Local breeding experience and the reproductive performance of Tree Swallows

ABSTRACT The potential advantages of repeated breeding at a particular location should improve reproductive performance in long‐lived species of birds. However, for short‐lived species, natural selection should favor individuals that most quickly develop competency in reproduction. Therefore, we hypothesized that local breeding experience beyond the first breeding attempt at a particular location would have little effect on subsequent reproductive performance of Tree Swallows (Tachycineta bicolor), a species where about 50% of adults breed only once in their lives. We tested this hypothesis using data collected from Tree Swallows in Michigan from 1993 to 2002. Because we were specifically interested in examining the effects of local breeding experience on reproductive performance, we restricted our analyses to after‐second‐year (ASY) females and their mates that we first encountered as breeders. Consistent with our hypothesis, we found no relationship between repeated local breeding experience and the reproductive performance of ASY female Tree Swallows and their mates as measured by clutch size and number of fledged young. However, pairs with more combined total local experience tended to lay eggs earlier in the season. These results suggest that Tree Swallows may benefit from breeding site fidelity, not because repeated local experience improves reproductive performance as measured by the production of fledglings, but because returning individuals acquire nest cavities earlier and are able to begin breeding earlier, providing time to renest in case of early nest failure.     

Differences in size between first and replacement clutches match the seasonal decline in single clutches in Tree Swallows Tachycineta bicolor

The seasonal decline in clutch size in birds can be a response to the environmentally conditioned decrease in prospects for offspring or a consequence of a lower physical ability of late‐breeding females. To find out which of the explanations apply in Tree Swallows Tachycineta bicolor, we assessed whether replacement clutch size in this species is affected by an individual female's ability to lay a certain number of eggs. To do this, we measured the decline in clutch size as a function of laying date between first and replacement clutches in individuals that re‐nested following natural failure, and compared this with the rate of decline in clutch size with laying date for Tree Swallows that laid only a single clutch in that season. Additionally, we assessed whether the clutch size and the rate of its seasonal decline varied across years. We accounted for the truncated and under‐dispersed nature of clutch size data by using a Bayesian approach in the analysis. We found little variation in the rate of clutch size decline across years at our breeding site. Accounting for this seasonal decline in clutch size, mean clutch size was similar between single‐time breeding females and those that laid replacement clutches, implying that the number of eggs laid on the second attempt by female Tree Swallows is determined by laying date, rather than by the female's physical ability to produce a clutch of a certain size.     

Breeding success and chronology of Wood Storks Mycteria americana in northern and central Florida, U.S.A.

The breeding success and chronology of Wood Storks Mycteria americana were studied at eight colonies in northern and central Florida during 1981–1985. Mean ± s.d. clutch size for all colony-years was 3.07 ± 0.56 (n = 2694 nests), with three-egg clutches (72%) most frequent. Mean clutch size among all colonies and years ranged from 2.73 ± 0.55 to 3.41 ± 0.61. Many colonies exhibited significant negative trends in clutch size with, hatching date because of a proportional decrease in four-egg clutches later in the season. Mean colony clutch size was not correlated with nest numbers, nesting density or mean hatching date within most years. Mean ± s.d. number of fledglings for all colonies and years was 1.29 ± 1.16 fledglings per nest (n = 2812 nests). Mean annual fledging rates in colonies ranged from 0 (colony failed) to 2.66 fledglings per nest. Most breeding failure occurred prior to egg hatching, and the second highest mortality occurred between hatching and 2 weeks of age. Four-egg clutches fledged more storks than three-egg clutches, which in turn were more successful than two-egg clutches. However, all clutch sizes showed similar fledgling per egg rates. The seasonal decline in productivity was associated proportionally with smaller clutch sizes later in the breeding season. An increase in mean hatching date was correlated with an increase in latitude. There was greater within-year breeding synchrony among colonies than interyear breeding synchrony within each colony. Breeding synchrony was not correlated with mean hatching date, latitude, longitude, nest numbers or nesting density.     

The breeding success of the Indian Cattle Egret Ardeola ibis in eastern Australia

Over a three year study the Cattle Egret's mean clutch size was 3.6 eggs and 81% of nests with eggs produced one or more fledglings. Breeding success averaged 2.1 chicks to fledgling stage, and between 1.5 and 1 8 fledged per nesting pair. These data illustrate the breeding performance of a population in an expansion phase of its history. The determinants of growth of immigrant Cattle Egret populations are sought in a comparison of its breeding parameters in different lands. A significant reduction in clutch size occurred after about two-thirds of clutches had been laid each year. One-year-old birds nesting for the first time are implicated in this reduction.     

Does prior breeding experience improve reproductive success in collared flycatcher females?

In order to assess the importance of breeding experience in shaping age-specific patterns of reproduction an experiment on collared flycatchers (Ficedula albicollis) was performed. Experimental females were restrained from reproduction by destroying their nests after prolonged incubation. Unmanipulated females were used as controls. Only young, first time breeders were considered. A year after manipulation, experimental (inexperienced) and control (experienced) females were compared for a number of breeding traits. They did not differ with respect to laying date, clutch size, number of fledglings and number of local recruits. However, fledglings from broods of inexperienced females were significantly lighter and had shorter tarsi. This suggests that prior experience of feeding young may indeed influence the future reproductive performance in altricial bird species such as the collared flycatcher.     

Food abundance and clutch size of Tree Swallows Tachycineta bicolor

We studied Tree Swallows Tachycineta bicolor during eight breeding seasons (1977-84) at three sites that differed in food availability, which we measured as an index of the biomass of flying insects derived from daily catches in suspended nets. Mean clutch size differed significantly between sites (6.2, 5.8, and 5.6 eggs) and was related to average food abundance at the site. Within two sites, clutch sizes were significantly larger in years when food was more abundant during egg-laying ( r = 0.83 and 0.97 at the high and low food abundance sites, respectively; r = 0.63, not significant, at the medium abundance site). Both clutch size and laying season food abundance were positively correlated with food abundance during the nestling stage at the 'high' site, but not at the other two sites. Our results support the hypothesis that female Tree Swallows respond proximately to abundant food during egg-laying by laying more eggs, but they do not elucidate the ultimate evolutionary factors involved in determining clutch size.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

Family break-up in Black and Red Kites Milvus migrans and M. milvus: is time of independence an offspring decision?

I observed the natural process of family break-up in 13 families of Black Kites Milvus migrans and five families of Red Kites M. milvus in which fledglings had been individually marked. In other broods, I performed experiments which modified the parental investment fledglings received by supplementing nests of both species with food or transferring Black Kite chicks to nests with younger or older chicks of the same species.
The time of family break-up in the Black Kite is mainly an offspring decision which is not affected by an artificial increase of parental investment. The duration of the post-fledging period was not increased in Black Kites that were given supplementary food. Chicks transferred to nests with a younger chick did not extend the post-fledging period, nor did chicks transferred to nests with an older chick shorten the post-fledging period. In Red Kites, parental investment seemed to have more influence on the timing of the family break-up. Red Kite adults invested less as parents than did Black Kites during the post-fledging dependence period, and in nests where supplementary food was given, Red Kite fledglings stayed attached for a longer period.     

Breeding success of a Lesser Spotted Woodpecker Dendrocopos minor population

We made intensive observations of 50 pairs of Lesser Spotted Woodpecker Dendrocopos minor in southern Sweden from 1989 to 1992. Between 7% and 22% of the pairs each year defended a territory but laid no eggs. Overall, 76% of the pairs laying eggs raised fledglings, but this proportion was only 33% in 1991 when the weather was cold and wet. The mean clutch size (5.9) and brood size (4.5) in successful nests did not vary significantly between years. Incubation time was 1 day longer in 1991 (11–12 days) than in 1990 and 1992 (10–11 days). Number of young fledged per nesting attempt (3.5) and per pair (2.7) in the population also varied significantly between years. The initiation of egg laying varied between years but coincided each year remarkably well with the date when oak came into leaf. Breeding success is discussed, and comparisons are made with other hole-nesting species.     

Nest predation reduces benefits to early clutch initiation in northern cardinals Cardinalis cardinalis

Life history theory and empirical studies suggest that early breeding confers higher reproductive success, but the extent to which this advantage can be generalized to human‐dominated systems and across species is less well understood. We studied the fitness consequences of clutch initiation for 181 female northern cardinals Cardinalis cardinalis and 1228 nests in forests within urban and rural landscapes of Ohio, USA between 2004–2007. Cardinals that bred earlier made significantly more nesting attempts, but cumulative number of young fledged was similar to that of later‐breeding individuals. The expected number of fledglings produced per successful nest was unrelated to date and remained ~1.8 fledglings across the season, despite the fact that nest survival rates improved dramatically as the season progressed. Because the probability of resighting breeding individuals in subsequent years was unrelated to first clutch initiation date, we have no evidence that clutch initiation affected adult survival. The absence of a clear benefit to early breeding appears to be a consequence of high rates of nest predation early in the breeding season.     

Colony size, reproductive success, and colony choice in Cave Swallows Petrochelidon fulva

We investigated how annual reproductive success, as measured by the number of nestlings surviving to day 10 and the percentage of nests that were successful, varied with colony size of the Cave Swallow Petrochelidon fulva in south central Texas. We also studied whether Cave Swallows chose colonies, in part, on the basis of reproductive success at a site the previous year. Neither measure of reproductive success varied significantly with colony size for either first-wave or second-wave nestings. Mean clutch size per colony did not vary significantly with colony size. Mean nestling body mass, an index of parental foraging efficiency, was unrelated to colony size, except for broods of five, in which nestling mass declined significantly with colony size. Colony size was not significantly affected by reproductive success at the site the previous year, although sites with more successful nests during the first wave declined less in size during the second wave within the same season than did sites that had fewer successful first-wave nests. Unlike the closely related Cliff Swallow Petrochelidon pyrrhonota , Cave Swallows did not use breeding performance of conspecifics in choosing nest-sites, because they did not preferentially aggregate at sites that were the most successful the previous year. Coloniality in Cave Swallows did not appear to have a net negative effect on annual fitness, suggesting that colonial nesting was not solely a response to nest-site limitation, but the benefits of breeding colonially (if any) were unclear.     

青海省海北地区高寒草甸雀形目鸟类的繁殖生产力

通过对高寒草甸１０种雀形目鸟类繁殖生产力研究表明,高寒草甸鸟类的繁殖生产与鸟类栖息地所处的地理环境、营巢类型和繁殖方式密切相关。其中繁殖季节长度和窝卵数是影响高寒草甸鸟类繁殖生产力的两个最重要的因子（Ｐ〈０．０５）。鸟类平均繁殖生产力为２．８０（只／对．年）。     

Unpredictable food supply modifies costs of reproduction and hampers individual optimization

Investment into the current reproductive attempt is thought to be at the expense of survival and/or future reproduction. Individuals are therefore expected to adjust their decisions to their physiological state and predictable aspects of environmental quality. The main predictions of the individual optimization hypothesis for bird clutch sizes are: (1) an increase in the number of recruits with an increasing number of eggs in natural broods, with no corresponding impairment of parental survival or future reproduction, and (2) a decrease in the fitness of parents in response to both negative and positive brood size manipulation, as a result of a low number of recruits, poor future reproduction of parents, or both. We analysed environmental influences on costs and optimization of reproduction on 6 years of natural and experimentally manipulated broods in a Central European population of the collared flycatcher. Based on dramatic differences in caterpillar availability, we classified breeding seasons as average and rich food years. The categorization was substantiated by the majority of present and future fitness components of adults and offspring. Neither observational nor experimental data supported the individual optimization hypothesis, in contrast to a Scandinavian population of the species. The quality of fledglings deteriorated, and the number of recruits did not increase with natural clutch size. Manipulation revealed significant costs of reproduction to female parents in terms of future reproductive potential. However, the influence of manipulation on recruitment was linear, with no significant polynomial effect. The number of recruits increased with manipulation in rich food years and tended to decrease in average years, so control broods did not recruit more young than manipulated broods in any of the year types. This indicates that females did not optimize their clutch size, and that they generally laid fewer eggs than optimal in rich food years. Mean yearly clutch size did not follow food availability, which suggests that females cannot predict food supply of the brood-rearing period at the beginning of the season. This lack of information on future food conditions seems to prevent them from accurately estimating their optimal clutch size for each season. Our results suggest that individual optimization may not be a general pattern even within a species, and alternative mechanisms are needed to explain clutch size variation.     

THE BIRDS OF LANGEBAAN LAGOON

Data are presented on breeding success of Red Bishops (Euplectes orix) collected over four breeding seasons at a colony in the Addo Elephant National Park, Eastern Cape, South Africa. Overall hatching and fledging success were 53.8% and 26.0% of all eggs laid, respectively, and the overall mean number of fledglings per breeding attempt was 0.77. Hatching and fledging success varied significantly among seasons, with both clutch and brood losses due to predation being the main reason for the observed differences. Hatching success also differed significantly among clutch sizes, being highest for four-egg clutches (63.2%), intermediate for three-egg clutches (55.5%) and lowest for two-egg clutches and five-egg clutches (33.2% and 34.3%, respectively). However, fledging success was not significantly different among clutch sizes. The mean number of fledglings per breeding attempt was 0.44 for two-egg clutches, 0.80 for three-egg clutches, 1.10 for four-egg clutches, and 0.57 for five-egg clutches. The height of accepted nests (i.e.nests in which at least one egg was laid) was significantly lower than the height of nests not accepted. In addition, accepted nests in which eggs hatched and young fledged were significantly lower than accepted nests in which no eggs hatched and no young fledged. These overall effects of nest height on nest acceptance and hatching and fledging success were, however, due only to nests built above water, since no such effects were found when nests built above ground (i.e.on dry land) were analysed separately. I detected no effect of nest coverage on the probability of a nest being accepted, nor was there any effect of nest coverage on hatching or fledging success. Nests above water were significantly more likely to be accepted than nests above ground; however, hatching and fledging success of nests that were accepted did not differ significantly between nests built above water and those built above ground.     

Comparative demography of Burrowing Owls in agricultural and urban landscapes in southeastern Washington

ABSTRACT.   Anecdotal evidence suggests that Burrowing Owls have declined in the state of Washington. We examined the status of these owls in agricultural and urban habitats to better understand the underlying causes of these declines. Nest density was higher in the area dominated by agriculture (0.67 nests/km²) than in the urban area (0.28 nests/km²), and re-use of nest burrows was more common in the agricultural area. We found no difference in mean clutch size between the two areas, but nesting success was higher in the agricultural area. The mean number of fledglings per nesting attempt was higher in the agricultural area (2.02 vs. 1.47), but we found no difference between the two areas in the mean number of fledglings per successful nest (3.2 vs. 3.1). Both natal recruitment (4% vs. 8%) and annual return rate of adults (30% vs. 39%) were lower in the agricultural area than in the urban area, suggesting that the owl population in the agricultural area may not be stable and may be a "sink" population. Due to high burrow fidelity from year to year, and the tendency of some owls in Washington to overwinter, we recommend that legal protection of nest burrows be extended to the nonbreeding season.     

Reproductive trade‐offs in a long‐lived bird species: condition‐dependent reproductive allocation maintains female survival and offspring quality

Life history theory is an essential framework to understand the evolution of reproductive allocation. It predicts that individuals of long‐lived species favour their own survival over current reproduction, leading individuals to refrain from reproducing under harsh conditions. Here we test this prediction in a long‐lived bird species, the Siberian jay Perisoreus infaustus. Long‐term data revealed that females rarely refrain from breeding, but lay smaller clutches in unfavourable years. Neither offspring body size, female survival nor offspring survival until the next year was influenced by annual condition, habitat quality, clutch size, female age or female phenotype. Given that many nests failed due to nest predation, the variance in the number of fledglings was higher than the variance in the number of eggs and female survival. An experimental challenge with a novel pathogen before egg laying largely replicated these patterns in two consecutive years with contrasting conditions. Challenged females refrained from breeding only in the unfavourable year, but no downstream effects were found in either year. Taken together, these findings demonstrate that condition‐dependent reproductive allocation may serve to maintain female survival and offspring quality, supporting patterns found in long‐lived mammals. We discuss avenues to develop life history theory concerning strategies to offset reproductive costs.     

Growth, fledging success and post-fledging survival of juvenile Oystercatchers Haematopus ostralegus

We studied the consequences of differences in growth rate on the subsequent survival of Oystercatcher Haematopus ostralegus chicks. Fledging success increased sharply with growth rate, from zero in chicks growing at less than 6 g per day to about 85% in chicks growing at more than 10 g per day. The age at which chicks fledged varied from 27 to 52 days. Chicks which fledged at an early age displayed a much faster growth rate than later fledging chicks. Although slow growth resulted in a considerable prolongation of the period before fledging, slow-growing chicks fledged at a smaller size and with a lower body-weight than fast-growing chicks. After fledging, all chicks remained almost completely dependent on their parents up to an age of 3 months and often longer.
Almost 40% of the fledglings eventually returned to the breeding area. This figure probably reflects post-fledging survival. Age and size at fledging had no effect on a chick's probability of return. Body-weight at fledging had a small positive correlation with the return probability, but this was not statistically significant. We conclude that although slow growth severely reduces a chick's chance of fledging, it probably does not result in irreversible damage causing an increased risk of mortality during the first years after fledging. Apparently, any possible disadvantage associated with small size or low body-weight could be compensated for after fledging.     

Post-fledging interactions between the Common Cuckoo Cuculus canorus and its cavity-nesting Common Redstart Phoenicurus phoenicurus host

Brood parasite–host interactions during the incubation and nestling stages have been well studied, but the post-fledging period remains virtually unknown. Using radiotracking, we provide the first detailed data on post-fledging interactions between the Common Cuckoo Cuculus canorus and its only regular cavity-nesting host, the Common Redstart Phoenicurus phoenicurus. Cuckoos raised alone (‘solitary’) fledged at higher mass, with higher wing and tarsus length and started to fly at a younger age than Cuckoos raised alongside young Redstarts (‘mixed’). However, a further 23 fledging and post-fledging parameters measured at five pre-determined times (fledging, first-flight, predation, starvation, independence) did not differ between solitary and mixed Cuckoos. In addition, none of the parameters measured during the post-fledging period (growth, dispersal distances, number of flights) differed between solitary and mixed Cuckoos. Redstart fledglings from non-parasitized broods (‘solitary’) showed generally similar fledging and post-fledging parameters to fledglings reared alongside a Cuckoo (‘mixed’). Surprisingly, there were no significant differences in post-fledging predation rate, starvation or overall survival rates between mixed and solitary Cuckoos or mixed and solitary Redstarts. Thus, during the post-fledging period, mixed Cuckoo fledglings successfully compensated for the poorer performance experienced during the nestling stage whereas mixed and solitary Redstarts did not differ in any measured parameters. This suggests that the regular occurrence of mixed broods in this host–parasite system – which is unique among the many Cuckoo hosts – is evolutionarily stable for both hosts and parasites.     

Annual and seasonal reproductive trends in the Lesser Spotted Woodpecker Dendrocopos minor

Annual and seasonal variation in reproductive timing and performance were studied in a population of the Lesser Spotted Woodpecker Dendrocopos minor over 10 years in southern Sweden. The median laying date of the first egg varied by up to 17 days between years, being generally larger than the variation of laying dates within years. Neither clutch size, brood size in successful nests, fledging success in successful nests nor mean nestling weight differed significantly between years. There was no trend for mean clutch size to vary between early and late years. In spite of a more than threefold variation in population size, no reproductive variable demonstrated an apparent density-dependence. Within the season, clutch size declined steeply with increasing clutch initiation date, whereas fledging success and nesting success did not, leading to a trend in brood size almost identical to the trend in clutch size. The survival prospects of fledged young declined with increasing clutch initiation date, and it is argued that the clutch size laid is a strategic adjustment to laying date. Out of 124 breeding attempts, 34% did not produce fledged young. In 9% of the breeding attempts, pairs laid no eggs. At least 20% of the breeding attempts failed after egg-laying. The most common cause of breeding failure was loss of the breeding partner followed by nest abandonment (40% of the failures). Only 16–28% of the failures were due to predation on the nest. Most complete failures, and also partial losses from nests, occurred at the early breeding stages. It is argued that the early nestling phase may be a critical stage, which the woodpeckers adjust to coincide with the seasonal food peak, explaining the strikingly late breeding season compared with other non-migrant species.     

Brood parasitism of White-rumped Swallows by Shiny Cowbirds

ABSTRACT.   Swallows and martins are infrequent hosts of the generalist brood parasite Shiny Cowbird ( Molothrus bonariensis ). We monitored 50 nesting attempts by White-rumped Swallows ( Tachycineta leucorrhoa ) over a two-year period in Argentina and detected low rates of brood parasitism (three nests, or 6%). Of the three nests parasitized, cowbirds ( N = 1 per nest) successfully fledged from two. Eight of 13 swallow eggs in these three nests were punctured by cowbirds, and all but one swallow nesting starved at the two parasitized nests. At least two factors may contribute to the low frequency of parasitism of White-rumped Swallows by Shiny Cowbirds, including the inability of the larger cowbirds to enter some nest cavities and the aggressive nest defense behavior of adult swallows.