Haemosporidian parasitism in the blackcap Sylvia atricapilla in relation to spring arrival and body condition

Parasites can exert strong selection on hosts. Throughout the year migrants are exposed to different sets of parasites, which may affect life history traits such as migratory schedules. Here, we studied the relationship between parasite infection and arrival date of blackcaps Sylvia atricapilla to their breeding grounds in Germany throughout a period of six years (2007–2012). We used two data sets, one that included all blackcaps and one that included only recaptured birds. We assesed whether parasites influence spring arrival to breeding grounds, and for the recaptured data set, we analysed temporal variation in parasitism (i.e. infection status and parasitaemia) throughout the breeding season. We used both microscopy and PCR (a fragment of ? 479 bp of the mtDNA cyt b) to determine haemosporidian infection. Blackcaps were mostly infected with Haemoproteus parabelopolskyi (lineages SYAT01 and SYAT02). Infection status, but not parasitaemia, was constant through time for individual birds; meaning that once a bird is infected, it most likely will retain the infection for life. We found that infection by haemosporidian parasites has no relationship to arrival date in this blackcap population; however, infection by H. parabelopolskyi has a marginally significant effect on arrival date of recaptured blackcaps, somewhat delaying their arrival to breeding grounds. Birds captured later in the season were more likely to be infected than those from early spring, and parasitaemia was frequently lower in birds captured earlier in the season compared to those captured later (summer).     

Isolation of 10 tetranucleotide microsatellite loci in the blackcap (Sylvia atricapilla)

We isolated 10 polymorphic microsatellite DNA loci from the blackcap (Sylvia atricapilla) and optimized them for future studies of population differentiation in populations with different migration strategies in southwestern Germany. The loci were screened for polymorphism using 178 individuals from two populations in Germany and Spain. The primers amplified highly variable loci characterized by two to 19 alleles per locus and their observed and expected heterozygosities range from 0.47 to 0.81 and from 0.50 to 0.91, respectively.     

Effect of Haemoproteus belopolskyi (Haemosporida: Haemoproteidae) on body mass of the blackcap Sylvia atricapilla

The effect of initial Haemoproteus belopolskyi infection on the weight of its natural host, the blackcap Sylvia atricapilla, was investigated. Fourteen blackcap nestlings were taken at the age of 4-5 days and raised by hand in the laboratory. They were free of blood parasites. Seven 20- to 21-day-old nestlings were infected experimentally by inoculation in their pectoral muscle with approximately 45 sporozoites, which had developed in the experimentally infected biting midge Culicoides impunctatus. Seven nestlings were used as negative controls. Parasitemia developed in 6 inoculated nestlings, with a prepatent period of 11-12 days. No infections were detected in the controls during this study. The weight of experimentally infected and control birds was measured 2 days before parasitemia became patent and for a 45-day period after patency. Blood smears were prepared from all birds on the days when they were weighed. When compared with controls, there was a significant weight loss of experimentally infected blackcaps during 6 days after the decline of parasitemia at 10-16 days of patency, indicating a short-term influence of the infection on the birds' body mass. Clinical symptoms of the infection were not recorded. All birds from both groups survived until the end of the experiment.     

Ecophysiological aspects of rapid population growth in a novel migratory blackcap (Sylvia atricapilla) population: an experimental approach

Summary Blackcaps (Sylvia atricapilla) that breed in central Europe have usually migrated to Mediterranean or African wintering grounds. In the past several decades, a portion of this breeding population has started migrating to the British Isles to overwinter and this population has increased dramatically. Several factors, including higher annual survivorship (due to supplemental feeding and reduced migratiry distance), assortative mating, and enhanced reproductive success may be involved in this rapid population growth. As part of an intensive, long-term study of this population, we tested the hypothesis that the differences in photoperiod experienced by British-wintering versus Mediterranean-wintering blackcaps might lead to relatively early vernal (i.e., migratory and/or reproductive) physiological condition in members of the former group. We found that birds exposed to photoperiodic conditions that simulated migration to Britain to overwinter generally initiated vernal migratory activity earlier than birds held under conditions simulating migration to traditional wintering areas in central Spain. This difference, coupled with the shorter migratory distance to the British Isles, leads to significantly earlier estimated arrival dates for blackcaps that winter in Britain compared to central Spain. Bimodality in arrival times suggests that assortative mating on central European breeding grounds might occur between members of the different wintering populations. Males exposed to British-winter photoperiods showed significantly earlier testicular development than males kept under Spanish-winter photoperiods. Early arrival on the breeding grounds, coupled with accelerated reproductive condition, should lead to a relatively early reproductive effort, perhaps increasing average reproductive success. In general, these results support the hypothesis that differences in photoperiod on the wintering grounds may play an important role in the dynamic state of this population.     

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     

Mechanical and structural adaptations to migration in the flight feathers of a Palaearctic passerine

Current avian migration patterns in temperate regions have been developed during the glacial retreat and subsequent colonization of the ice‐free areas during the Holocene. This process resulted in a geographic gradient of greater seasonality as latitude increased that favoured migration‐related morphological and physiological (co)adaptations. Most evidence of avian morphological adaptations to migration comes from the analysis of variation in the length and shape of the wings, but the existence of intra‐feather structural adjustments has been greatly overlooked despite their potential to be under natural selection. To shed some light on this question, we used data from European robins Erithacus rubecula overwintering in Campo de Gibraltar (Southern Iberia), where sedentary robins coexist during winter with conspecifics showing a broad range of breeding origins and, hence, migration distances. We explicitly explored how wing length and shape, as well as several functional (bending stiffness), developmental (feather growth rate) and structural (size and complexity of feather components) characteristics of flight feathers, varied in relation to migration distance, which was estimated from the hydrogen stable isotope ratios of the summer‐produced tail feathers. Our results revealed that migration distance not only favoured longer and more concave wings, but also promoted primaries with a thicker dorsoventral rachis and shorter barb lengths, which, in turn, conferred more bending stiffness to these feathers. We suggest that these intra‐feather structural adjustments could be an additional, largely unnoticed, adaptation within the avian migratory syndrome that might have the potential to evolve relatively quickly to facilitate the occupation of seasonal environments.     

Blackcaps Sylvia atricapilla stopping over at the desert edge; inter- and intra-sexual differences in spring and autumn migration

Spring and autumn Palaearctic-African migration patterns of Blackcaps Sylvia atricapilla during stopover at Elat, Israel, showed that males appeared significantly earlier than females during spring but not during autumn migration, suggesting that in males there is a stronger drive to reaching breeding territories early. The difference in mean appearance dates between sexes in spring tended to be greatest in years when the males appeared earliest. Longer spread of passage (the dates between which the central 50% of individuals were captured) for each sex in spring was found in years with an early mean passage datebut was significant only for females. These observations suggest that the timing of Blackcap migration is governed not only by endogenous factors but also by exogenous factors, and when the environmental conditions are unfavourable, the differences in passage dates between sexes decrease and the passage lengths shorten. The early individuals (both males and females) that stopped over at Elat in spring were those with relatively small body size (as indicated by relatively short wings) and relatively large fat reserves and in good body condition (as indicated from fat score and body mass/wing-length ratio). No differences in body size between early and late transients were detected during the autumn migration, but late birds of both sexes carried larger fat reserves. These phenomena may be explained either by leap-frog migration or by differential fitness among wintering males and females or both, with only the fittest Blackcaps being capable of an early departure. These individuals probably face much less intensive intra- and interspecific competition with residents and other transients in stopover sites than do later transients.     

Ecomorphological patterns in the Blackcap Sylvia atricapilla: insular versus mainland populations

Capsule Blackcap Sylvia atricapilla populations from the Azores archipelago show morphological differences to continental birds which are consistent with the ‘Island Rule’.

Aims The morphology of insular vertebrates is usually the result of the evolution in their particular environment and leads to predictable morphological patterns, according to the Island Rule. We test the predictions of the Island Rule, using the Blackcap of the Azores archipelago as our model.

Methods We compared morphological variation (body size and wing shape) of populations from the nine islands of the Azores to continental birds, using multivariate indexes. Also, we looked at the relationship between these patterns and possible insular ecological drivers of morphological divergence.

Results Our findings are concordant with Island Rule predictions, as in general birds from the Azores are larger than continental populations, especially birds from the most distant islands. Wing shape also differs significantly, as Azorean Blackcaps tend to have rounder wings than continental birds with a migratory-like phenotype.

Conclusion Overall, we conclude that the observed morphological patterns in Blackcap in the Azores conform in general to the Island Rule predictions.     

Wing moult in relation to autumn migration in adult Common Whitethroats Sylvia communis communis

Most long-distance passerine migrants in Sweden moult on breeding grounds before leaving on autumn migration to winter quarters. However, birds laying second or replacement clutches, or just breeding late, have too little time for a normal moult on the breeding grounds. When time is limited the birds may respond by making various adjustments to the moult, for example by moulting more quickly or by suspending the moult. In this study, the relationship between the performance of post-nuptial remex moult in Common Whitethroats breeding on Gotland, southeast Sweden, and autumn migration departure was investigated. The majority (77%) of the birds had interrupted moult in either the primaries or secondaries. Interruption of moult was more common among birds with a later onset date, as was asymmetry in moult between wings. The interruption of moult led to a significant time gain and moult completion was, consequently, more synchronized than moult onset. The results from this study indicate, in accordance with other data, that an early start of autumn migration is important. An early start may be crucial to facilitate the crossing of the Sahara Desert once the dry season has begun.     

Resistance of flight feathers to mechanical fatigue covaries with moult strategy in two warbler species

Flight feather moult in small passerines is realized in several ways. Some species moult once after breeding or once on their wintering grounds; others even moult twice. The adaptive significance of this diversity is still largely unknown. We compared the resistance to mechanical fatigue of flight feathers from the chiffchaff Phylloscopus collybita, a migratory species moulting once on its breeding grounds, with feathers from the willow warbler Phylloscopus trochilus, a migratory species moulting in both its breeding and wintering grounds. We found that flight feathers of willow warblers, which have a shaft with a comparatively large diameter, become fatigued much faster than feathers of chiffchaffs under an artificial cyclic bending regime. We propose that willow warblers may strengthen their flight feathers by increasing the diameter of the shaft, which may lead to a more rapid accumulation of damage in willow warblers than in chiffchaffs.     

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.     

Effect of photoperiod on incubation period in a wild passerine,Sylvia atricapilla

Time required for avian embryos to develop is influenced by incubation temperature and the amount of time adults incubate eggs. Experiments on poultry indicate that photoacceleration, the light‐induced stimulation of embryonic development, decreases the length of the incubation period as embryos receive more light. We hypothesized that eggs of wild birds exposed to longer periods of light should also have shorter incubation periods. We tested whether photoacceleration would occur in a species of open‐cup nesting passerine, the blackcap Sylvia atricapilla. We artificially incubated blackcap eggs under four different photoperiods, four hours of light (4L) and 20 h of dark (20D), 12L:12D, 20L:4D, and a skeleton photoperiod (1 h light, 2 times per day) that framed a 20 h day. While incubation periods were accelerated with increasing photoperiod length, the differences among photoperiods of 4, 12 and 20L were weak. Embryos exposed to skeleton photoperiods developed as fast as those exposed to 20L and significantly faster than those exposed to 4L and 12L treatments. Skeleton photoperiods may most closely approximate natural patterns of light exposure that embryos experience during dawn and dusk incubation recesses typically associated with adult foraging. If our results from this species also occur in other wild birds, exposure to different day lengths may help explain some of the variation in the observed seasonal and latitudinal trends in avian incubation period.     

Blackcaps (Sylvia atricapilla) increase the whistle part of their song in response to simulated territorial intrusion

Bird song is a sexually selected signal that serves two main functions, attracting a mate and deterring rivals. Different signal parameters may be important in advertising to females compared to advertising to rival males. Species solve the problem of this dual function in a variety of ways, one of which may be to have separate parts of song directed at male and female receivers. The blackcap song has two distinct parts, a complex warble, assumed to be directed at female receivers, followed by a louder and more stereotyped whistle putatively directed at males. We simulated territorial intrusions by broadcasting blackcap song in territories. Comparing songs sung prior, with those produced in response to playback, showed that the proportion of the whistle component of songs increased, but not the warble. This study thus provides empirical evidence that the whistle component of the blackcap song plays a prominent role in male–male competition. The warble component of the blackcap song may be directed at females, but this requires further testing.     

Consistency in Egg Rejection Behaviour: Responses to Repeated Brood Parasitism in the Blackcap (Sylvia atricapilla)

To evaluate host responses towards repeated brood parasitism we experimentally parasitized and continuously videotaped blackcap nests in two consecutive trials. The ejection of a foreign egg was the most common response (94.5%) in both trials, but desertion (4.1%) and acceptance (1.4%) also occurred. The general method of ejection was puncturing. In 9.8% of identified ejections, already punctured eggs stuck to the abdominal feathers of the incubating bird and were carried out of the nest. Females were responsible for the majority of ejections in both trials and their response time was significantly shorter than that of males. Blackcaps exhibited consistency in the sex responsible for egg ejection over the two trials; but in five (20.8%) experiments individuals changed their behaviour. Repeatability for host responses within the nest was very high. In ejections accomplished by the same bird, the response was significantly quicker in the second trial, indicating the presence of certain learning abilities. Our results suggest that cuckoo hosts are quite consistent in their responses towards parasitic eggs when parasitized repeatedly within one breeding attempt.     

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.     

Migratory Orientation of Blackcaps (Sylvia atricapilla): Population-specific Shifts of Direction during the Autumn

The population-specific orientation of two groups of blackcaps (Sylvia atricapilla), one from southwest Germany, the other from easternmost Austria, was studied outdoors in Emlen funnels. We investigated whether a seasonal shift in the migratory direction — as expected for the Austrian birds from ringing recoveries — occurs under experimental conditions and in a seasonally constant magnetic field. The West German birds, for which no shift was expected, oriented southwest during the entire season. The Austrian birds oriented southeast in October and southsouthwest in November. The clockwise shift by about 60° occurred within a 10-day period. The results indicate that in this species seasonal changes of migratory direction are probably based on an endogenous program, occur without the birds actually migrating and are independent of changes in the magnetic field. Our results provide further evidence that directional shifts in Sylvia warblers may be controlled by a different mechanism than in pied flycatchers (Ficedula hypoleuca).     

Lamellar orientation in human cornea in relation to mechanical properties

We have applied wide-angle X-ray scattering to the human cornea in order to quantify the relative number of stromal collagen fibrils directed along the two preferred corneal lamellar directions: superior-inferior and nasal-temporal. The data suggest that, on average, the two directions are populated in equal proportion at the corneal centre. Here, approximately one-third of the fibrils throughout the stromal depth tend to lie within a 45 degrees sector of the superior-inferior meridian, and similarly for the nasal-temporal direction. However, in some eyes we have measured significant differences between the two preferential fibril populations, with some corneas exhibiting as much as 25% more collagen in one direction than the other. Based on these findings, a mechanical model of the normal cornea may be envisaged, whereby the fibril tension in the underlying "background" of isotropically arranged collagen helps to balance the intraocular pressure; while the extra preferentially aligned fibrils take up the additional tensile stress along the superior-inferior and nasal-temporal meridians exerted by the rectus muscles and the orbicularis. It is possible that, through a direct impact on the elastic modulus of the tissue, an imbalance of superior-inferior and nasal-temporal fibrils in some eyes might affect corneal shape.     

Contrasting Patterns of Genetic Differentiation among Blackcaps (Sylvia atricapilla) with Divergent Migratory Orientations in Europe

Migratory divides are thought to facilitate behavioral, ecological, and genetic divergence among populations with different migratory routes. However, it is currently contentious how much genetic divergence is needed to maintain distinct migratory behavior across migratory divides. Here we investigate patterns of neutral genetic differentiation among Blackcap (Sylvia atricapilla) populations with different migratory strategies across Europe. We compare the level of genetic divergence of populations migrating to southwestern (SW) or southeastern (SE) wintering areas with birds wintering in the British Isles following a recently established northwesterly (NW) migration route. The migratory divide between SW and SE wintering areas can be interpreted as a result of a re-colonization process after the last glaciation. Thus we predicted greater levels of genetic differentiation among the SW/SE populations. However, a lack of genetic differentiation was found between SW and SE populations, suggesting that interbreeding likely occurs among Blackcaps with different migratory orientations across a large area; therefore the SW/SE migratory divide can be seen as diffuse, broad band and is, at best, a weak isolating barrier. Conversely, weak, albeit significant genetic differentiation was evident between NW and SW migrants breeding sympatrically in southern Germany, suggesting a stronger isolating mechanism may be acting in this population. Populations located within/near the SW/SE contact zone were the least genetically divergent from NW migrants, confirming NW migrants likely originated from within the contact zone. Significant isolation-by-distance was found among eastern Blackcap populations (i.e. SE migrants), but not among western populations (i.e. NW and SW migrants), revealing different patterns of genetic divergence among Blackcap populations in Europe. We discuss possible explanations for the genetic structure of European Blackcaps and how gene flow influences the persistence of divergent migratory behaviors.