirical support for the importance of one component of this model--individual predation risk--in predicting partial migration in wild populations of bream Abramis brama, a freshwater fish. Smaller, high-risk individuals migrate with a higher probability than larger, low-risk individuals, and we suggest that predation risk maintains size-dependent partial migration in this system.     

Evolutionary aspects of migratory behavior in European warblers

During the past 20 years, European Sylvia warblers have been used for a model study of the control mechanisms of bird migration and of evolutionary aspects of migratory behavior. Endogenous annual rhythms (‘circannual’ rhythms) and photo-period have proved to be the essential internal and external controlling factors. It is unknown whether this basic system, that also controls migration in other bird species, is currently evolutionarily stable or is instead adapting birds to the present slightly changing environmental conditions. Using the Blackcap, the control of partial migration in a bird species was analyzed. Two-way selective breeding experiments demonstrated a large selection response and high heritability values. These experiments have also indicated that a partially migratory population can become either almost completely migratory, or sedentary, in two to five generations. Hence, genetic influences are very important and presumably dominant over environmental factors in the expression of migratory or sedentary behavior. The large selection response implies a strikingly high evolutionary potential with respect to strong selection pressures. Further, in the Blackcap, migratory orientation behavior (in addition to migratory activity) was immediately transmitted into a F₁-generation when a cross-breeding experiment was performed using birds from a migratory and a resident population. The hybrids displayed their migratory activity along an axis that is used by their migratory parents. Finally, a rapidly developing novel migratory habit (new migratory direction to new wintering areas) in the Blackcap is discussed with respect to a positive feedback-mechanism, possibly including a series of advantages leading to above-average fitness.     

Predicting conditions for migration: effects of density dependence and habitat quality

Migration is widespread among animals, but the factors that influence the decision to migrate are poorly understood. Within a single species, populations may be completely migratory, completely sedentary or partially migratory. We use a population model to derive conditions for migration and demonstrate how migratory survival, habitat quality and density dependence on both the breeding and non-breeding grounds influence conditions for migration and the proportion of migrants within a population. Density dependence during the season in which migratory and sedentary individuals use separate sites is necessary for partial migration. High levels of density dependence at the non-shared sites widen the range of survival values within which we predict partial migration, whereas increasing the strength of density dependence at the shared sites narrows the range of survival values within which we predict partial migration. Our results have important implications for predicting how contemporary populations with variable migration strategies may respond to changes in the quality or quantity of habitat.     

Postglacial northward expansion and genetic differentiation between migratory and sedentary populations of the broad‐tailed hummingbird (Selasphorus platycercus)

Unlike other migratory hummingbirds in North America, the broad‐tailed hummingbird (Selasphorus platycercus) exhibits both long‐distance migratory behaviour in the USA and sedentary behaviour in Mexico and Guatemala. We examined the evolution of migration linked to its northward expansion using a multiperspective approach. We analysed variation in morphology, mitochondrial and nuclear DNA, estimated migration rates between migratory and sedentary populations, compared divergence times with the occurrence of Quaternary climate events and constructed species distribution models to predict where migratory and sedentary populations resided during the Last Glacial Maximum (LGM) and Last Interglacial (LIG) events. Our results are consistent with a recent northward population expansion driven by migration from southern sedentary populations. Phylogeographical analyses and population genetics methods revealed that migratory populations in the USA and sedentary populations in Mexico of the platycercus subspecies form one admixed population, and that sedentary populations from southern Mexico and Guatemala (guatemalae) undertook independent evolutionary trajectories. Species distribution modelling revealed that the species is a niche tracker and that the climate conditions associated with modern obligate migrants in the USA were not present during the LIG, which provides indirect evidence for recent migratory behaviour in broad‐tailed hummingbirds on the temporal scale of glacial cycles. The finding that platycercus hummingbirds form one genetic population and that suitable habitat for migratory populations was observed in eastern Mexico during the LIG also suggests that the conservation of overwintering sites is crucial for obligate migratory populations currently facing climate change effects.     

Settling down of seasonal migrants promotes bird diversification

How seasonal migration originated and impacted diversification in birds remains largely unknown. Although migratory behaviour is likely to affect bird diversification, previous studies have not detected any effect. Here, we infer ancestral migratory behaviour and the effect of seasonal migration on speciation and extinction dynamics using a complete bird tree of life. Our analyses infer that sedentary behaviour is ancestral, and that migratory behaviour evolved independently multiple times during the evolutionary history of birds. Speciation of a sedentary species into two sedentary daughter species is more frequent than speciation of a migratory species into two migratory daughter species. However, migratory species often diversify by generating a sedentary daughter species in addition to the ancestral migratory one. This leads to an overall higher migratory speciation rate. Migratory species also experience lower extinction rates. Hence, although migratory species represent a minority (18.5%) of all extant birds, they have a higher net diversification rate than sedentary species. These results suggest that the evolution of seasonal migration in birds has facilitated diversification through the divergence of migratory subpopulations that become sedentary, and illustrate asymmetrical diversification as a mechanism by which diversification rates are decoupled from species richness.     

Ancestry and evolution of seasonal migration in the Parulidae

Seasonal migration in birds is known to be highly labile and subject to rapid change in response to selection, such that researchers have hypothesized that phylogenetic relationships should neither predict nor constrain the migratory behaviour of a species. Many theories on the evolution of bird migration assume a framework that extant migratory species have evolved repeatedly and relatively recently from sedentary tropical or subtropical ancestors. We performed ancestral state reconstructions of migratory behaviour using a comprehensive, well-supported phylogeny of the Parulidae (the 'wood-warblers'), a large family of Neotropical and Nearctic migratory and sedentary songbirds, and examined the rates of gain and loss of migration throughout the Parulidae. Counter to traditional hypotheses, our results suggest that the ancestral wood-warbler was migratory and that losses of migration have been at least as prevalent as gains throughout the history of Parulidae. Therefore, extant sedentary tropical radiations in the Parulidae represent losses of latitudinal migration and colonization of the tropics from temperate regions. We also tested for phylogenetic signal in migratory behaviour, and our results indicate that although migratory behaviour is variable within some wood-warbler species and clades, phylogeny significantly predicts the migratory distance of species in the Parulidae.     

Partial migration in fishes: definitions, methodologies and taxonomic distribution

Partial migration, where populations are composed of both migratory and resident individuals, is extremely widespread across the animal kingdom. Researchers studying fish movements have long recognized that many fishes are partial migrants, however, no detailed taxonomic review has ever been published. In addition, previous work and synthesis has been hampered by a varied lexicon associated with this phenomenon in fishes. In this review, definitions and important concepts in partial migration research are discussed, and a classification system of the different forms of partial migration in fishes introduced. Next, a detailed taxonomic overview of partial migration in this group is considered. Finally, methodological approaches that ichthyologists can use to study this fascinating phenomenon are reviewed. Partial migration is more widespread amongst fishes than previously thought, and given the array of techniques available to fish biologists to study migratory variation the future of the field looks promising.     

The evolution of avian migration

The study of avian migration has reached sophisticated levels in many areas, including ecology, behaviour, and physiology. Traditional discussions of the evolution of migration, however, have been compromised for several reasons. Previous ideas concerning the ancestral home of migrant species, southern or northern, and whether a partially migratory stage always precedes a fully migratory stage, were not expressed as testable hypotheses. Plotting migratory behaviour on phylogenetic trees has become commonplace and allows tests of traditional hypotheses. Some of these studies are reviewed, lending some support for almost all of the previous ideas. Although phylogenetic mapping helps to frame questions about the evolution of migration in a testable framework, there are two serious issues. First, experimental and observational studies reveal that the expression of migratory behaviour can change rapidly within a lineage, which can violate assumptions of character mapping. In addition, a species distribution model is used to show that current conditions for obligate migratory populations of the chipping sparrow were much restricted at the Last Glacial Maximum, and that the species might have been considered a partial migrant at that time. The expression of migratory behaviour in an extant species might be an artefact of the current inter‐glacial period. Only if the rate of gains and losses of migratory behaviour can be incorporated into a phylogenetic mapping exercise will the actual evolutionary pattern of migration be revealed. For example, reconstruction of the ancestral area and the evolutionary history of migratory categories in a clade of New World warblers depended on the assumptions of character state transitions. A second concern is that the trait ‘migratory’ is too broad for evolutionary analysis and that, if possible, the expression of hyperphagia, Zugunruhe, and navigation could be mapped individually. Loss or suppression of any of these components can lead to sedentary populations, revealing how migratory behaviour can appear and disappear rapidly. A report of low levels of Zugunruhe in a sedentary bird, Saxicola torquata, is reconstructed as derived in a clade of otherwise migratory populations, suggesting that the loss of migration was a result of suppression (but not elimination) of Zugunruhe. When researchers mention the independent origin of migration in a clade, they are most likely referring to the gain or loss of the expression of the ancestral migratory programme, not the de novo evolution of migration per se. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 237–250.     

Correlates of alternative migratory strategies in western bluebirds

Partial migration occurs when only some animals in a population migrate. While evidence suggests that migratory strategies are partially controlled by genes, individual and environmental conditions which alter the cost‐benefit trade‐off of migration among individuals are also likely to play a role. Three hypotheses have been advanced to explain condition‐dependent partial migration: the arrival time, dominance and body size hypotheses. In this study, we asked whether these hypotheses explained differences in migratory strategy among individuals in a partially migratory population of western bluebirds Sialia mexicana breeding in southern British Columbia, Canada. We used stable hydrogen isotope signatures in claw tissue to determine migratory strategy of individual bluebirds, and examined patterns of migration at both individual and population levels. The proportion of resident bluebirds varied significantly over the three years of the study, and across study sites. Several migrants switched to the resident strategy between years; however, we found no evidence of strategy switching in the opposite direction. Young birds were significantly more likely to be resident than older birds, a pattern which could arise if early arrival is particularly important for birds obtaining a territory for the first time. Furthermore, young females were the most likely of all sex–age classes to be resident, which may reflect a survival advantage of residency for young females. Finally, birds mated assortatively by migratory strategy and isotopic evidence suggests that members of a pair often wintered in the same place. Our results provided no support for the dominance or body size hypotheses, and only limited support for the arrival time hypothesis in bluebirds. However, taken together, we suggest that our findings indicate that social factors may influence migratory strategies in this system.     

Phylogeography of the Ryukyu robin (Erithacus komadori): population subdivision in land-bridge islands in relation to the shift in migratory habit

We analysed the mitochondrial control region sequences of 150 Ryukyu robins (Erithacus komadori) from five migratory and three sedentary populations. E. komadori is endemic to the subtropical Asian islands, is well differentiated among populations in morphology and migratory behaviour within its narrow habitat range, and is ideal for examining the effects of altered migratory habits on population differentiation. Maximum-likelihood analysis among the haplotypes resulted in the generation of a single tree in which two phylogenetic clades corresponding to the two subspecies were evident. Within the northern lineage, three distinctive subgroups of populations (one migratory and two sedentary groups) were observed by population genetic analysis. The migration-related wing morphology and molecular data were then compared among groups. The wing shape of the northern lineage was significantly more pointed than that of the southern sedentary lineage, and they were exclusively discriminated from each other. The difference was not so apparent between the sedentary and migratory groups within the northern lineage, suggesting that the present northern subspecies is primarily a migratory lineage. The different migratory behaviours of the robin played an apparent role in maintaining the genetic structure at two different levels--one between the southern and northern lineages and the other between the sedentary and migratory groups within the northern lineage. While gene flow had long been maintained among the islands occupied by migratory individuals, migrants have been unable to contribute to the gene pool of the sedentary populations despite passing through the breeding range of such populations twice each year.     

The need for integrative approaches to understand and conserve migratory ungulates

Over the last two centuries overhunting, anthropogenic barriers and habitat loss have disrupted many ungulate migrations. We review the literature on ungulate migration disruptions and find that for many species the disruption of migratory routes causes a rapid population collapse. Previous research has focused on the proximal ecological factors that might favour migration, particularly spatiotemporal variation in resources and predation. However, this does not provide an adequate basis for understanding and mitigating anthropogenic effects on migratory populations. Migration is a complex behaviour and we advocate an integrative approach that incorporates population dynamics, evolution, genetics, behaviour and physiology, and that borrows insights and approaches from research on other taxa. We draw upon research on avian migration to illustrate research approaches that might also be fruitful in ungulates. In particular, we suggest that the migratory cycle should be evaluated in the context of seasonal population limitation, an approach we highlight with a preliminary demographic perturbation analysis of the Serengeti wildebeest ( Connochaetes taurinus ) population. We provide suggestions for avenues of future research and highlight areas where we believe rapid progress can be made by applying recent advances in theory, technology and analytical approaches.     

Coalescent analyses of multiple loci support a new route to speciation in birds

A widely accepted paradigm is that sedentary Neotropical bird species are a reservoir that gives rise to temperate-tropical migratory species. Recently, an alternative theory has been proposed, that developmental plasticity can allow some individuals within a migratory species to establish a disjunct breeding range through loss of migration, thus facilitating the founding of a new sedentary species. We used mtDNA and two nuclear introns to perform coalescent analyses for two closely related New World oriole species, one a long-distance temperate-tropical migrant and the other a short-distance intratropical migrant. Our results suggest that the short-distance migrant recently diverged from the long-distance migrant via a founder event. In this species pair, the widely accepted paradigm is not supported. These results are consistent with a model of speciation through reduction of migratory distance.     

Migration costs drive convergence of threshold traits for migratory tactics

Partial migration of some, but not all, members of a population is a common form of migration. We evaluated how migration costs influence which members migrate in 10 populations of two salmonid species. The migratory patterns of both species were evaluated based on the size at maturity for resident males, which is the threshold trait that determines the migratory tactics used within a population. In both species, this size was smaller in males located further from the sea, where migration costs are presumably higher. Moreover, the threshold sizes at maturity in males were correlated between both species. Our results suggest that migration costs are a significant convergent selective force on migratory tactics and life-history traits in nature.     

Postglacial population expansion drives the evolution of long-distance migration in a songbird

The evolution of long-distance migratory behavior from sedentary populations is a central problem in studies of animal migration. Three crucial issues that remain unresolved are: (1) the biotic and abiotic factors promoting evolution of migratory behavior, (2) the geographic origin of ancestral sedentary populations, and (3) the time scale over which migration evolves. We test the role of postglacial population expansions during the Quaternary in driving the evolution of songbird migration against prevailing views favoring the role of intraspecific competition. In contrast to previous attempts to investigate these questions using interspecific phylogenies, we adopt an intraspecific approach and examine the phylogeography of a North American songbird, the chipping sparrow (Spizella passerina), which exhibits both long-distance migratory behavior in temperate North America and sedentary behavior in Mexico and Central America. We show that migratory populations descend from sedentary populations in southern Mexico and that migration has evolved as a result of a northward population expansion into temperate North America since the last glacial maximum 18,000 years ago. Migration appears to have evolved rapidly in some species as populations colonized areas of high seasonality in the temperate zone. The phylogeography of the yellow-eyed junco (Junco phaeonotus), a strictly sedentary species, provides a null model supporting the view that northward range expansions were driven solely by environmental factors and not by a predisposition to evolve migratory behavior. These results provide the strongest evidence to date that historical climate patterns can drive the rapid evolution of avian migration in natural populations, and they suggest a general mechanism for the repeated evolution of migration within and across bird lineages.     

Acoustic but no genetic divergence in migratory and sedentary populations of blackcaps,Sylvia atricapilla

In songbirds, territorial songs are key regulators of sexual selection and are learned from conspecifics. The cultural transmission of songs leads to divergence in song characteristics within populations, which can ultimately lead to speciation. Many songbirds also migrate, and individual differences in migratory behaviours can influence population genetic structure and local song differentiation. Blackcaps, Sylvia atricapilla, exhibit versatile territorial songs and show diversity in migration behaviours. They therefore comprise a good model for investigating the relationships between migratory patterns, song variation, and genetic diversity. We studied a migratory population (two groups near Paris) and a sedentary population (three groups in Corsica). All of the birds were ringed and blood sampled to investigate genetic relatedness using 17 microsatellite loci. A detailed song analysis showed that this species has a complex repertoire (> 100 syllables), which required the development of a semi‐supervised method to classify different categories of syllables and compare sequences of syllables. Our analysis showed no genetic structuring among populations: individuals belonging to the same group were not genetically closer than those from different groups. However, we found a strong wingsize difference between sedentary and migratory populations. We also showed that geographical variations in songs rely at least on both syllable and sequence content. Unexpectedly, despite a higher turnover of individuals, migratory groups share as many syllables and sequences as sedentary groups, which raises interesting issues on song learning and the maintenance of dialects in migratory birds.     

Migration strategy and divergent sexual selection on bird song

Migratory birds are assumed to be under stronger sexual selection pressure than sedentary populations, and the fact that their song is more complex has been taken as confirmation of this fact. However, this assumes that sexual selection pressure due to both male competition and female choice increase together. A further issue is that, in many species, songs become less complex during competitive encounters; in contrast, female choice selects for more complex song, so the two selection pressures may drive song evolution in different directions. We analysed song in two sedentary and two migratory populations of blackcaps (Sylvia atricapilla), a species in which different song parts are directed to males and females. We found that migratory populations produce longer, female-directed warbles, indicating sexual selection through female choice is the strongest in these populations. However, the part of the song directed towards males is shorter and more repetitive (as observed in individual competitive encounters between males) in non-migratory populations, indicating sedentary populations, are under stronger selection due to male competition. We show for the first time that the intensity of selection pressure from male competition and female choice varies independently between populations with different migratory behaviours. Rapid alterations in the migration patterns of species are thus likely to lead to unexpected consequences for the costs and benefits of sexual signals.     

Caribou movement as a correlated random walk

Movement is a primary mechanism coupling animals to their environment, yet there exists little empirical analysis to test our theoretical knowledge of this basic process. We used correlated random walk (CRW) models and satellite telemetry to investigate long-distance movements of caribou, the most vagile, non-volant terrestrial vertebrate in the world. Individual paths of migratory and sedentary female caribou were quantified using measures of mean move length and angle, and net squared displacements at each successive move were compared to predictions from the models. Movements were modelled at two temporal scales. For paths recorded through one annual cycle, the CRW model overpredicted net displacement of caribou through time. For paths recorded over shorter intervals delineated by seasonal behavioural changes of caribou, there was excellent correspondence between model predictions and observations for most periods for both migratory and sedentary caribou. On the smallest temporal scale, a CRW model significantly overpredicted displacements of migratory caribou during 3 months following calving; this was also the case for sedentary caribou in late summer, and in late winter. In all cases of overprediction there was significant positive autocorrelation in turn direction, indicating that movements were more tortuous than expected. In one case of underprediction, significant negative autocorrelation of sequential turn direction was evident, indicating that migratory caribou moved in straightened paths during spring migration to calving grounds. Results are discussed in light of known migration patterns and possible limiting factors for caribou, and indicate the applicability of CRW models to animal movement at vast spatial and temporal scales, thus assisting in future development of more sophisticated models of population spread and redistribution for vertebrates. Received: 14 July 1999 / Accepted: 15 November 1999     

Genomic legacy of migration in endangered caribou

Wide-ranging animals, including migratory species, are significantly threatened by the effects of habitat fragmentation and habitat loss. In the case of terrestrial mammals, this results in nearly a quarter of species being at risk of extinction. Caribou are one such example of a wide-ranging, migratory, terrestrial, and endangered mammal. In populations of caribou, the proportion of individuals considered as “migrants” can vary dramatically. There is therefore a possibility that, under the condition that migratory behavior is genetically determined, those individuals or populations that are migratory will be further impacted by humans, and this impact could result in the permanent loss of the migratory trait in some populations. However, genetic determination of migration has not previously been studied in an endangered terrestrial mammal. We examined migratory behavior of 139 GPS-collared endangered caribou in western North America and carried out genomic scans for the same individuals. Here we determine a genetic subdivision of caribou into a Northern and a Southern genetic cluster. We also detect >50 SNPs associated with migratory behavior, which are in genes with hypothesized roles in determining migration in other organisms. Furthermore, we determine that propensity to migrate depends upon the proportion of ancestry in individual caribou, and thus on the evolutionary history of its migratory and sedentary subspecies. If, as we report, migratory behavior is influenced by genes, caribou could be further impacted by the loss of the migratory trait in some isolated populations already at low numbers. Our results indicating an ancestral genetic component also suggest that the migratory trait and their associated genetic mutations could not be easily re-established when lost in a population.     

Migratory diversity predicts population declines in birds

Declines in migratory species are a pressing concern worldwide, but the mechanisms underpinning these declines are not fully understood. We hypothesised that species with greater within‐population variability in migratory movements and destinations, here termed ‘migratory diversity’, might be more resilient to environmental change. To test this, we related map‐based metrics of migratory diversity to recent population trends for 340 European breeding birds. Species that occupy larger non‐breeding ranges relative to breeding, a characteristic we term ‘migratory dispersion’, were less likely to be declining than those with more restricted non‐breeding ranges. Species with partial migration strategies (i.e. overlapping breeding and non‐breeding ranges) were also less likely to be declining than full migrants or full residents, an effect that was independent of migration distance. Recent rates of advancement in Europe‐wide spring arrival date were greater for partial migrants than full migrants, suggesting that migratory diversity may also help facilitate species responses to climate change.