Age-related patterns of reproductive success in house sparrows Passer domesticus

A common life history pattern in many organisms is that reproductive success increases with age. We report a similar pattern in house sparrows Passer domesticus , older individuals performed better than yearlings for most measures of reproductive success. Older males and females began breeding earlier in a given season and fledged more young than their yearling counterparts. Individual males also fledged more young in their second breeding season than they did in their first, but individual females did not show consistent improvement in reproductive success from year one to two. A path analysis indicated that age in both sexes acted primarily through the timing of breeding; earlier nesters laid more eggs and hence fledged more young but did not have more nesting attempts. We tested whether the increased reproductive success with age arose from high quality individuals surviving to be older (selection hypothesis). In contrast to the main prediction of this hypothesis that reproductive success and survival should be positively related, we found that survival from one year of age to two years of age was negatively related to reproductive success in the first year for males and females combined. Additionally, individuals that survived to breed as two-year-olds did not differ in total young fledged in their first year from those that did not survive to their second season of breeding. Our results indicate that fledgling production increases with age due to improvements in timing of breeding, particularly in females, and not because of the loss of poor breeders or increased output. Mechanisms producing age-related differences in timing of breeding warrant further study.     

Starting over: experimental effects of breeding delay on reproductive success in early‐arriving male American redstarts

Birds that arrive and breed early often have higher reproductive success than late individuals, either as a consequence of timing‐specific advantages (the timing hypothesis) or because these individuals and/or their resources are of higher quality (the quality hypothesis). In this study, we examined the potential influence of several factors affecting reproductive success by experimentally delaying breeding of early‐arriving male American redstarts Setophaga ruticilla, a species for which early male arrival is strongly related to increased reproductive success. Our manipulation involved the capture, holding, and release of males following pairing and territory establishment, resulting in the majority of subjects (67%) losing their initial mate (47%) or mate and territory (20%) and forcing them to start over approximately 12 d after their initial arrival. Males forced to start over (i.e. those losing their first territory and/or mate) did not experience any decrease in body condition, nor did their reproductive behaviour differ from that of early‐arriving control males. We found that naturally early‐arriving but experimentally manipulated males suffered reduced fledging success in comparison to early‐arriving males that bred early or late, but equivalent success in comparison to males that arrived and bred late. Based on our results, we propose that the relationship between early arrival and higher reproductive success in this species is mediated not simply by individual male quality or absolute arrival timing alone, but rather some other aspect of resource quality is likely important. We discuss and present evidence for two alternative explanations under the quality hypothesis: female quality and territory quality. To our knowledge, our study is the first to investigate the effects of experimentally delaying male breeding time, strengthening previous correlational evidence for resource quality as a potentially important selective agent driving early arrival in migratory birds.     

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.     

Sex-differentiated migration patterns, protandry and phenology in North European songbird populations

This study aims to investigate causes and mechanisms controlling protandrous migration patterns (the earlier breeding area arrival of males relative to females) and inter-sexual differences in timing of migration in relation to the recent climate-driven changes in phenology. Using standardised ringing data from a single site for eight North European migratory passerines collected throughout 22 years, we analysed sex-differentiated migration patterns, protandry and phenology of the entire populations. Our results show protandrous patterns for the first as well as later arriving individuals for all studied species. Males show more synchronous migration patterns compared to females and, hence, first arriving females followed males more closely than later arriving individuals. However, we found no inter-sexual differences in arrival trends as both sexes advance spring arrival over time with the largest change for the first arriving individuals. These findings seem in support of the “mate opportunity” hypothesis, as the arrival of males and females is strongly coupled and both sexes seem to compete for early arrival. Changes in timing of arrival in males and females as a response to climatic changes may influence subsequent mating decisions, with subsequent feedbacks on population dynamics such as reproductive success and individual fitness. However, during decades of consistent earlier spring arrival in all phases of migration we found no evidence of inter-sexual phenological differences.     

Protandrous migration and variation in morphological characters in Emberiza buntings at an East Asian stopover site

The effect of the timing of spring migration on reproductive success differs between the sexes. As a consequence, various sex‐specific tactics relating to the timing of migration have evolved in migratory avian groups. Various hypotheses have been proposed to explain differential migration to breeding or wintering grounds, and inter‐ and intrasexual size differences are often considered one of the proximate mechanisms. We investigated arrival patterns in the spring by individuals of each sex, sexual size dimorphism and related morphological variables, and the relationship between size variation and arrival date in five bunting species that passed through an East Asian migratory flyway stopover site in 2006–08. Males of all the study species arrived before females, and significant sexual dimorphism was observed. Several morphological characters, including total length, wing‐length and tail‐length, contributed to the size variation. Although larger males arrived earlier, there was no relationship between arrival date and size in females. Our study confirmed that East Asian buntings display a discriminated protandrous migration pattern at the stopover site as well as at the breeding grounds. This is consistent with the view that larger body size in males is favoured due to its association with early arrival to help ensure access to the best resources and hence enhanced mating success.     

Does the expression of a male plumage trait in female Kentish plovers (Charadrius alexandrinus) signal individual quality?

A small percentage (4.5%) of females Kentish plovers (Charadrius alexandrinus) observed breeding in southern Spain expressed a plumage trait characteristic of males: the forecrown (frontal) black bar. The expression of this male trait may be age-related, and reversible, i.e., within individuals it may be present in some years but not in other years. Females with frontal bars were not in better condition and nor did they initiate breeding earlier than in years when they lacked frontal bars. Nevertheless, when females had frontal bars they laid larger eggs. I suggest that the frontal bar on females could be an epiphenomenon resulting from hormonal imbalances of females as they age.     

Antioxidants and condition-dependence of arrival date in a migratory passerine

Early arrival at the breeding area after spring migration determines several fitness benefits to migratory birds, including an early start of reproduction and a greater seasonal reproductive success. Environmental conditions determine the onset of migration at the population level, while physiological costs associated with migration and imposed on individuals determine the level of asynchrony in the pattern of arrival. These costs include several physiological changes, suppression of immune response, and an increased risk of oxidation by production of free radicals due to energetically expensive activities. According to the condition-dependent hypothesis of arrival date, only healthy individuals in prime condition can afford to arrive and thus reproduce early in the season.
We investigated condition-dependence of arrival date in male barn swallow ( Hirundo rustica L.), a migratory passerine, in two years that differed in environmental condition, and the role of carotenoids in mediating arrival date. We show that male tail length, the main secondary sexual character, body condition, and hematocrit were negatively correlated with arrival date in the first year, indicating better quality of early arriving males. In the second year, better environmental conditions advanced arrival of the entire population by weeks. In this season we could show no relationship between arrival and tail length or body condition. Moreover, a lower value of sedimentation rate and a brighter colour of the red throat feathers indicated better health status of the population. Arrival date is a condition-dependent character in barn swallows, since we found high repeatability of arrival date, lower values of carotenoids in blood and a lower depletion of carotenoids in blood shortly after arrival in early arriving males. This suggests that early arriving males either use fewer antioxidants for free radical scavenging, or they have differential access to antioxidants.     

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.     

Age-related change in breeding performance in early life is associated with an increase in competence in the migratory barn swallow Hirundo rustica

1. We investigated age-related changes in two reproductive traits (laying date and annual fecundity) in barn swallows Hirundo rustica L. using a mixed model approach to di-stinguish among between- and within-individual changes in breeding performance with age. 2. We tested predictions of age-related improvements of competence (i.e. constraint hypothesis) and age-related progressive disappearance of poor-quality breeders (i.e. selection hypothesis) to explain age-related increase in breeding performance in early life. 3. Reproductive success increased in early life, reaching a plateau at middle age (e.g. at 3 years of age) and decreasing at older age (> 4 years). Age-related changes in breeding success were due mainly to an effect of female age. 4. Age of both female and male affected timing of reproduction. Final linear mixed effect models (LME) for laying date included main and quadratic terms for female and male age, suggesting a deterioration in reproductive performance at older age for both males and females. 5. We found evidence supporting the constraints hypothesis that increases in competence within individuals, with ageing being the most probable cause of the observed increase in breeding performance with age in early life. Two mechanisms were implicated: (1) advance in male arrival date with age provided middle-aged males with better access to mates. Yearling males arrived later to the breeding grounds and therefore had limited access to high-quality mates. (2) Breeding pairs maintaining bonds for 2 consecutive years (experienced pairs) had higher fecundity than newly formed inexperienced breeding pairs. 6. There was no support for the selection hypothesis because breeding performance was not correlated with life span. 7. We found a within-individual deterioration in breeding and migratory performance (arrival date) in the oldest age-classes consistent with senescence in these reproductive and migratory traits.     

Patterns of territory settlement and consequences for breeding success in the Northern Wheatear Oenanthe oenanthe

We examined the pattern of territory settlement and its consequences for breeding success in the Northern Wheatear Oenanthe oenanthe on Bardsey Island, Wales, during the breeding seasons of 1991-93. Males returned earlier than females, and older males returned earlier than first-year breeders. Although their boundaries shifted between years, the general location of territories was consistent during the three-year study. There was a high degree of fidelity to area and territory between years for both sexes. The order of territory settlement, from which a territory rank was calculated, was highly consistent for males between years irrespective of individual settlement patterns and territory fidelity. Patterns of territory settlement were less predictable for females, although there was a significant correlation between the mean territory ranks of paired males and females. There was a male-biased sex ratio in each year, and between 5% and 26% of males remained unpaired throughout the breeding season. Male mating status and breeding success were dependent on arrival date, territory rank and breeding density. Early-arriving, usually older, males were able to settle on territories first and were more likely to pair, while later-arriving individuals were more likely to remain unmated. These effects were consistent between years, and consequently territories could be classified as either preferred (accounting for proportionately more breeding attempts) or non-preferred. Territory quality as opposed to individual quality appeared to explain much of the variation in breeding success, and both sexes benefited by breeding on preferred territories through enhanced breeding success and an increased probability that their offspring would be recruited to the population.     

A ptilochronological study of carry‐over effects of conditions during wintering on breeding performance in the barn swallow Hirundo rustica

The ecological conditions that a bird experiences during any stage of its life cycle may have consequences that become manifested at later life stages, and these ‘carry‐over effects’ may be major components of variance in individual performance. Condition‐dependent feather growth rate, as assessed by growth bars width (GBW), provides a unique, though largely under‐exploited tool to investigate carry‐over effects of ecological conditions and individual physiological state during molt. In this study of breeding barn swallows Hirundo rustica, which undergo a single complete annual molt of tail and wing feathers during wintering in sub‐Saharan Africa, we first show that old (≥ 2 yr) females have larger GBW than old males and yearlings. GBW was smaller with larger infestations by common ectoparasites of barn swallows, independent of the swallows’ age or sex, and larger GBW was associated with a higher index of body condition during the breeding season in males. Larger GBW predicted higher seasonal reproductive output of older males, but not reproductive output of younger males or females of any age class, with this higher reproductive output of older males mediated by higher offspring fledging success. However, no relationship with GBW was observed for seasonal reproductive output of males in the spring preceding the winter when the feathers were grown. Hence, this study suggests that the analysis of the rate of feathers growth (‘ptilochronology’) has a larger potential to serve as a powerful tool in the study of carry‐over effects than has been appreciated to date. Specifically, the present results support the idea that conditions experienced during wintering in Africa and proximately reflected by GBW have carry‐over effects on body condition and breeding success. These effects are sex and age specific, being more pronounced in older males, possibly as a consequence of differences in annual time routines and susceptibility to extrinsic factors among sex and age classes.     

Arrival date, age and breeding success in white stork Ciconia ciconia

Early arrival to breeding grounds is a life history trait in birds that can result in fitness benefits. We studied the relationship between arrival date and breeding success of individuals in a central Iberian population of white stork Ciconia ciconia , between 1999 and 2005, and the ways in which other potential factors, such as age or sex, affect this relationship. Our results showed that age was the factor most closely related to arrival date and breeding success. Older individuals returned earlier to the breeding grounds, achieved larger clutch sizes and produced more chicks than younger birds. After controlling statistically for age effect, breeding probability (laid eggs or not) and laying date were still significantly explained by arrival date. A higher probability of failure to reproduce (no eggs laid) was found in birds arriving later than in those arriving early. However, clutch size and nestling success (number of nestlings in the nest 40 days after hatching) were not correlated with arrival date. Food availability in the study area throughout the breeding cycle, due to a nearby rubbish dump, could be the factor mitigating differences in clutch size and nestling success related to individual arrival date.     

Arrival times,laying dates,and reproductive success of Snowy Plovers in two habitats in coastal northern California

ABSTRACT Habitat quality, as indexed by the reproductive success of individuals, can greatly influence population growth, especially for rare species near the limits of their range. Along the Pacific coast, the Snowy Plover (Charadrius alexandrinus nivosus) is a threatened species that, in recent years, has been breeding on both riverine gravel bars and ocean beaches in northern California. From 2001 to 2009, we compared the habitat characteristics, breeding phenology, reproductive success, and abundance of Western Snowy Plovers occupying these two habitats. Similar percentages of yearling and adult plovers returned to gravel bars and beaches, but plovers breeding on gravel bars arrived and initiated first clutches 2–3 weeks later than those breeding on beaches. Despite this delay, however, the mean annual fledging success of plovers on gravel bars (1.4 ± 0.4 [SD]) was double that on beaches (0.7 ± 0.3). Differences in cumulative reproductive success produced a stronger pattern. By their sixth year, males on gravel bars had fledged 14.5 ± 2.1 chicks, more than four times the number of young fledged by males on beaches (3.3 ± 3.1). Over 9 years, local population size decreased by about 75%, coincident with a shift in breeding distribution away from high‐quality gravel bars to ocean beaches. This unexpected population decline and shift to poorer quality beaches may have been related to occasional low survival of plovers that over‐winter exclusively on beaches in our study area. Consistently low productivity of plovers breeding on ocean beaches suggests the need for intensified management to ameliorate the negative impacts of predation and human activity on the recovery of this population.     

Pay-offs and penalties of competing migratory schedules

We relate variation in the timing of arrival by migrating birds breeding at northerly latitudes to individual differences in the prior accumulation of energy stores. Balancing starvation risks early in the season against the almost universal declining trend in reproductive prospects with advancing date is seen as an individual decision with fitness consequences. We review three studies implicating events at the staging sites or in winter in setting the individual migratory schedule. Climate change influences the timetable of a pied flycatcher (Ficedula hypoleuca) population breeding in The Netherlands and wintering in West Africa, followed since 1960. Mean air temperature in the period mid April‐mid May (arrival and laying) increased and laying date advanced by 10 days. Still, in recent years most birds did not lay early enough to maximise fitness (determined by recruitment and parental survival) whereas many parents achieved this goal in 1980–1985. As the flycatchers have not started to arrive earlier, some ecological constraint further upstream is postulated (possibly the hurdle of the crossing of Sahara and Mediterranean). The ability to follow individual migrants provides a second avenue to assess the fitness implications of migratory schedules. Thus, brightly coloured male bar‐tailed godwits (Limosa lapponica) captured in the Dutch Wadden Sea (the intermediate staging site linking a West African wintering area with breeding sites in arctic Russia) and traced with miniature radio‐transmitters did not depart early. The ‘best’ males (with bright breeding plumage) were picked up by the listening stations in Sweden 650 km further along the migratory route ten days later than the paler individuals. If early arrival confers the competitive advantage of prior occupancy but increases mortality, the ‘best’ males may be able to afford arriving later and thus avoid some of the survival costs. Return rate of the ‘bright’ males to the staging site in later seasons was indeed higher than for the ‘pale’, early males. Intensive observation of pink‐footed geese (Anser brachyrhynchus) fitted with coded neck‐collars substantiate the tight relationship between energy stores (fat) accumulated up to final departure from the final staging site (Vesterålen, N. Norway) en route to the nesting grounds (Spitsbergen) and subsequent success. The breeding outcome of individual parents (accompanied by juveniles or not) could be related to observations of body condition before departure (visual ‘abdominal profile index’). Recently, perceived conflicts with agriculture have resulted in widespread harassment by humans. The geese have: drastically shortened their stay on the Vesterålen, fail to achieve the body condition usual a decade ago and reproductive output has fallen. Although the geese are currently pioneering new staging sites, an adequate alternative has not materialised, underlining the critical role of the final take‐off site.     

Interactions between a Candidate Gene for Migration (ADCYAP1), Morphology and Sex Predict Spring Arrival in Blackcap Populations

Avian research has begun to reveal associations between candidate genes and migratory behaviors of captive birds, yet few studies utilize genotypic, morphometric, and phenological data from wild individuals. Previous studies have identified an association between ADCYAP1 polymorphism and autumn migratory behavior (restlessness, or zugunruhe), but little is known about the relationship between ADCYAP1 and spring migratory behavior. The timing of spring migration and arrival to the breeding ground are phenological traits which could be particularly favorable for establishing territories and acquiring mates, thus important to fitness and reproductive success. Here, we investigated how individual genotypic ADCYAP1 variation and phenotypic variation (wing length and shape) of blackcaps (Sylvia atricapilla) affect spring arrival date across nine natural populations in Europe. We hypothesized that longer alleles should be associated with earlier spring arrival dates and expected the effect on arrival date to be stronger for males as they arrive earlier. However, we found that longer wings were associated with earlier spring arrival to the breeding grounds for females, but not for males. Another female-specific effect indicated an interaction between ADCYAP1 allele size and wing pointedness on the response of spring arrival: greater allele size had a positive effect on spring arrival date for females with rounder wings, while a negative effect was apparent for females with more pointed wings. Also, female heterozygotes with pointed wing tips arrived significantly earlier than both homozygotes with pointed wings and heterozygotes with round wings. Stable isotope ratios (δ ² H) of a subset of blackcaps captured in Freiburg in 2011 allowed us also to assign individuals to their main overwintering areas in northwest (NW) and southwest (SW) Europe. NW males arrived significantly earlier to the Freiburg breeding site than both SW males and females in 2011. NW females had more pointed wing tips compared to SW females, but no difference in ADCYAP1 allele size was found between the different migration routes.     

HIGHER FITNESS FOR PHILOPATRIC THAN FOR IMMIGRANT MALES IN A SEMI-ISOLATED POPULATION OF GREAT REED WARBLERS

To compare the fitness of philopatric and immigrant individuals we examined the lifetime reproductive success of 116 male and 137 female great reed warblers. The study was carried out in a semi-isolated population in Sweden and covered breeding adults hatched between 1985 and 1993. Lifetime fitness, measured as life time number of fledglings and offspring recruits, was lower for immigrant than for philopatric males. We found no such relationships for females. The difference in reproductive success could not be explained by immigrant males having lower phenotypic quality because they had similar life span, spring arrival date, and territory quality as philopatric males. The lower lifetime fitness among immigrant than philopatric males appeared to result from reduced mating success. This suggests that females are reluctant to mate with immigrant males despite their apparently similar phenotypic quality. Though it is not known whether females gain in fitness by avoiding matings with immigrant males, it is notable that immigrant males have smaller song repertoires than philopatric males. Large repertoires, previously shown to sexually arouse great reed warbler females, correlate with the occurrence of extrapair paternity and postfledging survival of offspring in our population.     

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.     

Seasonal patterns in reproductive success of temperate‐breeding birds: Experimental tests of the date and quality hypotheses

For organisms in seasonal environments, individuals that breed earlier in the season regularly attain higher fitness than their late‐breeding counterparts. Two primary hypotheses have been proposed to explain these patterns: The quality hypothesis contends that early breeders are of better phenotypic quality or breed on higher quality territories, whereas the date hypothesis predicts that seasonally declining reproductive success is a response to a seasonal deterioration in environmental quality. In birds, food availability is thought to drive deteriorating environmental conditions, but few experimental studies have demonstrated its importance while also controlling for parental quality. We tested predictions of the date hypothesis in tree swallows (Tachycineta bicolor) over two breeding seasons and in two locations within their breeding range in Canada. Nests were paired by clutch initiation date to control for parental quality, and we delayed the hatching date of one nest within each pair. Subsequently, brood sizes were manipulated to mimic changes in per capita food abundance, and we examined the effects of manipulations, as well as indices of environmental and parental quality, on nestling quality, fledging success, and return rates. Reduced reproductive success of late‐breeding individuals was causally related to a seasonal decline in environmental quality. Declining insect biomass and enlarged brood sizes resulted in nestlings that were lighter, in poorer body condition, structurally smaller, had shorter and slower growing flight feathers and were less likely to survive to fledge. Our results provide evidence for the importance of food resources in mediating seasonal declines in offspring quality and survival.     

Arrival timing and hematological parameters in Gray Catbirds (Dumetella carolinensis)

Early arrival at the breeding grounds for migratory birds is associated with greater reproductive success. According to the condition-dependent arrival hypothesis, only those individuals in superior physiological condition are able to bear the costs (e.g., poor environmental conditions, limited food availability) of early arrival. Condition has usually been measured in terms of energy reserves or mass but other physiological measures of condition such as hematocrit and immune function have been gaining attention. We examined several measures of condition and their association with date of first capture in Gray Catbirds (Dumetella carolinensis) arriving at breeding grounds in northeastern Pennsylvania. Earlier arrivals had higher hematocrit and H/L ratios and lower lymphocyte counts. Arrival date was also negatively associated with fat score. Fat score was positively related to hematocrit, total number of leukocytes, and number of lymphocytes, but the other hematological parameters were not associated with traditional measures of condition (keel score, fat score, or a body condition index). Our results provide some support for the condition-dependent arrival hypothesis and suggest that there may be immunological differences between early- and late-arriving birds.     

Examining carry‐over effects of winter habitat on breeding phenology and reproductive success in prairie warblers Setophaga discolor

Winter habitat quality can influence breeding phenology and reproductive success of migratory birds. Using stable isotope ratios of carbon (δ¹³C) from bird claws and red blood cells collected in Massachusetts, USA, we assessed if winter habitat occupancy carried over to affect prairie warbler Setophaga discolor breeding arrival dates, body condition upon arrival, pairing success, first‐egg dates and reproductive success. In two of three years (in 2011 and 2012, but not in 2013), after‐second‐year (ASY) males wintering in drier habitat, as indicated by enriched δ¹³C values, arrived later on the breeding grounds. Based on the North Atlantic Oscillation index, there was likely less rainfall in the Caribbean wintering grounds during the winters of 2011 and 2012 compared to the winter of 2013, suggesting increased winter rainfall in 2013 may have diminished the influence of winter habitat occupancy on arrival date. We did not find any effects of winter habitat on breeding season phenomena for second‐year (SY) males or females, but our sample sizes for these age/sex classes were relatively low. Although winter habitat quality influenced arrival dates of ASY males, there was no evidence that it affected reproductive performance, perhaps because of high rates of nest depredation in our system. Our study adds to a growing body of research that shows the influence of carry‐over effects can differ among species and within populations, and also can be modulated by other environmental conditions. This information enriches our understanding of the role of carry‐over effects in population limitation for migratory birds.