The demography of Yellow Wagtails Motacilla flava on abandoned fields in northern European Russia

Capsule Apparent survival rates of Yellow Wagtails breeding in abandoned fields in Russia are determined by previous breeding success.

Aims To examine apparent survival and its link to previous breeding success in Yellow Wagtails breeding in abandoned fields in the Vologda region, northern European Russia.

Methods We ringed and measured apparent survival of Yellow Wagtails at two abandoned agricultural sites over eight years (2005–2012). We modelled the impact of age, nest stage, and time of season on daily nest survival rates.

Results Predation was the main cause of nest failure. Nest daily survival rate was highest at the beginning of the breeding season. Overall nest survival probability was 0.40?±?0.02. Adult apparent survival after successful breeding was 0.42?±?0.06 and after unsuccessful breeding this was 0.13?±?0.06.

Conclusion Reproductive success can be regarded as the crucial demographic parameter of the local Yellow Wagtail population in northern European Russia. Apparent survival after successful breeding is significantly higher than after unsuccessful breeding, because unsuccessful breeders probably move to new breeding sites the following year. High adult survival may be particularly important to Yellow Wagtail population dynamics in the study region, because second breeding attempts are apparently unusual.     

Nesting success in Redshank Tringa totanus breeding on coastal meadows and the importance of habitat features used as perches by avian predators

Capsule Nest survival rates could not be explained by distance to habitat edges or other features used by predators.

Aims To investigate if predation on Redshank nests was affected by habitat characteristics at a local scale.

Methods We examined survival rates of Redshank nests on coastal meadows on the Baltic island of Gotland, Sweden, over two breeding seasons. We analysed nest survival rates in relation to several habitat characteristics that may benefit predators searching for nests. We examined existing studies concerning predation rates on wader nests in relation to edges and habitat features potentially used by avian predators.

Results We found no significant effects of distance to habitat edge or to nearest potential lookout for avian predators or to shoreline. Abundance of Lapwings Vanellus vanellus, an aggressive species with active nest-defence, did not have any significant effect on nest survival rate, nor did vegetation concealment of nests. Nest survival rates were significantly different between years and lower later in the season.

Conclusions There is only weak support for general effects on wader nest predation rates of proximity to edges and features used by avian predators. Simple mechanical management actions such as removal of trees and bushes on coastal meadows may not directly, and by itself, result in higher reproductive success of waders. Further understanding is needed of the behaviour of predators and the composition of the predator community in different landscapes in order to increase the efficiency of management actions to remove threats to vulnerable species on coastal meadows.     

Nest survival of Woodpigeons (Columba palumbus) in North African forests

Capsule The nesting success of the Woodpigeon Columba palumbus in oak forests in Morocco is affected by human disturbance and nest-site characteristics.

Aims To determine factors affecting the nesting success of Woodpigeons subject to two levels of disturbance in a holm oak Middle Atlas forest.

Methods Woodpigeon nests were monitored in highly disturbed (n?=?30) and less-disturbed sites (n?=?60) over two years (2010–11). Logistic-exposure models were used to estimate daily nest-survival rate and evaluate mechanisms (altitude, temporal factors, nest location, nest-site habitat and degree of human influence) affecting nest survival.

Results Daily nest-survival rate was higher in less-disturbed sites (0.987; 95% CL: 0.980–0.991) than in high-disturbance sites (0.967; 95% CL: 0.949–0.978). Increasing density of trees surrounding nest sites and distance from the nest tree to the closest track increased daily nest-survival rate.

Conclusions Less-disturbed sites confer better nesting conditions than highly disturbed ones. Management measures should control the access of vehicles, especially trucks, in the forest of the Middle Atlas and introduce a grazing regime that will allow the forest regeneration, which would improve Woodpigeon nesting success by increasing the density of holm oak trees.     

Breeding ecology of the Little Bittern Ixobrychus minutus in northeast Algeria

Capsule This is the first study of the reproductive ecology of the Little Bittern Ixobrychus minutus in North Africa. During the study period, clutch size exhibited a sharp drop concurrent with a steady human encroachment and 30% habitat loss.

Aim To investigate the status, nest-site selection, breeding parameters, chick development and chick diet of Little Bittern in Algeria, under deteriorating habitat conditions

Methods Data were collected during surveys of all major Algerian wetlands (2002–08) and monitoring of breeding at Boussedra, a freshwater marsh in northeast Algeria, during the years 2005 and 2008.

Results Egg-laying spanned the period of 10 May to 30 June with one recorded case of a double brood in early July (n = 30). The mean clutch size in 2005 was 6.2?±?0.4 sd eggs per nest (n = 6 clutches), which was significantly higher than that of 2008, which amounted to 4.3?±?0.5 sd eggs per nest (n = 6 clutches), and this decline was associated with increased disturbance and loss of habitat. Overall, clutches suffered from a low rate of predation (17.6%) and displayed a high hatching rate (85.7%).

Conclusion Breeding parameters of the Little Bittern in North Africa are similar to elsewhere in Europe.     

Habitat Selection and Reproductive Success of Lewis's Woodpecker (Melanerpes lewis) at Its Northern Limit

Lewis''s Woodpecker (Melanerpes lewis) has experienced population declines in both Canada and the United States and in 2010 was assigned a national listing of threatened in Canada. We conducted a two-year study (2004–2005) of this species at its northern range limit, the South Okanagan Valley in British Columbia, Canada. Our main objective was to determine whether the habitat features that influenced nest-site selection also predicted nest success, or whether other factors (e.g. cavity dimensions, clutch initiation date or time of season) were more important. Nest tree decay class, density of suitable cavities and total basal area of large trees were the best predictors of nest-site selection, but these factors were unrelated to nesting success. Estimates of demographic parameters (mean ± SE) included daily nest survival rate (0.988±0.003, years combined), nest success (0.52±0.08), clutch size (5.00±0.14 eggs), female fledglings per successful nest (1.31±0.11), and annual productivity (0.68±0.12 female fledglings per nest per year). Although higher nest survival was associated with both early and late initiated clutches, early-initiated clutches allowed birds to gain the highest annual productivity as early clutches were larger. Nests in deep cavities with small entrances experienced lower predation risk especially during the peak period of nest predation. We concluded that nest-site selection can be predicted by a number of easily measured habitat variables, whereas nest success depended on complicated ecological interactions among nest predators, breeding behaviors, and cavity features. Thus, habitat-based conservation strategies should also consider ecological factors that may not be well predicted by habitat.     

Interactive effects of time and vegetation on reproduction of redshanks (Tringa totanus) breeding in Wadden Sea salt marshes

As shown for various species, nesting waders are non-randomly distributed on wetlands and preferentially select riparian nest-sites adjacent to limnic or marine waterbodies. Studying the redshank Tringa totanus, we tested the hypotheses that, in a coastal wader species which conceals its clutch in vegetation, predation and hatching success are affected by vegetation zonation, and that breeding in lower salt marsh areas has negative consequences for reproduction. We further predicted effects of timing of breeding and breeding experience/age of adults potentially reflected by egg biometrics both on nest-site selection and reproduction. Effects of vegetation, space, time and individual quality on hatching success of redshanks were studied in the German part of the Wadden Sea. Dominant plant species, vertical vegetation structure and nest concealment varied significantly between nests. Variation in nest concealment was relatively low: about 90% of clutches were classified as being well concealed. This variation was explainable by vegetation structure but not by vegetation composition at the nest-site, distance to shoreline, and time of clutch initiation. Vertical vegetation structure varied by dominant plant species but not by distance to shoreline and time of clutch initiation. Hatching success of clutches was low (10.6%) due to high predation (daily predation rate: 7.4%). Hatching success and duration of clutch survival were negatively and predation positively related to the date of clutch initiation. Furthermore, negative relationships were found between egg size and predation and duration of survival, respectively. We assume that concealed nests, early breeding and breeding experience diminish predation in salt marsh breeding redshanks. Thus, redshank reproduction appears to be affected by interactive effects of timing of breeding and vegetation facilitating early breeding. In contrast to open-nesting species, breeding in riparian habitats next to waterbodies may be disadvantageous for species breeding concealed in vegetation if these are covered by less structured vegetation.     

The reproductive biology of the Rock Partridge Alectoris graeca saxatilis in the southern French Alps: first evidence of double-nesting behaviour

Capsule: Double-nesting occurs frequently in Rock Partridges Alectoris graeca living in the southern French Alps.

Aims: To investigate reproductive parameters of a Rock Partridge Alectoris graeca population.

Methods: The reproductive behaviour of 62 radio-tagged birds was monitored during the breeding seasons 2012–16, to record breeding phenology, clutch sizes, hatching success, nest survival and parental care.

Results: Double-nesting behaviour in the Rock Partridge was confirmed for the first time. Participation in incubation was similar for males and females (86% versus 70%). Clutch size was larger for nests incubated by males (11.0?±?1.6 eggs) than by females (9.5?±?1.2). Male nest survival rate (0.37) tended to be lower than female nest survival rate (0.62), although the difference was not significant. No significant difference was detected between male and female hatching success. Nest failures were caused by mammals taking the eggs (78%) or predation of the incubating parent (13%). Some circumstantial evidence suggests that occurrence of double-nesting behaviour could depend on previous winter and spring weather influencing the body condition of females. The reproductive biology of the Rock Partridge contrasts in some breeding traits with Red-legged Partridge and natural partridge hybrids, possibly due to climatic differences between habitats.

Conclusion: Confirmation of double-nesting in Rock Partridges indicates that climatic constraints inherent to its mountain environment do not act as an impediment to this behaviour. Variation in weather conditions between years could influence the annual occurrence of double-nesting.     

Light data from geolocation reveal patterns of nest visit frequency and suitable conditions for efficient nest site monitoring in Common Swifts Apus apus

ABSTRACT

Capsule: We developed a protocol for efficient monitoring of potential Common Swift Apus apus nest sites which considers variation in nest visit frequency across the breeding season and in relation to time of day and weather.

Aims: To investigate patterns of nest visit frequency in Common Swifts in order to improve the efficiency and reliability of the monitoring of nest sites threatened by building renovations.

Methods: We derived information on nest attendance from light data recorded by geolocators from ten adult Common Swifts during three breeding seasons (n?=?686 individual sampling days) and analysed how nest visit frequency varied across the breeding season and in relation to time of day and weather.

Results: The mean nest visit frequency was 5.63 visits per bird per day (0.32 visits per hour of daylight). The daily number of visits was highest at the beginning of July during chick-rearing. Moreover, it was positively correlated with temperature and negatively correlated with rainfall and wind speed. Nest visit frequency showed a distinct peak around sunset, while also being relatively high in the morning and around noon.

Conclusion: We recommend monitoring potential Common Swift nest sites in Central Europe between the end of June and mid-July during good weather between 0.50 and 7.75?h after sunrise or between 3.00?h before sunset and sunset, when observation bouts of 0.5–2.0?h provide an encounter probability greater than 90%. Our study shows that repurposing geolocator light data – usually used to study bird migration – for investigating nest attendance in cavity-breeding birds can provide important information for bird conservation.     

Golden Plover Pluvialis apricaria breeding success on a moor managed for shooting Red Grouse Lagopus lagopus

Capsule The best estimate of breeding success was a mean of 0.57 fledglings per pair, which when combined with adult survival rates, successfully explained the observed population trend.

Aims To quantify Golden Plover breeding success on a moor managed for shooting Red Grouse Lagopus lagopus.

Methods An intensive study recorded the fate of individual Golden Plover nests and, using radiotelemetry, chicks. The factors associated with mortality were examined, allowing the construction of a model of breeding success. Adult survival was estimated from return rates of colour-ringed birds.

Results Estimated rates of daily nest survival during laying (0.8636) were significantly lower than during incubation (0.9913). The daily survival rate of chicks less than nine days (0.8868) was significantly lower than for older chicks (0.9792). A population model based on these parameters overestimated the rate of nest losses, but accurately described brood survival and fledging success. Although predation rates were low, poor survival of young chicks through starvation or exposure suggest other factors were limiting breeding success at the study site.

Conclusions Predation rates of Golden Plover nests and chicks can be low on moorlands managed for shooting Red Grouse. However, in the absence of predation, other factors may still reduce chick survival and limit breeding success.     

New is not always better: low breeding success and different occupancy patterns in newly built nests of a long-lived species,the white stork Ciconia ciconia

Capsule An increase in new nest building in a white stork population revealed that they were built further from human settlement and on non-typical structures; such nests had lower breeding success resulting from later breeding.

Aim To determine why some birds build new nests rather than occupy older ones, and how new nests affect breeding performance compared to old nests, in a long-lived bird, the white stork.

Methods We compared new nest construction in 2010 with a long-term data set on white stork in Western Poland from 1974 to 2009. For data from 2010, we analysed nest location and breeding biology in detail.

Results Since 1974, the proportion of new build nests was ca. 1.6%; in 2010 this was 13.2%. Pairs in new nests bred later than pairs in old, and had smaller clutches and lower breeding success. New nests were located further from settlements and tended to be built on different structures. A significantly lower proportion of new nests were re-occupied in subsequent years.

Conclusions Pairs may build new nests to gain experience in nest building, cooperation and foraging for subsequent seasons or because of competitive pressure when the environment is close to carrying capacity. Breeding success can be initially very low.     

Week-end bias at observatories

Capsule Although subject to human disturbance Turtle Doves do nest successfully in these olive and orange orchards.

Aim To investigate the breeding ecology of Turtle Doves in a man‐made agricultural habitat in central Morocco.

Methods Turtle Dove nests were monitored in orange and olive orchards over three years (2006, 2007 and 2008). Nest abundance, nest location, egg‐laying chronology, clutch size, nest survival rates and breeding success were determined and compared between orchard types.

Results The Moroccan population of Turtle Doves start breeding earlier than European populations. Clutch size, nest survival rates and breeding success were similar in orange and olive orchards. Nest location differed between orange and olive trees. Nest densities were 16 nests/ha in olive orchards and 45 nests/ha in orange orchards. Nest success rate averaged 48%. Daily nest survival rates did not vary according to orchard types, year and date. In the two orchards, no nest position variables were significant predictors of nesting success.

Conclusion Although highly frequented by people, fruit orchards seem to be suitable breeding habitats for Turtle Doves in this region.     

The impact of livestock on Lapwing Vanellus vanellus breeding densities and performance on coastal grazing marsh

Capsule Even at very low stocking densities, livestock reduce breeding densities of adult Lapwings and increase the risk of nest loss due to predation.

Aims To assess the effects of livestock on Lapwings breeding on coastal grazing marshes.

Methods Densities of breeding adults, clutch sizes, laying dates, incubation schedules, clutch and chick survival were compared between marshes grazed at low stocking densities (0.2–0.51 livestock units/ha) and marshes where livestock had been excluded. Repeated measurements of sward heights were also made.

Results. Breeding densities in 1995 and 1997, but not 1996, were negatively correlated with the presence of livestock. Though few nests were trampled, livestock disrupted incubation schedules and increased the risk of nest predation. Clutches were smaller on grazed marshes than on ungrazed marshes, while more clutches were also laid later on grazed marshes. Grazed swards remained shorter, and more suitable for nesting, longer than ungrazed swards but clutches laid later in the season were more likely to be predated.

Conclusion The exclusion of livestock from selected areas to increase the nesting success of lapwings is a desirable option on coastal grazing marshes where the rate of grass growth is slow in spring. Grazing regimes are suggested that would maintain relatively short swards, provide refuge to Lapwings from livestock during the peak nesting period, and allow grazers to exploit all of their marshes.     

Artificial nests for Barn Swallows Hirundo rustica: a conservation option for a declining passerine?

Capsule: Barn swallows Hirundo rustica using artificial nest cups had greater reproductive success than those building and using natural nests.

Aims: To quantify reproductive success of Barn Swallows breeding in artificial man-made nests and compare with those using natural nests.

Methods: In 2012–16, the breeding attempts of 231 pairs of Barn Swallows were monitored in artificial and natural nests in an urban habitat in Denmark.

Results: Pairs breeding in old natural nests and artificial nests bred earlier, laid more eggs, and produced more nestlings and fledglings than pairs breeding in new natural nests. Inter-clutch intervals were shorter in Barn Swallow pairs using artificial nests and old natural nests. Nest cup volume and nest volume of newly built nests were significantly larger and more variable than artificial nests excluding an effect of nest size as an explanation for the larger clutches laid in artificial nests. Increased energy and time costs of nest construction may have reduced reproductive success for pairs building natural nests.

Conclusion: Artificial nests were readily accepted by Barn Swallows, they had a similar low predation rate as natural nests, and they proved to be a possible conservation option for this declining passerine breeding in urban habitats as evidenced by their 75% higher reproductive success in artificial nests.     

Testing the importance of individual nest-site selection for a social and group-living vulture

Nest-site selection by species is expected to be adaptive and lead to improved breeding productivity, but in some settings, there exist mismatches between preferred nesting habitat and breeding productivity. We tested the expectation that nest-site selection is adaptive in a sample of 63 nests of a long-lived social species that breeds and forages in groups: the critically endangered white-backed vulture (Gyps africanus). By studying breeding groups in the same area, we controlled for landscape-level effects on habitat selection and investigated how fine-scale nest-site characteristics affect breeding productivity. We developed models to assess how nine characteristics of nest sites selected by breeding vultures compared with 70 random trees and tested associations between these characteristics and breeding productivity. White-backed vultures selected nest sites in taller trees (>7 m), but neither tree height nor any other nest-site characteristics had a clear effect on breeding productivity. Vultures selected nest trees closer to each other than random trees, and the associations between nest density, nearest neighbour distance and breeding success were all positive. These positive associations and the absence of an observable effect between nest-site characteristics and breeding productivity suggest that for this semi-colonial breeder, the social imperative of proximity to conspecifics (i.e., nesting near other vultures and group foraging) may be more important than individual nest-site selection.     

Why do earlier‐arriving migratory birds have better breeding success?

In migratory birds, early arrival on breeding sites is typically associated with greater breeding success, but the mechanisms driving these benefits are rarely known. One mechanism through which greater breeding success among early arrivers can potentially be achieved is the increased time available for replacement clutches following nest loss. However, the contribution of replacement clutches to breeding success will depend on seasonal variation in nest survival rates, and the consequences for juvenile recruitment of hatching at different times in the season. In particular, lower recruitment rates of late‐hatched chicks could offset the benefits to early arrivers of being able to lay replacement clutches, which would reduce the likelihood of replacement clutch opportunities influencing selection on migratory timings. Using a simulation model of time‐constrained capacity for replacement clutches, paramaterized with empirically‐derived estimates from avian migratory systems, we show that greater reproductive success among early‐arriving individuals can arise solely through the greater time capacity for replacement clutches among early arrivers, even when later renesting attempts contribute fewer recruits to the population. However, these relationships vary depending on the seasonal pattern of nest survival. The benefits of early arrival are greatest when nest survival rates are constant or decline seasonally, and early arrival is least beneficial when nest success rates increase over the breeding season, although replacement clutches can mitigate this effect. The time benefits of early arrival facilitating replacement clutches following nest loss may therefore be an important but overlooked source of selection on migratory timings. Empirical measures of seasonal variation in nest survival, renesting, and juvenile recruitment rates are therefore needed in order to identify the costs and benefits associated with individual migration phenology, the selection pressures influencing migratory timings, and the implications for ongoing shifts in migration and breeding phenology.     

Breeding biology of the endangered Dupont’s Lark Chersophilus duponti in two separate Spanish shrub-steppes

Capsule: The Dupont’s Lark Chersophilus duponti in Iberia has relatively high breeding success in both core and fragmented habitats, so population declines are more likely to be the result of low juvenile or adult survival.

Aims: To measure important aspects of the reproductive biology of one of the most endangered and least known larks: the Dupont’s Lark C. duponti.

Methods: We monitored 36 nests in 2 Spanish shrub-steppes, one holding one of the largest European populations (250 pairs) and one composed by fragmented habitat patches holding a smaller population (50 pairs).

Results: The breeding season went from late-March to early July. Overall mean (±sd) clutch size was 3.47?±?0.56, and the number of fledglings per successful nest was 3.0?±?1.15. Mean nestling period was short (8.2 days). Nests showed similar daily survival rate during the incubation period (0.9750?±?0.0110) as during the nestling period (0.9545?±?0.0168), with a mean breeding success of 50%. Predation was the main cause of complete nest failure (83.3% of failed nests in both localities).

Conclusion: Breeding parameters did show no significant variation between populations. Breeding success in both sites was generally higher than recorded in previous studies of this and most other lark species, which suggests that breeding success does not compromise long-term viability of these populations. The decline of the studied populations should be explained by other causes, such as a general decrease in habitat quality, habitat loss or habitat fragmentation.     

Nest location, clutch size and nest success in the Scarlet Ibis Eudocimus ruber

Breeding success and nest-site characteristics were studied during the 1996–1997 breeding season in a colony of Scarlet Ibises Eudocimus ruber in south-eastern Brazil to test the hypothesis that nest-site characteristics and clutch size affect nest success. Two nesting pulses produced young, the earlier being more successful. Predation accounted for most failures during the first pulse, wind destruction during the second. A third pulse with few nests produced no young. Adult Ibises abandoned nests when they lost sight of other incubating birds. Logistic regression analysis indicated that nest success during the first pulse was positively related to clutch size, number of nests in the nest tree and in the nearest tree, and negatively to the distance to the nearest neighbour. During the second pulse there were significant negative associations between success, nest height and distance to the fourth nearest nest, and a positive association between success and nest cover. The results agree with the 'selfish herd' hypothesis, indicating that nest aggregation may increase breeding success, but the nest-site characteristics affecting success can differ over the course of one breeding season.     

Selection through predation, snowfall and microclimate on nest-site preferences in the Water Pipit Anthus spinoletta

Nest-site characteristics can have a strong impact on reproductive success in birds. Nest sites should simultaneously protect from predators, offer shelter and provide a favourable microclimate. We studied the relationship between three agents of natural selection (predators [i.e. Adders and birds/mammals], snowfall and microclimate), nest-site characteristics and reproductive success to determine whether these influenced preference for specific nest-site characteristics in the Water Pipit Anthus spinoletta . Pooled over all nests, the relative importance as agents of natural selection decreased from mammalian/avian predation (15% of all nests) through Adder predation (12%) to snowfall (7%), but there were clear differences in space and time. Predation by Adders selected for nest sites surrounded by few medium-sized shrubs. Selection by mammalian and avian predators favoured no specific nest-site characteristics. Protection from snowfall was best in nests surrounded by relatively few medium-sized shrubs. Microclimate had a strong influence on nestling survival and duration of nestling period. In nests on ENE-facing slopes, where maximum temperatures were reached in the morning, nestling survival was higher than on WSW-facing slopes, where temperature maxima occurred in the afternoon. Our results indicate that weak, but significant, directional selection is acting on preference for certain nest-site characteristics through effects on survival and development of nestlings. As predation and snowfall are unpredictable, the evolution of an optimal nest placement strategy is unlikely on a small scale. On a larger scale, however, choice of one breeding area over another may be favoured because of predictable differences between locations in terms of survival and nestling development.     

DO NEST SITE CHARACTERISTICS AFFECT THE BREEDING SUCCESS OF RED BISHOPS EUPLECTES ORIX?

Summary

Ferguson, J.W.H. 1994. Do nest site characteristics affect the breeding success of Red Bishops Euplectes orix? Ostrich 65:274-280.

A colony of Red Bishop birds Euplectes orix in a reed bed near Pretoria was studied during two breeding seasons. Breeding activity was greater during the second year, following increased rainfall. Predation was the most important source of mortality with 25–30% of the nests yielding fledglings. Between 7 and 11% of the nests were parasitised by Diederik Cuckoos. Reed height and distance from the edge of the reed bed were strongly correlated with nest density. From a temporal perspective breeding success was highest in late January following the peak in breeding activity. These observations suggest that overall breeding success is higher when nest densities are low. The mean fledging rate per nest was lower in areas of high nest density. This was, however, a statistical artifact. Multivariate analyses could not identify any environmental factors as predictors of breeding success within a colony.     

Drivers of low breeding success in the Lesser Spotted Woodpecker Dendrocopos minor in England: testing hypotheses for the decline

Capsule The breeding success of Lesser Spotted Woodpeckers Dendrocopos minor is now lower in England than previously reported and also lower than found in studies elsewhere in Europe.

Aims To quantify the breeding success and identify the causes of nest failure. To test the hypotheses that breeding success is related to aspects of food limitation and parental care, and inclement weather during the nesting period, or to interactions with Great Spotted Woodpeckers.

Methods Nests were monitored in three regions of England, recording survival and causes of failure. We measured aspects of food limitation and parental care, rainfall and Great Spotted Woodpecker interactions at nests, to explore whether there was any evidence that these factors were related to breeding success. We compared results to other studies from the UK and continental Europe.

Results Nest survival was 52%. The average number of chicks produced from successful nests was 2.8. Chick-stage daily nest survival was positively related to provisioning rates, indicating that food supply may be limiting. The most common cause of nest failure was presumed starvation of chicks after the disappearance of an adult. Some females ceased visiting nests, leaving provisioning solely to the male. This behaviour has been reported elsewhere in Europe, but in the present study males were unable to compensate fully by increasing their provisioning rates, leading to poor nest survival. Provisioning rates and chick-stage daily nest survival were negatively associated with rainfall. Nest predation by Great Spotted Woodpeckers occurred but was a less frequent cause of failure. Aggressive interactions were recorded between the two woodpecker species but these were unrelated to breeding parameters.

Conclusions Low breeding success is most probably related to food shortages in the breeding period. Simple population modelling using parameters from the present study and from published work shows that if the low productivity that we have observed is replicated throughout Britain, it would be sufficient to account for the observed population decline. However, the possibility that survival rates are also low cannot be ruled out.