Heritability of arrival date in a migratory bird

The behaviour of long-distance migratory birds is assumed to partly be under the influence of genes, as demonstrated by selection experiments. Furthermore, competition for early arrival among males may lead to condition-dependent migration associated with fitness benefits of early arrival achieved by individuals in prime condition. Here I present field data on the repeatability and the heritability of arrival date in a trans-equatorial migratory bird, the barn swallow Hirundo rustica, and I test for a genetic correlation between arrival date and the expression of a condition-dependent secondary sexual character. The repeatability was statistically significant and the heritability of arrival date was estimated to be 0.54 (s.e. = 0.15). There was no significant evidence of this estimate being inflated by environmental or maternal condition during rearing. Arrival date and migration are condition dependent in the barn swallow, with males with the most exaggerated secondary sexual characters also arriving the earliest. There was a significant genetic correlation between arrival date and tail length in male barn swallows, providing indirect evidence for a genetic basis of this condition dependence. Given the high level of heritability, arrival date could readily respond to selection caused by environmental change.     

Pairing context determines condition-dependence of song rate in a monogamous passerine bird

Condition-dependence of male ornaments is thought to provide honest signals on which females can base their sexual choice for genetic quality. Recent studies show that condition-dependence patterns can vary within populations. Although long-term association is thought to promote honest signalling, no study has explored the influence of pairing context on the condition-dependence of male ornaments. In this study, we assessed the influence of natural variation in body condition on song rate in zebra finches (Taeniopygia guttata) in three different situations: during short and long encounters with an unfamiliar female, and within heterosexual mated pairs. We found consistent individual differences in male directed and undirected song rate. Moreover, body condition had a positive effect on song rate in paired males. However, male song rate was not influenced by body condition during short or long encounters with unfamiliar females. Song rate appears to be an unreliable signal of condition to prospective females as even poor-condition birds can cheat and sing at a high rate. By contrast, paired females can reliably use song rate to assess their mate''s body condition, and possibly the genetic quality. We propose that species'' characteristics, such as mating system, should be systematically taken into account to generate relevant hypotheses about the evolution of condition-dependent male ornaments.     

Extrapair paternity in relation to sexual ornamentation, arrival date, and condition in a migratory bird

We tested the novel hypothesis that arrival date in migratorybirds represents a reliable indicator of male quality that canbe used by females as a cue in extrapair mating decisions. Secondarysexual characters are often condition-dependent, and competitionfor early arrival leads to condition-dependent migration. Hence,both secondary sexual characters and arrival date are predictedto be condition-dependent indicators of male phenotypic quality.We studied the relationship between expression of a secondarysexual character, arrival date, and condition, respectively,and extrapair paternity in a Spanish population of barn swallows,Hirundo rustica. By using microsatellite markers to determinepaternity, we showed that 17.8% of all offspring (N = 674) and32.4% of all broods (N = 170) were due to extrapair paternity.Quasi-parasitism (in which the male nest owner fathered theoffspring, but the eggs were laid by another female) occurredin 2.6% of all nestlings and 2.9% of all broods. Individualswere consistent in the frequency of extrapair paternity amongfirst, second, and third broods. Males with long outermost tailfeathers, arriving early and in prime body condition, had littleextrapair paternity in their nests. This was also the case whencontrolling for the confounding effects of male age. Partialcorrelation analysis was used to investigate the direct andindirect effects of tail length, arrival date, and body conditionon extrapair paternity. Body condition accounted for most ofthe variance in extrapair paternity, whereas tail length andarrival date accounted for a smaller proportion of the variance.Body condition was strongly correlated with tail length andarrival date. However, because females cannot directly assesscondition or arrival date (males arrive before females), femalesmay obtain an indirect measure of condition and migration abilityfrom tail length and other phenotypic traits of males. Thissuggests that extrapair paternity depends on the effects ofcondition, through its indirect effects on arrival date, taillength, and other variables.     

Ecological conditions during winter predict arrival date at the breeding quarters in a trans-Saharan migratory bird

We studied variation in arrival date to the breeding colonies in Italy of a trans‐Saharan migratory bird, the barn swallow Hirundo rustica, in relation to variation in ecological conditions, as reflected by the normalized difference vegetation index (NDVI), in the winter quarters. Arrival date of old but not young individuals captured during consecutive breeding seasons was earlier after winters with favourable conditions. Change in arrival date in relation to change in NDVI was similar in the two sexes. Change in arrival date significantly and positively predicted change in breeding date. As a result of increased frequency of second broods determined by earlier arrival, the number of fledged offspring per season was larger after African winters with good in comparison to poor ecological conditions for barn swallows. This is the first study demonstrating phenotypic plasticity in migration phenology of a long‐distance migratory bird in relation to ecological conditions during wintering.     

Variability,heritability and condition-dependence of the multidimensional male colour phenotype in a passerine bird

Elaborate ornamental traits are commonly assumed to be honest signals of individual quality, owing to the presumed costs involved in their production and/or maintenance. Such traits are often highly variable, possibly because of condition-dependence and/or high underlying genetic variation, and it has been suggested that their expression should be more sensitive to condition and/or more heritable than non-ornamental traits. Many bird species display colourful plumage with multiple distinct patches of different developmental origins, forming complex colour phenotypes. Despite this complexity, colourful ornaments are often studied in isolation, without comparison to suitable non-ornamental controls. Based on plumage reflectance data collected over 8 years, we assessed the signalling potential of the multidimensional male colour phenotype in a tropical bird: the purple-crowned fairy-wren Malurus coronatus. Specifically, we tested the predictions that the expression of putative ornamental colours (purple and black – the breeding colours – and blue) is (1) more variable, (2) more heritable and (3) more condition-dependent compared to year-round non-ornamental colours (buff-white and brown). Our results show that ornamental colours exhibit greater levels of variability, and some chromatic components of purple and blue colouration appear slightly heritable (h² = 0.19–0.30). However, contrary to predictions of heightened condition-dependence in ornaments, only brightness of the buff-white and brown colouration increased with male body condition, although brightness of the purple colouration was related to male age as expected. Despite partial support for predictions, the lack of consistent patterns illustrates the complexity of visual signals and highlights the need to study colour phenotypes in their entirety.Subject terms: Behavioural ecology, Evolutionary ecology, Ecological genetics, Quantitative trait, Sexual selection     

A matter of timing: changes in the first date of arrival and last date of departure of Australian migratory birds

Although there is substantial evidence that Northern Hemisphere species have responded to climatic change over the last few decades, there is little documented evidence that Southern Hemisphere species have responded in the same way. Here, we report that Australian migratory birds have undergone changes in the first arrival date (FAD) and last date of departure (LDD) of a similar magnitude as species from the Northern Hemisphere. We compiled data on arrival and departure of migratory birds in south‐east Australia since 1960 from the published literature, Bird Observer Reports, and personal observations from bird watchers. Data on the FAD for 24 species and the LDD for 12 species were analyzed. Sixteen species were short‐ to middle‐distance species arriving at their breeding grounds, seven were long‐distance migrants arriving at their nonbreeding grounds, and one was a middle‐distance migrant also arriving at its nonbreeding ground. For 12 species, we gathered data from more than one location, enabling us to assess the consistency of intraspecific trends at different locations. Regressions of climate variables against year show that across south‐east Australia average annual maximum and minimum temperatures have increased by 0.17°C and 0.13°C decade^?1 since 1960, respectively. Over this period there has been an average advance in arrival of 3.5 days decade^?1; 16 of the 45 time‐series (representing 12 of the 24 species studied) showed a significant trend toward earlier arrival, while only one time‐series showed a significant delay. Conversely, there has been an average delay in departure of 5.1 days decade^?1; four of the 21 departure time‐series (four species) showed a significant trend toward later departure, while one species showed a significant trend toward earlier departure. However, differences emerge between the arrival and departure of short‐ to middle‐distance species visiting south‐east Australia to breed compared with long‐distance species that spend their nonbreeding period here. On average, short‐ to middle‐distance migrants have arrived at their breeding grounds 3.1 days decade^?1 earlier and delayed departure by 8.1 days decade^?1, thus extending the time spent in their breeding grounds by ～11 days decade^?1. The average advance in arrival at the nonbreeding grounds of long‐distance migrants is 6.8 days decade^?1. These species, however, have also advanced departure by an average of 6.9 days decade^?1. Hence, the length of stay has not changed but rather, the timing of events has advanced. The patterns of change in FAD and LDD of Australian migratory birds are of a similar magnitude to changes undergone by Northern Hemisphere species, and add further evidence that the modest warming experienced over the past few decades has already had significant biological impacts on a global scale.     

Extraordinary high natal philopatry in a migratory passerine

Natal dispersal is a key component of population dynamics in birds. It guarantees the genetic exchange of populations, favours range expansions and reduces intraspecific competition. In general, natal philopatry of migratory passerines is quite low (0–13.5%). In this study, we give evidence that, under specific conditions, migratory populations of passerine birds may show a considerably elevated natal philopatry. In a Linnet Carduelis cannabina population on the remote island of Helgoland in the North Sea, we found an extraordinary high return rate of yearlings (38%), which corresponds exactly to the annual survival rate of the species. Despite being completely migratory, the Linnets of Helgoland apparently return to a large extent to their native area and consequently might support the population maintenance on the island. Further studies are needed to reveal if this high natal philopatry is only an unusual 1-year event or a general characteristic of this partially isolated island population.     

Food availability during migratory stopover affects testis growth and reproductive behaviour in a migratory passerine

Long-distance migratory passerines initiate testicular recrudescence during spring migration to meet the demands of timely reproduction upon immediate arrival on the breeding grounds. The degree of testicular development is known to depend on environmental factors like stopover habitat quality; reproductive performance may be strongly impacted by testicular maturation upon arrival on the breeding grounds. We investigated the effect of stopover food availability on subsequent reproductive performance in garden warblers (Sylvia borin). Spring migration was simulated by repeated food deprivation and re-feeding to imitate the alternation of flight and stopover periods. During the two final stopover periods, males were either kept under ad libitum food (ad libitum males) or under limited food conditions (limited males). After simulated arrival in the breeding area, manipulation of previous stopover food availability resulted in significantly slower testicular recrudescence (p < 0.001) and decreased plasma testosterone (p < 0.01) in limited males compared to ad libitum males. Body mass change was not significantly different between the two groups (p = 0.38). Limited males also exhibited reduced performance in reproductive behaviours employed in territorial and sexual contexts. Limited males had a longer ‘freezing’ interval (p < 0.05) and decreased activity (p < 0.01) when challenged with a live male decoy. In direct confrontation between limited and ad libitum males in the presence of a female, limited males exhibited significantly fewer behavioural traits in sexual context, i.e. directed to the female (p < 0.001). Therefore, we suggest that conditions encountered during previous migratory stopover may affect subsequent annual reproductive success by influencing key reproductive behaviours.     

Maladaptive habitat selection of a migratory passerine bird in a human-modified landscape

In human-altered environments, organisms may preferentially settle in poor-quality habitats where fitness returns are lower relative to available higher-quality habitats. Such ecological trapping is due to a mismatch between the cues used during habitat selection and the habitat quality. Maladaptive settlement decisions may occur when organisms are time-constrained and have to rapidly evaluate habitat quality based on incomplete knowledge of the resources and conditions that will be available later in the season. During a three-year study, we examined settlement decision-making in the long-distance migratory, open-habitat bird, the Red-backed shrike (Lanius collurio), as a response to recent land-use changes. In Northwest Europe, the shrikes typically breed in open areas under a management regime of extensive farming. In recent decades, Spruce forests have been increasingly managed with large-size cutblocks in even-aged plantations, thereby producing early-successional vegetation areas that are also colonised by the species. Farmland and open areas in forests create mosaics of two different types of habitats that are now occupied by the shrikes. We examined redundant measures of habitat preference (order of settlement after migration and distribution of dominant individuals) and several reproductive performance parameters in both habitat types to investigate whether habitat preference is in line with habitat quality. Territorial males exhibited a clear preference for the recently created open areas in forests with higher-quality males settling in this habitat type earlier. Reproductive performance was, however, higher in farmland, with higher nest success, offspring quantity, and quality compared to open areas in forests. The results showed strong among-year consistency and we can therefore exclude a transient situation. This study demonstrates a case of maladaptive habitat selection in a farmland bird expanding its breeding range to human-created open habitats in plantations. We discuss the reasons that could explain this decision-making and the possible consequences for the population dynamics and persistence.     

Reduced compensatory growth capacity in mistimed broods of a migratory passerine

Phenotypic plasticity has recently been proposed to increase population viability when rapid anthropogenic environmental changes cannot be tracked by means of evolution. This assumes that environmental changes do not constrain phenotypic plasticity itself, which has rarely been examined in natural populations. In areas of climate warming, many long-distance migratory birds breed increasingly late relative to the period of peak food supply, and the temporal mismatch may constrain plastic life-history traits such as nestling growth. We combined 23 years of food availability and breeding data with a 3-year experimental manipulation of nestling growth trajectories in a Central European population of collared flycatchers (Ficedula albicollis) to examine the potential impact of climate-related mistimed breeding on nestling developmental plasticity. Timing of the food peak was predicted by winter climate, and the median hatching date of broods was earlier in springs with earlier food peaks. However, the adjustment of hatching date was incomplete and the population largely missed the food peak in years with very early food peaks. After imposing a temporary, experimental food shortage on nestlings, the extent of compensatory growth in body mass differed among years, and this difference was apparently related to the distance of hatching dates from the yearly food peak. Growth compensation declined with distance from the peak. These results suggest that mistimed phenology may not only create permanently adverse conditions for migratory species but it may also constrain the plastic responses of individuals to temporary disturbances. Therefore, climate change may not only favour but also restrict phenotypic plasticity.     

Reverse turning and angle compensation in a migratory passerine,passerculus sandwichensis

Reverse turning behaviour was revealed in the savannah sparrow (Passerculus sandwichensis), a migratory passerine, using a T-shaped runway system. Compensatory reverse turns were recorded for experimental birds which were forced to make 90° turns either to the left or right before proceeding to a choice point. Controls ran a straight runway from start to choice point and were as likely to turn left as right. A second experiment revealed an ability to compensate for angular deviations in course. Experimentals were forced to make turns of less than 90° before entering a semi-circular arena. They responded with a compensatory turn of the appropriate sign (left or right) and magnitude. An idiothetic mechanism is probably involved in the observed angle sense (Winkelsinn). The results are broadly relevant to animal orientation.     

Impact of density and environmental factors on population fluctuations in a migratory passerine

1. Populations of plants and animals typically fluctuate because of the combined effects of density-dependent and density-independent processes. The study of these processes is complicated by the fact that population sizes are typically not known exactly, because population counts are subject to sampling variance. Although the existence of sampling variance is broadly acknowledged, relatively few studies on time-series data have accounted for it, which can result in wrong inferences about population processes. 2. To increase our understanding of population dynamics, we analysed time series from six Central European populations of the migratory red-backed shrike Lanius collurio by simultaneously assessing the strength of density dependence, process and sampling variance. In addition, we evaluated hypotheses predicting effects of factors presumed to operate on the breeding grounds, at stopover sites in eastern Africa during fall and spring migration and in the wintering grounds in southern Africa. We used both simple and state-space formulations of the Gompertz equation to model population size. 3. Across populations and modelling approaches, we found consistent evidence for negative density-dependent population regulation. Further, process variance contributed substantially to variation in population size, while sampling variance did not. Environmental conditions in eastern and southern Africa appear to influence breeding population size, as rainfall in the Sahel during fall migration and in the south African wintering areas were positively related to population size in the following spring in four of six populations. In contrast, environmental conditions in the breeding grounds were not related to population size. 4. Our findings suggest negative density-dependent regulation of red-backed shrike breeding populations and are consistent with the long-standing hypothesis that conditions in the African staging and wintering areas influence population numbers of species breeding in Europe. 5. This study highlights the importance of jointly investigating density-dependent and density-independent processes to improve our understanding of factors influencing population fluctuations in space and time.     

Blood parasites shape extreme major histocompatibility complex diversity in a migratory passerine

Pathogens are one of the main forces driving the evolution and maintenance of the highly polymorphic genes of the vertebrate major histocompatibility complex (MHC). Although MHC proteins are crucial in pathogen recognition, it is still poorly understood how pathogen‐mediated selection promotes and maintains MHC diversity, and especially so in host species with highly duplicated MHC genes. Sedge warblers (Acrocephalus schoenobaenus) have highly duplicated MHC genes, and using data from high‐throughput MHC genotyping, we were able to investigate to what extent avian malaria parasites explain temporal MHC class I supertype fluctuations in a long‐term study population. We investigated infection status and infection intensities of two different strains of Haemoproteus, that is avian malaria parasites that are known to have significant fitness consequences in sedge warblers. We found that prevalence of avian malaria in carriers of specific MHC class I supertypes was a significant predictor of their frequency changes between years. This finding suggests that avian malaria infections partly drive the temporal fluctuations of the MHC class I supertypes. Furthermore, we found that individuals with a large number of different supertypes had higher resistance to avian malaria, but there was no evidence for an optimal MHC class I diversity. Thus, the two studied malaria parasite strains appear to select for a high MHC class I supertype diversity. Such selection may explain the maintenance of the extremely high number of MHC class I gene copies in sedge warblers and possibly also in other passerines where avian malaria is a common disease.     

Parasite assemblages distinguish populations of a migratory passerine on its breeding grounds

We attempted to establish migratory connectivity patterns for American redstarts Setophaga ruticilla between their breeding and wintering grounds by characterizing the composition of their haematozoan parasite assemblages in different parts of their range. We detected significant but limited geographic structuring of haematozoan parasite lineages across the breeding range of the redstart. We found that redstarts from the south-eastern (SE) region of the breeding range had a significantly different haematozoan parasite assemblage compared with populations sampled throughout the rest of the breeding range. Evidence using stable isotopes from feathers previously demonstrated that redstarts from the SE of the breeding range also have a unique and separate wintering range. Thus, although two methods of estimating migratory connectivity have now both shown the SE US breeding sub-population of redstarts to be distinct from other populations on both the breeding and wintering grounds, conclusive migratory connectivity for this species as a whole could not be established.     

Population-scale drivers of individual arrival times in migratory birds

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.