Is the later arrival of young male red-breasted flycatchers (Ficedula parva) related to their physical condition?

Intraspecific variation in the arrival time of migratory birds to breeding grounds is common. Although this phenomenon has been explained in various ways, the condition-dependency of arrival is often evoked. I analyzed the arrival time of male red-breasted flycatchers Ficedula parva, a long-distance migratory passerine, in relation to age, body size and body condition. Data were collected over the course of 5 years in the primeval Białowieża Forest of northeast Poland. Each of these years, young males arrived at the breeding grounds significantly later in the year than did older males. Young males also exhibited significantly shorter wings and lower body condition than older males. Settlement speed was significantly related to age, but not to wing length or body condition. Only the arrival time of old males was related to the body mass and condition. Later arrival of young males could be explained by the lack of experience or by avoidance of both aggression and competition from older males, and both explanations are thought to conserve energy for breeding. Young male red-breasted flycatchers exhibit delayed plumage maturation, and this duller plumage supports the strategy of restraint in the arrival time of young males in this species.     

Advancement of spring arrival in a long-term study of a passerine bird: sex,age and environmental effects

In migratory birds, mistimed arrival might have negative consequences for individual fitness, causing population declines. This may happen if arrival time is not synchronized with breeding time, especially when earlier springs favour earlier reproduction. We studied spring arrival time to the breeding areas in a pied flycatcher Ficedula hypoleuca population in southern Norway during a 30-year period (1985–2014). We investigated trends in arrival both for the entire population and for different population fractions (e.g. early vs. late arrivals). We also studied sex and age class differences, along with repeatability of arrival. Finally, we explored how arrival is influenced by environmental conditions at the areas birds use throughout the year, using mixed-effects models and quantile regressions with individual-based data. Spring arrival advanced over five days, at a similar rate through the entire population. Males and adult birds arrived earlier than females and yearlings. Arrival was significantly repeatable for males and females. Birds arrived earlier in years with high temperature and rainfall at the breeding grounds, and low NDVI both on the Iberian Peninsula and in central Europe. Later fractions of the population showed a steeper response to these environmental variables. This intra-population heterogeneity in the responses to the environment probably stems from a combination between the different selection pressures individuals are subject to and their age-related experience. Our results highlight the importance of studying how migration phenology is affected by the environment not only on the breeding grounds but also on the other areas birds use throughout the year.     

Blood parasites and male fitness in the pied flycatcher

In vertebrates the effect of parasites on host ecology has almost been ignored. Recently the view that well-adapted parasites do not harm their hosts has been challenged and there is growing evidence that parasites do have a present-day effect on a great variety of host fitness components. The pied flycatcher is a small migratory passcrine bird. Any decrease in condition caused by disease should affect its ability to cope with physical demands of migration. Here we examine whether blood parasites have any effect on male arrival time. Males infected with Trypanosoma arrived on average 2 days later than males with no Trypanosoma infection. Infected males also had shorted tails and tended to have shorter wings. By contrast, there was no difference in male arrival time between males infected with Haemoproteus and healthy males. It seems that Trypanosoma infection lowered male condition and consequently the ability to moult and migrate. The difference in length of feathers may have generated the difference in arrival times. Early arrival is highly important for males, since only the first males become polygynous and breeding prospects deteriorate rapidly with any delay in egg laying. Estimated reduction in breeding success for infected males was about 20%.     

Relationship between arrival timing and breeding success of intra‐tropical migratory Fork‐tailed Flycatchers (Tyrannus savana)

For migratory birds, early arrival at breeding areas has many benefits, such as acquisition of better territories and mates. This strategy has been found in numerous species breeding at north‐temperate latitudes, but has not been yet reported for intra‐tropical migratory species. We evaluated the relationship between arrival date, initiation of breeding, and breeding success of Fork‐tailed Flycatchers (Tyrannus savana) breeding in southeastern Brazil and overwintering in northern South America. We color‐banded adult flycatchers during three breeding seasons and searched for them during the following breeding seasons. We also monitored nests from construction until either failure or fledging of young. We found that: (1) male Fork‐tailed Flycatchers arrived at the breeding site earlier than females, (2) males that arrived earlier had greater breeding success, and (3) nests where eggs were laid earlier in the breeding season were more likely to be successful than those where eggs were laid later. Male Fork‐tailed Flycatchers appeared to benefit from early arrival at a tropical breeding site, potentially mediated by their ability to acquire a high‐quality territory and mate as early as possible, and by the ability of their mate to begin breeding as early as possible. Breeding success for female Fork‐tailed Flycatchers may be determined primarily by a combination of the arrival date of their mate and how quickly they can begin breeding. Our results suggest that protandry occurs in an intra‐tropical migratory bird and that early arrival of males and early initiation of reproduction by females results in greater reproductive success. A better understanding of the underlying mechanisms that control the timing of migration and reproduction of this and other intra‐tropical migratory species is important for evaluating the challenges they face in light of current and future rapid environmental changes.     

Is Divorce in Young Common Terns,Sterna hirundo,after Recruitment just a Question of Timing?

The probability of divorce in birds has been linked with age, breeding experience, reproductive output and synchrony in return. Here, we investigate the consequences of first breeding attempts in common terns for mating in the subsequent season. Nearly 20% of all first‐time breeders disappeared or skipped at least one season after recruitment. In 84 pairs, which consisted of at least one recruit and of which both partners returned to the colony, the divorce rate was 45%. We compared reproductive success, arrival dates, and asynchrony in arrival dates of pairs of the first breeding season against the second season, for both reunited and divorced pairs and males and females separately. First, in pairs of which both members came back to the colony, we found an increase of reproductive success most pronounced in males. In the second season reproductive success of divorced compared with reunited pairs was higher, as only divorced pairs significantly improved the number of fledglings, and again this relation was stronger in males. Secondly, females of reunited pairs arrived significantly earlier from the first to the second season and by far more days than their males. However, in divorced pairs former mates did not differ in the number of days they advance their arrivals. Finally, divorced males arrived on average 4 d earlier than their former mates, whereas divorced females arrived 5 d later compared with their former mates of the recruitment season. Contradictory to nearly all other divorce studies in birds so far, we found a clear fitness gain in divorced males. We suggest that the improvement in reproductive success of young males stems from a side‐effect of the birds’ quality and ability to reach the breeding site in appropriate time and earlier as potential competitors. In long‐lived bird species the heterogeneity among young individuals in the timing of arrival at the colony seems to explain why former recruit‐pairs reunite or split. For young males we suggest as best explanation of divorce that they profit from ‘pushing for an empty chair’, while females seem to profit from their choosiness and may actively decide between former and other mates.     

Fat stores of American redstarts Setophaga ruticilla arriving at northerly breeding grounds

Many long-distance passerine migrants arrive with more fat stores than necessary to have reached their northerly breeding grounds. Researchers have argued for adaptive advantages associated with arriving with 'surplus' migratory fat, including increased breeding performance and insurance against adverse weather, and reduced food resources during the days following arrival. The objectives of this work were to document fat stores in American redstarts Setophaga ruticilla arriving to breed in northern Michigan and to test predictions associated with the hypothesis that arrival fat serves an insurance role. Results suggest that redstarts arrived in northern Michigan with fat stores sufficient to have continued migrating an additional distance greater than 1000 km. Significant yearly variation in arrival fat corresponded in part with environmental conditions measured at the breeding grounds. Birds arrived with the most fat in the year with the coolest temperatures and the lowest food abundance at the time of arrival to the site. Further, an inverse relationship between arrival fat and arrival day in males indicated that early arrivals carried more fat than later arrivals. Birds that arrived early faced cooler temperatures and lower densities of terrestrial invertebrates, and arrival fat may have provided a mechanism to overcome poor early season foraging conditions. However, our results are not entirely consistent with the hypothesis that arrival fat serves only as insurance. Arrival fat appeared important even during the most benign year of this study. Further, evidence suggests that females may have derived more benefit from arrival fat than males. These results highlight the connection between phases of a bird's annual cycle. Migrants that do well en route , arrive at breeding grounds in better condition, which may contribute to survival and reproductive success.     

EVOLUTION OF PLUMAGE COLOR IN MALE PIED FLYCATCHERS (FICEDULA HYPOLEUCA): EVIDENCE FOR FEMALE MIMICRY

We present the first evidence for sexual deception by female mimicry in birds. Using live, caged birds we show that territorial male pied flycatchers behave aggressively toward bright-colored males but display sexually toward female-like male intruders. We also show that the males that are fooled are those that lack recent sexual experience. All male pied flycatchers are dull-colored in winter. It is possible that young males are more constrained during the spring molt than older males since the former are more dull-colored in spring. According to the molt-constraints hypothesis a subadult plumage would be maladaptive in the breeding season. Analysis of male settling pattern at breeding sites in spring suggests that brownish males are allowed to settle closer to already-established males than dark-colored males. This result suggests an adaptive value of having a subadult plumage color, in particular for young males arriving late from spring migration. However, we also show that mimicry incurs a cost, that of increased aggression from females, which may explain why female-like males have reduced mating success.     

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.     

Lifetime patterns of pairing success in male Ortolan Buntings Emberiza hortulana

Analyses of lifetime fitness in birds are typically based on estimates of breeding success, in particular the number of offspring fledged. Small and isolated bird populations often have a male‐skewed adult sex ratio, so that male lifetime productivity depends to a large degree on pairing success, but few studies have focused on patterns of lifetime pairing success. The Norwegian population of Ortolan Buntings Emberiza hortulana is strongly male‐skewed, such that in any year about half of all males are unpaired. Pairing success of first‐year males (16–44%) was significantly lower than for older males (52–89%). Lifetime pairing success was correlated with lifespan and was strongly skewed, with a majority of males being paired only once or never, and only 11% paired three or more times despite a stable lifetime annual survival rate of 63%. Males that were paired in one year were more likely to be paired the next year than males that were unpaired in the previous year. The shortage of females caused even the older males to have a substantial probability of becoming unpaired, and 49% of long‐lived males (known as adults for at least 4 years) were unpaired after years in which they were paired. Pairing success in the Ortolan Bunting therefore follows similar age‐related and lifetime patterns in breeding success documented in other species. However, even the older males ran a high risk of not being paired, contrasting with earlier distinctions between pre‐breeding and breeding lifespans. I discuss the importance of knowledge of pairing success for the management of endangered and declining populations.     

Age at pollination modifies relative male and female reproductive success in a monoecious fig tree

Plants that depend on a single species of insect pollinator must often contend with infrequent and unpredictable visitation. Prolongation of floral receptivity comes at the cost of reduced male and/or female reproductive success among older flowers. Fig trees (Ficus spp.) have a highly specific pollination symbiosis and individual inflorescences (syconia) that remain receptive for days or weeks. Reproductive success in monoecious fig trees involves production of both seeds and fig wasp offspring. We assessed whether the reproductive output of individual syconia changes with the length of time they waited for pollination, and whether the relative female and male reproductive success also changes. A pollination experiment was conducted in an SE Asian monoecious fig tree Ficus curtipes, in which receptive syconia were covered with mesh bags to exclude wasps and pollinated by single pollinators of this fig tree at their different receptive ages. When the syconia matured their size and contents were recorded. Seed quality was also assessed. The results showed that pollinators entered syconia that had been waiting for up to 36?days. The frequencies of abortions among syconia pollinated at different ages were low throughout. The number of un-utilised flowers increased progressively in older syconia. Seed production was highest in syconia entered on the first day of receptivity, whereas pollinator production peaked in syconia pollinated on day 12, then declined in older syconia. Consequently, overall reproductive efficiency declined with syconium age and floral sex allocation became more male-biased in older syconia. Older syconia also produced lighter seeds. These results suggest that un-pollinated syconia of F. curtipes can remain receptive for several weeks. This makes pollination of each syconium more likely, but at the cost of reduced productivity and with more ovules allocated to male function. However, the prolongation of floral receptivity has significance for the co-adaptation between syconia and fig wasps and for the evolution of the fig tree-fig wasp symbiosis.     

Plumage colour and sexual selection in the pied flycatcher Ficedula hypoleuca

The plumage colour of male pied flycatchers varies on a continuum from brownish and female-like to a conspicuous black-and-white. Males have a conspicuous plumage only during the breeding season, when the mortality rate was low and not correlated with plumage colour. Brownish males breed successfully, and breeding males had no lower annual rate of return than non-breeders. Experiments on male intrusion showed that possession of a cryptic, female-like plumage did not facilitate nest site settlement by such males. Dark males were on average older, arrived earlier, and defended more nestboxes than brown males. However, experiments showed that plumage colour was not a reliable indicator of the outcome of competitive combats between males. These facts argue against several existing hypotheses about variation in male plumage colour. A new explanation is proposed: a bright colour serves to signal presence and thereby reduces territory holding costs. The brownish males arrive only shortly before the females and thus will not benefit as much by being conspicuous as do the black-and-white males that arrive earlier in spring.     

Variation with age in the timing and duration of spawning in American plaice

Spawning time of female American plaice Hippoglossoides platessoides varied significantly with age. This effect, however, was not consistent across years. Generally, younger fish spawned earlier than older fish, but in the 1990s they spawned later than older fish. Spawning duration also varied with age, with younger fish generally having a longer spawning period than older fish. Changes in depth distribution were related to changes in both spawning time and duration. As depth increased, spawning time became later and duration became shorter. Fish <11 years old showed the largest change in both spawning time and depth distribution.     

Age-specific patterns of infection with haemosporidians and trypanosomes in a warbler: implications for sexual selection

Although the selective loss of individuals susceptible to disease can favor the evolution of female preference for older males, the interrelationship between age, infection, longevity, and mating success remains poorly characterized in natural populations. In a longitudinal study of 61 male common yellowthroats (Geothlypis trichas), we found that the probability of infection with hematozoa increased as males aged from 1 to 5 years. Despite a significant, negative association between infection and longevity that partially masked age-effects, the odds that a male was infected with Trypanosoma, Plasmodium, or Leucocytozoon increased 71–212% per year. Nearly 75% of males in their first breeding season were either uninfected or infected with only a single parasite, while 50% of older males were infected with at least two parasites and 16% were infected with all three. No males escaped infection after their second breeding season. Older males were also more likely to sire extra-pair young (EPY) and, as a consequence, infection with multiple parasites was associated with a fourfold increase in the odds of producing EPY. Unlike younger males, 80% of the oldest males had a history of either surviving chronic infection or recovering. Combined with previous work showing higher diversity at the major histocompatibility complex among older males, our results suggest that the song and plumage traits that signal male age in common yellowthroats also, perforce, signal resistance to parasites. By preferring older males, females may obtain good genes for disease resistance even in the absence of any effect of infection on male ornamentation.     

Different reproductive tactics in male collared flycatchers signalled by size of secondary sexual character

Most studies of variation in male reproductive tactics have focused on conspicuous categorical differences in mating behaviour (i.e. variation in mating strategies). However, in the presence of trade-offs between investment in competition over matings, parental care and survival, a male''s optimal allocation rule might vary according to his physiological condition and social or ecological environment. Thus, there may also be more subtle variation in male reproductive tactics. Here, I show that the reproductive effort (estimated as residual change in condition) of male collared flycatchers was affected by the size of their forehead patch (a secondary sexual character), age and date of arrival at the breeding grounds. Among early males (i.e. males with a high likelihood of both attracting more than one female and obtaining extra-pair copulations), large-patched males made a relatively large reproductive effort and as a result were in worse condition at the time of feeding offspring as compared to small-patched males. Furthermore, among early breeders, young males and males with experimentally increased forehead patch size made a relatively high effort. By contrast, regardless of age and badge size, there were no such patterns observed among late breeders. These results suggest that collared flycatchers use different reproductive tactics depending on both internal and external factors, and that the size of a secondary sexual trait may not only indicate variation in individual condition but also predict how resources will be allocated between pre- and post-mating reproductive activities.     

Spatial patterns of white stork (Ciconia ciconia) migratory phenology in the Iberian Peninsula

In contrast to the attention attracted by temporal trends of phenology, the spatial patterns of arrivals, departures or stays of trans-Saharan birds are still nowadays largely unknown in most of their European breeding areas. In the case of the white stork (Ciconia ciconia), some studies have attempted to describe its migratory patterns throughout some European countries but, to our knowledge, no one has related these patterns to some kind of explanatory variable which offers an ecologically-based explanation for the heterogeneous phenology observable among populations. Here, arrivals, departures and stays of this species, recorded in hundreds of Spanish localities, were related to a set of environmental, geographical, biological and spatial predictors, and modeled by multiple regression. The best model for arrival dates accounted for up to 34% of variability of data and pointed towards an earlier arrival in those populations located in south-western Iberia and with higher population densities. This last relationship is probably due to the competition for nest-site fidelity maintenance. However, no variable was able to explain properly the blurred spatial pattern recorded for departure dates. Departure decisions are strongly influenced by social behavior in this species and depend on collective decisions influenced by peculiar local environmental conditions of each year rather than macrogeographical gradients. Environmental, geographical or spatial variables also did not capture much of the observed variability in the length of the stays among populations. However, this variable was strongly related to the arrival and departure dates of populations. White storks stay longer in localities with earlier arrivals and, especially, later departures.     

Annual variation in the timing of reproduction in Antarctic fur seals, Arctocephalus gazella, at Bird Island, South Georgia

The arrival of Antarctic fur seals at a breeding beach on Bird Island, South Georgia, was studied over five consecutive breeding seasons, 1983 to 1987. Experienced bulls arrived first and established breeding territories on the beaches in anticipation of the arrival of the cows. Male arrival, which is less synchronous within years than female arrival, was significantly later in 1987 than in any other year. Female arrival, estimated by pup birth date where necessary, was highly synchronous; it usually started when 80% or more of potential territory sites were occupied by males. Cows arrived significantly later in 1984 and 1987 than in 1983, 1985 or 1986. The late arrival of both males and females in 1987 is attributed to unusually severe climatic conditions during the preceding winter. The late arrival and reduced fecundity of females in 1984 is attributed to markedly reduced food availability during the austral winter and summer of 1983. Males were not affected in 1984 because they could move away from the area of reduced food availability earlier than females and because they have a more varied diet. Factors influencing the winter distribution, the timing and pattern of arrival and the breeding of male and female Antarctic fur seals are discussed.     

A comparison of the annual cycles in testicular size and moult in captive European starlings Sturnus vulgaris during their first and second years

There is considerable evidence that reproductive success improves with age in birds. It is often suggested that improved performance of older birds is because they are more experienced. Less consideration has been given to the possibility that improvements may be a consequence of age-related changes in reproductive physiology. One factor that consistently changes with age is laying date – first year birds lay later than older birds. In European starlings Sturnus vulgaris , older males begin reproductive activity earlier than first year males and dominate available nest sites. I monitored changes in testicular volume in captive starlings exposed to natural changes in day length and temperature, from their first autumn through the next two annual cycles. Testicular maturation was advanced by 3–4 weeks in birds during their second year compared to the first, and testicular regression occurred about 2 weeks later. The period of full sexual maturation was 50% longer during the second year. The timing of the post-nuptial moult was the same. A possible physiological mechanism to explain this is discussed. The results show that earlier reproductive activity in older birds can be explained, at least in part, by intrinsic physiological mechanisms. This does not preclude additional effects of experience of an individual in the improvement in reproductive performance between first and second year birds.     

Positive feedback between ecological and reproductive character displacement in a young avian hybrid zone

Character displacement can reduce costly interspecific interactions between young species. We investigated the mechanisms behind divergence in three key traits-breeding habitat choice, timing of breeding, and plumage coloration-in Ficedula flycatchers. We found that male pied flycatchers became expelled from the preferred deciduous habitat into mixed forest as the superior competitor, collared flycatchers, increased in numbers. The peak in food abundance differs between habitats, and the spatial segregation was paralleled by an increased divergence in timing of breeding between the two species. Male pied flycatchers vary from brown to black with brown coloration being more frequent in sympatry with collared flycatchers, a pattern often proposed to result from selection against hybridization, that is, reinforcement. In contrast to this view, we show that brown male pied flycatchers more often hybridize than black males. Male pied flycatcher plumage coloration influenced the territory obtained in areas of co-occurrence with collared flycatchers, and brown male pied flycatchers experienced higher relative fitness than black males when faced with heterospecific competition. We suggest that allopatric divergence in resource defense ability causes a feedback loop at secondary contact where male pied flycatchers with the most divergent strategy compared to collared flycatchers are favored by selection.     

Factors influencing plasticity in the arrival‐breeding interval in a migratory species reacting to climate change

Climate change is profoundly affecting the phenology of many species. In migratory birds, there is evidence for advances in their arrival time at the breeding ground and their timing of breeding, yet empirical studies examining the interdependence between arrival and breeding time are lacking. Hence, evidence is scarce regarding how breeding time may be adjusted via the arrival‐breeding interval to help local populations adapt to local conditions or climate change. We used long‐term data from an intensively monitored population of the northern wheatear (Oenanthe oenanthe) to examine the factors related to the length of 734 separate arrival‐to‐breeding events from 549 individual females. From 1993 to 2017, the mean arrival and egg‐laying dates advanced by approximately the same amount (~5–6 days), with considerable between‐individual variation in the arrival‐breeding interval. The arrival‐breeding interval was shorter for: (a) individuals that arrived later in the season compared to early‐arriving birds, (b) for experienced females compared to first‐year breeders, (c) as spring progressed, and (d) in later years compared to earlier ones. The influence of these factors was much larger for birds arriving earlier in the season compared to later arriving birds, with most effects on variation in the arrival‐breeding interval being absent in late‐arriving birds. Thus, in this population it appears that the timing of breeding is not constrained by arrival for early‐ to midarriving birds, but instead is dependent on local conditions after arrival. For late‐arriving birds, however, the timing of breeding appears to be influenced by arrival constraints. Hence, impacts of climate change on arrival dates and local conditions are expected to vary for different parts of the population, with potential negative impacts associated with these factors likely to differ for early‐ versus late‐arriving birds.     

The effects of climate change on the phenology of winter birds in Yokohama, Japan

Observations made largely from summer breeding sites in Europe and North America have been used to document the effects of climate change on many bird species. We extend these studies by examining 23 years of observations between 1986 and 2008 of six winter bird species made by citizens at a city park in Yokohama, Japan. Bird species arrive in autumn and spend the winter in the area, before departing in the late winter or spring. On average, birds species are arriving 9 days later than in the past and are departing on average 21 days earlier, meaning that the average duration of their stay in Yokohama is about 1 month shorter now than in the past. Patterns of changes over time varied among species, but departure dates changed for more species than did arrival dates. Dates of departure and arrival were sometimes correlated with monthly average temperatures—later arrivals and earlier departures were associated with warmer temperatures. In addition, interannual variation in arrival and departure dates were strongly correlated across species, suggesting that species were responding to the same or similar environmental cues. This study provides a clear demonstration of the value of using citizens to make observations that contribute to research in climate change biology.