Seasonal decline in provisioning effort and nestling mass of Blue Tits Cyanistes caeruleus: experimental support for the parent quality hypothesis

A common feature of many birds breeding in seasonal environments is that fitness‐related parameters such as nestling mass or survival decline as the breeding season progresses. Consequently, there is a tendency for early breeders to have better reproductive performance than individuals breeding later in the season. This variation could be caused by factors associated with the date of laying, such as changing environmental conditions (date hypothesis), or by differences in parental quality between early and late breeders (parent quality hypothesis). To evaluate the relative importance of both hypotheses, we manipulated hatch dates of Blue Tits Cyanistes caeruleus by exchanging clutches with different incubation stages and assessed the impact on nestling mass, nestling diet and provisioning rates. Mean nestling mass declined significantly over the season. This was the combined result of differences in parental quality, which dominated in the early part of the season, and the influence of hatching date (date effect per se), which prevailed later in the season. Nestling diet composition was apparently uninfluenced by the manipulation, suggesting that deteriorating food supplies are the primary reason for the seasonal variation in the nestling diet. Counter to the date hypothesis, delayed parents did not feed their young less than control pairs did, but in fact exhibited higher provisioning rates. Our results suggest that in this population, parental quality seems to constrain post‐hatching reproductive performance and such intrinsic limitations may help to explain why certain individuals breed later.     

The quality and the timing hypotheses evaluated using data on great reed warblers

The seasonal decline in reproductive success observed in many animal species may be caused by timing per se (timing hypothesis) or by variation in phenotypic quality between early and late breeding females (quality hypothesis). To distinguish between these two hypotheses, several studies of birds have used clutch removal experiments to manipulate breeding date. However, removal experiments also increase the females' previous reproductive effort due to the production of an extra clutch and a longer incubation period. According to life-history theory an increase in reproductive effort lowers future reproduction. Hence, life-history theory predicts lowered success of replacement broods for other reasons than expected from the timing hypothesis. Female great reed warblers, Acrocephalus arundinaceus , studied in Sweden are frequently exposed to nest predation, after which many lay replacement clutches. In order to examine possible effects of previous reproductive effort on different fitness components, we analysed the re-laying frequency and the reproductive success of replacement broods in relation to time of the season and previous reproductive effort (measured as the length of the previous breeding attempt, LPB). In clutches of re-laying females both the number of fledglings and the proportion of recruits were negatively correlated with LPB, whereas re-laying frequency and clutch size were not related to LPB. We expect such relationships to be present also among other species. Consequently, the use of replacement clutches, as for example in clutch removal experiments, in evaluations of the cause of the often observed seasonal decline in various fitness components, might exaggerate the importance of the timing hypothesis over the quality hypothesis.     

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.     

Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

Fitness consequences of timing of breeding in birds: date effects in the course of a reproductive episode

In seasonal environments, avian reproductive performance almost generally declines in the course of the season. Quantifying the associated fitness consequences of timing of breeding, i.e. of date‐related factors, is important for understanding the evolution of temporal patterns in avian life‐histories and for predicting consequences of climate change. The seasonal decline can also be caused by an effect of parental quality: individuals with high phenotypic quality may breed early. The results of existing experimental studies investigating whether date or quality effects cause the seasonal decline are inconsistent, indicating that both mechanisms might be involved. However, it remains unclear to what extent the confounding effect of quality occurs and what the fitness consequences of timing per se over a whole breeding episode are. In a cross‐fostering experiment using the barn swallows’ second broods we evaluated the causes for the seasonal decline in reproductive performance for three distinct periods of a reproductive attempt, the early nestling period, the late nestling period and the post‐fledging period, and we assessed the overall fitness consequences of timing per se. A seasonal decline in juvenile feather growth rate was mainly due to date effects in the late nestling period, although we determined quality effects during early nestling development. Date effects on survival were present in the post‐fledging period, but not in the nestling period. The decline in feather length due to date effects in the nestling period accounted for 9% of the seasonal decline in post‐fledging survival, whereas date effects arising only in the post‐fledging period caused 91% of the decline. These results suggest that date effects increase in the course of a reproductive episode. Thus, the benefits of an early timing of breeding can be quantified only when considering also the post‐fledging period. We suggest that the timing of breeding evolved through a trade‐off between date‐related benefits and quality‐related costs of early breeding.     

Seasonal declines in reproductive success of the common tern Sterna hirundo: timing or parental quality?

Reproductive success declines over the course of the breeding season in many bird species. Two categories of hypothesis have been evoked to explain this decline. The “timing” hypothesis suggests that seasonal declines in breeding success are attributable to the date of laying. The “parental quality” hypothesis suggests that seasonal declines result from the fact that young, inexperienced, or low quality birds breed later in the season. To evaluate the relative importance of timing and parental quality, egg exchanges and removals were used to manipulate hatching dates of common terns Sterna hirundo. Indices of quality, attendance, provisioning rates, and reproductive success of birds in three experimental groups (delayed hatch pairs, advanced hatch pairs, and pairs induced to relay) were compared to those of date‐matched controls. Pairs that hatched chicks early raised more chicks than pairs hatching chicks late in the season, regardless of initial laying date. This suggests that hatching chicks early is advantageous in itself. Our results, however, also support the parental quality hypothesis. There was a significant negative relationship between natural laying date and fledging success, independent of hatching date. Differences in chick growth and survival suggest that higher quality adults may be able to compensate for the disadvantages of late hatching dates and achieve similar reproductive success to that of pairs hatching chicks early. We found that pairs hatching chicks late in the season were subject to more incidents of kleptoparasitism than those hatching chicks early. This may be a proximate factor contributing to seasonal declines in reproductive success for common terns, although such a mechanism would not be likely in non‐colonial species. Failure to control for egg quality may have biased the results of some prior egg exchange experiments. Additionally, altered cost of incubation may be an unavoidable confounding factor in studies designed to manipulate timing of breeding.     

Age-specific variation in reproduction is largely explained by the timing of territory establishment in the New Zealand stitchbird Notiomystis cincta

1. Using data from 327 nests over a consecutive 8-year period we examined age-specific variation in reproduction in a population of stitchbirds (or hihi) Notiomystis cincta and related how differences in reproductive performance were linked to the timing of territory establishment and breeding. 2. Across the population all reproductive parameters showed a quadratic relationship with an increase mainly between the first and second breeding season and a decline after the fourth year. A longitudinal analysis showed evidence of senescence by the sixth year in the numbers of chicks fledged and recruited. 3. Reproductive increases between years 1 and 2 were the result of poor-quality females dying after their first breeding season (differential selection hypothesis) in combination with surviving females showing improvements in reproduction in their second year (individual improvement/constraint hypothesis). 4. There was no effect of mate experience or territory quality on improvements in breeding between years. 5. The key variable influencing reproductive output was the timing of breeding. Birds that started laying earlier were more likely to lay multiple clutches in any given season. This was the main difference between first-year and older birds; generally first-year birds initiated egg laying later and consequently laid fewer clutches. 6. Approximately half of all first-year birds did not establish their territory until after the breeding season had begun. This delay in territory establishment resulted in these birds delaying breeding, which resulted in them having a lower reproductive output relative to all other birds. First-year birds that managed to establish their territory before breeding commenced, had similar rates of reproduction as older birds. 7. There was a positive relationship between the timing of territory establishment during a female's first year and her hatching date in the previous breeding season. We hypothesize that this was because late-hatched females were less able to effectively compete for territories against earlier-hatched members of their cohort, and this delayed their establishment and breeding in their first year. Thus, this social constraint is likely to be a major factor driving age-specific reproductive variation in this population.     

Experimental dissociation of individual quality, food and timing of breeding effects on double-brooding in a migratory songbird

Annual reproductive success in many species is influenced by the number of breeding attempts within a season. Although previous studies have shown isolated effects of female quality, food, and timing of breeding on the probability of female birds producing second broods, to our knowledge, none have tested the relative importance of multiple factors and their interactions using simultaneous manipulations within populations of free-living birds. In this study, we show that individual quality and timing of breeding interact to affect the probability of double-brooding in female mountain bluebirds (Sialia currucoides). High-quality females (those that naturally initiated clutches early in the season) were more likely to double-brood, regardless of whether their hatching date was advanced or delayed, whereas later breeding, lower quality females were much less likely to double-brood when their first attempt was delayed. This indicates that annual fecundity of poorer quality (or younger) female bluebirds may be more sensitive to seasonal variation in environmental conditions. In addition, birds that were provided with supplemental food throughout first breeding attempts were more likely to double-brood in one of the study years, suggesting that female bluebirds may be energetically limited in their capacity to initiate a second brood. Females that had their first brood delayed also had a shorter inter-brood interval and were moulting fewer feathers during second broods compared to controls, while females in better condition showed more advanced moult in second breeding attempts. Taken together, our results demonstrate the combined effects of age- or individual quality-mediated energetic trade-offs between current and future reproduction, and between investments in offspring and self-maintenance, on annual fecundity of female birds.     

Cavity size, clutch-size and the breeding ecology of Tree Swallows Tachycineta bicolor

Previous studies provide evidence that cavity size influences clutch-size and reproductive success in some hole-nesting birds, because overcrowding in cavities may cause brood mortality due to trampling or hyperthermia. We tested this hypothesis with two experiments at nestbox populations of Tree Swallows Tachycineta bicolor in southeastern Ontario. Female Tree Swallows showed a preference for nesting in large boxes over small ones in the first experiment, and they laid significantly smaller clutches in small nestboxes during both experiments. Differences in clutch–size between large and small nestboxes could not be attributed to other factors known to influence clutch-size in birds (e.g. parental quality, habitat quality, laying date). Reproductive success, however, did not differ between pairs using the two box types during either experiment, nor did it differ during within-clutch-size comparisons between box types. Some measures of nestling quality were significantly lower for broods in large boxes, but during most comparisons there were no differences. These results do not support the adaptive reason for why cavity size affects clutch-size. We suggest that broods in our experiments did not experience the microhabitat conditions necessary to induce the expected differences in brood mortality.     

Kittiwakes strategically reduce investment in replacement clutches

Many life-history traits are expressed interactively in life, but to a varying extent on different occasions. Changes in trait expression can be accounted for by differences in the quality of the environment ('environmental constraint' hypothesis) or by strategic adjustments, if the relative contribution of the trait to fitness varies with time ('strategic allocation' hypothesis). In birds, egg production is lower in replacement clutches than in first clutches, but it is unknown whether this reduction results from an environmental constraint (e.g. food being less available at the time when the replacement clutch is produced) or from a strategic allocation of resources between the two breeding attempts. To distinguish between these two hypotheses, we performed an experiment with black-legged kittiwakes (Rissa tridactyla). Pairs were either food-supplemented or not before the first clutch was laid onwards and we induced them to produce a replacement clutch by removing eggs once when the first clutch was complete. As predicted by the 'strategic allocation' hypothesis, egg production of food-supplemented and non-food-supplemented birds decreased between first and replacement clutches. This suggests that kittiwakes strategically reduce investment in egg production for their replacement clutches compared to first clutches.     

Passerine birds breeding under chronic noise experience reduced fitness

Background

Fitness in birds has been shown to be negatively associated with anthropogenic noise, but the underlying mechanisms remain obscure. It is however crucial to understand the mechanisms of how urban noise impinges on fitness to obtain a better understanding of the role of chronic noise in urban ecology. Here, we examine three hypotheses on how noise might reduce reproductive output in passerine birds: (H1) by impairing mate choice, (H2) by reducing territory quality and (H3) by impeding chick development.

Methodology/Principal Findings

We used long-term data from an island population of house sparrows, Passer domesticus, in which we can precisely estimate fitness. We found that nests in an area affected by the noise from large generators produced fewer young, of lower body mass, and fewer recruits, even when we corrected statistically for parental genetic quality using a cross-fostering set-up, supporting H3. Also, individual females provided their young with food less often when they bred in the noisy area compared to breeding attempts by the same females elsewhere. Furthermore, we show that females reacted flexibly to increased noise levels by adjusting their provisioning rate in the short term, which suggests that noise may be a causal factor that reduces reproductive output. We rejected H1 and H2 because nestbox occupancy, parental body mass, age and reproductive investment did not differ significantly between noisy and quiet areas.

Conclusions/Significance

Our results suggest a previously undescribed mechanism to explain how environmental noise can reduce fitness in passerine birds: by acoustically masking parent–offspring communication. More importantly, using a cross-fostering set-up, our results demonstrate that birds breeding in a noisy environment experience significant fitness costs. Chronic noise is omnipresent around human habitation and may produces similar fitness consequences in a wide range of urban bird species.     

Cold tolerance,and not earlier arrival on breeding grounds,explains why males winter further north in an Arctic‐breeding songbird

Sex biases in distributions of migratory birds during the non‐breeding season are widespread; however, the proximate mechanisms contributing to broad‐scale sex‐ratio variation are not well understood. We analyzed a long‐term winter‐banding dataset in combination with spring migration data from individuals tracked by using geolocators to test three hypotheses for observed variation in sex‐ratios in wintering flocks of snow buntings Plectrophenax nivalis. We quantified relevant weather conditions in winter (temperature, snowfall and snow depth) at each banding site each year and measured body size and condition (fat scores) of individual birds (n > 5500). We also directly measured spring migration distance for 17 individuals by using light‐level geolocators. If the distribution pattern of birds in winter is related to interactions between individual body size and thermoregulation, then larger bodied birds (males) should be found in colder sites (body size hypothesis). Males may also winter closer to breeding grounds to reduce migration distance for early arrival at breeding sites (arrival timing hypothesis). Finally, males may be socially dominant over females, and thus exclude females from high‐quality wintering sites (social dominance hypothesis). We found support for the body size hypothesis, in that colder and snowier weather predicted both larger body size and higher proportions of males banded. Direct tracking revealed that males did not winter significantly closer to their breeding site, despite being slightly further north on average than females from the same breeding population. We found some evidence for social dominance, in that females tended to carry more fat than males, potentially indicating lower habitat quality for females. Global climatic warming may reduce temperature constraints on females and smaller‐bodied males, resulting in broad‐scale changes in distributional patterns. Whether this has repercussions for individual fitness, and therefore population demography, is an important area of future research.     

Parental age and reproduction in the Marsh Tit Paras palustris

In the Marsh Tit Parus palustris , breeding performance improved with the age of both the male and the female parent—older birds produced larger clutches earlier in the season. The effect of male age, however, was mainly due to a positive association with the age of the female. When the effect of laying date was accounted for, there was no additional effect of female age on clutch-size. However, pairs with previous breeding experience laid larger clutches than newly formed pairs of the same age. There was some evidence that young females that laid late in the season were less likely to survive to breed again, leading to an age effect on reproductive performance. There was no evidence that individual birds laid earlier or produced larger clutches when they became older.     

Differential maternal investment counteracts for late breeding in magpies Pica pica: an experimental study

Reproductive success in many avian populations declines throughout the breeding season. Two hypotheses have gained attention to explain such a decline: the "timing" hypothesis proposes that deteriorating food availability causes the decline in reproductive success (causal effect of breeding time), whereas the "quality" hypothesis proposes that individuals of lower phenotypic quality reproduce at the end of the breeding season, causing the correlation between breeding time and breeding success. We tested both of these hypotheses in a monogamous single breeder, the magpie Pica pica , by experimentally inducing some pairs to lay a replacement clutch, after removal of the first clutch. The first clutch was left in the nest of another magpie pair (matched by laying date and clutch size), and incubated and raised by these foster parents. In this way we obtained two clutches from the same magpie pair with full siblings raised in conditions of the first and second reproductive attempts. High quality pairs (with laying dates in the first half of the breeding season) reached similar breeding success in replacement clutches as compared to first clutches of the same female. In addition, experimental pairs reared significantly more offspring of similar quality in their replacement clutches as compared to late-season first clutches, thereby suggesting that late season first clutches were produced by pairs of lower phenotypic quality. Results indicate that high quality pairs trade-off clutch size for larger eggs in replacement clutches, which could help magpie pairs to partly compensate for poorer environmental conditions associated with a delayed breeding attempt.     

Correlates of timing of spring migration in birds: a comparative study of trans-Saharan migrants

The evolution of migratory strategies in birds is likely to have been influenced by ecological as well as socio-sexual factors in both wintering and breeding areas. In this comparative study, we analysed timing of spring passage of 38 long-distance migratory bird species that winter south of the Sahara desert and breed in Europe, in relation to wintering and breeding latitudes, moult strategy, nesting site (open vs. cavity), and sexual dimorphism in size and coloration, which may reflect intensity of sexual selection. We employed a large data set consisting of more than 190 000 individuals ringed during spring migration in the Mediterranean Sea. We found that the species that migrated earlier were those wintering farther north, nesting in cavities and showing the largest degree of sexual size dimorphism (SSD). However, sexual dichromatism was not related to migration date. Among passerine species, moulting wing-feathers in Africa delayed migration. We found no support for the energetic constraint hypothesis, which proposes that early arrival selects for large male size, since early arriving species were not larger than late arriving ones. Thus, the observed associations suggest that variation in migration schedules at the interspecific level may have evolved in relation to ecological factors and SSD, possibly reflecting the intensity of mating competition.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 199–210.     

Offspring fitness and individual optimization of clutch size

Within-year variation in clutch size has been claimed to be an adaptation to variation in the individual capacity to raise offspring. We tested this hypothesis by manipulating brood size to one common size, and predicted that if clutch size is individually optimized, then birds with originally large clutches have a higher fitness than birds with originally small clutches. No evidence was found that fitness was related to the original clutch size, and in this population clutch size is thus not related to the parental capacity to raise offspring. However, offspring from larger original clutches recruited better than their nest mates that came from smaller original clutches. This suggests that early maternal or genetic variation in viability is related to clutch size.     

Migration phenology and winter habitat quality are related to circulating androgen in a long‐distance migratory bird

In migratory birds, the timing of departure from wintering grounds is often dependant on the quality of habitat on an individual's territory and may influence individual fitness, resulting in an interaction of life history stages across large geographical distances. American redstart Setophaga ruticilla males who overwinter in high quality habitats arrive early to breed and subsequently produce more offspring than late arrivers. Since many migratory species overlap vernal migration with the physiological transition to breeding, we examined if breeding preparation plays a role in this seasonal interaction. We tested the hypothesis that early arriving male redstarts from high quality winter habitats are in superior breeding condition by simultaneously measuring winter habitat quality (stable‐carbon isotopes) and breeding preparation (circulating androgen, cloacal protuberance (CP) diameter) upon arrival at breeding grounds. Compared with late arrivers, early arriving males were from higher quality winter habitats and had higher androgen, but smaller CPs. Males arriving with higher androgen were in more advanced physiological migratory condition, as measured by haematocrit. Early arrivers were more likely to successfully breed, but there was no significant relationship between androgen upon arrival and breeding success. One possible explanation for these relationships is that androgen measured during arrival is most relevant in a migratory context, such that birds with high androgen may benefit from effects on migratory condition, positively influencing fitness through earlier arrival.     

Temperature but not rainfall influences timing of breeding in a desert bird, the trumpeter finch ( Bucanetes githagineus )

Reproductive performance in birds depends on several factors, one of the most important being the time of breeding. Birds try to fit offspring birth and growth to peak vegetative production in order to assure fledgling survival. In arid environments, where weather conditions are often extreme, birds must face unpredictable abiotic conditions. This study uses a border population of the trumpeter finch (Bucanetes githagineus) as a model to test whether climate variables (rainfall and temperature) influence breeding parameters by comparing 2 years with very different weather. The study was carried out in the Tabernas desert (southeastern Spain) in 2004 and 2005. A comparison of laying dates in the 2 years shows a 40-day delay in the date of the first clutch in the coldest year (mean minimum temperature 3°C lower in 2005 than in 2004). However, once the breeding season started, the number of clutches, clutch size, duration of the incubation period, nestling phase, fledgling rates and productivity were similar. One likely explanation for this delay is that low temperatures did not allow the germination of Diplotaxis sp., a plant forming the bulk of the trumpeter finch diet during spring. Its absence could prevent onset of breeding, although other temperature-related factors could also be involved. Although rainfall has frequently been reported as a limiting factor for arid bird species, our 2-year study shows that temperature can also influence the breeding biology of arid bird species, by affecting its timing.     

Reduced immunocompetence of nestlings in replacement clutches of the European magpie (Pica pica)

Laying date is one of the most important determinants of reproductive success and recruitment probability in birds. Late breeders usually fledge fewer chicks than individuals with earlier breeding dates, and fledglings produced late in the season have high mortality rates. Food availability and nestling mass have been evoked as the principal mechanistic links between laying date and offspring survival. Here we suggest that another factor may actually account for the difference in survival rate between early and late offspring: immunocompetence. We predicted that nestlings produced later in the season or in replacement clutches should have lower immune responses when challenged with an antigen, than early nestlings or nestlings produced in first clutches. This hypothesis was tested in a population of magpies (Pica pica), in which we experimentally induced breeding failure in a group of nests and compared the immune response of nestlings in replacement clutches with the immune response of first clutch nestlings. Cellular immune response, as measured by wing web swelling (a correlate of T-lymphocyte production after injection of phytohaemagglutinin-P), significantly decreased with hatching date and was significantly lower in nestlings of replacement clutches. Furthermore, coefficients of intraclutch variation in immune response were higher in nestlings of replacement clutches. This experiment demonstrates an inverse relationship between immune responsiveness and breeding date, and reduced recruitment probability of late nestlings may be a direct consequence of their inability to cope with parasites.     

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.