Herring and Sprat Abundance Indices Predict Chick Growth and Reproductive Performance of Common Terns Breeding in the Wadden Sea

Herring (Clupea harengus) and sprat (Sprattus sprattus) are the key prey resources of common terns (Sterna hirundo) breeding in the Wadden Sea. Breeding success of the terns has been below average since 2002, coinciding with exceptionally low herring recruitment and sprat abundance. Time series of herring and sprat abundance in the North Sea and in the Wadden Sea were analyzed to explain long-term breeding success and chick development at two common tern breeding colonies. North Sea herring recruitment and sprat abundance in the Wadden Sea explained the largest part of common tern breeding success, both as single variables and in a multiple regression approach. Breeding success showed stronger correlations with herring recruitment indices derived from the North Sea region compared to the Wadden Sea. Also, herring and sprat abundance data explained more variability in breeding success than of more directly responding measures such as growth rate and maximum weight of chicks. Despite spatial and temporal incoherences between fish surveys and the common tern breeding season, breeding success of common terns reflected the abundance of their key prey fish beyond their foraging range and breeding season. We argue that the ecological connectivity between large- and small-scale herring abundance and the responsiveness of common tern breeding success is strong enough to establish a fish–seabird indicator system to be potentially valuable in monitoring and conservation.     

Modelling the occurrence and abundance of a colonial species,the arctic tern Sterna paradisaea in the archipelago of SW Finland

Knowledge of the habitat requirements and suitable breeding areas of sea birds is crucial for their management and conservation. However, there are still few studies that have modelled the breeding distribution and abundance of colonial sea birds. In this study, we created predictive distribution models for a colonial species, the arctic tern Sterna paradisaea, using 14 environmental variables calculated for 525 islands in the Archipelago Sea in SW Finland. We modelled the occurrence (presence‐absence) using generalised additive models (GAMs) and abundance (pair numbers/colony size) using hurdle models fitted with GAM. We tested for spatial autocorrelation in model residuals and evaluated the models on independent data. Critical factors influencing the occurrence of the arctic tern were the proportions of boulder or gravel and forest of island area, as well as island maximum elevation and area, such that the species seemed to prefer large and low islands with sparse vegetation. Abundance was influenced by the proportions of boulder or gravel and bare rock of island area, as well as exposure and island area. To some extent, different factors influenced the occurrence and the abundance. The evaluation results of the models were good, with an AUC value of 0.91 for the most accurate presence‐absence model and a Pearson's correlation coefficient of 0.60 for the most accurate hurdle model. The predictive ability of the models increased when we removed islands with single or few breeding pairs from the data set. Although the hurdle models did not produce accurate pair number estimates, they indicated which islands are suitable for larger colonies. Abundance is a crucial factor for colonial species. This modelling technique can therefore be of great value for the conservation and management of the arctic tern and similar colonial species.     

The importance of observer effort on the accuracy of breeding success estimates in the Common Guillemot Uria aalge

ABSTRACT

Capsule: A data-thinning approach was used to assess the effects of reducing the frequency of nest-checks on estimates of breeding success of Common Guillemots Uria aalge. Inter-year and inter-colony differences in fledging age and their implications for setting a minimum age after which a chick could be assumed to have fledged were evaluated.

Aims: To assess the consequences of reducing sampling frequency on the estimation of breeding success, and on the robustness of the assumption that breeding has been successful if a chick survives to 15 days old.

Methods: Breeding success, ages at fledging and loss of chicks were estimated from daily checks at two Scottish colonies over a six-year period. Data-thinning was used to assess the consequences of reducing checks from daily to every two or three days. Breeding success was recalculated assuming that all chicks surviving to 15 days fledged.

Results: Reducing the frequency of checks from daily to every second or third day resulted in a small but statistically significant increase in the estimate of success. Between 20% and 25% of chick losses occurred when chicks were ≥15 days old. Assuming that these chicks had fledged resulted in significant increases in breeding success.

Conclusion: Assumptions about whether or not a chick fledged had a greater impact on the estimate of breeding success than reducing the frequency of nest-checks from daily to every second or third day. There was no threshold age after which a chick could be assumed to have fledged. These findings are relevant to other monitoring schemes where there is pressure to reduce input. Sampling methods used in monitoring schemes need to be clearly stated and changes in protocols documented so that sampling effects can be incorporated into future analyses.     

Factors affecting breeding distribution of Storm-petrels Hydrobates pelagicus in Orkney and Shetland

Capsule The main factors are past and present human activities, especially the introduction of rats to islands.

Aims To assess factors that influence breeding distribution and abundance of Storm-petrel.

Methods We used a database for 142 islands in Shetland and Orkney. Breeding status of Storm-petrel was related to data for each island on introduced and indigenous predators, other human-related features, and aspects of island geography.

Results Although 92% of the total land area of the archipelagos comprised islands with rats present, Storm-petrel colonies were almost totally restricted to rat-free islands. They also occurred more frequently on islands with cliffs, far from neighbouring islands with humans, and on islands with a low rate of human visits. Colony size was smaller on the smallest occupied islands. Breeding numbers of Great Skuas Stercorarius skua, Great Black-backed Gulls Larus marinus, and Storm-petrels all correlated, as each increased with island size.

Conclusions The presence or absence of rats is the single most important influence on Storm-petrel breeding distribution in Orkney and Shetland. However, geographical and human-related effects, such as the presence of cliffs or the occurrence of human visits, also appear to influence the distribution of Storm-petrels, whereas avian predators appear to have had little effect until now.     

Nestbox orientation: a species-specific influence on occupation and breeding success in woodland passerines

Capsule Nestbox orientation has species-specific influences on nestbox occupation and breeding success for woodland passerines.

Aims To determine if nestbox orientation had an influence upon nestbox selection or breeding success for three co-occurring woodland passerines.

Methods We analysed 15 consecutive years of breeding data (1990–2004) from 295 nestboxes in the UK using circular statistical analyses to examine the influence of orientation upon nestbox occupation and breeding success for three species, Blue Tit Cyanistes caeruleus, Great Tit Parus major and Pied Flycatcher Ficedula hypoleuca.

Results The three species used nestboxes of all orientations during the 15-year period. The frequency of nestbox occupation by Great Tits correlated with orientation (the mean number of nests in boxes oriented south-southwest was lower than the mean number of nests in boxes facing other directions). There was no such relationship for Blue Tits or Pied Flycatchers. Nestbox orientation influenced the breeding success of Pied Flycatcher (the mean number of young to fledge from boxes oriented south-southwest was lower than from boxes facing other directions). There was no such relationship for Blue or Great Tits.

Conclusion Nestbox orientation can be an important influence on occupation and breeding success, but this differed between species. Intriguingly, although the directionality reduced nestbox occupation (Great Tit) and breeding success (Pied Flycatcher) was the same (south-southwest), there was a disparity in the influence of orientation for Great Tit (orientation influenced the frequency of occupation but not success) and Pied Flycatcher (orientation did not influence occupation but did affect success). We discuss these disparities, considering the possible influences of mating strategy, breeding phenology, nestbox microclimate and offspring quality.     

The breeding population of Peregrine Falcon Falco peregrinus in the United Kingdom,Isle of Man and Channel Islands in 2014

Capsule: Overall numbers of Peregrines are higher than previously recorded, due primarily to the growth of lowland populations, despite declines in many upland areas.

Aims: To estimate the breeding population of Peregrines, and changes in this population over time, as well as to relate variation in trends and observed breeding success to variation in region, breeding site type and other ecological influences.

Methods: Two types of survey methods were used. Site-based visits focussed on locations where Peregrines were known to have bred previously, while area-based searches aimed to find all Peregrines breeding in randomly chosen 5?km?×?5?km squares. Findings from each method were used to estimate population size in different parts of the survey area. Breeding outcome and site type were assessed for most of the pairs found during either survey.

Results: In 2014, the breeding population of Peregrines in the UK, Isle of Man and Channel Islands was estimated at 1769 pairs. This is 22% larger than the population estimate from the previous survey in 2002. Most of this increase is accounted for by increases in lowland England. Populations in some upland areas have declined.

Conclusions: The gap between the fortunes of lowland and upland Peregrines has continued to grow, along with the overall UK population. Likely reasons for the continued success of Peregrines in the lowlands include increasing uptake of breeding sites on human structures, abundant availability of prey in many lowland situations and, in many areas, a relative lack of conflict with humans. Factors likely to be limiting upland Peregrine populations vary between different regions, and include ongoing illegal killing and deliberate disturbance, and food supply.     

Climate fluctuation effects on breeding of blue penguins (Eudyptula minor)

Abstract

El Niño and La Niña climate perturbations alter sea currents and food availability for seabirds in many areas of the world. This changes their breeding success and mortality. Blue penguin (Eudyptula minor) breeding success is dependent upon whether one or two clutches per season are laid, and the hatching and fledging success of these clutches. This study uses six years of data from five blue penguin breeding colonies, three from Taiaroa Head, Otago Peninsula and two from Oamaru, to examine whether annual variation in breeding success correlates with El Niño/La Niña perturbations. When La Niña conditions prevailed, penguins started breeding later, and there was a lower proportion of double breeders than in El Niño and normal years. The probability of a newly hatched chick surviving to fledging was also dependent on whether large‐scale climatic conditions prevailed, whereas hatching success and overall breeding success (number of fledged chicks per breeding pair) showed no correlation with climate perturbations.     

Home ranges of introduced rats on Christmas Island: A pilot study

Understanding the spatial ecology of invasive rats (Rattus spp.) is necessary to inform management actions to reduce their impact on native flora and fauna. This study investigates home range sizes of exotic rats around seabird colonies and urban areas on Christmas Island, where rat predation is suspected to be adversely affecting fledgling success among local seabirds. It was hypothesised that rat home range sizes would be smaller in urban areas owing to more consistent food availability. Home ranges of male rats were significantly larger compared with their female counterparts, with male rats maintaining larger home ranges in urban areas compared with seabird colonies. Conversely, female rats had smaller home ranges in urban areas compared with seabird colonies. Our findings suggest a possible correlation between the spatial distribution of food resources and home range size. Additionally, the spatial distribution of breeding females across the landscape had a significant influence on the home ranges of male rats. These findings have important implications for proposed efforts to manage rat populations on Christmas Island, while also providing valuable information regarding the ecology of invasive rats on tropical islands.     

Changes in sooty shearwater (Puffinus griseus) abundance and harvesting on the Rakiura Titi Islands

Abstract

We estimated the change in abundance of sooty shearwater (titi, Puffinus griseus) at six Rakiura Titi Islands, New Zealand, by comparing historical and recent surveys of the density of entrances to breeding burrows. We found evidence that entrance density between 1994 and 2006 was lower than it was between 1961 and 1976. Our overall estimate of the annual rate of change in burrow entrance density is ‐1.0% (95% CI ‐2.3 to ‐ 0.1%). Declines have been slower on four islands where Rakiura Maori maintain a traditional harvest of sooty shearwater chicks ("muttonbirding") compared with three unharvested islands. Density‐dependent population processes may explain this difference: rates of decline have been faster in areas of relatively high initial entrance density, and historically the harvested islands have had lower initial density. There was a strong, apparently linear, relationship between entrance density and chick density on breeding colonies, so changes in entrance density probably do indicate a real population decline. The western side of Taukihepa, the largestof the Titi Islands, first became accessible for muttonbirding with the advent of helicopters in the 1970s, but it is unknown whether this has caused an increase in the number of sooty shearwaters harvested by Rakiura Maori.     

Population trends and reproductive success of the European shag Phalacrocorax aristotelis on the Iberian Peninsula following the Prestige oil spill

In 2003, immediately following the Prestige oil spill in Galicia, Spain, we studied the population trends and reproductive performance of European shags (Phalacrocorax aristotelis) at oiled and unoiled colonies. This bird is an important member of the nearshore marine community, breeding in the area affected by the Prestige oil spill. The European shag feeds around the breeding colonies throughout the year, making it a useful indicator of environmental change. Before the oil spill, population trends were similar between oiled and unoiled colonies. Nevertheless, colonies located within the path of the oil suffered greater declines (ca. 10%) compared with pre-spill trends and with population trends at unoiled colonies. In 2003, the breeding success was 50% lower in oiled colonies compared with unoiled colonies. The data available from pre-spill years suggest that the annual reproductive success did not differ among colonies before the impact. European shags breeding at colonies affected by oil showed a negative initial impact from the Prestige oil spill. The reduction in reproductive success at oiled colonies may be due to sub-lethal effects of oil exposure or low food availability after the oil spill.     

Effect of nest-site microclimatic conditions on nesting success in the Lesser Kestrel Falco naumanni

Capsule: Microclimatic conditions in the nests of the Lesser Kestrel Falco naumanni, particularly the percentage of time of extremely low humidity, affect breeding success.

Aim: To study the effect of within-nest temperature and humidity on nest productivity, and the correlation between nest productivity and the order of dates on which nests were occupied by the parents. To compare microclimatic conditions in the nest, breeding success and order of occupation between nests under tile roofs and in artificial nest boxes.

Methods: Three different Lesser Kestrel colonies were monitored in Israel – one rural, one urban and one in an open country habitat. Data loggers were placed in 39 nests for the entire breeding period to measure temperature and humidity. The number of fledglings was recorded for each nest, as well as the date of occupation.

Results: Full microclimatic data from 35 nests suggest that percentage of time of extremely low humidity is the major predictor of nest productivity – low humidity is negatively correlated with nest productivity. Sites of more successful nests were occupied earlier. Considering only successful nests, the urban colony had the lowest breeding success of the three colonies. There was no significant difference in mean productivity between nests in roofs and nest boxes, but nests in roofs were occupied earlier.

Conclusion: Nest microclimate, particularly low humidity, affects nesting success in addition to colony location.     

Breeding performance and timing of breeding of inland and coastal breeding Cormorants Phalacrocorax carbo in England and Wales

Capsule Breeding performance was higher at inland colonies than on the coast due to higher nestling survival during late development.

Aims To compare breeding performance of inland and coastal breeding Cormorants in England and Wales and to provide breeding performance data for Cormorants for future demographic studies.

Methods Breeding performance and timing of breeding was monitored at six inland and four coastal colonies during 1997 and 1998. We compare clutch and brood size and egg and nestling survival.

Results Breeding performance was higher at inland colonies because of higher nestling survival during the later stages of nestling development, from 15–56 days.

Conclusions Greater and earlier food availability at inland colonies, resulting in earlier and more protracted breeding is the most probable explanation for differences in breeding performance. A more protracted breeding season would reduce competition for food and enhance breeding performance, which may be especially apparent during late chick development when energy demands are greatest.     

Reproductive measures and chick provisioning of Cory's Shearwater Calonectris diomedea borealis in the Azores

Capsule Timing of breeding influenced wing-length at fledging, and egg size may be an indicator of fledging weight and the amount of food received by chicks.

Aims To investigate chick growth, temporal patterns of chick food provisioning and the importance of indices of parental condition or quality, egg size and hatching date, to predict nestling body mass and wing-length at fledging, and compare breeding and chick feeding characteristics between colonies in the northeast Atlantic.

Methods A survey of Cory's Shearwater nests was carried out at Vila islet. A sample of 52 chicks, ringed and weighed at hatching, was selected to study chick growth and food provisioning.

Results Hatching success (51%) was much lower than fledging success (87%). Both hatching date and egg size contributed to explain wing-length at fledging, but hatching date, which was negatively correlated with wing-length at fledging, had the most important contribution (22%). There was some indication that egg size may explain variation in fledging weight and the amount of food received by chicks. Food delivery and feeding frequency of chicks varied throughout the chick development stage and three phases were distinguished: (1) 0–29 days, the highest feeding frequency values and a linear increase in food delivery; (2) 30–69 days, an oscillation in food delivery and medium feeding frequencies; (3) 70–90+ days, a sharp decrease in both food delivery and feeding frequency.

Conclusion Variation in food availability did not seem sufficient to override the overall importance of indices of parental quality in determining reproductive measures and chick provisioning. Breeding and feeding characteristics were similar between colonies in the northeast Atlantic, with variability in chick provisioning higher further south.     

SELECTION OF COLONY SITES AND NEST SITES BY COMMON TERNS STERNA HIRUNDO IN OCEAN COUNTY, NEW JERSEY

Common Terns nested on 34 of 259 saltmarsh islands along 46 miles of Ocean County coastline, New Jersey. They nested on low islands of Spartina with less than 12%, windrow ranging in size from 0.6 to 108 acres. All islands selected by terns faced at least two miles of open water from at least one direction. Selection of islands seemed a response to vegetation, size, distance to the nearest island, distance to the nearest shore and exposure to open water. Of the 225 islands without nesting terns, only three fulfilled appropriate criteria. Eighty per cent of nests were situated on windrow. Terns laid larger clutches on windrow compated to those nesting in Spartina. Nearest neighbour distance on islands with low predation varied from 85 to 485 cm and was a function of space, vegetation, and the size of the island. Terns nested closer together on windrow compared with Spartina. Considering all colonies, the mean nearest neighbour distance negatively correlated with the number of nests. Skimmers, Oystercatchers, Laughing Gulls, and Herring Gulls nested in some of the tern colonies. All tern colonies on islands also occupied by nesting Herring Gulls suffered over 10% egg-loss by predation. Proportionally more nests were preyed upon in windrow than in Spartina. Clutch sizes in nests in windrow on islands with predation were significantly lower than those in windrow on islands without predation, even though clutch sizes in Spartina were similar on islands with and without predation. Flooding by exceptional high tides destroyed significantly more nests in Spartina than in windrow. The nests that survived in Spartina were built deeper, and their rim heights were higher than nests destroyed by tides. Choice of colony and nest sites is discussed in terms of the balance of two selection pressures: predation and tidal flooding. Tern nests on windrow are more exposed to predators, but safer from flooding, whereas those nests in Spartina are more susceptible to flooding and less vulnerable to predators. The recent invasion of Herring Gulls into Ocean County has significantly increased the rate of nest predation among affected colonies of Common Terns. It is possible that in future years the terns' behaviour may be modified in response to this new selective pressure.     

Factors influencing Hen Harrier Circus cyaneus territory site selection and breeding success

ABSTRACT

Capsule: Our findings regarding Hen Harrier Circus cyaneus territory site selection and breeding success in Ireland offer an opportunity for the development of initiatives and conservation actions aimed at enhancing the suitability of upland areas for breeding Hen Harriers and ensuring the long-term persistence of the species.

Aims: To investigate landscape-scale associations between habitat composition and Hen Harrier territory site selection, and to explore the influence of habitat and climate on breeding success.

Methods: We used multi-model inference from generalized linear models and Euclidean distance analyses to explore the influence of habitat, topographic, anthropogenic and climatic factors on Hen Harrier territory selection and breeding success in Ireland, based on data from national breeding surveys in 2010 and 2015.

Results: Hen Harrier territories were associated with heath/shrub, bog and pre-thicket coniferous forests. Comparisons between territories and randomly generated pseudo-absences (upland and lowland) showed that breeding pairs preferentially select for these habitats. Breeding success was negatively influenced by rainfall early in the breeding season and by climatic instability, and was positively influenced by the presence of heath/shrub and bog.

Conclusions: The results suggest that Hen Harrier breeding success is compromised by the synergistic effects of climate, landscape composition and management. Effective conservation of Hen Harriers in Ireland will therefore rely on landscape-scale initiatives.     

Low breeding success of the little egret (Egretta garzetta) near residential areas and in colonies exposed to gales: a comparison of colony in Sichuan,Southwest China,with literature

The breeding biology of the little egret (Egretta garzetta) was studied in 20 nests within the mixed-species breeding colonies at Nanchong, Sichuan, Southwest China, in 2006. By measuring a set of physical characteristics of vegetation at the nests and at a set of 20 randomly chosen sites we showed that birds preferentially used taller trees in areas with fewer shrubs of higher species diversity. Nests at lower locations in trees had marginally lower hatching success due to their destruction by humans; this destruction contributed marginally significantly to lowering of the total nesting success in all studied nests. Although gale winds also had a negative effect on breeding success, the anthropogenic influences were a greater factor in reproductive failure. We found similar effects in our review of literature on breeding success of the little egret from various geographical areas. Our results may be of use by conservation organizations in their actions to protect colonies of the little egret.     

Estimating regional population size and annual harvest intensity of the sooty shearwater in New Zealand

Abstract

Recent comprehensive survey data from multiple New Zealand offshore islands were combined with demographic population models to produce the first formal estimate of the total population of sooty shearwaters within New Zealand territory. We estimated the total population over 1994–2005 to be 21.3 (19.0–23.6) million individual birds in the New Zealand region. This population consisted of 12.8 (12.0–13.6) million adults, 2.8 (2.5–3.1) million chicks, and 4.4 (4.2–4.7) million breeding pairs. Breeding sooty shearwaters were concentrated primarily around the southern islands of New Zealand, with 53% breeding in the Titi Islands surrounding Rakiura (Stewart Island). Rakiura Maori muttonbirders were estimated to harvest 360 000 (320 000–400 000) sooty shearwaters per year, equivalent to 18% of the chicks produced in the harvested areas and 13% of chicks in the New Zealand region. Overall, 11% of the chicks within the Titi Islands live on unharvested ground. Systematic and widespread surveys of breeding colonies in South America are needed before a reliable global sooty shearwater population estimate can be calculated.     

Behavioural plasticity in nest-site selection of a colonial seabird in response to an invasive carnivore

Non-native invasive species are one of the most serious threats to biodiversity and are considered the leading cause of extinction of several bird taxa, including seabirds. Introduced American mink (Neovison vison) have caused devastating effects on island populations of several colonial seabird species. In this study, we investigated the direct and indirect effects of mink on population dynamics and reproductive success of European shags (Phalacrocorax aristotelis) breeding at Illas Cíes, one of the most important colonies of the species in Southern Europe. A severe episode of mink predation on adult shags occurred in the year when mink arrived in the breeding colonies, though the number of shags killed dropped abruptly in subsequent years. We found that, after the arrival of mink, shags moved to nest-sites that afforded greater protection from carnivores. This shift caused a substantial reduction in mortality by predation, but probably entailed a cost in terms of their reproductive success because sites with lower levels of predation risk showed a higher risk of egg loss by nest flooding due to poorer drainage. Our study highlights that behavioural plasticity may allow shags to cope with invasive predators.     

Nest-site selection and characteristics in a mixed-species colony of Avocets Recurvirostra avosetta and Black-winged Stilts Himantopus himantopus

Capsule Causes of breeding failure determined optimal nest location, Black-winged Stilts being better adapted than Avocets to nest close to water.

Aims To compare nest size and location between the two species and to test whether there were inter-species differences that might suggest specific nesting adaptations, and to investigate possible correlates between hatching success and nest location.

Methods Nest-site selection, characteristics and hatching success were studied during 1989 in a large mixed-species colony located in southern Spain.

Results Black-winged Stilt nests differed in composition and size depending on distance to water (nests touching the water were larger and included mud), but Avocet nests did not. Avocets nests were more aggregated and central within the colony than those of Black-winged Stilts. Causes of breeding failure were flooding and rat predation. Successful or unsuccessful Black-winged Stilt nests did not differ significantly in characteristics or location. Successful Avocet nests were further from water than flooded nests and further from dry land than depredated ones. Distance to the centre of the colony did not affect hatching success.

Conclusion Black-winged Stilts were better adapted than Avocets to nest near water, due to their greater plasticity in nest-building behaviour. Causes of breeding failure determined optimal nest location.     

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.