Temporal changes in the breeding ecology of European Turtle Doves Streptopelia turtur in Britain, and implications for conservation

The migratory European Turtle Dove Streptopelia turtur has undergone a 69% decline in population size and a 25% contraction in breeding range in Britain over the last 30 years. An investigation of the breeding ecology of this summer visitor was undertaken in 1998–2000 at two study sites in East Anglia, England. The only previous study of Turtle Dove reproduction in Britain provided pre-decline data for comparison with the current situation in a modern agricultural environment. Territory sizes ranged from 1.91 to 3.08 ha, were established in areas with scrub, hedges and woodland and contained less cropped land than expected from its availability. The majority of nests were sited 1–3 m above ground level within thorny bushes, particularly Hawthorn Crataegus monogyna . Nests found by radiotelemetry were significantly higher above ground level and were found in greater numbers than expected in woodland and coniferous trees than those found by cold searching, which were lower and found predominantly in hedges and thorny bushes. Turtle Dove nest success rate averaged 53% during incubation and 65% during the nestling stage, so that only 35% of nests successfully produced young. A comparison with data collected during the 1960s showed that Turtle Doves today have a shorter breeding season and consequently produce about half the number of clutches and young per pair than formerly. A simple simulation model suggested that the reduction in productivity alone would lead to a population decline of 17% per annum. This study suggests that the recovery of Turtle Doves in Britain is dependent upon the provision and sympathetic management of nesting and foraging habitats. The current arrangements for set-aside and agri-environmental schemes provide the framework for delivering these requirements.     

Breeding season food limitation drives population decline of the Little Owl Athene noctua in Denmark

Many farmland bird species have declined markedly in Europe in recent decades because of changes in agricultural practice. The specific causes vary and are poorly known for many species. The Little Owl, which feeds extensively on large invertebrates and is strongly associated with the agricultural landscape, has declined over most of northwestern Europe, including Denmark. We investigated the likely reasons for the population decline in Denmark by identifying patterns of local extinction (scale, 5 × 5 km²) and estimating demographic parameters affecting local survival, focusing on changes over time and their relationship to habitat characteristics. The distribution of the Little Owl in Denmark contracted considerably between 1972–74 and 1993–96. The extent of contraction varied across the country, and the only habitat correlate was that local disappearance was associated with smaller amounts of agricultural land. Analyses of ring recovery data suggested a constant annual adult survival rate of 61% from 1920 to 2002, which is similar to estimates from countries with stable populations. First‐year annual survival rates were much lower than values previously reported. From the 1970s into the 21st century, the mean number of fledglings declined from around 3 to < 2 young per territory, but the decline in clutch size was considerably less. Reproductive parameters were higher closer to habitat types known to be important foraging habitats for Little Owls, and were also positively correlated with the amount of seasonally changing land cover (mostly farmland) within a 1‐km radius around nests as well as temperatures before and during the breeding season. Experimental food supplementation to breeding pairs increased the proportion of eggs that resulted in fledged young from 27 to 79%, supporting the hypothesis that the main proximate reason underlying the ongoing population decline is reduced productivity induced by energetic constraints after egg‐laying. Conservation efforts should target enhancement of food availability during the breeding season. Other farmland species dependent on large invertebrates are likely to share the problems that Little Owls face in modern agricultural landscapes.     

Breeding success of Spotted Flycatchers Muscicapa striata in southern England – is woodland a good habitat for this species?

The UK population of the Spotted Flycatcher Muscicapa striata has declined markedly in the last 30 years but there have been few recent studies of the species. This study examined the relationship between nest success and the predominant habitat type around Spotted Flycatcher nests in two contrasting areas of England. A breeding population in eastern England, a region where numbers of Spotted Flycatchers are known to have decreased dramatically in recent decades, was compared with another in southwest England, where numbers have remained stable or even increased. Whilst there was no difference in breeding success between the two study areas, there were significant differences between habitats, with garden nests more successful than those in farmland or woodland, at both egg and chick stages. Estimates of productivity per nesting attempt were also lower in farmland and woodland, with nests in gardens fledging twice as many chicks as those in either woodland or farmland. The proximate cause of lower success in farmland and woodland was higher nest predation rates during both egg and chick stages. In terms of nesting success, farmland and woodland appear to be similar in quality for this species, but both appear to be suboptimal habitats when compared with gardens, providing evidence of a problem on the breeding grounds for this species, in at least these two habitats.     

Survival of British Sedge Warblers Acrocephalus schoenobaenus in relation to west African rainfall

Fluctuations in the population levels and annual adult survival rates of British Sedge Warblers Acrocephalus schoenobaenus since the late 1960s are strongly correlated with indices of wet season rainfall in the west African winter quarters. Population changes are unrelated to estimates of breeding productivity in the previous year. Habitat availability in the winter quarters has probably been the main factor limiting the size of the Sedge Warbler population in Britain during the period of study.     

The breeding biology of Black Guillemots Cepphus grylle in Shetland

The breeding biology of Black Guillemots Cepphus grylle was studied from 1982 to 1984 in Shetland, on islands lacking introduced mammalian predators. Adult survival (94%), clutch size (1.75-1.92) and breeding output (1.05-1.34 young fledged per breeding pair) were very high, indicating that conditions were relatively favourable on the islands studied. The timing of breeding varied significantly between years and was somewhat later than in other temperate parts of the range. In 1984 food was probably more readily available (possibly due to better weather conditions) in the pre-breeding period, which enabled more inexperienced birds to reach the condition threshold for breeding. It is likely that these birds tended to lay single-egg clutches in more open sites which were more susceptible to egg predation by crows. Nest-site quality and food availability were the key determinants of breeding productivity.     

Habitat associations and breeding success of yellowhammers on lowland farmland

1.  Yellowhammers began to decline on British lowland farmland in the late 1980s and losses are presently 10% per year. This study examined variation in the habitat selection and breeding success of yellowhammers, allowing an evaluation of whether Britain's yellowhammer decline might have been caused by recent changes in agriculture.
2.  Yellowhammer territories were associated with hedgerows, vegetated ditches and wide uncultivated grassy margins around fields. Pasture and silage leys were avoided. Nests were built among herbaceous vegetation in ditches or in the shrubby vegetation of hedgerows.
3.  Breeding started slightly earlier on organic farms than on intensively managed farms, but no measure of breeding success differed between farm types. Predation was the cause of most (64%) nest failures. A maximum of three breeding attempts (two successful) was observed per pair, with a mean clutch size of 3·3, a Mayfield nest success rate of 0·46, and 2·6 nestlings fledged per successful brood. These data, together with published estimates of adult yellowhammer survival and of post-fledging survival among other passerines, suggest that breeding productivity is too low to maintain a stable population.
4.  The removal of hedgerows or abandonment of hedge management, filling or clearing of ditches, intensification of grassland management and cropping or grazing right up to the field edge, are all likely to have adversely affected yellowhammers on lowland farmland in southern England. Policy reforms that redirect subsidy support to environmentally beneficial management of field margin habitats and retention of winter-feeding sites such as stubbles should assist in restoring populations of breeding yellowhammers on lowland farmland.
5.  Our data expand further the array of farmland bird species for which interactions between agricultural change and population change are increasingly understood.     

The breeding distribution and abundance of the rook (Corvus frugilegus L.) in Great Britain since the 1920s

A dramatic decline in the breeding density of the rook ( Corvus frugilegus ) took place in Great Britain between 1944/6 and 1975. This was evident from the results of two national surveys conducted in these years and other more local censuses undertaken in the intervening period. The most notable decreases occurred in the eastern counties of England: Cambridgeshire (68%) and Lincolnshire (72%). Data from local surveys indicated that the decrease in rook breeding densities began in the late 1950s and continued into the 1960s. There were regional variations in the timing of this decline.
Regional variation in the breeding density of the rook and rookery distribution was analysed for the two national censuses. The pattern was shown to be similar for both periods, despite the fall in overall densities. Rooks occupied lowland (< 300 m) agricultural land and avoided areas of high human population. Higher rook densities and rookery sizes were consistently found in Scotland compared with England and Wales.     

Inter‐annual variability and long‐term trends in breeding success in a declining population of migratory swans

Population declines among migratory Arctic‐breeding birds are a growing concern for conservationists. To inform the conservation of these declining populations, we need to understand how demographic rates such as breeding success are influenced by combinations of extrinsic and intrinsic factors. In this study we examined inter‐annual variation and long‐term trends in two aspects of the breeding success of a migratory herbivore, the Bewick's swan Cygnus columbianus bewickii, which is currently undergoing a population decline: 1) the percentage of young within the wintering population and 2) mean brood size. We used an information‐theoretic approach to test how these two measures of productivity were influenced over a 26 yr period by 12 potential explanatory variables, encompassing both environmental (e.g. temperature) and intrinsic (e.g. pair‐bond duration) factors. Swan productivity exhibited sensitivity to both types of explanatory variable. Fewer young were observed on the wintering grounds in years in which the breeding period (May to September) was colder and predator (Arctic fox) abundance was higher. The percentage of young within the wintering population also showed negative density‐dependence. Inter‐annual variance in mean swan brood size was best explained by a model comprised of the negative degree days during the swan breeding period, mean pair‐bond duration of all paired swans (i.e. mean pair duration), and an interaction between these two variables. In particular, mean pair duration had a strong positive effect on mean brood size. However, we found no long‐term directional trend in either measure of breeding success, despite the recent decline in the NW European population. Our results highlight that inter‐annual variability in breeding success is sensitive to the combined effects of both intrinsic and extrinsic factors.     

The influence of habitat management on the breeding success of the Great Bittern Botaurus stellaris in Britain

Great Bitterns Botaurus stellaris have experienced a population decline in the UK, such that in 1997 the total number of breeding males was just 11. This study aimed to identify factors affecting productivity, and how management could be used to manipulate this. An intensive study of Great Bittern breeding success was conducted between 1997 and 2001. The date that males established their booming territories was closely correlated with when females started nesting. Wetter sites with greater fish densities had males that established their booming territories earlier in the season. However, only the date that males started booming determined when females started nesting. The mean clutch size of Great Bitterns was four and the only cause of nest failure was predation. Of eight nests suspected to be second attempts the mean interval between these first and second attempts was 12.25 ± 0.88 (sd) days (range 8–15). The fate of radiotagged chicks followed to fledging revealed that the overall probability of a chick surviving to fledge was 39.1%. Daily losses of Great Bittern chicks due to starvation/exposure accounted for 76.25% and predation 21.25%. The youngest chicks in poor condition were most likely to die, particularly in periods of high rainfall. A simulated renesting model allowed estimation of Great Bittern productivity as 1.24 chicks per female and 1.52 nesting attempts per female. Habitat management, or lack of water control that resulted in sites being drier in spring, delayed nesting, although statistically there was no difference in productivity compared with wetter sites.     

Temporal changes in the migration phenology of turtle doves Streptopelia turtur in Britain, based on sightings from coastal bird observatories

Using daily counts of birds seen at six coastal bird observatories in southern and eastern England, we explored the migration phenology of turtle doves during the period 1963 to 2000. Annual totals increased threefold up to the late 1970s then decreased again, in accordance with the BTO Common Birds Census (CBC) index of abundance. Median annual spring arrival and autumn departure dates of turtle doves were not related to abundance (CBC index) or mean temperature in spring or summer respectively. Although median annual spring arrival date has not altered over the 38-year period, median annual autumn departure date has become earlier by 8 days. This has resulted in a shortening of the breeding season by 12 days, which ties in with a reduction in average number of nesting attempts per pair observed by a recent autecological study. It is possible that breeding turtle doves are now out of phase with peaks in food availability. This may have resulted in reduced breeding performance and earlier termination of the breeding season, and may have partly contributed to the decline of the species.     

Population declines and reproductive performance of Skylarks Alauda arvensis in different regions and habitats of the United Kingdom

Declines in the number of breeding Skylarks Alauda aruensis and changes in their reproductive performance were analysed using data from two long-running surveys co-ordinated by the British Trust for Ornithology: the Common Birds Census and the Nest Record Scheme. In the UK, the number of breeding Skylarks declined by approximately 55% between 1975 and 1994. This decline was steepest in agricultural habitats and in regions associated with intensive agriculture. In contrast, Skylark reproductive performance per nest, in terms of clutch size, brood size and post-hatching survival rate of nests, showed a general improvement over time. This improvement was greatest in intensively farmed agricultural habitats. Therefore changes in reproductive performance per nesting attempt were probably not responsible for the decline in numbers. It is inferred that possible causes of the decline of the Skylark are: reductions in the number of breeding attempts per pair per season, reductions in the proportion of birds attempting to breed, and increased mortality outside the breeding season.     

Is the large‐scale decline of the starling related to local changes in demography?

The decline of one farmland bird, the migratory European starling, has been attributed to both agricultural intensification and farmland abandonment and to factors operating both during the winter and during the breeding season. We analysed population data from thirty‐three Swedish nestbox colonies over more than two decades to determine if the national decline was caused by a common factor affecting all colonies or by local changes in the breeding grounds affecting starling colonies. We found that numbers of breeding starling had declined significantly, but at different rates in different colonies. The local population sizes were affected by previous years’ productivity at both national and local scales, suggesting that changes in habitat quality at both scales could affect local population trends. There were no long‐term trends in reproductive output, but fledgling production was lowest at intermediate years. The local population changes were positively related to local changes in reproductive output, but only when including complete nest‐failures. A relationship between population declines and low mean local productivity was the result of the association between population sizes and reproductive success over time, since decline rates of starlings were not related to the average success during the first part of the study, but to the average success during the later part of the study. The relationship between population change and changes in reproductive output was evident, but fledgling production showed negative density‐dependence. In conclusion this study suggests that the decline of the starling population in Sweden has been affected by processes at small spatial scales during the breeding season affecting reproductive success, but does not exclude an additional role for processes at large spatial scales or outside the breeding season.     

Characterising demographic contributions to observed population change in a declining migrant bird

Populations of Afro‐Palearctic migrant birds have shown severe declines in recent decades. To identify the causes of these declines, accurate measures of both demographic rates (seasonal productivity, apparent survival, immigration) and environmental parameters will allow conservation and research actions to be targeted effectively. We used detailed observations of marked breeding birds from a ‘stronghold’ population of whinchats Saxicola rubetra in England (stable against the declining European trend) to reveal both on‐site and external mechanisms that contribute to population change. From field data, a population model was developed based on demographic rates from 2011 to 2014. Observed population trends were compared to the predicted population trends to assess model‐accuracy and the influence of outside factors, such as immigration. The sensitivity of the projected population growth rate to relative change in each demographic rate was also explored. Against expectations of high productivity, we identified low seasonal breeding success due to nocturnal predation and low apparent first‐year survival, which led to a projected population growth rate (λ) of 0.818, indicating a declining trend. However, this trend was not reflected in the census counts, suggesting that high immigration was probably responsible for buffering against this decline. Elasticity analysis indicated λ was most sensitive to changes in adult survival but with covariance between demographic rates accounted for, most temporal variation in λ was due to variation in productivity. Our study demonstrates that high quality breeding habitat can buffer against population decline but high immigration and low productivity will expose even such stronghold populations to potential decline or abandonment if either factor is unsustainable. First‐year survival also appeared low, however this result is potentially confounded by high natal dispersal. First‐year survival and/or dispersal remains a significant knowledge gap that potentially undermines local solutions aimed at counteracting low productivity.     

POPULATION,DIET AND THE ANNUAL CYCLE OF THE LAUGHING DOVE AT BARBERSPAN,PART 4. BREEDING DATA AND POPULATION ESTLMATES

Dean, W. R. J. 1980. Population, diet and the annual cycle of the Laughing Dove at Barber-span, Part 4: Breeding data and population estimates. Ostrich 51:80-91.

From 1974–1976, Laughing Doves Streptopelia senegalensis raised an observed total of 436 young in 619 nest attempts, giving a breeding success of 0,70 young per pair nest attempt, or an overall success of 0,35 young per egg. Breeding success, estimated by computing the probability of survival of an egg through the incubation period, and of a chick through the nestling period suggest that the breeding success is about 0,33 young per egg. Nests were found in every month of the year, though there was a tendency for breeding to be concentrated during the late rainy season and the dry season. The mean annual population size of the Laughing Dove in the study area was estimated at 221 adults and juveniles by one method and at 237 adults and juveniles by another method. The number of young produced each year is correlated with the estimated population size for each year.     

El Niño in the warm tropics: local sea temperature predicts breeding parameters and growth of blue-footed boobies

1. There is increasing interest in the impacts of El Ni?o Southern Oscillation (ENSO) on reproduction of apical predators such as seabirds and marine mammals. Long-term studies documenting ENSO effects on reproduction of seabirds in the warm tropics are scarce, and differential sensitivity of breeding parameters to ENSO has rarely been explored. 2. Analysis of 18 years of breeding data from a colony of the blue-footed booby Sula nebouxii (Milne-Edwards) showed a delay in onset of breeding when the global Southern Oscillation Index was negative; each unit of the atmospheric pressure differential (hPa) across the Pacific Ocean meant a delay of 7 days. 3. ENSO conditions also produced declines in breeding participation, clutch size, brood size, hatching success and fledging success, especially when surface waters surrounding the colony were warmer during winter and spring. Each additional degree (°C) of water temperature produced a reduction of 0.45 fledglings per nest. Different breeding parameters were sensitive to ENSO indices in different blocks of months. 4. Warming of local waters during the winter was associated with decline in ocean productivity in the current year and the following year, consistent with ENSO impacts on breeding parameters being mediated by effects on local productivity and prey availability. However, there was no evidence of lagged effects of ENSO on any breeding parameter. 5. Comparison of 5 years revealed that when local surface waters were warm, chicks grew more slowly, but no effects of ENSO on weight and size of eggs were evident in data of 9 and 7 years, respectively. 6. Our findings extend evidence of impacts of ENSO on seabird reproduction to the eastern tropical Pacific and indicate that several breeding parameters of blue-footed boobies (but not egg size) are affected in the short term by ENSO conditions, particularly by local anomalies in sea surface temperature associated with decline in ocean productivity.     

The use of predator-exclusion fencing as a management tool improves the breeding success of waders on lowland wet grassland

Waders breeding on lowland wet grassland have undergone dramatic declines across Europe in recent decades. Few species now achieve the levels of breeding success required for population stability and recovery, with predation from large mammals acting as a key compounding factor limiting nest survival and productivity. Predator management through lethal control is often controversial, yet alternative non-lethal methods are little tested in the context of grassland breeding waders. Excluding predators through the use of electric fences has led to improvements in nest and chick survival in other habitats. To test the applicability of this method to lowland wet grassland we constructed predator-exclusion fences on sites across the UK and, with Lapwing Vanellus vanellus as a study species, used historical and contemporary data to test whether excluding large mammalian predators leads to an increase in wader nest survival and productivity, and whether effects differ between fence designs. Lapwing nest survival was significantly higher in the presence of any type of predator-exclusion fence, with significantly fewer nests predated each day. Overall productivity also improved, with significantly higher numbers of chicks fledged per pair in years when fences were operational. Different designs and methods of powering fences resulted in different levels of success, with combination design fences and those powered by mains electricity performing best. Excluding large mammalian predators from areas of lowland wet grassland with predator-exclusion fencing successfully improves Lapwing nest survival and productivity, allowing breeding success to exceed the levels required for population recovery. Other wader species breeding in the same habitat are also likely to benefit from the increased protection from predation provided by fences. Predator-exclusion fencing is therefore an effective management tool for protecting restricted and declining populations of breeding waders on lowland wet grassland.     

Reproduction of the white-tailed sea eagle Haliaeetus albicilla in Sweden

Sixty-eight white-tailed sea eagle territories were studied on the Swedish Baltic coast in 1964–1982 and 22 territories in Swedish Lapland in 1976–1982. Older data were assembled for comparison. The breeding frequency of adult pairs was high in both study areas (ca. 90%). Median egg-laying dates were 17 March (Baltic) and 7 April (Lapland). Average clutch size and egg volume did not differ between samples from the Baltic taken between 1856 and 1982. Average clutch size was significantly smaller in Lapland (1.6 eggs) than on the Baltic coast (2.1 eggs). Clutch size in Lapland was positively related to carrion availability during March. Nestling survival was significantly lower in Lapland, due to starvation in broods of more than one young. Brood loss was significantly more frequent in Lapland. At the Baltic, nest success decreased from 75 to 22%, and brood size from 1.8 to 1.2, from the early 1950's to the mid 1960's. Nest success in 1976-1982 was significantly lower at the Baltic (26%) than in Lapland (45%), with a much larger proportion of consistently failing pairs. The proportion of eggs laid that resulted in fledged young was estimated at 16% at the Baltic and 39% in Lapland. It is concluded that egg production and nestling survival were suppressed by limited food supply and nest success by human interference in Lapland, and that the reproductive impairment of the Baltic population was due primarily to contamination with toxic substances, mainly DDE.     

Breeding biology and breeding success of the Lesser Grey Shrike Lanius minor in a stable and dense population

The Lesser Grey Shrike Lanius minor is highly endangered throughout Europe, having declined markedly in abundance and range. Long-term changes in climate and agricultural practices have been identified as the main reasons for its decline. To determine which factors influence short-term changes in breeding success, we examined several aspects of its breeding biology. Our investigation revealed that our study area bears a large and stable population of this species. In 1996 and 1997, we recorded 84 and 77 breeding pairs in an area of 20 km², with an average of 4.20 and 3.85 pairs/km² respectively. Data on breeding density, clutch size and fledging success from 1989 to 1997 (excluding 1992) indicate a stable breeding population with a constant high breeding success. Reproductive success declined through the season, mainly through seasonal variation in clutch size rather than chick mortality. However, breeding success was generally high (69% and 79% of the nests produced chicks], with low hatching failure and few nest losses. The main cause of breeding failure was nest predation (at least 50% of nest losses), mainly by magpies (at least 66% of depredated nests). Although in this population the Lesser Grey Shrike tends to aggregate in clusters, breeding density had no obvious effect on breeding success and nest predation.     

Density-dependent decline of breeding success in an introduced, increasing mute swan Cygus olor population

An increasing population of the mute swan Cygnus olor , was studied from its very establishment in 1976 until 1998. As the number of pairs increased, there was a decline in all production parameters, including the average number of clutches per pair, the average number of broods per clutch, and the average number of fledged young per brood. In a multiple regression analysis covering the whole breeding season, the number of pairs and the average number of clutches per pair explained 71% of the variation in the average number of fledglings per pair. During the brood season, the average number of fledglings per brood was an additionally important parameter in explaining fledgling production. Habitat quality seemed to affect breeding: in places occupied earlier more cygnets fledged from a clutch than in habitats inhabited later. The decreased production of young very likely reflects density-dependent effects on reproduction. This density dependence seems to operate on the breeding grounds, since winter harshness did not affect breeding success. Density-dependent processes started acting already when pairs were beginning to nest, and continued during the brood period. Density dependence has apparently not been detected at the pair stage in earlier studies of bird populations.     

First pairing in Snow Geese Anser caerulescens: at what age and at what time of year does it occur?

In migratory birds, the place and time of pair formation are important parameters for population structure and dynamics. Geese are not only migratory but also exhibit long-term monogamy, and therefore the first pairing event in a bird's lifetime is of particular importance. Through behavioural observations of young, known-age, marked birds conducted on the wintering grounds during three winter seasons we investigated two aspects of the timing of first pair formation in the Wrangel Island population of Snow Geese Anser caerulescens : (1) the age at which birds first form pair bonds, and (2) the seasonal pattern of first pair formation. Wrangel Island Snow Geese paired considerably later in life than Snow Geese from a low-Arctic population: almost none of the birds formed pairs in their second winter, and many were still in sibling groups for at least part of that season. The proportion of birds in pairs continued to increase until at least 5 years of age. Most pairing took place during the observation periods, and in general the proportion of birds in pair bonds increased gradually throughout the winter season. The amount of pairing during spring migration or summer varied annually and among cohorts, indicating that even very young birds may be able to form pair bonds quickly if conditions on the breeding grounds are unusually favourable. Pairing later and remaining in family groups longer may be a response to breeding conditions in this high-Arctic colony. Here, productivity is typically low due to harsh weather and predation, whereas Snow Geese breeding in the low Arctic are less restricted and form pairs and start to breed when younger. The fact that most, but not all, pairing takes place on the wintering grounds helps explain why a previous study found a certain amount of gene flow between the two Wrangel Island subpopulations with separate wintering grounds.