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.     

The significance of clutch overlap in Great Tits Parus major

We studied the effects of manipulation of the size of first broods in the Great Tit Parus major on the size and breeding success of second clutches and its relation to the degree of clutch overlap. The rearing of first brood fledglings always overlapped with the laying of the second clutch and in most cases also with the incubation period of the latter. The degree of clutch overlap depended on the size of the first brood, being less when the first brood was large. Clutch overlap also increased with season. Mechanisms affecting the timing of laying of second clutches are discussed. A large first brood imposed reproductive costs. It affected the size of the second clutch by causing it to be delayed; second clutch size decreases with season. It affected the post-fledging survival of second brood young as, in this population, this decreases with fledging date. The breeding success of second clutches was, however, not affected by the size of the first brood, but instead by the weight of the female when rearing the first brood.     

Landscape heterogeneity drives intra‐population niche variation and reproduction in an arctic top predator

While intra‐population variability in resource use is ubiquitous, little is known of how this measure of niche diversity varies in space and its role in population dynamics. Here we examined how heterogeneous breeding environments can structure intra‐population niche variation in both resource use and reproductive output. We investigated intra‐population niche variation in the Arctic tundra ecosystem, studying peregrine falcon (Falco peregrinus tundrius, White) breeding within a terrestrial‐marine gradient near Rankin Inlet, Nunavut, Canada. Using stable isotope analysis, we found that intra‐population niches varied at the individual level; we examined within‐nest and among‐nest variation, though only the latter varied along the terrestrial‐marine gradient (i.e., increased among‐nest variability among birds nesting within the marine environment, indicating higher degree of specialization). Terrestrial prey species (small herbivores and insectivores) were consumed by virtually all falcons. Falcons nesting within the marine environment made use of marine prey (sea birds), but depended heavily on terrestrial prey (up to 90% of the diet). Using 28‐years of peregrine falcon nesting data, we found a positive relationship between the proportion of terrestrial habitat surrounding nest sites and annual nestling production, but no relationship with the likelihood of successfully rearing at least one nestling reaching 25 days old. Annually, successful inland breeders raised 0.47 more young on average compared to offshore breeders, which yields potential fitness consequences for this long‐living species. The analyses of niche and reproductive success suggest a potential breeding cost for accessing distant terrestrial prey, perhaps due to additional traveling costs, for those individuals with marine nest site locations. Our study indicates how landscape heterogeneity can generate proximate (niche variation) and ultimate (reproduction) consequences on a population of generalist predator. We also show that within‐individual and among‐individual variation are not mutually exclusive, but can simultaneously arise and structure intra‐population niche variation.     

Climatic effects on the breeding phenology and reproductive success of an arctic-nesting goose species

Climate warming is pronounced in the Arctic and migratory birds are expected to be among the most affected species. We examined the effects of local and regional climatic variations on the breeding phenology and reproductive success of greater snow geese ( Chen caerulescens atlantica ), a migratory species nesting in the Canadian Arctic. We used a long-term dataset based on the monitoring of 5447 nests and the measurements of 19 234 goslings over 16 years (1989–2004) on Bylot Island. About 50% of variation in the reproductive phenology of individuals was explained by spring climatic factors. High mean temperatures and, to a lesser extent, low snow cover in spring were associated with an increase in nest density and early egg-laying and hatching dates. High temperature in spring and high early summer rainfall were positively related to nesting success. These effects may result from a reduction in egg predation rate when the density of nesting geese is high and when increased water availability allows females to stay close to their nest during incubation recesses. Summer brood loss and production of young at the end of the summer increased when values of the summer Arctic Oscillation (AO) index were either very positive (low temperatures) or very negative (high temperatures), indicating that these components of the breeding success were most influenced by the regional summer climate. Gosling mass and size near fledging were reduced in years with high spring temperatures. This effect is likely due to a reduced availability of high quality food in years with early spring, either due to food depletion resulting from high brood density or a mismatch between hatching date of goslings and the timing of the peak of plant quality. Our analysis suggests that climate warming should advance the reproductive phenology of geese, but that high spring temperatures and extreme values of the summer AO index may decrease their reproductive success up to fledging.     

Combining Site Occupancy,Breeding Population Sizes and Reproductive Success to Calculate Time-Averaged Reproductive Output of Different Habitat Types: An Application to Tricolored Blackbirds

In metapopulations in which habitat patches vary in quality and occupancy it can be complicated to calculate the net time-averaged contribution to reproduction of particular populations. Surprisingly, few indices have been proposed for this purpose. We combined occupancy, abundance, frequency of occurrence, and reproductive success to determine the net value of different sites through time and applied this method to a bird of conservation concern. The Tricolored Blackbird (Agelaius tricolor) has experienced large population declines, is the most colonial songbird in North America, is largely confined to California, and breeds itinerantly in multiple habitat types. It has had chronically low reproductive success in recent years. Although young produced per nest have previously been compared across habitats, no study has simultaneously considered site occupancy and reproductive success. Combining occupancy, abundance, frequency of occurrence, reproductive success and nest failure rate we found that that large colonies in grain fields fail frequently because of nest destruction due to harvest prior to fledging. Consequently, net time-averaged reproductive output is low compared to colonies in non-native Himalayan blackberry or thistles, and native stinging nettles. Cattail marshes have intermediate reproductive output, but their reproductive output might be improved by active management. Harvest of grain-field colonies necessitates either promoting delay of harvest or creating alternative, more secure nesting habitats. Stinging nettle and marsh colonies offer the main potential sources for restoration or native habitat creation. From 2005–2011 breeding site occupancy declined 3x faster than new breeding colonies were formed, indicating a rapid decline in occupancy. Total abundance showed a similar decline. Causes of variation in the value for reproduction of nesting substrates and factors behind continuing population declines merit urgent investigation. The method we employ should be useful in other metapopulation studies for calculating time-averaged reproductive output for different sites.     

Female brood desertion increases with number of available mates in the Rock Sparrow

We studied the reproductive strategy of a Rock Sparrow Petronia petronia population, breeding in nest boxes in the Western Alps (Italy). Over seven years of study (1991–1997) 19% of the females laid second clutches after successfully fledging the first one. Among these, about 50% deserted the first nest when nestlings were 14.3 d old (range=8–19 d), 3.6 d before fledging (range=1–8 d). In all these cases the primary male mate took over all parental duties and successfully reared the young. Inter-clutch time of deserting females was 8.1 d shorter than that of non-deserting double-brooded females. The breeding success of deserting females was significantly greater than that of both single-brooded females and double-brooded females that did not desert their first brood. The fledging success of the second clutches depended on the status of the secondary male: females paired with previously unpaired males had a higher fledging success than those that paired with a polygynous male. The frequency of deserting females varied among years from 0 to 16%, and was significantly and positively correlated with the frequency of males available as mates at the time of desertion. In this study we showed that sequential polyandry with brood desertion is a regularly occurring strategy in the female Rock Sparrow.     

The breeding biology of the White-spectacled Bulbul,Pycnonotus xanthopygos,at the northwestern edge of its distribution range

The White-spectacled Bulbul, Pycnonotus xanthopygos, is an abundant and possibly invasive species in Turkey, where it has gradually expanded its distribution and breeding range in both western and southeastern directions. This study focused on its breeding biology, which is still poorly known. The breeding activity extends from February until September. The preferred nesting areas are mainly gardens and maquis groves, where 24 different nesting tree species have been identified. The clutch size is 3.3 ± 0.8 eggs per pair, nesting success 68%, hatching success 94%, fledgling success 95%, and overall breeding success 89%. While nesting success differs significantly between the years, we found no significant differences in hatching, fledging, and overall breeding success between the years studied. Despite favourable climatic conditions in the Mediterranean region, the species makes only one brood per year in a relatively extended breeding season extending over seven months, and has a relatively a high reproduction rate per nest.     

Long‐term relationship between diet breadth and breeding success in a declining population of Egyptian Vultures Neophron percnopterus

Between 2000 and 2009 we studied the diet and breeding success of Egyptian Vultures Neophron percnopterus in southern Spain. Wild species accounted for 74.9% of prey items (n = 1071) with a predominance of mammals (62.3%), followed by birds (20.8%) and reptiles (13.1%). Spatially, the diet was highly varied and not restricted to carcasses of livestock; wild Rabbits Oryctolagus cuniculus accounted for 54% of the overall remains. The spatial variability may reflect regional and local disparity in the availability of main prey. The temporal relationship between variation in trophic diversity and Vulture nesting productivity (both values showing a long‐term decrease) might suggest a causal link between variation in diet and reproductive output. We hypothesize that high turnover rates could explain productivity variation as a consequence of the recruitment of less experienced individuals to the breeding population. This could in turn generate covariation between diet and reproductive output.     

Nosy neighbours: large broods attract more visitors. A field experiment in the pied flycatcher, <Emphasis Type="Italic">Ficedula hypoleuca</Emphasis>

Life is uncertain. To reduce uncertainty and make adaptive decisions, individuals need to collect information. Individuals often visit the breeding sites of their conspecifics (i.e., “prospect”), likely to assess conspecifics’ reproductive success and to use such information to identify high-quality spots for future breeding. We investigated whether visitation rate by prospectors and success of visited sites are causally linked. We manipulated the reproductive success (enlarged, reduced, and control broods) in a nest-box population of migratory pied flycatchers, Ficedula hypoleuca, in Finland. We measured the visitation rates of prospectors at 87 nest-boxes continuously from manipulation (day 3 after hatching) to fledging. 302 adult pied flycatchers prospected 9194 times on these manipulated nests (at least 78% of detected prospectors were successful breeders). While the number of visitors and visits was not influenced by the relative change in brood size we induced, the resulting absolute brood size predicted the prospecting behaviour: the larger the brood size after manipulation, the more visitors and visits a nest had. The parental provisioning rate at a nest and brood size pre-manipulation did not predict the number of visitors or visits post-manipulation. More visitors, however, inspected early than late nests and broods in good condition. Our study suggests that individuals collect social information when visiting conspecific nests during breeding and provides evidence that large broods attract more visitors than small broods. We discuss the results in light of individual decision-making by animals in their natural environments.     

Within-season divorce in Blue Tits (Cyanistes caeruleus)

Divorce between breeding seasons, i.e. mate change while the old breeding partner is still alive, has been well-studied in several bird species. It can be viewed either as the result of reproductive decisions of individuals to maximize their own fitness or as the outcome of intra-sexual competition for mating opportunities. In contrast, divorce within a breeding season—also referred to as rapid mate switching—has received much less attention. Within-season divorce may occur after sudden changes in environmental conditions, such as those causing the disappearance or devaluation of nesting sites or territories, or when better breeding partners become available. Within-season divorce can be initiated by a member of the pair, or it can be the result of a take-over by an unpaired individual that competes for access to the resource. During a field experiment investigating the effects of limiting nesting sites on reproductive behaviour in Blue Tits, we recorded several cases of within-season divorce. The rate of divorce was not related to the experimental nest-site limitation, and pairs that changed their partner suffered reduced reproductive success compared to faithful pairs. Although there were no differences in the timing of breeding, clutch size or hatching success, pairs with a new partner also suffered a reduced fledging success, which was partly explained by complete brood failures. This study highlights that the pair bond prior to egg laying can be unstable when conditions force individuals to compete for a new partner or nest site and indicates the importance of the correct timing of divorce within the breeding cycle.     

The social system of the Pale Chanting Goshawk Melierax canorus; monogamy v polyandry and delayed dispersal

The social organization and reproductive strategy of the Pale Chanting Goshawk Melierax canorus were investigated in four southern African study areas. Territories supporting adults additional to the breeding pair were observed in two study areas. Within one of these areas, these additional adults were either non-breeders staying in their natal territory but actively excluded from the nesting area during the breeding season or a male cobreeder participating fully in reproductive activities with the breeding pair. Polyandrous trios were, however, observed in only one vegetation type in this area, Broken Veld. The annual number of offspring fledged per group did not differ significantly between vegetation types nor between polyandrous trios and monogamous pairs within Broken Veld. Pale Chanting Goshawks lay predominantly two-egg clutches. In 2 out of 5 years, breeding groups succeeded in laying, hatching and even fledging a second brood of young after successfully fledging their first brood. Double brooding occurred more frequently in Broken Veld and most frequently in polyandrous trios in this vegetation type. To accommodate their relatively long breeding cycle (>115 days) in this temperate study area with its limiting summer breeding period, double-brooding Pale Chanting Goshawks laid the first clutch in midwinter and the second, on average, 24 days after the offspring from the first brood left the nest.     

The reproductive capacity of Monk Parakeets Myiopsitta monachus is higher in their invasive range

Breeding parameters for Monk Parakeets Myiopsitta monachus nesting in Barcelona, Spain, were collected for 651 nests over five breeding seasons. This invasive population has a high reproductive capacity compared with the species in the native range: fledging success was double, the percentage of pairs attempting second broods three times higher and 55% of first-year birds bred compared with almost zero in South America.     

Male brood provisioning rates provide evidence for inter‐age competition for mates in female Cooper's Hawks Accipiter cooperii

Life history theory predicts that individuals should maximize lifetime reproductive success (LRS) by breeding as soon as they reach sexual maturity, yet many species delay breeding, either because there are insufficient available mates or breeding sites, or because delayed breeding yields higher LRS. Accipitriform species, such as Cooper's Hawk Accipiter cooperii, exhibit both delayed breeding and delayed plumage maturation. However, in certain circumstances, first‐year females in non‐definitive plumage do breed and apparently compete with older females for high‐quality breeding territories. We predicted that these young females are at a competitive disadvantage compared with older females and that older females would have both higher reproductive success and be able to acquire higher quality nesting territories. We conducted brood counts and measured prey delivery rates by male Cooper's Hawks in an expanding urban population located in Albuquerque, New Mexico (USA), to assess our prediction. We found that older females had higher reproductive success, fledging 1.6 more offspring than younger females, and that they occupied territories where males provisioned at higher rates of 0.37 more prey items per 2‐h period. Our results showed that older females fared better than first‐year females but it is unclear if this is the result of passive or active competition. Older females initiated nesting 14.3 days sooner than first‐year females and thus may have filled vacant, high‐quality territories before first‐year females began seeking mates. Additionally, first‐year females were never observed persistently to confront older females for breeding territories, but they did actively compete against each other. First‐year females may defer to older females who, in a direct competitive interaction, would be most likely to prevail. Thus, delayed plumage maturation in Cooper's Hawks may serve to focus competition for nesting territories within age classes.     

Predicting reproductive success from hormone concentrations in the common tern (<Emphasis Type="Italic">Sterna hirundo</Emphasis>) while considering food abundance

In birds, reproductive success is mainly a function of skill or environmental conditions, but it can also be linked to hormone concentrations due to their effect on behavior and individual decisions made during reproduction. For example, a high prolactin concentration is required to express parental behaviors such as incubation or guarding and feeding the young. Corticosterone level, on the other hand, is related to energy allocation or stress and foraging or provisioning effort. In this study, we measured individual baseline prolactin and corticosterone between 2006 and 2012 in breeding common terns (Sterna hirundo) using blood-sucking bugs. Reproductive parameters as well as prey abundance on a local and a wider scale were also determined during this period. Baseline prolactin and corticosterone varied significantly between years, as did breeding success. At the individual level, prolactin was positively and corticosterone was negatively linked to herring and sprat abundance. At the population level, we also found a negative link between corticosterone and prey abundance, probably reflecting overall foraging conditions. High prolactin during incubation was mainly predictive of increased hatching success, potentially by supporting more constant incubation and nest-guarding behavior. It was also positively linked to a lesser extent with fledging success, which could indicate a high feeding rate of young. Corticosterone concentration was positively related to high breeding success, which may be due to increased foraging activity and feeding of young. In general, our study shows that baseline prolactin and corticosterone levels during incubation can predict reproductive success, despite the presence of an interval between sampling and hatching or fledging of young.     

Radio‐transmitters do not affect seasonal productivity of female Golden‐winged Warblers

Investigating the potential effects of handling and marking techniques on study animals is important for correct interpretation of research results and to effect progress in data‐collection methods. Few investigators have compared the reproductive output of radio‐tagged and non‐radio‐tagged songbirds, and no one to date has examined the possible effect of radio‐tagging adult songbirds on the survival of their fledglings. In 2011 and 2012, we compared several parameters of reproductive output of two groups of female Golden‐winged Warblers (Vermivora chrysoptera) breeding in Minnesota, including 45 females with radio‐transmitters and 73 females we did not capture, handle, or mark. We found no difference between groups in clutch sizes, hatching success, brood sizes, length of incubation and nestling stages, fledging success, number of fledglings, or survival of fledglings to independence. Thus, radio‐tags had no measurable impact on the productivity of female Golden‐winged Warblers. Our results build upon previous studies where investigators have reported no effects of radio‐tagging on the breeding parameters of songbirds by also demonstrating no effect of radio‐tagging through the post‐fledging period and, therefore, the entire breeding season.     

Double brooding and polygyny in Sedge Warblers Acrocephalus schoenobaenus breeding in north-west England

Sedge Warbler breeding was studied at a site in Greater Manchester, UK, during 1991 and 1993. Birds were individually colour-ringed and, in the 2 years of the study, 18 pairs and 38 nesting attempts were observed. In both years, the majority of pairs (> 90%) attempted a second brood after successfully raising their first. Overall nest survival was 44% and the median reproductive output of individual males was 6.5 fledglings in each year (range 0–10 and 0–15 in 1991 and 1993, respectively). After fledging, one or both parents continued to feed the young for up to 15 days (median 11, range 0–15 days). One male was apparently bigamous in 1991 and in 1993, raising four broods successfully in 1993 and 25 fledglings over the 2 years. These observations suggest that double brooding is the normal strategy for Sedge Warblers in this population and that polygyny may be common, although easily overlooked.     

Brood desertion and polygamous breeding in the Kentish Plover Charadrius alexandrinus

In the Kentish Plover Charadrius alexandrinus one of the adults, typically the female, deserts the brood when the chicks are a few days old. Once parental care is terminated, adults may initiate a second nesting attempt if sufficient time remains within the season. For these nests, individuals pair with different mates from those of the first nesting attempt, thus becoming sequentially polygamous. In a small population of Kentish Plovers in Fuente de Piedra lake (southern Spain), the duration of biparental care of broods was longer than in other localities. It also showed considerable variation between years that was evidently related to Gull-billed Tern Gelochelidon nilotica predation pressure on the chicks. There was year-to-year variation in the number of polygamous matings. Both the duration of the breeding season and nesting success in the first half of the season limited the occurrence of polygamy. Despite females deserting broods earlier than males, the interval between the first and second nesting of polyandrous females and polygynous males was similar. The interval was not affected by the body condition of females after the first nesting attempt, nor by problems related to egg formation ability, but was probably due to the availability of potential mates. More females than males initiated second nests, suggesting that polygamous opportunities were more limited for males than for females. In terms of delayed breeding, reduced survivorship or reduced breeding opportunities in years following polygamous breeding, polygamous individuals did not have greater costs than non-polygamous ones. Females with second nests did not seem to be selective in mate choice, mating with any available male. Mates for second nests may therefore be of lower quality than those for first nests, as judged by male plumage characteristics. Clutch sizes and egg characteristics of polyandrous females were similar in first and second nests. Nest success of second nests was only 40% of that of first ones, with nest desertion accounting for 60% of the losses. As the costs of polygamy are apparently low and as breeding success is very variable among years, polygamous breeding of the long-lived Kentish Plover may be an important breeding strategy with which to increase individual lifetime reproductive success.     

The most common diet results in low reproduction in a generalist seabird

Dietary specialization has been described across a wide range of taxa in the animal kingdom. Fitness consequences are, however, not well documented. We examined the reproductive consequences of different dietary specializations in the herring gull Larus argentatus, an omnivorous seabird, using an extensive dataset which includes breeding and dietary data of 10 successive years. We hypothesized that pairs that focused on prey of higher energetic value would yield higher fledging rates. An alternative hypothesis is that pairs that relied on more resources simultaneously would reproduce better. The novelty of this study is that we used continuous measurements representing dietary composition and degree of specialization rather than restricting our analysis to predefined categories. By relating these two continuous measurements for diet to several proxies for reproductive success, we show clear consequences of dietary choice. Most pairs concentrated on bivalves, a prey type not particularly rich in energy. Pairs feeding on energy‐rich prey (e.g., “domestic refuse and fishery discards”) during chick rearing were found to have a higher reproductive success, supporting the first hypothesis. Pairs that used more resources did not clearly have a higher reproductive success. The majority of the pairs did not switch to energy‐rich prey during chick rearing, despite low breeding outcome. We discuss how trade‐offs between factors such as resource availability, predictability, and the time and energy needed to obtain certain prey species may influence resource selection.     

Evolution of single-chick broods in the Swallow-tailed Gull Creagrus furcatus

Swallow-tailed Gulls lay single-egg clutches, and so raise single-chick broods. As they are pelagic seabirds, this small brood size is expected to relate to proximate food limitation owing to infrequent food deliveries. However, a previous brood doubling experiment detected an 82% increase in fledging success from experimentally doubled broods compared to controls. We repeated the brood doubling experiment, and found that none of 50 enlarged broods produced more than one independent offspring. Control and experimental parents produced fledglings of similar body size, which also had indistinguishable rates of fledging and subsequent survival and reproduction. A variety of parameters estimating survival and breeding costs of reproduction showed no treatment effect. Since two-chick broods yield dramatically higher fledging rates at some times, apparently without excess costs of reproduction, selection on brood size appears to favour a two-chick brood. However, selection may not favour a two-egg clutch if egg production is very costly. Additionally, our estimates of reproductive success do not incorporate the performance of experimental and control offspring as adults, which could differ, since growth of chicks differed slightly by treatment.     

Reproductive success depends on the quality of helpers in the endangered,cooperative El Oro parakeet (Pyrrhura orcesi)

In cooperative species, helping behaviour and reproductive success can be correlated, but understanding this correlation is often impaired by the difficulty to correctly infer causation. While helpers can incur costs by participating in brood care, it is yet unclear if their help depends on their individual quality. We address these questions in the previously unknown cooperative breeding system of the endangered El Oro parakeet (Pyrrhura orcesi). Specifically, we ask (i) whether breeders benefit directly from helpers by an enhanced reproductive success and if so, (ii) whether the amount of this potential benefit is regulated by the quality of contributing group members. Groups consist of a dominant breeding pair accompanied by helpers, but cooperation is not obligate. Microsatellite heterozygosity was used to assess individual quality; its suitability as indicator of quality was reflected in the positive relationship between offspring heterozygosity and recruitment into the population. The reproductive success of breeding pairs depended on helper (genetic) quality and the number of helpers. This relationship occurred on two different levels: clutch size and fledging success, indicating (i) that females profit from high‐quality helpers and probably adjust clutch size accordingly and (ii) that the helpers increase fledging success. Congruently, we found that offspring body condition is positively affected by helper quality, which is most probably explained by the increased feeding rates when helpers are present. We suggest a causal link between cooperation and reproductive success in this frugivorous, endangered parakeet. Further, helper (genetic) quality can be a relevant factor for determining reproductive fitness in cooperative species, particularly in small and bottlenecked populations.