Intra-population variation in the diet of an avian top predator: generalist and specialist foraging in Great Black-backed Gulls Larus marinus

ABSTRACT

Capsule: Great Black-backed Gulls Larus marinus breeding on Skokholm, UK, fed predominantly on seabirds, rabbits, refuse, and marine prey, with the majority of pairs being dietary generalists, but with some specialist pairs.

Aims: To understand the significance of Great Black-backed Gulls as top predators on a small offshore island with internationally important numbers of breeding seabirds (Skokholm, UK) by quantifying their diet and to determine how this varies within the breeding season, to test for pair-level dietary specialization and to examine the consequences of dietary differences for reproductive performance.

Methods: Regurgitated pellets were collected and analysed from 26 breeding pairs on Skokholm during 2017 and related to breeding success.

Results: Analysis of 1035 pellets revealed that, overall, Great Black-backed Gulls fed on seabirds (48% – mostly Manx Shearwaters Puffinus puffinus), mammals (38% – mostly European Rabbits Oryctolagus cuniculus), anthropogenic waste (7%), and marine prey (7%). Diet varied among pairs with 18 (73%) generalist pairs and 7 (27%) specialist pairs (of which, 5 were bird specialists and 2 were mammal specialists). Diet also varied seasonally, but pair-level dietary diversity was repeatable through the breeding season. Dietary diversity did not covary with breeding success.

Conclusion: Great Black-backed Gulls are top predators on Skokholm. Variation in diet among pairs emphasizes that not all individuals contribute equally in terms of predation. Understanding the incidence of this variation has important ecological implications, particularly where apex predators may exert a strong top-down influence.     

Breeding biology of the Little Egret Egretta garzetta on the southern coast of the Bay of Biscay

ABSTRACT

Capsule: Annual reproductive parameters of Little Egrets Egretta garzetta in an Atlantic coastal colony showed strong variation in a 20-year study, mainly due to extreme events.

Aims: To describe the breeding biology of Little Egrets in the Bay of Biscay and compare it with that of the Mediterranean basin. Also, to explore relationships between breeding parameters and colony size with some climatic indicators.

Methods: Phenology, number of nests, clutch size, number of hatchlings, brood size, and hatching and breeding success were recorded over 20 years.

Results: Median laying date of 214 nests was 1 May (range: 1 April–25 June) and 54% of the clutches were laid in the second half of April and the first half of May. Over the 20-year study period mean clutch size of 270 nests was 4.0, mean number of hatchlings was 3.1, and mean brood size was 2.3. Hatching success ranged from 46.1% to 100% and breeding success from zero to 100%. Number of nests was negatively associated with clutch and brood size. The highest clutch size and lowest brood size were recorded at the beginning of the season. Significant relationships were found between the number of hatchlings and the rainfall during the pre-breeding season, and between brood size and the summer rainfall.

Conclusions: Reproductive parameters showed significant variation over the study period, which highlights the importance of using long-term data sets. Breeding occurred one month later than in natural colonies of the Mediterranean basin. Negative relationships between the number of nests and clutch and brood size suggest some degree of density-dependent effects. Ranges of clutch size, number of hatchlings, and brood size were within those reported in Mediterranean populations. The effect of rainfall on reproductive parameters was weak. Extreme weather and predation events caused low rates of hatching and breeding success that affected the growth of the colony.     

THE INFLUENCE OF THE ENVIRONMENT ON BREEDING SUCCESS OF A SUBURBAN POPULATION OF CRESTED BARBETS TRACHYPHONUS VAILLANTII

Van Zyl, A.J. 1994. The influence of the environment on the breeding success of a suburban population of Crested Barbets Trachyphonus vaillantii. Ostrich 65: 291–296.

I studied the breeding biology of the Crested Barbet Trachyphonus vaillantii in Colbyn, a suburb east of Pretoria, South Africa, for nine breeding seasons from 1981 to 1989 to examine patterns in annual breeding success, breeding attempt success in multiple broods, and rainfall. The modal incubation period was 14 days and the nestling period ranged from 28 to 31 days. Average clutch size for all the years was 3,3 eggs/clutch and there was no significant difference in clutch size or number of young fledged/nest between years. On average, Crested Barbet pairs made 2,4 breeding attempts/season. There was no difference in clutch size or breeding success between the breeding attempts. Crested Barbets nesting in natural nests laid on average larger clutches than those in artificial nestboxes, but had non-signficantly lower breeding success. Failure to raise Crested Barbet chicks was attributed to parasitism by Lesser Honeyguides Indicator minor, bee swarms occupying nestboxes, and flooding of natural nests. Breeding performance was not correlated with rainfall or adult body size. The suburban environment may be less variable than a natural environment, resulting in a stable breeding Crested Barbet population.     

Against the trend: increasing numbers of breeding Northern Lapwings Vanellus vanellus and Black-tailed Godwits Limosa limosa limosa on a German Wadden Sea island

Capsule The increase in population sizes over the last 30 years cannot be explained by reproductive success.

Aims To establish whether the positive population trends are due to increasing and self-sustaining populations or to immigration.

Methods We studied the population development of breeding lapwings from 1971 until 2005 and of godwits from 1977 until 2005 on Wangerooge, a German Wadden Sea island. Both species increased significantly during the last three decades. For each species we used a logistic growth model to derive the reproductive output required to explain the past population development without assuming immigration. We compare the values derived by this model with empirical findings of reproductive output of the respective populations.

Results For neither lapwings nor godwits can empirical reproductive success explain the observed population development.

Conclusion Our results imply that the increase in breeding pairs of Northern Lapwing and Black-tailed Godwit on Wangerooge Island is not due to reproductive output. We propose that it is mainly caused by immigration onto the island.     

The Crested Lark Galerida cristata as an example of a bird species that benefits from agricultural management in western Poland

ABSTRACT

Capsule: The Crested Lark Galerida cristata as an example of a species which selects habitats which have undergone agricultural intensification.

Aims: To describe habitat preferences of the Crested Lark in western Poland.

Methods: We surveyed 30 randomly chosen study plots (1?km²) in western Poland to detect breeding pairs of Crested Larks. Sentinel and direct observations in the field were used to provide habitat data. Generalized linear mixed models were used to determine what type of environments habitats were selected by the Crested Larks.

Results: We recorded a total of 106 pairs of Crested Larks in 30 study plots. Birds inhabited farmland environments, where their preferred habitats included buildings, silage, and arable land with cereals.

Conclusion: Silage, which is associated with agricultural intensification within agricultural landscapes, increased the chances of the occurrence of Crested Larks.     

Age-related effects on breeding phenology and success of Common Guillemots Uria aalge at a North Sea colony

Capsule: Common Guillemots Uria aalge show delayed breeding and marked age-related changes in reproductive success consistent with improved performance with experience.

Aims: To determine age of first breeding and age-related effects on breeding phenology and success of Common Guillemots.

Methods: Resighting data from a long-term colour-ringing study of Common Guillemot chicks were combined with observations of breeding phenology and success to follow the recruitment process, breeding phenology and success of 62 birds at a major North Sea colony over a 30-year period.

Results: The median age of first breeding of Common Guillemots was 6.6 years. There were no detectable costs of first breeding on return rates or the likelihood of breeding the next season but first time breeders bred later and less successfully. Age of first breeding and lifetime breeding success both varied among individuals but there was no clear optimal age of first breeding and early first breeding was not associated with higher lifetime breeding success.

Conclusions: Common Guillemots in the Isle of May population delayed breeding for 3–4 years beyond physiological maturity. The marked increase in breeding success with age was consistent with improved performance with experience rather than selection for higher quality individuals. Findings from this study will inform population models by providing improved estimates of age of first breeding and age-related changes in reproductive performance.     

The breeding biology of Rose-ringed Parakeets Psittacula krameri in England during a period of rapid population expansion

Capsule The reproductive rate of Rose-ringed Parakeets in the UK was higher in 2001-2003 than previously estimated.

Aim To measure reproductive rate and the factors affecting this in Rose-ringed Parakeets in England during a period of rapid expansion in population of this potential invasive species.

Methods During 2001–2003, 108 nests were located and monitored in southwest London, southeast London and the Isle of Thanet. Nest survival using the Mayfield method, fledging success and the characteristics of the nest and surrounding vegetation were measured.

Results The mean date of first egg was 26 March?±?1.3 days and the median clutch size was 4 eggs. Mayfield nest survival rate was 72%. Reproductive success was 1.4?±?0.3 young fledged per nest. Parakeets bred predominantly in ash (Fraxinus) and oak (Quercus). Nest trees had a diameter at breast height (DBH) of 73.7?±?4.7?cm (mean?±?se).

Conclusion Reproductive success in this parakeet was higher than previously estimated (0.8 young fledged per nest prior to 1999). This reproductive output is sufficient to explain the rapid increase in Rose-ringed Parakeet numbers (from an estimated 1500 birds in 1996 to nearly 10,000 birds by 2004) if Rose-ringed Parakeets have a low death rate similar to other parrots.     

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.     

Improvement of reproductive performance with age and breeding experience depends on recruitment age in a long‐lived seabird

Reproductive success increases with age in many species, however, the underlying mechanisms remain unclear and several hypotheses have been proposed to explain age‐related improvements in reproductive output. In this contribution we investigated the effects of age, recruitment age, breeding experience and sex on reproductive performance during the early breeding career in the common tern Sterna hirundo using long‐term individual‐based data. We used measurements of performance, which spanned the entire breeding process: clutch size, hatching success, fledging success and fledglings per pair. Longitudinal analyses within individuals showed a clear increase with age in all performance measures. Furthermore, a significant change in reproductive performance was found between first time‐ and experienced breeders. Recruitment age had a strong influence on hatching and fledging success: two‐year‐old recruits had significantly lower reproductive success than birds which recruited at older ages, but the increase in breeding performance with experience was stronger in young recruits. Comparing age and experience effects, age effects were more pronounced during the first breeding attempts, whereas experience effects were also visible in subsequent breeding attempts. The degree of intra‐individual improvements in reproductive performance is due to a complex interplay of age at first breeding and experience. The results strongly support the constraint hypothesis.     

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.     

Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

Increased hunting effort buffers against vole scarcity in an urban Kestrel Falco tinnunculus population

Capsule In years with low vole abundance birds visited hunting grounds more frequently and for longer.

Aims To describe diet composition, hunting behaviour, habitat choice and reproductive success of urban Kestrels during changing vole abundance.

Methods For five years, we studied the hunting effort of Kestrels in a medium-sized city during the breeding season. Pitfall traps were used for determining vole abundance. Kestrel diet composition was determined from pellet analyses. The number of eggs and offspring was recorded during at least two consecutive visits for each nest and each breeding stage.

Results In contrast to larger European cities, the Common Vole Microtus arvalis was a key part of the Kestrels' diet and did not fluctuate significantly according to vole availability. Reproductive success was quite high and stable throughout years with different vole abundance. In years of low vole abundance, the arrival frequency at hunting grounds and time spent there increased. During years with a low vole population, Kestrels had less hunting success and the rate of successful visits decreased. Therefore, Kestrels probably had to change hunting grounds more frequently. During low vole years Kestrels used less demanding techniques, e.g. perching, despite the lower success of these hunting techniques, to avoid extremely high energetic costs.

Conclusions An increase in hunting helps to maintain a proper diet and consequently reproductive success. Vole abundance did not change dramatically during the study period, as reported by studies from western and northern Europe. The proportion of ruderal habitats on the city periphery is higher than in more monotonous farmland habitats. Ruderal habitats can be important when Kestrels look for mammals other than voles, especially during vole scarcity.     

Population trend and breeding biology of Montagu's Harrier Circus pygargus in a natural vegetation site in northeast Spain

Capsule Reproductive output in a natural habitat was higher than in birds breeding in cereal crops, highlighting the importance of natural habitats for the species.

Aims To evaluate breeding in a natural habitat in inland Castellon province, Spain, and compare breeding parameters with other European populations.

Methods Breeding population size in inland Castellon was recorded between 1981 and 2003. A logistic growth model was used to describe population increase. We also calculated clutch size, brood size, productivity, percentage of successful nests and fledging success for the period 1989–2003.

Results Population size increased from three pairs in 1981 to 98 pairs in 2003. During 1989–2003, productivity (2.74 ± 1.49 fledglings/pair) and percentage of successful nests (84.%) were higher than observed in other European populations.

Conclusions Our results show the importance of our study area for the conservation of this species in the Iberian Peninsula, as it might act as a source of colonists for other areas. Food and protection against predators might explain the high breeding output, which may in turn explain the rapid population growth in the area. The study also suggests that natural habitats might be important for the species elsewhere.     

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.     

Population size,breeding performance and territory quality of Bonelli's Eagle Hieraaetus fasciatus in eastern Spain

Capsule A five-year monitoring study is described of overall population stability and differential reproductive success in relation to habitat heterogeneity.

Aims To assess the effect of the altitude and orientation on laying date and breeding performance, and analyse the effect of territory quality as a likely factor that could be regulating the population.

Methods We monitored a population of 28–33 pairs, from 2002 to 2006, counting a total of 131 breeding attempts.

Results Territories located at lower altitude showed higher mean fecundity than those located at higher altitude. The mean laying date was February 18 ± 16 days. Laying date was positively correlated with nest altitude, the coastal pairs laying earlier than those in mountainous regions. Pairs located at lower altitudes showed higher mean fecundity than those located at higher altitudes. There was no preference in mean orientation either in breeding performance or in relationship to nest altitude. We did not find a difference in breeding performance between territories classed as being at high density and those classed as being at low density.

Conclusion The population has remained stable since the first national census was conducted 17 years ago. Our results could be explained in the light of the Habitat Heterogeneity Hypothesis. We suggest a differential reproductive success in relation to habitat heterogeneity.     

北戴河朱鹮野化训练种群的繁殖和婚外配北大核心CSCD

为了扩大朱鹮(Nipponia nippon)种群的数量和分布区,于2018年7月从洋县引进20只朱鹮在北戴河建立了朱鹮野化种群。2020年繁殖期,北戴河共有22只朱鹮,其中成年朱鹮17只,雌雄性比为1.1;实际繁殖密度为37.8只/hm2;人工巢的密度为40.0巢/hm2,人工巢筐内径为50 cm。2020年繁殖期,北戴河朱鹮共配对8对,其中6对繁殖成功,营巢成功率为75%;共产卵33枚,平均窝卵数为(4.1±1.8)枚;出壳18只,孵化率54.5%;出飞13只,出飞率72.2%,繁殖成功率为39.4%,繁殖生产力为2.2±1.2。与洋县饲养种群相比,北戴河种群首枚卵的产卵时间晚17 d,与两地温差相吻合。北戴河朱鹮的窝卵数显著高于洋县种群,这可能是由种内巢寄生所致。监控录像表明,北戴河1巢朱鹮的窝卵数高达6枚,超过野生种群平均窝卵数(2或3枚)的2倍,推测发生了种内巢寄生行为。此外,还观察到8号巢朱鹮同时发生了婚外配和种内巢寄生行为,婚外配雌鸟将卵产在该雄鸟的巢中,使得窝卵数高达7枚。此后在该巢中发现3只朱鹮轮流孵卵,以及2只雌性朱鹮并排孵于同一巢中的异常情况。北戴河朱鹮的种内巢寄生行为可能是因网笼内人工巢址密度较高但隐蔽性较低所致,而婚外配行为可能与种群密度和繁殖密度过高有关。本文有关朱鹮异常繁殖行为的研究结果可为野化网笼内人工巢筐的设置和野化种源的选择提供参考,并提示我们进一步关注朱鹮在环境压力下的表型可塑性和生态适应能力。     

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.     

Reproductive consequences of natal dispersal in a highly philopatric seabird

Natal and breeding dispersal have a major impact on gene flowand population structure. We examined the consequences of nataldispersal on the reproductive success (proportion of pairs rearingchicks) of colonial-breeding Thick-billed murres (Uria lomvia).Reproductive success increased with distance dispersed for thefirst and second breeding attempt. The increase in breedingsuccess leveled off at natal dispersal distances above 7 m.Our results were consistent with the idea that the relationshipbetween dispersal and reproductive success is caused by siteavailability and mate choice as birds willing to disperse fartherhad a greater choice of potential sites and mates. This hypothesiswas supported by the fact that birds dispersing farther weremore likely to pair with an experienced breeder, which increasesthe likelihood of breeding success for young breeders. Explanationsfor increasing breeding success with increased dispersal basedon inbreeding effects were unlikely because most breeding failureswere caused by egg loss rather than infertility or nestlingdeath. However, we could not explain why >50% of birds returnwithin 3 m of the natal site, despite having an up to 50% lowerreproductive success than birds dispersing 7 m or more.     

Annual productivity and individual female reproductive success in a Great Bustard Otis tarda population

The reproductive success of Great Bustards Otis tarda in north-western Spain was studied between 1987 and 1998, both at the population ( c . 700 adult females breeding in our study area) and the individual level (sample of 32 marked females). Overall productivity was low, with a population mean of only 0.14 chicks reared per adult female, and an average breeding success of 0.15 chicks per year in the sample of marked females, but interannual variability was high (0.04–0.29). Population productivity was positively correlated with winter (October–March) precipitation prior to each breeding season, and negatively correlated with the number of days of rain during the hatching period. High annual productivity resulted from a high proportion of females rearing two chicks. Reproductive success was higher in females older than 6 years than in younger birds. The proportion of females in the marked sample that failed in breeding after having bred successfully the previous season was significantly higher than the proportion of those that did not. Finally, females with a higher than average breeding success tended to breed successfully in years of both low and high population productivity, whereas those with lower than average breeding success did so only in years of high productivity.